2677-oikonomoy et al-fish greece
TRANSCRIPT
91Medit. Mar. Sci., 8/1, 2007, 91-166
Mediterranean Marine ScienceVolume 8/1, 2007, 91-166
The freshwater ichthyofauna of Greece - an updatebased 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 speciesfrom 105 hydrographic basin areas were compiled and analysed in order to develop a nation-wide inven-tory (including transboundary river basins). Overall, 162 species, including diadromous and euryhaline,with documented occurrence records in freshwaters, and taxa of unclarified taxonomic status, are accom-modated in the distributional compilation. An annotated checklist summarises the confirmed ichthy-ofauna of Greek freshwaters (161 species); a provisional supplementary list contains species recorded inbrackish waters (55 species). In comparison to the last published (1991) checklist of freshwater fish ofGreece, the present checklist shows an increase in species number of 53% (56 species). This increase hasresulted mainly from taxonomic re-evaluations of existing taxa on the basis of new information and adop-tion of a new systematic concept. The current trend, as reflected in recent ichthyological publications, istowards abandonment of the biological species concept (BSC) and adoption of the phylogenetic speciesconcept (PSC) for the delineation of species boundaries. The practical implications of the change inspecies concept on biodiversity conservation and watershed management are discussed. An overview ofthe composition and characteristics of the freshwater fish fauna of Greece is provided, especially withregard to the native and introduced status of species, and the spatial patterns of species richness andendemism. This systematic inventory may assist in efforts to develop nation-wide surface water bioassess-ment tools within the demands of the Water Framework Directive (WFD); it may further promote biodi-versity conservation and biologically-orientated fishery management approaches.
Keywords: Freshwater ichthyofauna; Fish distribution; River basin area; Endemism; Biodiversity; Greece.
Review Article
Introduction
Freshwater fish represent an impor-tant component of the aquatic ecosystemand are highly valued for their economic,social and aesthetic importance. Fish arealready involved in environmental policiesas biodiversity and ecological quality indi-cators (KESTEMONT et al., 2000;SCHMUTZ et al., 2007) and they havebeen used successfully in biogeographicalstudies (B N RESCU, 2004), ecore-gion delineations (HAWKES et al., 1986;ABELL et al., 2002), conservation evalua-tions (MOYLE & RANDALL, 1998) andassessments of ecologically acceptablewater regime management (JOWETT,1997). Greece has diverse inland waterresources and hosts one of the richestfreshwater ichthyofaunas in Europe. Thenumber of species occurring in Greekwaters is still an issue of active investiga-tion. The last published checklist of thefreshwater fish of Greece contains 105native and introduced species (includingeuryhaline) plus five of doubtful occur-rence (ECONOMIDIS, 1991). A recentre-evaluation gives 135 species, of which89 are native exclusively freshwater, and 54are endemic to the country or to the south-ern Balkans (BOBORI &ECONOMIDIS, 2006). However, a com-plete nation-wide inventory of fish occur-rences by river basin has never been pub-lished.
A review of publications concerningthe freshwater fish of Greece was recentlyundertaken with the scope of assessing theutility of available ichthyological informa-tion for the implementation of the EUWater Framework Directive(ECONOMOU et al., 2004a). One of theoutcomes of this review is that there hasbeen a change in the thematic focus of
ichthyological research over the pastdecades, from distributional surveys andtaxonomic work based on morphology, tostudies of fish physiology, ecology, biologyand genetics. This change had a negativeside-effect on the availability of fish distri-butional data. Large-scale distributionalsurvey work was abandoned in favour ofregional or local scientific investigations,often focusing on single species or singleareas. It is characteristic that most of theinformation used to produce the checklistof freshwater fish of Greece published byECONOMIDIS (1991) was derived fromhis earlier published catalogue of fish(ECONOMIDIS, 1973), which is based onresults of investigations undertaken beforethe 1970s.
According to the aforementionedreview (ECONOMOU et al., 2004a) valu-able ichthyological information exists forlarge rivers and lakes, however, the fishfauna of small aquatic systems has beenpoorly investigated. For some areas noinformation at all could be found, whereasfor other areas there are data shortages formost, except the larger water bodies. Anassociated problem is that the distributionof alien fish is poorly known since speciesintroductions or translocations are rarelyannounced, and when they are, the dataare not always accessible. Quite often,information on the occurrence of non-native species can be obtained only fromgrey literature or local authorities. A riskof misreporting and misidentification isinevitably inherent in this approach as thetaxonomy of the introduced species is notoften verified by experts. Moreover, muchof the available information on species dis-tributions is spread out in various, oftenobscure, publications which are not readi-ly accessible to the scientific community,especially to non Greek-speaking scien-
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tists. This holds particularly true for disser-tation theses, works published in older orlocal journals, national conference pro-ceedings and project reports. It is obviousthat a synthesis and compilation of rele-vant information is required to facilitateknowledge access and sharing.
Deficiencies in distributional dataavailability, coupled with taxonomic ambi-guities, have a negative impact on scientif-ic research and management applicationsdependent on fish data. For example, plan-ning for biodiversity conservation inGreece is often hindered by lack of knowl-edge of the distributional ranges of endan-gered species and vagueness in ichthyolog-ical nomenclature. These problems reflectthe fact that the list of protected fishspecies for Greece included in the Annex-es of the Habitats Directive (92/43/EC)contains species listed under invalidnames, while many really threatenedspecies are not listed. Recent taxonomicrevisions undertaken on a European scaleand the application of new criteria fordefining species (KOTTELAT, 1997;KOTTELAT & FREYHOFF, 2007a)have attempted to resolve problems relat-ed to nomenclature and systematics but atthe same time they have complicated theichthyological faunal list of Greece byrenaming and ‘splitting’ many species.Thus, much confusion may be generatedamong environmental managers and non-specialist users of ichthyological data, assome species appear under new names,several new species have emerged, and thedistributional ranges of many species arenot clearly defined. This again reiteratesthe need for a systematic treatment ofexisting fish distributional records to pro-duce hydrographic basin area compilationsincorporating these changes.
The aim of the present work is to
develop an updated and, to the degree thatit is possible, complete inventory of thedistribution of freshwater fish over Greekterritory. Using a combination of datasources and historic or contemporaryaccounts, we compiled lists of the freshwa-ter fish occurring in 105 Greek hydro-graphic basin areas and we characterizedtheir provenance status (i.e. native orintroduced in each basin area). The hydro-graphic basin area was chosen as the spa-tial unit for this inventory because of itsrelevance to biogeography. Indeed, no sin-gle factor is more important in explaininglarge-scale distributional patterns of fresh-water fish than hydrographic basin limits(GILBERT, 1980). River basins have longbeen considered as the operational unit forthe biogeographical analysis of fishthroughout the world (e.g. ABELL et al.,2002; HIGGINS et al., 2005; THIEME etal., 2005; REYJOL et al., 2006). This isbecause both primary and many secondaryfreshwater fish are freshwater-obligateorganisms that cannot readily disperseacross terrestrial boundaries or marineareas. Effectively, the freshwater fish pop-ulations within hydrological basins arereproductively isolated entities adapted totheir basin-scale ecosystems. River basinsoften exhibit a distinct fish compositionand can be seen as ‘biogeographicalislands’ containing a specific pool ofspecies (HUGUENY, 1989). This island-like character also renders the basin areaan appropriate scale for studies ofendemism and speciation (PETER, 2006).
The main product of this work is ahydrographic basin area-based compila-tion of fish that live in the freshwaters ofGreece. These data may assist in under-standing regional assemblage structureand other faunistic attributes of relevanceto biogeography. In addition, the data are
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of potential utility in resource and envi-ronmental management, especially withregard to ecological quality assessments,biodiversity conservation and fisheryexploitation.
From the perspective of ecologicalquality assessment, interest is centered onthe successful implementation of the EUWater Framework Directive 2000/60/EC(WFD). The WFD establishes an ecosys-tem-based policy framework for watermanagement and protection in Europe atthe river basin scale. The Directivedemands, among others, that the memberstates should develop monitoring pro-grammes and ecological status classifica-tion systems within river basin districtsusing fish and other organisms as biologi-cal indicators. An important requirementfor the establishment of the monitoringprogrammes is that reliable informationon freshwater fish assemblages for eachriver basin area is available, both for thecharacterization of the undisturbed, type-specific ‘reference conditions’ and for theselection of the appropriate biologicalmetrics with which to measure ecologicaldegradation (ECONOMOU, 2002;ECONOMOU et al., 2003).
With regard to biodiversity conserva-tion, the emphasis is on identifying areashosting species in need of protectionaccording to the EU Habitats Directive(92/43/EC). Incomplete and/or inaccessi-ble information on fish species’ distribu-tions has hindered the timely evaluation ofareas of high conservation value and theirinclusion in the NATURA 2000 protected-area framework. This is unfortunate, asGreece is one of Europe’s biodiversityhotspots in terms of freshwater fishendemism (MAITLAND & CRIVELLI,1996). The country has already lost nearly75% of its natural wetland areas during the
last century (OECD, 2000) and manyaquatic ecosystems are threatened even indesignated protected-areas. Data showthat several fish populations have becomethreatened or extirpated (ECONOMIDIS,1995, 1999, 2002; ECONOMIDIS et al.,1996; ECONOMOU et al., 1999;STOUMBOUDI et al., 2002;PERDIKARIS et al., 2005) and manymore may face threats in the near futuredue to escalating pressures upon surfacefreshwater resources. Organization of theichthyological information on a basin areascale provides a practical approach for theevaluation of conservation priorities andthe design of restoration measures. Knowl-edge of fish distributions may also facili-tate the monitoring of changes in the fishfaunas in relation to climatic change.
Lastly, freshwater fishing has a strongsocio-economic dimension, especially inrural areas, where alternative employmentpossibilities are limited or highly seasonal.The freshwater fisheries’ resources arethreatened by destructive or illegal fishingand, increasingly in recent years, by theuncontrolled introduction of alien species(CRIVELLI et al., 1997; ECONOMIDISet al., 2000a; ECONOMOU et al., 2001a).The present compilation may provide pol-icy-relevant information on the distribu-tion status of native, exotic and translocat-ed species that may help in designing bio-logically-based fisheries management.
Study Area
Greece’s land area (132 000 km2) sup-ports remarkably varied inland water fea-tures. The country has a highly fragmentedgeography and various parts of the territo-ry have very different environmental con-ditions and biogeographical histories.65.4% of the land exceeds 200 m in alti-
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tude (CATSADORAKIS, 2003). Thecoastal morphology both on the continentand on the islands is diverse but it usuallyconsists of a highly fragmented, narrowcoastal zone with a varied relief. The mostextensive lowland areas exist in northernGreece where relatively large alluvialplains are drained by the rivers Evros,Nestos, Strymon, Axios and Aliakmon.
In terms of inland water aquatic biota,Greece is at a biological crossroads amongMediterranean, temperate European,Danubian-Black Sea, and Anatolian influ-ences (B N RESCU, 2004). Althoughthe biotic influences due to the country’sgeography are unique, important biogeo-graphic barriers criss-cross the country andcreate even more heterogeneity. Longmountain chains, climatic rain-shadows,wide marine gulfs and deep seas that werenot drained due to sea level lowering at theglacial maxima, create inland water isola-tion which has remained rather stable formillennia. More geographic idiosyncrasiesare created by the country’s geologicalfragmentation and dynamics, its extensivecoastline (16000 km), and a variety of cli-matic zones. Climatically, Greece main-tains remarkable extremes ranging fromhigh-rainfall mountains receiving 2400 mmprecipitation per year (Central Pindos), toseasonally semi-arid areas (southern Atti-ki) receiving less than 400 mm. Rain-shad-ow areas in the southeast mainland andthe Cyclades islands create pockets of sea-sonally arid conditions with high evapo-transpiration rates and a long summerdrought. Unlike most European countriesGreece’s inland aquatic ecosystems arestrongly characterized by geographical iso-lation, both spatially as small river basinareas, and temporally as long-standing rel-atively stable waterbodies which createdbiotic refugia when much of the continent
was being affected by the Pleistoceneglaciations (SKOULIKIDIS et al., 1998;PERISSORATIS & CONISPOLIATIS,2003).
A proper classification of Greece’ssurface water systems has never been com-pleted and the variety of aquatic ecosys-tems and habitat types is certainlyimmense. These water features range fromlarge transboundary rivers, medium andsmall perennial and intermittent streamsto small endorheic karstic streams, springsystems, inland lakes, coastal lagoons,swamps and marshes. Water features havebeen poorly inventoried for their aquaticbiota and little published informationexists on regional and local scale speciesdistributions; data for invertebrates andlower vertebrates is especially lacking(LEGAKIS, 2004). Until recently, formalinventory procedures have catalogued onlythe large wetland sites; one such compila-tion lists 378 wetland sites coveringapproximately 2000 km2 in the entirecountry (ZALIDIS & MANTZAVELAS,1994). Unfortunately, these published cat-alogues of sites are a poor and incompleterepresentation of the total number ofaquatic sites in the country (e.g.CATSADORAKIS & PARAGAMIAN,2006). In a recent regional-based survey,CATSADORAKIS & PARAGAMIAN(2007) describe 352 wetland areas solely inthe Aegean islands (excluding Crete).
Profile of Greek hydrographic basins
It has been recently stated that limnol-ogy and freshwater ecology in the Mediter-ranean should not be based on temperateEuropean paradigms, patterns and gener-alizations (ALVAREZ-COBELAS et al.,2005). Greek river basins differ markedlyfrom most temperate European ones.
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They usually comprise isolated hydro-graphic basins, characterized by short,steep fluvial systems that exhibit very ero-sive behavior, flashy, irregular flowregimes and are influenced by varied geo-graphical, geological and climatic condi-tions. Most rivers run through narrowmountain valleys and descend abruptly tothe coast, usually lacking extensive lowlandsections and floodplain habitats. There areonly eight large rivers in the country (withdrainage areas larger than 6000 km2),including five transboundary ones(SKOULIKIDIS et al., 1998). The numberof smaller rivers or streams with perennialflowing segments is certainly in the hun-dreds; a complete inventory of all hydro-graphic basin delineations does not exist,despite attempts in the recent past (e.g.NTUA, 1994). ‘Lowland’ fluvial habitatconditions are mainly encountered in theplateaus and inland plains of the largerrivers, which create unique biotic assem-blages since their access to the river’s mainstem may be blocked by gorges or othernatural barriers including karstic phenom-ena. The deltas of Greek rivers are oftenextensive, although they are not affectedby estuarine tidal regimes (TSIOURIS &GERAKIS, 1991).
Most lakes in Greece are located with-in river basins or have a historic relationwith their wider river basins. Some of theolder lakes have a unique history of isola-tion and are centers of endemism (e.g.Lakes Prespa, Vegoritis, Pamvotis, Doira-ni) (FROGLEY et al., 2001; GRIFFITHSet al., 2002; FROGLEY & PREECE,2004; ALBRECHT et al., 2007). Although56 major natural lakes have beendescribed (ZALIDIS & MANTZAVE-LAS, 1994; ZACHARIAS et al., 2002),Greece has many smaller lentic bodies,such as various wetlands, ponds and coastal
lagoons; the number of these small waterfeatures is certainly in the thousands. Someof these water features are associated withlowland rivers and lowland or plateau lake-swamp environments. In contrast, manysmall karstic spring-fed lentic systems espe-cially in the limestone-dominated southernand western parts of the country are oftentotally isolated from other surface-flowingwaters in the wider landscape.
Methods
Our inventory approach takes the fol-lowing steps: a) selection of adequately-studied hydrographic basin areas (andother ‘isolated’ aquatic sites); b) compila-tion of fish species occurrence for eacharea; and, c) compilation of a provisionalannotated freshwater fish species checklistand a supplementary list of species record-ed in transitional waters.
Definition of studied basin areas
The basic geographical unit to assem-ble our distributional compilation isdefined as the hydrographic basin area.Only basin areas where documented fishspecies occurrences have led to a ‘near-complete’ ichthyofauna list are included inthe compilation. The size of each basinarea was retrieved largely from the Hydro-scope database of the National TechnicalUniversity of Athens (NTUA, 1994). Thisgeo-database delineates 737 river basinsand ‘wider river basin areas’ in Greece.The NTUA delineations do not includesome of the smaller hydrographic basinareas we provide data for; their area wasroughly estimated by us from maps (Table1). Each hydrographic basin area was cate-gorized into one of six general biogeo-graphical units, these roughly follow
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Table 1Hydrographic 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.
EstimatedNo Name Given Biogeo-graphical 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.)
3 Filiouri North Aegean 2107 Filiouris R., Bospos R., Mitrikou (or Ismarida) L.;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.
7 Laspias North Aegean 138 Laspias (or Laspopotamos) R., Avdira wetlands andsurrounding 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.
16 Volvi North Aegean 1903 Volvi L., Koronia (or Aghiou Vassileiou or Lagada) L.,tributary streams.
17 Doirani North Aegean 420 Doriani (Dorjan) L. (within the broader basin of the AxiosR.), 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.
22 Vegoritis North Aegean 752 Vegoritis L., Cheimaditis L., Petron L., Zazari L.,tributary streams.
23 Kastoria North Aegean 264 Kastoria (or Orestias) L. (within the broader basin of theAliakmon R.), tributary streams.
24 Aliakmon North Aegean 8677 Alkiakmon R., Almopeos R., Tripotamos R., reservoirs,Delta wetlands.
25 Mavroneri North Aegean 815 Mavroneri (or Itamos) R.26 Pinios The North Aegean 9500 Thessalian Pinios R.
27 Prespa South Adriatic 1383 Mikri and Megali Prespa L. (within the broader basin ofthe 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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Table 1 (continued)
EstimatedNo Name Given Biogeo-graphical Area Included water bodies
Area (km2)
34 Acheron Ionian 752 Acheron 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.
42 Acheloos Ionian 6329 Acheloos R., Agios Dimitrios (Lesini) R., lakes,reservoirs, Delta and lagoons.
43 Evinos Ionian 1112 Evinos R. and reservoir.44 Mornos Ionian 998 Mornos R., reservoir, Delta wetlands and Gouvos Spring.
45 Kerkyra Ionian Insular Kerkyra (Corfu) Island water features.drainages
46 Lefkas Ionian Insular 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 142 Glafkos R.58 Piros Ionian 577 Piros R.59 Tsivlos Ionian 10 Tsivlos L. (within the Krathis R. basin).
60 Prokopos Ionian 280 Prokopos Lagoon, Lamia Swamp and adjacent springs andstreams.
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 102 Taka L.73 Evrotas Ionian 1738 Evrotas R.
(continued)
B N RESCU (2004): 1) NorthernAegean (includes both the Thrace andMacedonia-Thessaly zoogeographicregions), 2) Southern Adriatic (represent-
ed here by the Aoos R. and the Lake Pres-pa), 3) Ionian (includes the Ionian islandsand nearly all of the Peloponnese exceptthe eastern part 4) East Peloponnese
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Table 1 (continued)
EstimatedNo Name Given Biogeo-graphical Area Included water bodies
Area (km2)
74 Vassilopotamos Ionian 14 Vassilopotamos canals within the Evrotas Delta Area.75 Smynous Ionian 192 Smynous (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.
90 Sperchios Attiko-Beotia 1828 Sperchios R., Gorgopotamos and other tributaries, Deltawetlands.
91 Cholorema Attiko-Beotia 192 Cholorema 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.
96 Samothraki Aegean islands Insular Samothraki Island water features.drainages
97 Lesvos Aegean islands Insular Lesvos Island water features.drainages
98 Samos Aegean islands Insular Samos Island water features.drainages
99 Rhodos Aegean islands Insular 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.
(rain-shadow coastal area only), 5) Attiko-Beotia (includes Euboea and Fthiotis ineastern central Greece), and 6) Aegeanislands (includes Crete).
The hydrographic basin areas cited aredefined by the traditional watershedboundaries of the entire catchment, takinginto account hydrographic idiosyncrasiesand historical drainage connections. In thiscontext, lakes with a present or past con-nection to large rivers were usually incor-porated into the wider river basin area.This also includes small rivers lying in thevicinity of the deltaic depositional zone oflarger rivers. For instance, the Acheloos R.basin area is defined here to include thedrainages of the natural lakes Trichonis,Lyssimachia, Ozeros and Amvrakia; theartificial reservoirs Kremasta, Kastraki,Stratos and Tavropos; the entire deltaicwetland area, including the small AghiosDimitrios (or Yeroporos) R. with whichthe Acheloos R. had a direct connectionbefore the draining of the deltaic LakeMeliti. The hydrographic basin areas ofhydrologically isolated lakes are consid-ered to contain the drainages of associatedsmaller lakes, as well as the drainages ofrivers discharging into them. For example,the hydrographic basin area of LakeVegoritis contains the watershed of LakesVegoritis, Cheimaditis, Zazaris and Petronand all streams discharging into theselakes. Expert judgment based on criteria ofgeographical distinctiveness and surfacehydrological connectedness was used todefine the areal extent of the hydrograph-ic basin area unit considered in this inven-tory. In certain cases (14 basin areas), sev-eral exceptions and violations to the use oftraditionally defined hydrographic basinareas were made:a. Certain older lakes or lake groups
within major river catchments are
included as ‘isolated’ water features,despite present or past drainage con-nections with wider river basins (e.g.Lakes Kastoria, Doirani and Pamvo-tis). In a few instances even very smalllakes, semi-isolated spring-fed wet-lands, and other karstic surface water-bodies are delineated in isolation,either because they represent uniquegeological features or they have othersurface water attributes which areworthy of particular conservationinterest (e.g. Lakes Tsivlos, Vouliag-meni and Zaravina, Kato AlmiriSpring).
b. Some assemblages of separate smallstream basins on the mainland and onthe islands are incorporated into singleartificially defined ‘basin areas’. This isdone in cases where complete infor-mation on species composition foreach separate stream basin is lacking,but incidental records among thegroup of proximate basins is consid-ered adequate to warrant the widerareas’ inclusion (i.e. SW Messiniaincludes several small streams south ofKyparissia-Messini, Peloponnese). Insome other cases, surface hydrology iscomplicated and semi-connected wet-land conditions or artificial canals blurwatershed boundaries; hence nearbystreams and wetlands are included(e.g. coastal wetland and isolatedcanal features such as the Laspias-Avdira wetlands near Xanthi, Thrace).In the case of small islands, the entirenetwork of an island’s inland waterfeatures is presented as an artificiallydefined ‘hydrographic basin’ area.This treatment is of course provisionaland was followed simply because inmost insular systems ichthyologicalresearch is still inadequate. Only for
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two of the largest Aegean islands(Crete and Euboea) are specific riverbasin area compilations provided.
Compilation of fish species occurrence foreach area
We compiled a species-hydrographicbasin area dataset hosting the freshwaterfish species occurring in Greece and, in thecase of transboundary rivers, in neighbour-ing southern Balkan countries (Albania,Bulgaria, FYR Macedonia, Turkey-in-Europe). This compilation is based on anextensive bibliographic study that began in2003 (ECONOMOU et al., 2004a, 2006).Unpublished survey data from HCMRfield surveys were also utilized when theavailable material was collected throughthe participation of the authors in fieldsampling projects. The authors have beeninvolved in wide-ranging site-based sam-pling surveys particularly in western,northwestern and southern Greece(including 73 of the presented hydrograph-ic basins areas); much of this work remainsunpublished in refereed journals (ECO-NOMOU et al., 1999, 2004a, 2007; ZO-GARIS et al., 2004, 2006).
Attention was paid to documentingand ascertaining the quality of each partic-ular record for each basin area. The occur-rence of a species in a basin area had to beprovided by at least one reliable source(publication, technical report) or to havebeen confirmed by the authors during fieldsurveys. Some information was receivedfrom competent ichthyologists throughpersonal communications. On some occa-sions substantiated uncertainty of the qual-ity of an accepted record is provided with aquestion mark notation beside the record.
Fish systematic taxonomy was basedon KOTTELAT & FREYHOF (2007a),
and all relevant publications up to mid2007. KOTTELAT & FREYHOF (2007a)provide a major revision of taxonomicunits in Europe and include informationon the distribution of the taxa occurring inGreece. Some of the taxa described arenot given separate species status while dis-tributional information for some species isvague or incomplete. For the sake of con-sistency and practical policy-relevantapplications, we adopt the taxonomicchanges given by KOTTELAT &FREYHOF (2007a) and we critically com-ment on particular difficulties in the taxon-omy and distribution of certain taxa in thechecklist (Appendix I). In some cases wecarefully document divergence in certaintaxa distributions from that presented bythe above authors.
The following species inclusion andnomenclature premises were made:a. All species, native or introduced, that
spend their entire lives or a significantportion of their life-cycle in freshwa-ters were considered in the basin areadistributional compilation, i.e. primaryfreshwater species (intolerant to salt-water) and secondary freshwaterspecies (species that now live exclu-sively in freshwater but were once ableto tolerate saltwater). Diadromousspecies and euryhaline species forwhich adequate distributional dataexist are included, but so-called ‘spo-radic species’ or marine stragglers thatseem indifferent to salinity or have adefinite marine life-history, are notincluded. Information is largely lack-ing in order to confirm the regularpresence of certain marine species infreshwaters, although proof of severalmarine species’ residence in brackishtransitional waters exists (e.g. KOU-TRAKIS et al., 2000).
Medit. Mar. Sci., 8/1, 2007, 91-166 101
b. Certain euryhaline species known tobe regularly present in freshwaters butfor which consistent records are miss-ing are excluded from the distribution-al compilation. However, these speciesare placed within the summary check-list of freshwater fish in the appendixand should be considered as importantelements of the freshwater fauna.
c. Valid species names are given but insome instances tentative operationaltaxa names are given for species thatpresent identification problems, strict-ly following KOTTELAT & FREY-HOFF (2007a). In addition, in five iso-lated cases, we provide only our ownoperational name for unidentifiedspecies known only to genus level (i.e.two Eudontomyzon spp., Knipowitschiasp. Squalius sp1, Squalius sp2). This isnot a diversion from any accepted tax-onomy, but it is required to show thepresence of an unidentified species ina particular basin area.
d. Species of undefined taxonomic statusare included with a notation to indi-cate taxonomic uncertainty. For exam-ple, KOTTELAT & FREYHOF(2007a) provide the controversialtaxon Salmo sp. Louros and weinclude this poorly described taxon inour compilation, but with a notation toshow ‘doubtful taxonomic status’. Fur-thermore there are several instanceswhere KOTTELAT & FREYHOFF(2007a) refer to species occupying par-ticular hydrographic basins but give noinformation on presumably the sameor closely related fish that exist innearby basins. In such cases of uncer-tainty we have used available distribu-tional data and expert judgment toassign these fish to a specific taxon.We used the acronym ‘cf’ (Latin for
confer) between the genus name andthe species name to show that thespecies in question is similar to anamed species but there is uncertaintyabout its taxonomic status, or that thisspecies may represent a distinctunnamed species. Our overall objec-tive was to produce an operational andcomplete distributional compilation ofthe freshwater fish of Greece ratherthan a taxonomically more accuratebut incomplete distributional account.
e. Notations are given where uncertaintyexists about the native or introducedstatus of a population and where thepopulation is presumed extirpated orpossibly extirpated.
Annotated freshwater species checklist (inAppendix)
A critical assessment of the ichthy-ological data assists in the production of anupdated checklist of species known toinhabit freshwaters in Greece. As previ-ously mentioned, certain migratory andtransient marine species regularly presentin freshwaters are included but mostmarine species are excluded, since evi-dence is lacking on how frequent thesespecies are in the freshwater parts of thebasin areas. This checklist is still a prelimi-nary contribution for use in practical con-servation-relevant applications. Sweepingtaxonomic name changes have drasticallyaltered previously given names and abol-ished the use of sub-species names, so apractical updated list linking the previousand the current taxonomy is needed toclarify confusion. In this respect, notationon previous species names and new addi-tions to Greece’s species list are given. Forconservation purposes the level of‘endemicity’ of each species according to
Medit. Mar. Sci., 8/1, 2007, 91-166102
the scale of the Greek territory is summa-rized. Endnotes present special taxonomicproblems and distributional uncertainties.It must be made clear that in some casesthe available data is often of poor qualityand uncertainties exist; consequently,unreliable records and poorly validatedtaxonomic problems are inevitable. Thechecklist provides notes concerning ourprofessional opinion on these difficulties.
Supplementary list of species recorded intransitional waters (in Appendix)
A separate supplementary list ofspecies recorded in transitional waterbod-ies is also attempted (this includes rivermouths, coastal lagoons and the brackishreaches of lowland rivers). This is certainlya very rough provisional contributionneeding further research. References aregiven for each species that is included inthe list. This list does not include speciescited in the aforementioned checklist offreshwater species; therefore, many fresh-water species that have been recorded intransitional waterbodies are not included.This supplementary list attempts to pro-vide a more complete picture of the hydro-graphic basin-based species assemblage,since transitional waters are definitely apart of the river basin area unit. Some ofthese species are potential candidates forinclusion in the freshwater checklist whenfurther evidence is gathered.
Results
Hydrographic basin areas considered inthe survey
Table 1 lists the hydrographic basinareas considered in this study along withtheir main synonyms, the waterbodies
included in each area and the estimatedtotal surface area. The basin areas arearranged numerically according to theirposition on the orientation map of Greecewhich is given in Figure 1. Most of thebasin areas covered in this survey lie on thewestern, more humid side of the country(50 areas). Northern Greece covers a verylarge region with the predominance oflarge or very large hydrographic basins (26areas). Relatively few basins from the drierrain-shadow parts of the country (Central-Eastern Greece, Eastern Peloponnese, theAegean islands) are accommodated in thedataset, reflecting both the scarcity of larg-er perennial waterbodies and poor sam-pling effort (29 areas).
Composition and characteristics of theichthyofauna of Greece
All species known from freshwaters,native or introduced, are recorded in thechecklist of the freshwater fish fauna ofGreece (Appendix I). The supplementarylist of other euryhaline and marine speciesthat have been recorded in the publishedliterature from transitional brackish watersis provided in Appendix II.
The distributional compilation ofspecies occurrence data in 105 hydrograph-ic basin areas is presented in Tables 2-6.The compilation includes all species con-tained in the checklist, plus seven speciesthat have been reported from sections oftransboundary rivers outside Greek territo-ry; however, it excludes 9 euryhalinespecies contained in the checklist for whichthe distributional accounts in freshwatersare incomplete. Among these, mugilids(five species) have a widespread occur-rence in estuaries and lowland sections ofrivers but their presence is not regularly
Medit. Mar. Sci., 8/1, 2007, 91-166 103
reported or given at species level. The sameholds for Atherina boyeri, which is com-monly found in coastal lagoons, estuariesand the lower reaches of rivers. Notably,there are records of three landlockedsandsmelt populations in Greece, in LakesVoulkaria, Kourna and Trichonis respec-tively (LEONARDOS, 2001; TIGILIS,
2001; ECONOMOU et al., 2001b). Twomore species excluded from the compila-tion are Dicentrarchus labrax and Zosteriss-esor ophiocephalus, which are not a fre-quent target of faunistic investigations, andtheir occurrence in transitional and adjoin-ing freshwater waterbodies often goesunreported.
Medit. Mar. Sci., 8/1, 2007, 91-166104
Fig.1: Map of Greece showing the location of the hydrographic basin areas considered in the ichthy-ofaunal distributional compilation.
Medit. M
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Number 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 26Abramis brama 1 2 1 1Acipenser gueldenstaedtii 2Acipenser stellatus 1b,c 2b,c
Acipenser sturio 1c 1c 1c 1c 1c
Alburnoides bipunctatus (1) 1 1 1 1 1 1 1 1 1Alburnus alburnus 1a 1a
Alburnus macedonicus 1Alburnus sp. Volvi 1 1 1Alburnus thessalicus (1)a 1 1 1 1 1Alburnus vistonicus 1 1 1 1Alburnus volviticus 1Alosa fallax 1 1 1 1 1 1? 1Alosa macedonica 1Alosa vistonica 1Anguilla anguilla 1 1? 1 1? 1 1 1? 1 1 1 1 1 1 1c 1 1 1 1 1 1 1 1Aphanius fasciatus 1 1 1 1 1 1 1Aspius aspius 1 1 1 1Barbatula barbatula (1?)a,b,c (2) 1 1Barbus balcanicus 1 1 1 1 1 1 1 1Barbus cyclolepis 1 1 1 1 1 1Barbus macedonicus 1 1 1 1Barbus sperchiensis 1Barbus strumicae 1 1 1 1 1 1 1 1Carassius auratus 2
Evro
s
Avas
Filio
uri
Kom
psat
os
Vist
onis
Kos
sint
hos
Lasp
ias
Nes
tos
Mar
mar
as
Nev
roko
pi
Stry
mon
Lado
pota
mos
Mav
rola
kas
Aspr
olak
as
Rih
ios
Volv
i
Doi
rani
Axio
s
Anth
emou
ntas
Gal
likos
Loud
ias
Vego
ritis
Kas
tori
a
Alia
kmon
Mav
rone
ri
Pini
os T
he
Table 2Fish faunistic lists for the hydrographic basins of Northern Greece. The position of the basins is shown in Figure 1.
(continued)
Medit. M
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Number 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 26Carassius carassius 1 2Carassius gibelio 1b 2 1?b 2 2 1b 2 2 2 2 2 2 2 2 2Chondrostoma vardarense 1 1 1 1 1 1 1 1 1Cobitis puncticulata 1Cobitis punctilineata 1Cobitis stephanidisi 1Cobitis strumicae 1 1 1 1 1 1 1 1 1 1 1Cobitis vardarensis 1 1 1 1 1 1 1 1Coregonus cf. albula (2)a
Coregonus cf. lavaretus 2a 2a
Coregonus cf. peled (2)a
Ctenopharyngodon idella 2 2 2Cyprinus carpio 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 2b 1Esox lucius 1 (1) 1 1 1 1 1 1 1b 2 1 1Eudontomyzon hellenicus 1Eudontomyzon sp. 1a
Gambusia holbrooki 2 2? 2 2 2 2 2 2 2 2 2 2 (2?) 2 2Gasterosteus gymnourus 1 1 1 1 1 1 1 1? 1Gobio bulgaricus 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1Gobio feraeensis 1Huso huso 2b,c
Hypophthalmichthys molitrix 2 2 2Ictalurus punctatus (2)Ictiobus sp. (2)Knipowitschia caucasica 1 1 1 1 1 1 1 1 1 1 1 1 1
Table 2 (continued)
Evro
s
Avas
Filio
uri
Kom
psat
os
Vist
onis
Kos
sint
hos
Lasp
ias
Nes
tos
Mar
mar
as
Nev
roko
pi
Stry
mon
Lado
pota
mos
Mav
rola
kas
Aspr
olak
as
Rih
ios
Volv
i
Doi
rani
Axio
s
Anth
emou
ntas
Gal
likos
Loud
ias
Vego
ritis
Kas
tori
a
Alia
kmon
Mav
rone
ri
Pini
os T
he
(continued)
Medit. M
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Number 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 26Knipowitschia thessala 1Lepomis gibbosus 2 2 2 2 2 2 2 2Leucaspius delineatus 1 1 1 1 1 1 1Misgurnus fossilis (2?) (2?) (2)Oncorhynchus kisutch 2Oncorhynchus mykiss 2 2 2 2 2 2 2Oxynoemacheilus bureschi 1 1 1 1 1 1Pachychilon macedonicum 1 1 1 1 1? 1 1 1Perca fluviatilis 1 2 1 1c 1 1 1 2 1 1Petroleuciscus borysthenicus 1 1 1? 1 1? 1 1 1Petromyzon marinus 1Phoxinus cf. phoxinus (1)a 1a 1a 1a 1a 1a 1a
Phoxinus strymonicus 1Proterorhinus semillunaris 1 1Pseudorasbora parva 2 2 2 2 2 2 2Pungitius platygaster 1 1 1Rhodeus amarus 1 1 1 1 1 1 1 1 1 1 1 1Rhodeus meridionalis 1 1 1 1 1 1 1 1Romanogobio elimeius 1 1 1 1Rutilus rutilus 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1Sabanejewia balcanica 1 1 1a 1 1 1 1 1 1Salaria fluviatilis 1 1 1 1 1 1 1 1 1 1 1Salmo cf. macedonicus 1a 1a 1a (1)Salmo farioides 2 2Salmo pelagonicus 1 1 1a,c
Table 2 (continued)
Evro
s
Avas
Filio
uri
Kom
psat
os
Vist
onis
Kos
sint
hos
Lasp
ias
Nes
tos
Mar
mar
as
Nev
roko
pi
Stry
mon
Lado
pota
mos
Mav
rola
kas
Aspr
olak
as
Rih
ios
Volv
i
Doi
rani
Axio
s
Anth
emou
ntas
Gal
likos
Loud
ias
Vego
ritis
Kas
tori
a
Alia
kmon
Mav
rone
ri
Pini
os T
he
(continued)
Medit. M
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Number 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 26Salmo cf. trutta 2a
Salvelinus fontinalis 2 2Sander lucioperca 1 2Scardinius erythrophthalmus 1 1 1 1 1 1 1 1 1? 1 1 1 1Silurus aristotelis 2Silurus glanis 1 1? (1?) 1 1 1 1 1 1 1 1 1 1Squalius orpheus 1 1 1 1 1 1 1 1 1 1 1a 1a
Squalius vardarensis 1 1 1 1 1 1a 1 1 1 1Tinca tinca 1 1 1 1 1 1c 1 1 1b 2 1 1Vimba melanops 1 1 1 1 1 1 1 1 1Zingel balcanicus 1SUM 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
Table 2 (continued)
Evro
s
Avas
Filio
uri
Kom
psat
os
Vist
onis
Kos
sint
hos
Lasp
ias
Nes
tos
Mar
mar
as
Nev
roko
pi
Stry
mon
Lado
pota
mos
Mav
rola
kas
Aspr
olak
as
Rih
ios
Volv
i
Doi
rani
Axio
s
Anth
emou
ntas
Gal
likos
Loud
ias
Vego
ritis
Kas
tori
a
Alia
kmon
Mav
rone
ri
Pini
os T
he
1 = Native, confirmed presence in river basin; 1? = Presumably native, reported but unconfirmed presence; 2 = Introduced to the basin; 2? = Reported but unconfirmed introduction. The above symbols placed in parenthesis indicate occurrence only in sections of transboundary rivers outside the Greek territory.Notations are further given where taxonomic and native status uncertainty exists or where the population may be presumed extirpated, as follows: a : Doubtful taxonomic status of population; b : Doubts on native or introduced status; c : Extirpated or possibly extirpated population.
Medit. M
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Number 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46Acipenser baeri 2 2Acipenser gueldenstaedtii 2 2Acipenser naccarii (1)c 1b,c
Acipenser ruthenus 2Acipenser sturio (1)c 1?c
Alburnoides bipunctatus 1 1?Alburnoides prespensis 1Alburnus belvica 1Alburnus cf. scoranza 1a
Alosa fallax (1) 1 1? 1? 1Anguilla anguilla 1 1 1 1? 1 1 1 1 1 1 1 1 1 1 1 1 1Aphanius fasciatus (1) 1 1 1 1 1 1 1 1Barbus peloponnesius 1 1 1 1? 1 1 1 1Barbus prespensis 1Barbus rebeli 1Carassius auratus 2?a 2 2Carassius gibelio 2 2 2 2 2 2Chondrostoma prespensis 1Chondrostoma vardarense 1Clarias gariepinus 2Cobitis arachthosensis 1a
Cobitis hellenica 1a 2a 1a
Cobitis meridionalis 1Cobitis ohridana 1
Pres
pa
Aoos
Kal
amas
Zara
vina
Pam
votis
Para
myt
hia
Kal
odik
i
Ache
ron
Ziro
s
Lour
os
Arac
htho
s
Vouv
os
Vlyc
hos
Voul
kari
a
Asta
kos
Ache
loos
Evin
os
Mor
nos
Ker
kyra
Lefk
as
Table 3Fish faunistic lists for the hydrographic basins of Western Greece. The position of the basins is shown in Figure 1.
(continued)
Medit. M
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Number 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46Cobitis trichonica 1Coregonus cf. lavaretus 2a
Ctenopharyngodon idella 2 2 2? 2Cyprinus carpio 1 (2?) 2 2 2 2 2 2 2 2Economidichthys pygmaeus 1 2 1 1 1 1 1 1 1 1 1Economidichthys trichonis 1Esox lucius 2Eudontomyzon sp. Louros 1a
Gambusia holbrooki 2 (2) 2 2 2 2 2 2 2 2 2 2? 2 2Gasterosteus gymnourus 1 1Gobio cf. skadarensis 1a
Hypophthalmichthys molitrix 2 2 2Hypophthalmichhtys nobilis 2Ictalurus punctatus 2?Knipowitschia goerneri 1c
Knipowitschia milleri 1Knipowitschia sp. 1 2 1 1? 1 1 1 1 1?Lampetra sp. (1)Lepomis gibbosus 2 2Luciobarbus albanicus 1 1 1 1b 1 1 1 1? 1 1 1Micropterus salmoides 2Oncorhynchus kisutch 2Oncorhynchus mykiss 2 2 2 2 2 2 2 2Oreochromis niloticus 2? 2Oxynoemacheilus pindus 1
Table 3 (continued)
Pres
pa
Aoos
Kal
amas
Zara
vina
Pam
votis
Para
myt
hia
Kal
odik
i
Ache
ron
Ziro
s
Lour
os
Arac
htho
s
Vouv
os
Vlyc
hos
Voul
kari
a
Asta
kos
Ache
loos
Evin
os
Mor
nos
Ker
kyra
Lefk
as
(continued)
Medit. M
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Number 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46Pachychilon pictum 1Parabramis pekinensis 2Pelasgus epiroticus 1?a 1 1a 1?a,c
Pelasgus prespensis 1Pelasgus stymphalicus 2a 1 1 1 1 1 1 1a
Pelasgus thesproticus 1 1 1 1 1 1Perca fluviatilis 2Petromyzon marinus 1Poecilia sp. (2?)Polyodon spathula 2?Pseudorasbora parva 2 2 2Rhodeus amarus 1a 2a
Rutilus ohridanus (1)a
Rutilus panosi 2 1?a,c 1Rutilus prespensis 1Salaria economidisi 1Salaria fluviatilis 1 1 1 1 1 1Salmo dentex 1a
Salmo farioides 1 1 1c 1 1 1 1Salmo letnica 2Salmo peristericus 1Salmo sp. Louros 1a
Salvelinus fontinalis 2Scardinius acarnanicus 2a,b 1Silurus aristotelis 2 1
Table 3 (continued)
Pres
pa
Aoos
Kal
amas
Zara
vina
Pam
votis
Para
myt
hia
Kal
odik
i
Ache
ron
Ziro
s
Lour
os
Arac
htho
s
Vouv
os
Vlyc
hos
Voul
kari
a
Asta
kos
Ache
loos
Evin
os
Mor
nos
Ker
kyra
Lefk
as
(continued)
Medit. M
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Number 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46Silurus glanis 2 2 2Squalius cf. peloponnensis 1a 1a 1?a,b 1a 1a 1a 1a 1a
Squalius pamvoticus 1Squalius prespensis 1Squalius sp. Aoos 1Squalius sp. Evinos 1a 1a 1a
Telestes pleurobipunctatus 1 1 1 1b 1 1 1 1 1 1Tinca tinca 2 2 2 2 2 2 2Tropidophoxinellus hellenicus 1Valencia letourneuxi 1 1 1 1 1 1 1 1 1c 1c
SUM 25 24 20 9 23 1 2 15 5 22 20 3 8 8 4 39 13 10 10 5
Table 3 (continued)
Pres
pa
Aoos
Kal
amas
Zara
vina
Pam
votis
Para
myt
hia
Kal
odik
i
Ache
ron
Ziro
s
Lour
os
Arac
htho
s
Vouv
os
Vlyc
hos
Voul
kari
a
Asta
kos
Ache
loos
Evin
os
Mor
nos
Ker
kyra
Lefk
as
Notes as in Table 2
Medit. M
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Number 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79Acipenser sturio 1?c
Anguilla anguilla 1? 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1? 1 1Aphanius almiriensis 1Aphanius fasciatus 1 1 1Barbus peloponnesius 1? 1 1 1 1 1? 1 1 1 1 1 1 1 1 1 1 1Carassius gibelio 2 2?Ctenopharyngodon idella 2 2 2Cyprinus carpio 2 2 2 2? 2 2 2Gambusia holbrooki 2 2 2 2 2 2 2 2 2 2 2 2Gasterosteus gymnourus 1 1Hypophthalmichthysmolitrix 2 2Knipowitschia sp. 1 1 1Leponis gibbosus 2?Luciobarbus albanicus 1 2?Oncorhynchus kisutch 2Oncorhynchus mykiss 2 2 2 2 2 2Pelasgus laconicus 1 1 1 1a 1a
Pelasgus stymphalicus 2 1 1 1 1 1 1 1 1 1 1 1 1a 1a
Perca fluviatilis 2?Petromyzon marinus 1?Salaria fluviatilis 1? 1 1 1? 1 1 1 1 1 1 1 1 1Salmo farioides 1Salmo cf. trutta 2Silurus glanis 2Squalius cf. peloponnensis 1c 1 1 1 1 1b 1 1 1 1c 1a 1a 1a 1 2a
Eras
sinos
Arg
Kato
Alm
yri
Lern
i
Adre
li
Smyn
ous
Vass
ilopo
tam
os
Evro
tas
Taka
Stym
phal
ia
Fene
os
Kand
ila
Pam
issos
SW M
essin
ia
Peris
tera
s
Yian
nous
agas
Neda
Alfio
s
Pini
os P
el
Koty
chi
Prok
opos
Tsivl
os
Piro
s
Glaf
kos
Volin
aios
Phoe
nix
Meg
aniti
s
Selin
ous
Kero
nitis
Vour
aiko
s
Krat
his
Krio
s
Derv
enio
s
Asso
pos P
el
Table 4Fish faunistic lists for the hydrographic basins of the Peloponnese. The position of the basins is shown in Figure 1.
(continued)
Medit. M
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Number 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79Squalius keadicus 1 1Squalius moreoticus 1c
Telestes pleurobipunctatus 1 1 1Tinca tinca 2?Tropidophoxinellushellenicus 1
Tropidophoxinellusspartiaticus 1 1 1 1 1 1 1Valencia letourneuxi 1 1c
SUM 1 1 3 4 4 1 4 2 1 1 1 4 5 5 7 13 22 5 5 6 5 12 5 2 4 3 7 6 2 2 5 2 3
Table 4 (continued)
Eras
sinos
Arg
Kato
Alm
yri
Lern
i
Adre
li
Smyn
ous
Vass
ilopo
tam
os
Evro
tas
Taka
Stym
phal
ia
Fene
os
Kand
ila
Pam
issos
SW M
essin
ia
Peris
tera
s
Yian
nous
agas
Neda
Alfio
s
Pini
os P
el
Koty
chi
Prok
opos
Tsivl
os
Piro
s
Glaf
kos
Volin
aios
Phoe
nix
Meg
aniti
s
Selin
ous
Kero
nitis
Vour
aiko
s
Krat
his
Krio
s
Derv
enio
s
Asso
pos P
el
Notes as in Table 2
The distributional compilation indi-cates the presence of 162 species (includingdoubtful occurrences) in 105 basin areas.Of these species, 155 have been recorded
from hydrographic basins within the Greekterritory. Five of the species present in theNorth Aegean region, all introduced, donot occur in Greece and are known only
Medit. Mar. Sci., 8/1, 2007, 91-166 115
Number 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95Alburnoides bipunctatus 1a
Anguilla anguilla 1 1 1 1 1 1 1 1 1 1 1 1Aphanius fasciatus 1Barbus euboicus 1Barbus sperchiensis 1 1 1Carassius gibelio 2 2Ctenopharyngodon idella 2? 2Cyprinus carpio 2 2 2 2Gambusia holbrooki 2 2 2 2 2 2 2 2Gasterosteus gymnourus 1Hypophthalmichthys molitrix 2Hypophthalmichthys nobilis 2Knipowitschia caucasica 1 1Luciobarbus graecus 1 1 1Oncorhynchus kisutch 2Oncorhynchus mykiss 2 2a
Oreochromis niloticus 2Pelasgus marathonicus 1c 1 1 1 1c 1 1 1 1Poecilia cf. latipinna 2a
Pungitius hellenicus 1Rutilus sp. Sperchios 1Rutilus ylikiensis 1 1Salaria fluviatilis 1Salmo salar 2?Scardinius erythrophthalmus 1a
Scardinius graecus 1 1Silurus glanis 1?b
Squalius sp.1 1c
Squalius sp. Evia 1a 1Squalius vardarensis 1a 1a
Telestes beoticus 1c 1 1SUM 2 3 1 3 3 3 4 11 11 1 17 4 3 3 2 3
Voul
iagm
eni
Eras
sinos
Vra
Rafin
a
Kato
Sou
li
Mar
atho
n
Kifis
sos A
tt
Asso
pos B
eo
Kifis
sos B
eo
Ylik
i
Ther
mop
yles
Sper
chio
s
Chol
orem
a
Kire
as
Man
ikio
tiko
Dysto
s
Rigia
Table 5Fish faunistic lists for the hydrographic basins of Central-Eastern Greece.
The position of the basins is shown in Figure 1.
Notes as in Table 2
from Bulgarian sections of the rivers Evros,Strymon and Nestos (Coregonus albula, C.peled, Ictalurus punctatus, Misgurnus fossilis,Ictiobus sp.). Some characteristics of thefish assemblages in the designated biogeo-graphic regions are summarised in Table7a. The North Aegean region has the high-est (85 species) and the East Peloponneseregion has the lowest (6 species) speciesrichness. The total number of nativespecies with confirmed occurrence in theexamined basins is 130. 25 species wereinvariably assigned as introduced in all
areas of their occurrence; these arereferred to as aliens to the country (anoth-er two species were reported but their pres-ence requires confirmation). Ten of thesealiens have been established through natu-ral reproduction; the presence of theremaining 14 species in the wild dependson stocking and aquaculture escapes.Finally, 19 species have native populationsin Greece but are introduced in one ormore hydrographic basins outside theirnative range. These species are referred toas translocated, though there are instances
Medit. Mar. Sci., 8/1, 2007, 91-166116
Number 96 97 98 99 100 101 102 103 104 105Alosa fallax 1Anguilla anguilla 1 1 1 1 1 1 1 1?Aphanius fasciatus 1 1Barbus pergamonensis 1Carassius auratus 2?a
Carassius gibelio 2Ctenopharyngodon idella 2 2Cyprinus carpio 2? 2Gambusia holbrooki 2 2 2 2? 2 2 2 2Hypophthalmichthys molitrix 2 2Knipowitschia caucasica 1 1 1Ladigesocypris ghigii 1Oncorhynchus kisutch 2Oncorhynchus mykiss 2 2 2 2Oxynoemacheilus theophilii 1Petroleuciscus smyrnaeus 1Salaria fluviatilis 1 1 1Salmo salar 2Squalius cf. cii 1Squalius sp.2 1SUM 2 10 5 7 4 4 2 6 2 3
Sam
othr
aki
Lesv
os
Sam
os
Rho
dos
Alm
yros
Kou
tsou
lidis
Kou
rtal
iotis
Kou
na
Agia
Tavr
oniti
s
Table 6Fish faunistic lists for the hydrographic basins of the Aegean Islands.
The position of the basins is shown in Figure 1.
Notes as in Table 2
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* 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 7aSummary data of the ichthyofauna of Greece.
Fish occurrences in the hydrographic basin areas included in the distributional compilation.
Attributes
Designated regions
North Aegean1 South Ionian3 East Attiko- Aegean All
Adriatic2 Peloponnese4 Beotia5 Islands regions
Number of basins areas 26 2 48 3 16 10 105Number of fish taxa (species) 85 45 65 6 31 20 162Species within Greek territory(confirmed occurrences) 80 37 61 6 29 19 155
Species recorded only inneighbouring countries 5 6 - - - - 7(confirmed occurrences)Native species(confirmed occurrences)* 65 29 37 5 19 11 130
Introduced species(confirmed occurrences)** 20 14 24 1 10 8 25
Regional endemics (confinedto the biogeographic region) 29 18 28 1 9 1 86
Proportion of regionalendemics to the native fish 44.6 62.1 75.7 20.0 47.4 9.1 66.1fauna of the region (in %)Average basin area speciesrichness (confirmed occurrences) 18.0 23.0 6.9 3.0 4.4 4.2 9.1
Average basin area native speciesrichness (confirmed occurrences) 14.0 15.0 4.8 2.3 3.1 2.3 6.9
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) 161Typical freshwater species (do not readily enter seawater conditions) (Appendix I) 138Brackish water or marine species that spend part of their lives in freshwater conditions (Appendix I) 23Number of Greek endemics (species confined exclusively to Greece) (Appendix I) 47Number of Balkan endemics (species with a distribution restricted south of the Danube R.) (Appendix I) 28Number of "near endemic" species (species found along the frontiers of Greece) (Appendix I) 14Total number of fish species recorded in transitional waters but not confirmedas residents of freshwaters (Appendix II) 55
Total number of fish species recorded in fresh and transitional waters (Appendices I+II) 216
where the introduced specimens wereimported from abroad, rather than beingtranslocated from another Greek basin.
Taking into account the distributionalranges of the native species, 86 species arerecorded as regional endemics (their dis-tribution is confined to only one biogeo-graphical region). Some endemics areknown from single or very few basin areas.This category of range-restricted endemicsincludes species of high conservation pri-ority, such as Aphanius almiriensis, Alosavistonica, Barbus euboicus, Cobitisstephanidisi, Eudontomyzon hellenicus,Pungitius hellenicus and Squalius keadicus.A number of species have a distributionconfined to the frontiers of Greece withneighbouring countries (Lake Doiraini:one species; Lake Prespa: nine species).All regional endemics of the East Pelo-ponnese, Attiko-Beotia and AegeanIslands are entirely within Greek territory.The Ionian region has the highestendemicity level (75.7%) and the AegeanIslands the lowest (9.1%).
Table 7b summarises the fish dataappearing in Appendices I and II. The totalnumber of fish species recorded in thefresh and transitional waters of Greece is216. The checklist of freshwater fish species(Appendix I) contains 161 species that livein freshwaters, arranged in 28 families. Ofthese, the family Cyprinidae strongly dom-inates with 80 species that comprise 49% ofthe total number of species. Another fivefamilies (Acipenseridae, Cobitidae, Sal-monidae, Mugilidae, and Gobidae) arerepresented by 5 or more species. Twelvefamilies are represented by single species.Judging from spatial occurrence informa-tion, 139 taxa are provisionally classified astypically freshwater species (recorded pri-marily from freshwaters) and 23 are classi-fied as euryhaline species with a confirmed
presence in freshwaters. The supplemen-tary list (Appendix II) contains 55 specieswith a recorded presence in transitionalwaters, arranged in 22 families.
Table 8 shows the ten native and theten introduced species with the highest fre-quency of occurrence in the examinedbasins. Of the native species, Anguillaanguilla is the most widespread, reportedfrom 74 basins, followed by Salaria fluvi-atilis, which is known from 32 basins. Firstin the list of introduced species are Gam-busia holbrooki, with confirmed occurrencein 52 basins and Oncorhynchus mykiss, withoccurrence in 27 basin areas. The latterspecies has not yet been reported as repro-ducing in Greece and its occurrencedepends on stocking programmes andaquaculture escapes. Two alien species,Pseudorasbora parva and Lepomis gibbosus,are highly invasive and their distributionalrange is expanding.
A comparison of fish assemblage com-position among the six defined biogeo-graphical regions reveals that only 15native species, of the 130 native speciesrecorded, have joint presence in two ormore regions (see Table 9). Seven of thesespecies (Aphanius fasciatus, Knipowitschiacaucasica, Salaria fluviatilis, Anguillaanguilla, Alosa falax, Gasterosteus gymnu-rus and Petromyzon marinus) are second-ary freshwater or peripheral fish with pre-sumed ability to utilise the marine routefor their dispersal. One species, Cyprinuscarpio, with paired presence in the Adriat-ic region (Lake Prespa) and the NorthEast Aegean regions has been ommittedbecause its native status in Lake Prespa isnot certain. The high degree of faunisticdissimilarity among regions can only partlybe attributed to the fine-level taxonomyadopted in this study. The most probablereason of the dissimilarity is the presence
Medit. Mar. Sci., 8/1, 2007, 91-166118
of geographic barriers preventing faunalexchanges among regions.
Fish species richness and endemism
Hydrographic basin area species rich-ness (number of fish species per area) isgenerally low in Greece. The number ofnative freshwater fish species per area var-ied from 0 to 32 (Evros) and that of intro-duced species varied from 0 to 18 (Pamvo-tis and Acheloos). Only 53 hydrographicbasins, mostly small, had no record ofintroduced species. The areas with thehighest total basin richness (number ofnative and introduced species) are Evros(41), Strymon (40), Acheloos (38) and Ali-akmon (38). Figure 2 shows the distribu-tion of richness in the basin areas exam-ined, where richness was calculated sepa-rately for native, introduced and allspecies. More than half of the examinedbasin areas (59) host up to five species,with only five basin areas hosting morethan 35 species. If only the native speciesare considered, the number of basin areashosting up to five species rises to 65. Onlyten basin areas host more than 20 native
species. Basin area species richness isdetermined by a multiplicity of factors rep-resenting local and regional scales. Localinfluences on species richness include fac-tors determining habitat diversity andenvironmental stability, such as basin sur-face area, discharge level and variability,mean elevation and slope, presence oflakes or floodplains in the basin, etc. Herewe restrict ourselves to the examination ofthe relationship between basin nativespecies richness and basin surface area(see Fig. 3). Despite the high scatter ofpoints, the data show a clear increase ofrichness with basin area. The small size ofmost basins explains, at least partly, the lowaverage richness in Greek waterbodies.
On the regional scale, basin areaspecies richness reflects the pool ofspecies that occur in a biogeographicregion, as determined by a combinationof historical factors and contemporaryenvironmental influences. Other condi-tions being equal (e.g. when basins of sim-ilar size are compared), basin speciesrichness is associated with regionalspecies richness (defined as the numberof fish species known to occur within a
Medit. Mar. Sci., 8/1, 2007, 91-166 119
Table 8The 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 nativein some hydrographic basins.
Top 10 Native species Top 10 Introduced species
Species Catchments % Species Catchments %
Anguilla anguilla 74 70.5 Gambusia holbrooki 52 49.5Salaria fluviatilis 32 30.5 Oncorhynchus mykiss 27 25.7Squalius cf. peloponnensis 21 20.0 Carassius gibelio 20 19.1Barbus peloponnesius 21 20.0 Cyprinus carpio 18 17.1Aphanius fasciatus 21 20.0 Ctenopharyngodon idella 11 11.4Pelasgus stymphalicus 20 19.1 Hypophthalmichthys molitrix 10 10.5Cyprinus carpio 19 18.1 Lepomis gibbosus 10 10.5Knipowitschia caucasica 18 17.1 Pseudorasbora parva 10 10.5Rutilus rutilus 16 15.2 Tinca tinca 7 6.7Gobio bulgaricus 15 14.3 Oncorhynchus kisutch 5 4.8
biogeographic region). Inspection of thedata shows that the basins in the NorthAegean contain on average more speciesthan basins of similar size in the otherregions. This is largely a reflection of thericher ichthyofauna in the North Aegeanregion in comparison to other regions(see Fig. 4). The regional species richnessis particularly low in the Aegean islands,which are comprised of very small insularbasins. Low annual precipitation and thefrequent occurrence of prolongeddroughts may have contributed to species
extinctions and the depauperation of thelocal fish communities.
Figure 5 shows the spatial distribu-tion of fish endemicity. Within Greeceendemicity is very high (47 species foundexclusively in Greece; or 35% of its nativefish fauna). In some basin areas (e.g. Ach-eloos, Evrotas, Beotian Kifissos), the pro-portion of endemic fish exceeds 75% ofthe total number of native fish. The gen-eral trend is towards an increase of theproportion of endemics westwards andsouthwards. This trend is opposite to that
Medit. Mar. Sci., 8/1, 2007, 91-166120
Fig. 2: Frequency distribution of species richness in Greek hydrographic basin areas (all species, nativespecies and introduced species).
of species richness, which increases east-wards and northwards. Another 14species (10.3%) are considered ‘near-endemic’ since they inhabit isolatedwaters on the borders of Greece (Prespaand Doirani) or their range extends
slightly beyond the Greek territory(specifically in the Butrint basin in Alba-nia). Finally 28 species (20.6%) have awider distribution in the Balkan Peninsu-la south of the Danube, considered hereas Balkan endemics.
Medit. Mar. Sci., 8/1, 2007, 91-166 121
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 bio-geographical regions. Figures above columns indicate number of basin areas.
Discussion
Sources of bias in compiling the basinarea species lists and checklists – dataavailability, knowledge gaps and unmetneeds
The freshwater fish of Greece havebeen studied for more than 150 years. His-torical ichthyological information andearly records of commercial catches (e.g.VALENCIENNES, 1844; HELDREICH,1878; APOSTOLIDIS, 1883, 1892;ATHANASSOPOULOS, 1917, 1923,1925; KOLLER, 1927) provide a soundbasis for ascertaining the native distribu-tion of many species. However, the distri-bution of some species is still insufficientlyknown and their native or introduced sta-tus in some waterbodies is uncertain.
Despite the relatively large number ofpublications dealing with freshwater fish(ECONOMOU et al., 2004a), few onlyprovide complete fish faunistic lists in indi-vidual drainages (e.g. KATTOULAS,
1972; ECONOMIDIS et al., 1981; ECO-NOMIDIS & SINIS, 1982; KOKKI-NAKIS et al., 1999; TIGILIS, 2000;KOUTRAKIS et al., 2000; DAOULAS etal., 2001; ECONOMOU et al., 2001b,2004b; TACHOS, 2003; KOKKINAKIS,2006; STOUMBOUDI et al., 2006;LEONARDOS et al., 2007) or describethe distributional ranges of species andspecies groups (e.g. ECONOMIDIS, 1989;ECONOMOU, 2000; ECONOMIDIS etal., 2000b; BOBORI et al., 2001;DAOULAS, 2003; KALOGIANNI et al.,2006). An even smaller number of publica-tions take a synthetic approach to fish dis-tribution treating all species and/or com-piling basin-specific species lists over widegeographical areas (STEPHANIDIS,1939, 1950, 1971; ECONOMIDIS, 1974;DAGET & ECONOMIDIS, 1975;ECONOMOU et al., 1999, 2001a;ECONOMIDIS et al., 2001; BARBIERI etal., 2002). Two of the aforementionedworks, namely the thesis dissertations ofSTEPHANIDIS (1939) and ECONO-
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Fig. 5: Endemicity of the fish fauna relative to the territorial boundaries of Greece. Endemics confinedto 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 forcodes descriptions.
MIDIS (1974), deserve special mentionfor comprehensive taxonomic work anddetailed accounts of species occurrences ina large number of freshwater bodies. Acatalogue of the fish of Greece producedby ECONOMIDIS (1973) forms a land-mark in the ichthyological research ofGreece for reporting all freshwater fishoccurrences known at that time. A popu-larized check-list was later published bythe same author (ECONOMIDIS, 1991).
It is inevitable that some of our basinarea’s fish compilations may have errors ofomission. It is remarkable that with fewexceptions many smaller isolated aquaticsites, especially along Greece’s Aegeancoastline and its islands, have never beenproperly surveyed for fish. For example,our compilation provides data for onlyeight islands although wetland habitatsexist on a very large number of Greekislands. In a recent inventory of wetlandsites (CATSADORAKIS & PARAGA-MIAN, 2007), fish presence was recordedin 72 of 352 small wetlands in the Aegeanislands; unfortunately, however, these fishwere rarely identified to species level. Astark example of the extent of the unex-plored areas is given by the extent of wet-land exploration on the large island ofEuboea (3.685 Km2), which hosts a uniquefreshwater fish fauna that includes twolocal endemics. CATSADORAKIS &PARAGAMIAN (2007) provide descrip-tions of 39 wetlands on Euboea and theynote that this number is far from a com-plete inventory; we compiled ichthyofau-nal data of only four Euboean wetlands,two of which are not listed in the afore-mentioned inventory. The number of wet-lands on Euboea is certainly very large,and sadly we have only the slightest knowl-edge of freshwater fish distributions onthis island.
Another problem with our dataset –that again addresses unmet data needs – isthe unresolved issue of each basin area’sfish assemblage completeness. The dataprovided here sometimes refer to verylarge basin areas, thus much local informa-tion has been amalgamated, and importantsite-specific data is not presented (i.e. fishassemblages along a particular river seg-ment). We are almost certain that somespecies, especially introduced ones, aremissing from even the larger river basinarea accounts in our dataset. Also, somesmaller river basins, such as the Acheron,Kalamas, Kireas, and Lerna, do not yethave complete species lists. This is eitherbecause sampling efforts have been few orthe sampling conditions are especially dif-ficult due to deep-water, non-wadeablereaches which have never been surveyedwith appropriate tools (i.e. boat-basedelectrofishing, multi-mesh gill-nets etc).Lastly, there has been no recent samplingin some basin areas. For example, the mostreliable account of the Mornos R. fishdates back to 1972 (KATTOULAS, 1972)and is based on data obtained with old-fashioned sampling techniques that mayhave been ineffective for small-bodiedspecies likely to occur in the lower parts ofthe river. In addition, this investigationtook place before the huge wave of alienspecies’ introductions that occurred fromthe 1970’s through the 1990’s. Thus, theapparent absence of introduced species inthis system may reflect lack of recent fau-nistic information.
Apart from these distributional ambi-guities, there are also problems with occur-rence records of some species appearing inold publications but are not verified byrecent surveys. For example, we decided tonote our doubt for the occurrence ofAlburnoides bipunctatus in the Acheron R.,
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despite the fact that its presence in thisriver has been mentioned by STE-PHANIDIS (1939) on the basis of a singlespecimen collection. This species has notbeen found in the Acheron R. since thenand, more importantly, it is absent from allother river basins of the Ionian region,which substantiates our doubt. However,the Acheron R. is far too deep to be sam-pled efficiently with conventional samplingtechniques, so the presence of A. bipuncta-tus in this river cannot be excluded withcertainty. On similar grounds, we decidedto express doubt for the occurrence of Bar-bus peloponnesius in the Louros R., men-tioned in older surveys, because thisspecies is persistently absent in the collec-tions of extensive recent surveys.
In addition, the bibliography is rifewith ambiguities concerning site-specificinformation that would be of use to practi-cal management and conservation applica-tions (ECONOMOU et al., 2006). Difficul-ties in ‘deciphering’ locality informationfrom the bibliography are widespread andthey may also be a major reason for poten-tial errors. There is even difficulty withinterpreting the basin-specific location ofdata for many species. For example,ECONOMIDIS (1991) mention the exis-tence of Pseudophoxinus (now Pelasgus)stymphalicus on the island of Euboea with-out giving specific locality information.Other survey compilations have alsoencountered difficulty with unconfirmed orerroneous identifications of fish(MAURAKIS et al., 2003). Unfortunatelyit is very difficult to verify distributionalinformation, and errors can easily creepinto a dataset if the inventoried data arenot confirmed with recent field observa-tions. This shows that more field work isneeded to confirm and monitor the pres-ence of species in many river basins or sites.
Moreover, there are unresolved taxo-nomic issues with respect to the identity ofsome populations. Due to the difficulties inresolving taxonomy rapidly, many pastrecords of occurrence are difficult to sub-stantiate. For example, the species Pseu-dophoxinus stymphalicus has recently beensplit to five species under the new genusname Pelasgus (see KOTTELAT &FREYHOFF, 2007b). All previous pub-lished records referring to the nominatespecies P. stymphalicus are difficult to beincluded in a geographical compilationbecause the distributional limits of thenewly described taxa are not yet complete-ly defined. Are the specimens from the east-ern coast of the Peloponnese P. stymphali-cus, P. laconicus or P. marathonicus? Also,what is the translocated Pelasgus species inLake Pamvotis? Challenging problemswith the systematics and the distribution ofmany species remain to be resolved, such asconfirming the taxonomic validity of someunnamed and undescribed taxa within thegenera Squalius and Alburnus for whichmorphological characters alone do notallow reliable identification. Molecularstudies have greatly contributed to unravel-ing the genetic and the underlying phyloge-netic relationships of some species. Ongo-ing genetic work and other biological inves-tigations within the next few years shouldfocus on sorting relationships in taxa withdisjunct or fragmented distributions.
Finally, there is a lack of publishedsite-based ichthyofaunal surveys on transi-tional waters. This is a serious problembecause a large number of species useestuarine areas, rivermouths, lagoons andbrackish waters in deltas as rearing areasor seasonally as transient habitats. Ourbibliographic search showed that manyspecies have been recorded in transitionalwaters but many surprising records of
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marine species need verification (includingmarine stragglers such as Epinephelusaeneus, Scorpaena scrofa and Gaidropsarusmediterraneaus). We must reiterate theprovisional and tentative nature of oursupplementary list of species recorded inbrackish waters since 36 (56%) of thespecies that we present had been submit-ted by a single published bibliographic ref-erence. On the other hand, our list con-tains several species which possibly enterfreshwaters regularly. In fact, some speciesin this list were located in the lists of‘freshwater’ species by ECONOMIDIS(1991) and BOBORI & ECONOMIDIS(2006). Pending accurate detailed infor-mation on these and other species’ resi-dence in freshwaters we retain them onlywithin this provisional supplementary list.
The current ichthyological picture ofGreece
The last published annotated checklistof the freshwater fish of Greece contained105 species, including introduced, diadro-mous and euryhaline species with a regularpresence in freshwaters, and also fivespecies of doubtful occurrence (ECONO-MIDIS, 1991). Since then the list ofspecies has expanded considerably as sev-eral species have been recently described(e.g. KOTTELAT, 2004; KOTTELAT &BARBIERI, 2004; ECONOMIDIS, 2005;KOTTELAT & ECONOMIDIS, 2006;BOGUTSKAYA & ILIADOU, 2006;STOUMBOUDI et al., 2006; KOTTE-LAT, 2007; KOTTELAT et al., 2007).Some more taxa were named and desig-nated as species by KOTTELAT &FREYHOFF (2007a) mostly on the basisof morphological data. Molecular datahave contributed to some taxonomic clari-fication, especially when morphological
differences of diagnostic importance couldnot be established. For instance,KOTTELAT & FREYHOFF (2007) uti-lized evidence of genetic distinctivenessprovided by BOHLEN et al. (2006) toestablish Rhodeus meridionalis that other-wise would not be easily recognized as adistinct species on the basis of classical fea-tures such as morphology. Another causeof this increase of species number has beenthe introduction of several alien species tomany waterbodies for aquaculture andfishery enhancement (e.g. CRIVELLI etal., 1997; ECONOMIDIS et al., 2000a;LEONARDOS et al., 2007).
The present checklist of the freshwaterfish of Greece (Appendix I) compiles thefish data in a standardized way comparableto ECONOMIDIS (1991), i.e. it includesintroduced, diadromous and some euryha-line species. In total, the checklist contains161 species, of which 14 are of unclarifiedtaxonomic status and are given provisionalor generic names. The great increase inspecies numbers (56 species, 53%) since1991 has resulted mainly from taxonomicre-evaluations of existing taxa, rather thanfrom the genuine discovery of new species.In fact, most of the new species containedin the checklist were known as biologicalentities in 1991 but were recognized assubspecies or were lumped with theirclosely related species.
The apparent tendency towardsspecies splitting is in accordance with thecurrently prevailing trend towards adop-tion of the Phylogenetic Species Concept(PSC) over the older Biological SpeciesConcept (BSC). KOTTELAT (1997) hasstrongly advocated the use of the PSC(considered as equivalent to the Evolu-tionary Species Concept) for speciesrecognition and employed this concept in afirst revision of the systematics and
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nomenclature of the European freshwaterfish. This revision resulted in a greatincrease in the number of species inEurope with well over a hundred morespecies than previously recognised.Recently KOTTELAT & FREYHOFF(2007a) produced a handbook that furtherrevises the systematics of European fresh-water fish. This second revision has radi-cally changed the ichthyofaunal list ofGreece by introducing new names forspecies and genera and raising many pop-ulations and subspecies to species rank. Infact, 29 species from Greek freshwaters(18%) are ‘new’ since the ECONOMIDIS(1991) checklist was published. With thedemise of the subspecies unit, many sub-specific taxa have been lost while othershave been awarded species status. Overall,only 41% (67 species) of the species namesgiven in the present list are identical tothose used by ECONOMIDIS (1991).
Judging from trends in recent taxo-nomic publications, the new species con-cept is accepted by several taxonomistsworking with Greek freshwater fish and islikely to become the dominant conceptguiding fish systematics in the future(MILLER, 1998). The changes of speciesnames and the emergence of new speciesmay create problems to users of the fishdata who do not have a background in sys-tematics. We have attempted to resolvesome of the anticipated problems by quot-ing in the checklist of freshwater species(in Appendix I) the previous speciesnames and indicating new additions. Giventhat both the checklist and the distribu-tional compilation are based on the taxon-omy proposed by KOTTELAT &FREYHOFF (2007a), we consider itimportant to clarify the meaning of speciesunder the two species concepts and toexplore what impact the acceptance of the
PSC might have on the users of fish data.We shall begin with the definition ofspecies and the criteria used to delineatespecies boundaries under the two speciesconcepts, namely the BSC and the PSC.We shall also discuss the strengths andweaknesses of the two concepts and thepotential implications of the change inspecies concept on the uses (and users) offish data. Next, we shall provide anoverview of the distributional status of thefreshwater fish of Greece, particularly withrespect to regional patterns of richness andendemism. Last, we shall examine somepolicy-relevant implications of the fishdataset presented in this paper and weshall explore areas of research and man-agement priority.
Species concepts - distinctiveness criteriaand utility relative to the needs of users
How to define ‘species’ is one of themost fundamental and controversial issuesin biology (KULLANDER, 1999; BAR-TON, 2001; MAYDEN, 2002; REYDON,2005). More than 20 species concepts havebeen put forward (MAYDEN, 2002) andconsiderable debate still exists about theirtheoretical basis, applicability, and conse-quences for studies of ecology and biodi-versity (MAYR, 1996; HENDRY et al.,2002; ISAAC et al., 2005; AGAPOW &SLUYS, 2005). We confine ourselves to abrief presentation of the two prevailingspecies concepts, the BSC and the PSC,focusing on topics of relevance to biodiver-sity conservation and watershed manage-ment from a fish-based perspective. Moredetailed presentations and arguments infavour or against their conceptual andmethodological basis can be found in theaforementioned publications and also inKOTTELAT (1997), TURNER (1999),
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CRANDALL et al. (2000), RUFFING etal. (2002) and AGAPOW et al. (2004).
Both the BSC and the PSC seek topartition the natural variability observed inbiological communities into distinguish-able components (species) but differ overthe partitioning criteria and the charactersused to delineate species boundaries. TheBSC emphasizes reproductive compatibili-ty among individuals within and amongpopulations and defines a species as ‘agroup of interbreeding natural popula-tions which is reproductively isolated fromother such groups’ (MAYR, 1940). Thisdefinition is straightforward and providesone definitive criterion for assessing dis-tinctness of species – inability to inter-breed. Different species maintain theirgenetic integrity because gene flowbetween species is prevented by reproduc-tive isolation. One of the major criticismsof this concept is that interbreeding capac-ity can only be assessed in sympatry(MAYDEN & WOOD, 1995). In organ-isms with disjunct distributions the inter-breeding criterion becomes non-opera-tional, since there is no practical way totest whether individuals belonging toallopatric populations would be able tomate and produce viable and fertile off-spring. Due to the inability of assessingreproductive isolation in allopatric taxa,the distinctiveness of species is usuallyinferred indirectly, e.g. from observationof morphology and distribution. Tradition-ally, species have been defined by morpho-logical traits under the assumption thatmorphological variation reflects geneticvariation which, when sufficiently high,may cause reproductive isolation. The con-ceptual problem inherent in this assump-tion is that morphological changes pro-duced by variation and selection do notnecessary correlate with the genetic
changes that produce reproductive incom-patibility. Eventually, genetically distincttaxa may look very similar and, contrarily,large morphological differences may existbetween very closely-related taxa. TheBSC has also been criticised for its difficul-ty in dealing with introgressive hybridiza-tion and its inability to cope with asexualreproduction (TURNER, 1999).
The PSC emphasises membership in aunique genealogical (phylogenetic) lineageand defines a species as ‘the smallest diag-nosable cluster of individual organismswithin which there is a parental pattern ofancestry and descent’ (CRACRAFT,1983). Under this concept, criteria forspecies distinctiveness are monophyly (allmembers of a group are descended from asingle common ancestor) and autapomor-phy (presence of genetically-based charac-teristics shared by all members of the groupand not found in other groups). Subspeciesdo not exist according to the PSC. In com-parison to the BSC, the criteria employedto delineate species are more functionaland testable – for example, the problemat-ic notion of having to demonstrate repro-ductive isolation is avoided, while the exis-tence of phenetic dissimilarity is not a pre-requisite for considering genetically dis-tinct taxa as separate species. Major advan-tages of the PSC are its potential to handleasexual organisms and to delineate andclassify allopatric taxa. Main grounds forcriticism are the difficulty in demonstratingmonophyly, the tendency of PSC applica-tions to produce excessive ‘splitting’ ofspecies, the recognition of species irrespec-tive of the degree of phenotypic or geneticdivergence between evolutionary lines, andthe absence of standardised approaches forselecting traits and/or determining levels oftrait discreteness needed for diagnosingspecies-level differences. In fact, two
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groups would qualify for the status of sepa-rate species simply on the basis of differ-ences in any single character, morphologi-cal or genetic, provided that all individualswithin a group share one evolutionary line-age. Questions may therefore arise as towhich traits are appropriate for establish-ing species, whether species can be delimit-ed on the basis of genetic differences alone,and how much difference in the selectedtrait(s) is enough for distinguishing species.
Another problem associated with thePSC is how to deal with genetic introgres-sion, given the tendency of freshwater fishto hybridise even at the generic level(SMITH, 1993). For instance, how will ataxon formerly characterised as a distinctspecies be defined and named if extensiveintrogressive hybridisation has occurreddue to repeated stockings? In a strictapplication of the PSC, hybrid populationscannot be classified as belonging to anyspecies because they are polyphyletic atthe species level. In practical applicationsof the PSC, it is left to the taxonomists toevaluate evidence for the monophyly oflineages and to decide whether or not thephyletic integrity of the taxon has been dis-rupted by stocking beyond acceptable lim-its. A last issue of concern relates to phe-notypic plasticity. Many fish displayremarkable morphological variation, andit is rarely clear what portion of this varia-tion is based on inheritable genetic varia-tion and what portion reflects local envi-ronmental features. Although plasticitymay posses adaptive value and be undersome genetic control (BAMBER &HENDERSON, 1988; JENNINGS &BEVERTON, 1991; SWAIN et al., 1991),an extensive expression of plasticity maymislead taxonomic identifications. Cau-tion is therefore warranted when interpret-ing the results of comparisons of allopatric
populations, especially when: (a) taxa fromcontrasting environments show trait differ-ences that correlate with environmentaldifferences; (b) the comparisons involve alimited number of individuals and fewcharacters; and (c) the examined traitsshow slight differences and/or overlappingvalues. Again, it is left to competent taxon-omists to judge if the kind, amount anddirection of observed differences inspecies traits permit discrimination inspecies-level identifications.
Overall, application of the BSCencounters methodological difficulties,particularly with regard to demonstratingreproductive isolation in allopatric popula-tions. The PSC is equally vulnerable to var-ious methodological difficulties, such assample data variability and reliability onstatistical grounds, and problems in assess-ing phylogenetic lineages. It is also open tocriticism for the objectivity with which thespecies-discrimination criteria can be cho-sen or applied. Taxonomies based on theBSC contain few and often ambiguouslydefined species. However, these taxono-mies tend to maintain their stability overtime, because persistent and multiple-character differences must be demonstrat-ed before altering the taxonomic status oftaxa. By contrast, taxonomies based on thePSC usually contain more species and aresensitive to revisions and changes, becausethe species are more loosely and variouslydefined.
LEE & WOLSAN (2002) attempted toreconcile the apparently contradictoryviews concerning the conceptual frame-work and operational applicability of thetwo concepts, arguing that the BSC refersexplicitly to synchronic species, while thePSC refers explicitly to diachronic lineages.Therefore a biological species (synchronicspecies) and a phylogenetic species
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(diachronic lineage) are ontologically dis-tinct entities. The authors proposed that itmight be appropriate to use the termspecies solely for a synchronic, integratedassemblage of organisms, as defined by theBSC, and to apply the term lineage for adiachronic, non-integrated assemblagewhich is the historical product of past evo-lution. Nonetheless, most authors perceivespecies as both synchronic and diachronicentities. Inevitably, the conflict over themerits and practicability of the BSC andthe PSC will continue to persist. In Greece,a trend towards acceptance of the PSC isevident in almost all recent publications.Below we shall discuss the consequences ofthis trend for some applied biological disci-plines. After all, an important factor to beconsidered in systematics is how useful aspecies concept is relative to the needs ofthe users of fish data.
Taxonomic standardisation and specieslists. Many users of fish data (e.g. fisheriesmanagers and administrative officersinvolved in commercial catch statistics orregional development programmes) arenot interested in taxonomy per se but onthe applications of taxonomy. For theseusers, one of the more serious implicationsof adopting the PSC is that the fish specieslists are large, differ among basin areasand must change constantly. An additionaldifficulty arises when the species containedin the lists have been defined under differ-ent species concepts and with a mixture ofcriteria. This may also be a cause of insta-bility since any re-evaluation of taxadefined and named under the BSC mayresult in the definition of new species andnames under the PSC. Such taxonomicchanges may have a disturbing effect onmost users of fish data, who would like tosee the species list remaining constant
rather than changing each time a taxo-nomic study is undertaken.
Biodiversity conservation. The freshwa-ter fish fauna of Greece is characterized bya high level of endemism, which leads to aconcomitant need for a conservation focus.Given the high number of species in needof protection and the relatively poorresources available for biodiversity conser-vation, biologists and environmental man-agers are faced with the problem of decid-ing which taxa warrant special protectionusing criteria such as the magnitude of theextinction threat, ecological value or role inthe ecosystem and biological distinctive-ness. Assessing ‘biological distinctness’ is achallenging issue with biodiversity conser-vation. Several authors have argued thatthe PSC may promote conservation effortsbetter than other concepts (e.g.KOTTELAT, 1997; GOLDSTEIN et al.,2000). On the one hand, the PSC allows therecognition of genetically distinct popula-tions as evolutionarily significant unitsregardless of morphological similarity orthe ability to interbreed. On the otherhand, the taxonomic identification of vul-nerable populations makes them a cleartarget for conservation effort on politicalgrounds. Indeed, policy-makers recognizeand give more value to ‘distinct’ namedspecies than to lower taxonomic categories.For example, the Aphanius populationinhabiting the spring of Kato Almiri (NEPeloponnese) is more likely to be strictlyprotected than other Aphanius popula-tions, simply because it has been given thename A. almiriensis. Opponents of this viewconsider that adoption of the PSC mayhave little effect on conservation. Reasonsinvoked include the species-centred wasteof limited resources and the difficulty ofdeciding what should be conserved, given
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the proliferation of species under this con-cept; and, also the huge bureaucraticprocesses triggered to update conservationpolicies (e.g. GARNETT & CHRISTIDIS,2007). Taking position, we recognise thatthe PSC has various operational limita-tions, such as the problem of handlinghybridisations, and also that taxonomicclassifications are unstable. However, wetake into account that (a) many freshwaterfish populations in Greece have a long his-tory of isolation, often dating back to theMiocene, and thus may represent uniquegenetic units, deserving immediate man-agement consideration; (b) only ‘species’have high prominence under the nationaland the EU conservation legislations, and(c) many sites harbouring threatened fishpopulations or species (some until recentlyrecognized only as subspecies) have notbeen incorporated in officially designatedprotected-areas (e.g. under the NATURA2000 framework). On these grounds, weconsider that the PSC may better assist inefforts for conserving Greek freshwaterfish, despite its various limitations.
Ecological quality assessments. Todayone of the most active areas of ichthyolog-ical research in Greece is the use of ichthy-ological indicators for the assessment ofthe ecological status of freshwater water-bodies, in accordance with the provisionsof the EU Water Framework Directive2000/60. The principle behind the applica-tion of fish-based bioassessment method-ologies is that freshwater fish assemblagesreflect structural and functional aspects ofaquatic ecosystems. Therefore, the impor-tant issue for applying such methodologiesis not the genetic discreteness of taxa, butrather their ecological properties and abil-ity to diagnose ecological degradation.Many closely related taxa that are recog-
nised as separate species under the PSCexhibit a similar range of ecological char-acteristics and tolerances to a variety ofanthropogenenic disturbances (ECONO-MOU et al., 2007). For example, all Squal-ius species inhabiting Greek freshwaters inWestern Greece have similar ecologicalrequirements with respect to rheophilyand habitat use. Likewise, all Salmospecies of Greece exhibit similar require-ments with respect to habitat, thermal tol-erance and oxygen demands. Therefore, inichthyological research aimed at ecologicalquality assessments, high-level classifica-tions based on the BSC are more mean-ingful than the less inclusive fine-level clas-sifications based on the PSC.
In conclusion, it appears no speciesconcept can fulfil all of the differentrequirements posed by the various disci-plines and applications. Each has itsstrengths and weaknesses and may workbetter in some situations, while the othermay apply better in other situations. Giventhe different demands of potential users offish data, a possible solution is to use stan-dardised lists of species incorporating finerand coarser taxonomies suitable for differ-ent applications, so that studies based onsuch lists can at least be consistent. Never-theless, caution is needed in applicationsof the PSC concept in species descriptions;otherwise the taxonomy will become un-manageable and vulnerable to disruptivechanges.
Distribution pattern of native freshwa-ter fish
Ancient arrivals, travel routes and barriersto dispersal
Two major and perhaps non-exclusiveexplanations have been proposed for the
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arrival and dispersion of freshwater fishin the Balkans. One postulates that Euro-Siberian and Palaearctic species reachedthe area during Oligocene and Miocenetimes through river captures (ECO-NOMIDIS & B N RESCU, 1991). Asecond wave of arrivals of central Euro-pean and Danubian species that intrudedinto the area from the Danube and theBlack Sea during Pliocene or Pleistocenetimes is also postulated by the aboveauthors. The alternative hypothesis holdsthat colonisation of freshwater speciesaround the circum-Mediterranean mayhave occurred during a short period ofthe late Miocene when the Mediter-ranean dried up completely and then waspartially refilled with freshwater from theParatethys (BIANCO, 1990). It seemsthat no single explanation can account forthe diversity of the Balkan ichthyofauna,and of the Greek ichthyofauna in particu-lar (B N RESCU, 2004). Differentspecies may have arrived in differenttimes and through different pathways,and may have experienced variousdegrees fragmentation and isolation(ZARDOYA et al., 1999). Isolation, com-bined with complex climatic events, pro-moted speciation and produced a greatvariety of endemic taxa. During periodsof intense tectonism and marine regres-sion secondary contacts of previously iso-lated populations may have occurred,resulting to hybridization and geneticintrogression (e.g. DURAND et al.,2003). The following vicariant and disper-sal events have been proposed to accountfor the structural diversity and highdegree of endemicity of the Greek ichthy-ofauna (see ECONOMIDIS, 1974;ECONOMIDIS & B N RESCU,1991; ZARDOYA et al., 1999;DURAND et al., 2003; B N RESCU,
2004; BOBORI & ECONOMIDIS, 2006;SKOULIKIDIS et al., 2008): (a) the grad-ual uplift of the Alps and the BalkanMountains from late Oligocene to theend of the Miocene isolated the Balkandrainages preventing faunal exchangeswith the rest of Europe; (b) the rise of thePindos mountain range created a north-west-southeast barrier for fish rangeexpansions, while the rise of the MountOthrys cut the connections of the rivers ofcentral-eastern Greece from those ofnorthern Greece; (c) at thePliocene–Pleistocene boundary, a com-munication of the NW Aegean drainageswith the Danube R. was temporarilyestablished through a river-captureinvolving the Morava R. and the AxiosR.; (d) at about the same time, a similarcommunication of the Adriatic drainageswith the Danube R. was establishedthrough a river capture involving theOhrid-Drim-Skadar system in the area ofKosovo; (e) also in Plio-Pleistocenetimes, intrusion of Black Sea waters (thena freshwater lake) into the Mediter-ranean through the former Aegeopota-mos R. permitted dispersal of Black Seafreshwater fish to the NE Aegeandrainages; and (e) sea-level regressions atthe glacial maxima of the Pleistocene hada homogenising effect on fish assem-blages allowing dispersal among neigh-bouring river basins.
During the Pleistocene glaciations theGreek rivers remained free of ice, servingas refugia for the preservation of ancestralelements of the European ichthyofauna,which were eradicated from most otherparts of Europe. In the post-glacial times,the Greek rivers did not contribute to therecolonization of the European rivers withfreshwater fish; consequently, the fish fau-nas of the Greek rivers have retained their
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unique endemic forms which thus repre-sent distinct taxononomic entities differentfrom the ones in the rest of Europe.
Regional fish assemblages and endemicitypatterns
The present-day fish composition hasbeen determined by a combination of vic-ariant and dispersion events and faunalrelaxation by extinction episodes. Theuplift of the Pindic Cordillera in the Mid-dle Miocene (DERMITZAKIS & PAPA-NIKOLAOU, 1981) acted as a major fac-tor for the faunal divergence of the west-ern and eastern Greece. Consequently,two major aquatic biogeographical divi-sions are unanimously recognised, definedwith different names by various authors,and referred to here as the Aegean and theIonian divisions. However, opinions differover the number of minor divisions andtheir boundaries (STEPHANIDIS, 1939;BIANCO, 1986, 1990; ECONOMIDIS &B N RESCU, 1991; MAURAKIS &ECONOMIDIS, 2001; MAURAKIS et al.,2001). ECONOMIDIS & B NRESCU (1991) distinguished four mainichthyogeographic divisions (regions) inthe Balkans of which three encompass theGreek territory: the Ponto-Aegean (con-taining the subdivisions Thracian-EastMacedonia and Macedonia-Thessaly), theAttiko-Beotia, and the South Adriatic-Ion-ian. B N RESCU (2004) retained thisichthyogeographic scheme with someslight modifications, e.g. he charted theEast Peloponnese as a separate entity.
The fish distributional data presentedin the present paper corroborate theabove biogeographical separation ofGreece indicating the presence of charac-teristic endemics in each region (Appen-dix I) and a low degree of faunistic simi-
larity among regions (Table 9). Speciesrichness and endemicity levels also differamong biogeographic regions. The west-ern, central and southern parts of Greece(Ionian and Attiko-Beotia regions) holdan old and long-isolated ichthyofauna,and present low species richness but ahigh degree of endemicity. The northernand eastern parts of the country presenthigher richness and lower endemism,most probably because these parts are ingreat proximity to the dispersal areas ofthe Danube and the Black Sea. Speciesdepauperation in East Peloponnese andthe Aegean Islands makes faunistic com-parisons difficult and particularly chal-lenging. Nonetheless, the data indicate afaunistic distinctness of the Aegean regionthat most likely has its explanation inindependent origins of species. So far ithas been difficult to determine the faunis-tic relationships of the East Peloponnesewith the other regions because only few,poorly studied fish are present in thisregion. A general difficulty in performingbiologically relevant comparisons amongregions is that many areas in central andsouth-eastern Greece and in the AegeanIslands are in a bioclimatic semi-aridzone, where few species have survivedprolonged drought episodes or recenthuman water abstraction impacts.
The North Aegean basins harbourmany species with Black Sea and Danu-bian affinities (B N RESCU, 2004).Despite many common faunistic elementsover the entire North Aegean region (e.g.Scardinius erythrophalmus, Rutilus rutilus,Silurus glanis) the Thracian rivers (east-wards of the Strymon R.) and the Mace-donian-Thessalian rivers show ichthyologi-cal distinctiveness that would justify theirplacement in different ichthyogeographicregions. As pointed out by ECONOMI-
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DIS & B N RESCU (1991) most fishoccurring in the Thracian rivers are inhab-itants of still or slow-flowing waters andmay have reached the area chiefly from theBlack Sea during its freshwater phase.Nonetheless, dispersal opportunities haveexisted at least since the Miocene(SAKIN & YALTIRAK, 2005). Barbusstrumicae and Squalius orpheus are two ofthe most characteristic endemics of theThracian ichthyofauna.
The fish inhabiting the Macedonianand Thessalian rivers show distant affini-ties with Danubian fish and may haveentered the area via the Axios R. in Plio-Pleistocene times, as previously discussed.Indeed, several species endemic to thisarea (e.g. Cobitis vardarensis, Squalius var-darensis, Pachychilon macedonicum,Rhodeus meridionalis and Zingel balcani-cus) have sister group relationships withDanubian fish species. However, the firstarrivals might have occurred quite earlier,as it seems that the area had hydrological
contact with the former Paratethys Sea inMiocene times (e.g. SONNENFELD,2005). All Macedonian-Thessalian riversshow remarkable faunistic similarities, asexpected, given that these rivers were con-nected during the last glacial maximum(LYKOUSIS et al., 2005).
The Ionian region is considered as oneof the most isolated zoogeographic units inEurope, since it is blocked from the rest ofthe Balkans by mountain ranges. Thisregion contains unique endemics that areoften confined to one or few drainages(TSIGENOPOULOS & KARAKOUSIS,1996; BARBIERI et al., 2002;KETMAIER et al., 2003; MILLER et al.,2004a; ECONOMOU et al., 2004c).Species of some genera (e.g. Squalius,Scardinius and Barbus) show deep geneticdivergence from their counterparts in theBalkans and the rest of Europe and oftenhave a basal or almost basal position inphylogenetic reconstructions (DOADRIO& CARMONA, 1998; KETMAIER et al.,
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Table 9Faunistic similarities among biogeographical regions
(fish species with joint presence in more than two regions).
Fish species North Attiko- Ionian South East AegeanAegean Beotia Adriatic Peloponnese Islands
Alburnoides bipunctatusAlosa fallaxAnguilla anguillaAphanius fasciatusBarbus sperchiensisChondrostoma vardarenseGasterosteus gymnourusKnipowitschia caucasicaPelasgus stymphalicusPetromyzon marinusRhodeus amarusSalaria fluviatilisSalmo farioidesScardinius erythrophthalmusSqualius vardarensis
1998, 2003; DURAND et al., 1999a, 1999b;TSIGENOPOULOS & BERREBI, 2000;TSIGENOPOULOS et al., 2002). The dis-tinctness and ancient origin of the Ionianichthyofauna is further indicated by (a) thepresence of endemic genera (Tropidophox-inellus and Economidichthys) (BIANCO etal., 1987; STEPHANIDIS, 1974) and (b)the absence of widespread European gen-era that are typically present in otherBalkan regions (Chondrostoma, Barbatula,Gobio, Alburnus, Alburnoides, Phoxinus,Cottus and Rhodeus). Interestingly, someIonian species show much closer relation-ships to Iberian species than to speciesinhabiting other Balkan regions (DO-ADRIO & CARMONA, 1998; ZAR-DOYA et al., 1999; PERDICES et al.,1996; SANJUR et al., 2003). Geologicalevents that have contributed to isolationand speciation in the Ionian regioninclude: the early isolation of the southernpart of Peloponnese by mountain barriersand deep seas; the separation of the Pelo-ponnese by the opening of the CorinthianGulf during the early part of the LatePliocene; the progressive uplift of the Ion-ian islands throughout the Pliocene; andthe entrance of seawater in the PatraikosGulf during the Holocene. The existenceof paleo-lakes in Epirus, Acarnania, Arca-dia and Laconia allowed the maintenanceof old fish lineages. The confluence of theEpirus rivers, and the similar confluencesof the Acarnanian rivers, permitted faunalexchanges in the Pleistocene.
The breadth of fish diversity in theIonian region is easily underestimated ifone counts only the number of speciespresent. A larger and yet poorly exploredamount of diversity exists below thespecies level, and is represented by uniquephenotypes and genetic profiles oftenshowing a north-south clination. For
instance, the present-day distribution ofSqualius keadicus is restricted to Laconia(Evrotas and Vassilopotamos Rivers).However, there is evidence from geneticstudies that the historic range of thisspecies was wider and included rivers ofsouth-west Peloponnese from which it wasextirpated by introgression with newSqualius invaders (DURAND et al., 2000).
The Attiko-Beotia region is a diversearea which seems to be a true ‘geneticcrossroad’, as species have presumablyemigrated in both from the north, west andeast, however they have been isolated longenough to show differentiation and specia-tion. The rivers of this region share fewonly species with the North Aegean andthe Ionian rivers (Table 9) and are inhab-ited by a depauperate freshwater fishfauna that includes distinctive endemics.The fluvio-lacustrines Luciobarbus grae-cus, Scardinius graecus and Rutilus ylikien-sis are confined in the Kifissos R. systemand are probably remnants of the ichthy-ofauna of the ancient lake Kopais, nowdrained. Several species of the Attiko-Beotia region (of the genera Luciobarbus,Rutilus, Pelasgus, Telestes and Scardinius)have sister species in the Ionian region,which may reflect the past hydrologicalconnection of the Sperchios basin with theAmvrakikos Gulf in Miocene times.Today’s ichthyofauna is only a relict of thepast. An unclarified cyprinid, identified asa Squalius taxon, geographically isolatedfrom other Squalius taxa, inhabited theBeotian Assopos (STEPHANIDIS, 1974).This species presumably disappearedbefore it was scientifically studied and sim-ilar extinctions may have occurred in riversof Attiki and Euboea, which have beenseverely impacted by human activities.
Although the knowledge of theAegean island’s fish fauna is still incom-
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plete, the available evidence suggests thatthe eastern islands show faunal affinitieswith Asia Minor. A distinctive endemic inthis region is Ladigesocypris ghigii, inhabit-ing streams of Rhodes Island, which haslost its connection with Anatolia in thePliocene (DERMITZAKIS, 1990). Bycontrast, other eastern Aegean islands,including Lesvos and Samos, remainedconnected to Anatolia until Pleistocenetimes, and fish colonisations from themainland, especially during marineregressions, were possible. The ichthy-ofauna of the East Peloponnese isextremely depauperate and not yet prop-erly studied. In the 1990s an Aphaniuspopulation was discovered in the KatoAlmiri spring (ECONOMOU et al., 1997).This population was provisionallyassigned to A. fasciatus, though theauthors noted its morphological andbehavioural distinctiveness from the latterspecies. Later, this taxon was described asA. almiriensis (KOTTELAT et al., 2007).Whereas in the 1990s this species wasmoderately abundant, recent investiga-tions have failed to show its persistence inthe site.
It is safe to conclude that the remark-able diversity of the fish assemblagesamong biogeographic regions has a histor-ical explanation (vicariance and isolation).It is interesting that many cases of sharedspecies presences indicated in Table 9 con-cern secondary freshwater or peripheralfish with the ability to utilise the marineroute for their dispersal (Aphanius fascia-tus, Knipowitschia caucasica, Salaria fluvi-atilis, Anguilla anguilla, Alosa falax, Gas-terosteus gymnurus and Petromyzon mari-nus). This is especially the case in theAegean Island region, where all speciesshared with other regions are peripheral orsecondary freshwater species.
Species richness and species – area rela-tionships
High species richness generally corre-lates negatively with the degree ofendemicity, except in the Aegean Islandsand the East Peloponnese, where fish fau-nal depauperation does not permit mean-ingful comparisons. This is particularlytrue when the fish faunas of the Ionian andthe North Aegean regions are compared.Lower richness in the Ionian region (Table7a; Fig. 4) may have a historical explana-tion: the Ionian fish faunas have probablyremained isolated since the Miocene, andtherefore may have been subjected toextinction processes for a longer time thanthe North Aegean fish faunas, which hadPliocene or more recent contacts with theDanubian and Black Sea faunas. However,this explanation should be considered withcaution because the low species richness inthe Ionian hydrographic basins may be dueto ecological or physiographic factors. Ourdata show an increase of species richnesswith increasing basin size (see Fig. 3), con-firming long-standing generalisations thatmore species exist in larger basins, eitherbecause such basins contain a wider arrayof habitat configurations, or because theprobability of extinction is more likely insmall basins (REYJOL et al., 2006). Giventhe positive correlation between speciesrichness and basin size, a logical explana-tion of the lower richness of the Ionianbasins is that their size is much smallerthan the size of the North Aegean basins.However, local ecological conditions andlandscape features often disrupt the rich-ness-basin area correlation. For instance,the spring-fed Louros R. hosts a largernumber of species (14 species) than theadjacent and much larger nivo-pluvialArachthos R. (11 species). Presumably,
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hydrological stability and the presence ofmore extensive floodplain habitats in theformer river have resulted in a greateravailability of habitat types and/or higherability of species to persist in the longterm.
That both explanations may hold is notcontroversial. Richness may depend onboth demographic processes (colonisationand extinction events) and ecological orphysiographic factors. When basins of sim-ilar size are compared, those of the Ionianregion contain fewer species than basins ofthe North Aegean region, suggesting thathistorical factors and demographicprocesses had an important structuringeffect on regional fish assemblages. Withinregions, however, physiographic factors,such as basin size and relief, may betterexplain the observed richness patterns. Forinstance, most rivers of North Pelopon-nese are small and have high slope, lackingfloodplains; these rivers host only one ortwo fish species that are tolerant to flashyand erosive stream conditions. The depau-peration of the fish faunas of East Pelo-ponnese and the Aegean Islands can simi-larly be attributed to small basin size.Their isolated fish communities are partic-ularly vulnerable to repeated dryingepisodes, and many historical extirpationsmay have taken place.
Policy relevant implications of the survey
River basin area management and WFDapplication
As previously expressed, ecosystemhealth assessment and monitoring is oneaspect of aquatic conservation where fishplay an important role as biological indica-tors. Greece is lagging very much behind inthe application of the WFD assessment
scheme and an important reason is theinformation deficit on the organisms to beused as biotic quality elements (ECO-NOMOU et al., 2006). As of late 2007,Greece had not yet delineated river basinareas or constituent waterbodies, so eventhe basic geographic framework for eco-logical quality assessments is still missing.The river basin area distributions providedhere can contribute to bioassessment tooldevelopment in various ways, e.g. byenabling the characterisation of historicalreference conditions and the selection ofappropriate metrics. They also reveal twokinds of difficulties in building robust fish-based bioassessment indices. The first kindrelates to the faunistic idiosyncrasies andheterogeneity that characterised theGreek hydrographic basins (low basinspecies richness, high degree of endemici-ty, and varied basin-specific taxa assem-blages). Such aspects of the ichthyofaunalassemblages are characteristic of the high-ly heterogeneous Mediterranean-climateenvironmental conditions (GASITH &RESH, 1999; FERREIRA et al., 2007)and, among others, restrict the number ofpotential metrics or prevent the applica-tion of a common metric system over wideareas. The second kind of difficulties aris-es from taxonomic uncertainties, insuffi-cient knowledge of the species’ distribu-tions, and the wide tolerance of manynative species to varying environmentalconditions. The adoption of the phyloge-netic species concept (PSC) has exacerbat-ed the problems of reference conditionsand metric development because theapplication of a finer taxonomy generatesan ‘apparent’ increase of the biologicalheterogeneity among hydrographic basins.However, bearing in mind that phyloge-netically closely related species are morelikely to be ecologically similar, a solution
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to these problems is to identify ecological-ly-equivalent species that respond to eco-logical degradation in a similar manner.Base-line research projects for fish assem-blage community ecology is vital for estab-lishing and standardising lists of ecologi-cally-similar species.
Species conservation
Species entities are basic units in bio-diversity research and conservation appli-cations. Experience has shown that themore ‘distinct’ a species is, in relation toother species, the greater the priority isgiven to its protection (MAITLAND,2004). At the Mediterranean scale thenumber of threatened species is very largeand encompasses about 56% of theendemic fish; this is one of the highest pro-portional assessments of threatenedspecies worldwide (see SMITH &DARWELL, 2006; PETER, 2006). Thedistributional ranges of species are a keyissue for the assessment of their vulnera-bility and the design of protective meas-ures. In Greece, little work has been doneto systematically document the distribu-tions of endemic species; most studies pro-vide anecdotal accounts of species rangesand are not quantitative with respect toabundance or density (ECONOMIDIS,1995). Problems of this kind have impededconservation efforts. As a consequence,conservation priorities are usually basedon fish assemblage data available at thenational level (i.e. check-lists) or regionallevel (i.e. protected-areas), rather than ondetailed basin area-based fish compila-tions. Therefore, it is not surprising thatmany endangered fish species are still notgiven appropriate legal protection, e.g.they are not present in Annex II of theHabitats Directive. Besides, with the
recent name changes, we are faced with aremarkable number of localized endemicspecies as well as of taxa of unclear taxon-omy. Our basin survey identifies a largenumber of range-restricted species thatneed conservation attention. For somespecies complete distributional data areprovided that can assist in conservation re-assessments. For a number of otherspecies, however, the survey indicates gapsin the knowledge of their distributionand/or problems with their taxonomic sta-tus. Distributional surveys and taxonomicwork are essential for planning conserva-tion-orientated research.
Habitat conservation
While it is widely accepted that savingthe sites where vulnerable species live is avery important aspect for their conserva-tion, current legislation does not includesatisfactory provisions for sites and aquat-ic habitat types. For example, the HabitatsDirective does not provide adequatedescriptions or classifications of the arrayof aquatic habitat types which exist ininland freshwaters, since mostly terrestrialhabitat types are listed and targeted forconservation (DIMOPOULOS et al.,2005). This is unfortunate, because it iswell known that freshwater habitats belongto the world’s most threatened ecosystems(SMITH & DARWELL, 2006). Theseimportant weaknesses in current legisla-tive coverage or enforcement of protectionof aquatic habitats and their biota haveresulted in the ‘exclusion’ of many smalland highly vulnerable sites from the exist-ing protected-area network of Greece.Our basin-based compilation of fish distri-butions indicates the occurrence of basinscontaining potentially important evolu-tionary units. However, the compilation
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does not detail the exact limits of the geo-graphic distribution of each unit withinbasins. Further analyses and screening ofsite-specific data will be required to ensurethe preservation of independent geneticpools and to highlight unmet conservationgaps on a nation-wide scale.
Fishery management
Inland waters fisheries are facing sig-nificant problems due to mismanagement,water quality problems and the effects ofinvasive alien fish species (CRIVELLI etal., 1997; LEONARDOS et al., 2007).Destructive fishing practices lead to dam-aged fish habitat and less fish, and speciesintroductions and translocations causegenetic pollution, representing a majorreason of the degradation of native genepools. Misinformed fishermen and fish-eries managers are definitely the largestcause of alien fish species spread inGreece (ECONOMIDIS et al., 2000a). Inplanning and enforcing fisheries manage-ment policies, it is important that reliabledata on the composition of fish assem-blages and the native ranges of species areavailable. The present dataset providesbaseline ichthyological information thatmay help to track the spread of alienspecies and to report vital ichthyologicaldata in a standaridised manner. However,commercial and sport fishing have majoreconomic and political implications thatalso need to be taken into account in thewatershed management plans. While it isclear that holistic approaches coveringsocial, economic, environmental and tech-nical aspects should be used to promotefishery management, biodiversity issuesshould not be sacrificed for the sake ofdevelopment; the conservation value ofspecies and habitats should be given at
least as much importance as economicand social factors.
Biogeography
Since freshwater fish are restricted topathways offered by hydrogeographic sys-tems, their distribution largely reflects his-torical patterns of drainage connections(e.g. VARGAS et al., 1998; REYJOL etal., 2006). Moreover, fish do have betterdocumented distributional informationthan do most other freshwater-obligateorganisms in Greece (i.e. invertebrates,amphibians); therefore, they are capableof supporting biogeographical analyses(LEGAKIS, 2004). The influence of his-toric drivers determining fish distributionis especially important in the speciesextinction-invasion process. Many biogeo-graphic applications can be developedwhen a complete inventory of fish speciesoccurring in the hydrographic basins ofGreece will be created (for instance,assessing the relative importance of his-torical factors and physiographic orhydrological characteristics in determin-ing basin area fish assemblages). Weacknowledge that our database still needsverification and completion; neverthelessit is a first step in developing a nation-wide basin-based inventory that can beused for conservation-relevant biogeo-graphical research.
Research and management priorities
Below we summarize five imperativeactions with respect to conservation-rele-vant ichthyological research in Greece:
1. Fish distributional surveys. Our distribu-tional data clearly suggest that conserva-tion and management programmes should
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refer to the geographic ranges of each tax-onomic unit to ensure the preservation ofindependent genetic pools. Distributionaldata are also critical for the implementa-tion of the WFD (establishment of refer-ence conditions, metric selection, site-based index development), the HabitatsDirective (site protection and monitoring,populations reporting, conservation man-agement) and basic environmental impactassessment. Likewise, fisheries manage-ment should be seen as a conservationissue that needs planning and enforcementon the basis of distributional and informa-tion assemblage structure. Ichthyologicalresearch is especially needed in smallwater features such as springs, wetlandsand coastal lagoons, as well as in deep sec-tions of large rivers, which have not beenadequately sampled. An important initia-tive to integrate the aforementioned needswould involve a coordinated atlas projectfor freshwater fish in Greece.
2. Fish taxonomy - genetic research. Taxon-omy and conservation must come togeth-er; and taxonomists who are motivated byconservation action must strive to producereliable standardized taxonomic units.Obviously, genetic research is critical intaxonomy. A community of taxonomistsmust develop and a forum should be creat-ed in order to help establish the validity oftaxa (CRIVELLI & MAITLAND, 1995).Genetic variation in any species being con-served or managed is very important tomonitor. Without this basic research it isimpossible to effectively manage species,populations or communities of fish. Someisolated populations are poorly studied,and yet they may represent cryptic endem-ic species that may be worth protecting assignificant evolutionary units (e.g.Alburnoides bipunctatus in the Sperchios
R. and various Knipowitschia populationsin western and central-eastern Greece).Particular attention is needed in theboundary areas between species wheregenetically distinct units can occur. Forexample, the progress of recent Squaliuspeloponnensis populations and the colo-nization of new aquatic systems might havetaken place in successive waves. Perhaps insome systems of southern Peloponnese(e.g. in the SW Messinia streams or in themontane plateau of the Lousios R.) thenew invaders found old local Squaliusstocks and either eliminated them ormated with them. Similar hybridisationzones may exist in the boundaries ofspecies belonging to the genera Salmo andPelasgus.
3. Aquatic habitat inventory. Habitat loss isthe most important conservation problemfor fish species, and this is especially acutein seasonally-semi-arid environmentswhere many small watersheds are vulnera-ble to human pressures. Greece and thewestern Balkans have one of the largestconcentrations of range-restricted species;many species are restricted to one or tworiver basin areas – some are confined tocertain river segments or special habitattypes (spring-fed wetlands, lakes, pondsetc). Coordinated efforts are needed todocument aquatic habitats in an inventoryand to create wide-ranging campaigns fortheir preservation and restoration. As stat-ed by CRIVELLI & MAITLAND (1995),the conservation of freshwater habitats ismore important than that of individualspecies; but to conserve these effectivelywe need: a) a list of all aquatic habitatareas, b) priority ranking and conservationevaluation of the habitat areas, c) conser-vation management plans for sites and thefish species accommodated within them.
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4. Alien species control. Alien species arewidely considered as the second mostimportant threat to aquatic biodiversityafter habitat loss. Awareness and enforce-ment of the control of the spread of intro-duced species is critical for conservation.We have provided evidence that 25 alienspecies have been introduced to Greece ofwhich some are spreading fast, seriouslyimpacting the natural biota. Translocationof fish among basins is an equally seriousproblem, since they are usually performedwithout any concern for the evolutionaryhistory of the species (for trout species: seeAPOSTOLIDIS et al., 1999). The impactof translocations on genetic diversity mayexceed the impact of alien species intro-ductions due to the high possibility ofintrogressive hybridization between popu-lations or closely related species. In thelight of the recorded human effect on thedistribution of the different species, it isdesirable that both the donor populationsand the indigenous populations in therecipient areas would be geneticallyscreened before any introduction. Publicawareness of this problem is extremelyimportant. A research priority is to identi-fy areas hosting unaffected remnantindigenous fish stocks that can be pre-served and used as a population source forrehabilitation projects.
5. River basin management plans. A multi-tiered approach to biota and habitat pro-tection must be incorporated in river basinmanagement plans. Integrated and holisticplanning is needed to co-ordinate waterresource exploitation, conservation andrestoration in basins; however, at thisscale, biodiversity is not often given appro-priate consideration. Indeed, as alreadystated, traditional protected-areas oftendisregard fish and other elements of the
aquatic biota. It is therefore important topromote legal protection schemes and spe-cial management initiatives to keep aquat-ic habitats in existence (APERGHIS &GAETHLICH, 2006). Carefully sited pro-tected-areas are needed in order to coverlinear aquatic features or focus on relative-ly small sites and threatened populationrefuges (e.g. MOYLE & YOSHIYAMA,1994). Micro-reserves may be effective as ashort-term direct protective measure.
Conclusion
Hydrographic basin areas are of highrelevance to current water policy and con-servation. Such areas have well definedboundaries (watershed limits) and arebecoming important for effective aquaticecosystem monitoring, assessment, report-ing and management. In this work, a basinarea survey method was employed to com-pile the best-available distributionalassemblage data for freshwater fish inGreece. Our work is a preliminary butwide-ranging attempt that may help toidentify: (a) unmet needs in our under-standing of freshwater species’ distribu-tions in Greece, (b) problems with thespecies’ recent taxonomic changes, and (c)basic gaps in science-based conservationwork on threatened species. Indeed, oneof the major obstacles in effectively assess-ing the conservation status of fish, theirhabitat needs, and the anthropogenic pres-sures they may face, is the large gap inknowledge of their geographical distribu-tions.
A coordinated effort is needed to pro-mote field ichthyology as a scientific enter-prise that is directly useful to nature con-servation. There is still poor baselineknowledge of Greece’s inland water aquat-ic ecosystems. The number of different
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aquatic habitat areas in Greece is large; asgeographic entities, aquatic habitat sitescertainly number in the thousands. Wehave very little information for many of thesmaller more isolated ones, and the quali-ty of the information for many parts of thelarger river basin areas is also poor.Reviews from other inland water assess-ments from the Mediterranean have simi-larly depicted the problem of importantgaps in the inventory of ecosystems andtheir aquatic biota (ALVAREZ-COBELAS et al., 2005), so this is not aproblem unique to Greece. However, theichthyofauna of Greece is especially rich inmany range-restricted species and needsimmediate attention. Particular problemsconcern conservation-relevant aspects ofits taxonomy, biology and ecology. Impor-tant problems also exist with informationmanagement and dissemination. Thiswork also underlines the urgent need forbuilding site-based inventories of fishassemblages in this country.
Acknowledgements
This work is a product of extensive col-laboration with many scientists and natu-ralists who have shared data and reviewedfish species lists with us during the last fiveyears. The authors have greatly benefitedfrom the landmark works of P.S. Econo-midis; constructive advice and discussionswith him are highly appreciated. Notableassistance was provided by D. Bobori, D.Mdrak, A. Flloko, A. Apostolou, A.Legakis, P. Dimopoulos, Th. Naziridis, K.Poirazidis, Y. Kazoglou, Ch. Papaioannou,Y. Leonardos, Y. Roussopoulos, C.G.Papaconstantinou, A. Crivelli, G. Catsado-rakis, A. Apostolidis, G. Tingilis, Y.Reyjol, M.T. Ferreira and A. Vlamis-Gardikas who contributed to the biblio-
graphic compilation with references, dis-cussions or fish data. Many colleagueshelped in fieldwork; we are indebted to U.Dussling, K. Blasel, G. Dal ge, R. Bjork-land, P.J. Miller, B. Zimmerman, M. Kot-telat and W. Beaumont for assistance inwide-ranging surveys. At HCMR we espe-cially thank colleagues Y. Chatzinikolaou,V. Tachos, E. Kalogianni, C. Daoulas, T.Psarras, N. Koutsikos and D. Kommatas,who participated in field surveys, and N.Skoulikidis, A. Zenetos, P. Drakopoulouand E. Moussoulis, who contributed inmany other ways. We express our thanksto E. Economou, who coordinated data-quality control and detailed referencecross-checking. We appreciate the graphicwork done by A. Vidalis. Thanks are alsodue to E. Tzovara for her time andpatience in editing this manuscript. Final-ly, we are grateful to W. Beaumont forconstructive comments and editing and toB. Zimmerman and Z. Kaczkowski forreviewing drafts of this paper. Obviously,any errors during this compilation and itspresentation are the responsibility of theauthors or are based on their opinion atthe time of writing. This work was partial-ly funded through a project of the HellenicMinistry of Development and we aregrateful to M. Ghini for her assistance inthis cooperation. Lastly, the Departmentof Environment and Natural ResourcesManagement at the University of Ioanninasupported electrofishing field survey workinvolving the last author.
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Medit. Mar. Sci., 8/1, 2007, 91-166154
Accepted in 2007
Medit. Mar. Sci., 8/1, 2007, 91-166 155
12
34
56
Nam
eAu
thor
ityPr
evio
us N
omen
clat
ure
Fres
hwat
erEn
dem
icPe
trom
yzon
idae
1Eu
dont
omyz
on h
ellen
icus
1V
lady
kov,
Ren
aud,
Kot
t &
NO
CH
AN
GE
1EN
NO
Econ
omid
is, 1
982
2Eu
dont
omyz
on sp
.Lou
ros2
Und
escr
ibed
Eudo
ntom
yzon
hell
enic
us1
(EN
WE)
3Pe
trom
yzon
mar
inus
Linn
aeus
, 175
8N
O C
HA
NG
E0
OTH
ERAc
ipen
seri
dae
4A
cipe
nser
bae
ri Br
andt
, 186
9N
O C
HA
NG
E(1
)A
LN5
Aci
pens
er g
ueld
ensta
edtii
Br
andt
& R
atze
burg
, 183
3N
O C
HA
NG
E0
ALN
6A
cipe
nser
nac
carii
Bona
part
e, 1
836
NO
CH
AN
GE
0O
THER
7A
cipe
nser
ruth
enus
Linn
aeus
, 175
8N
O C
HA
NG
E(1
)A
LN8
Aci
pens
er st
ellat
us3
Palla
s, 17
71N
O C
HA
NG
E0
OTH
ER9
Aci
pens
er st
urio
Linn
aeus
, 175
8N
O C
HA
NG
E0
OTH
ER10
Hus
o hu
so4
(Lin
naeu
s, 17
58)
NO
CH
AN
GE
0(A
LN)
Poly
odon
tidae
11Po
lyodo
n sp
athu
la(W
alba
um, 1
792)
NEW
1A
LNAn
guill
idae
12A
ngui
lla a
ngui
lla(L
inna
eus,
1758
)N
O C
HA
NG
E0
OTH
ERC
lupe
idae
13A
losa
falla
x(L
a C
epèd
e, 1
803)
Alo
sa fa
llax
nilo
tica
0O
THER
14A
losa
mac
edon
ica
(Vin
cigu
erra
, 192
1)A
losa
(Cas
pial
osa)
mac
edon
ica
1EN
NO
15A
losa
vist
onic
aEc
onom
idis
& S
inis,
198
6A
losa
(Cas
pial
osa)
cas
pia
visto
nica
0
ENN
OC
ypri
nida
e16
Abr
amis
bram
a(L
inna
eus,
1758
)N
O C
HA
NG
E1
OTH
ER17
Alb
urno
ides
bip
unct
atus
(Blo
ch, 1
782)
NO
CH
AN
GE
1O
THER
18A
lbur
noid
es p
resp
ensis
(Kar
aman
, 192
4)A
lbur
noid
es b
ipun
ctat
us p
resp
ensis
1N
ENN
O
19A
lbur
nus a
lbur
nus
(Lin
naeu
s, 17
58)
Alb
urnu
s alb
urnu
s the
ssal
icus
, Alb
urnu
s alb
urnu
sm
aced
onic
us1
OTH
ER
20A
lbur
nus b
elvic
aK
aram
an, 1
924
Cha
lcal
burn
us b
elvic
a1
NEN
NO
21A
lbur
nus c
f.sc
oran
za5
Bona
part
e, 1
845
NEW
1EN
BAL
App
endi
x I.
Ann
otat
ed C
heck
list
of F
resh
wat
er F
ishe
s of
Gre
ece.
(con
tinu
ed)
Medit. Mar. Sci., 8/1, 2007, 91-166156
12
34
56
Nam
eAu
thor
ityPr
evio
us N
omen
clat
ure
Fres
hwat
erEn
dem
ic22
Alb
urnu
s mac
edon
icus
Kar
aman
, 192
8A
lbur
nus a
lbur
nus m
aced
onic
us1
NEN
NO
23A
lbur
nuss
p. V
olvi
6In
K&
F200
7N
EW1
(EN
NO
)24
Alb
urnu
s the
ssal
icus
7St
epha
nidi
s, 19
50A
lbur
nus a
lbur
nus t
hess
alic
us1
ENBA
L25
Alb
urnu
s vist
onic
usFr
eyho
f & K
otte
lat,
2007
Cha
lcal
burn
us c
halc
oide
s mac
edon
icus
1EN
NO
26A
lbur
nus v
olvi
ticus
Frey
hof &
Kot
tela
t, 20
07C
halc
albu
rnus
cha
lcoi
des m
aced
onic
us1
ENN
O27
Asp
ius a
spiu
s(L
inna
eus,
1758
)N
O C
HA
NG
E1
OTH
ER
28Ba
rbus
bal
cani
cus8
Kot
lik,
Tsig
enop
oulo
s, R
a’ b &
Ba
rbus
pelo
ponn
esiu
s pet
enyi
1O
THER
Berr
ebi,
2002
29Ba
rbus
cyc
lolep
isH
ecke
l, 18
37Ba
rbus
cyc
lolep
is cy
clol
epis
1EN
BAL
30Ba
rbus
eub
oicu
sSt
epha
nidi
s, 19
50N
O C
HA
NG
E1
ENC
E31
Barb
us m
aced
onic
usK
aram
an, 1
928
Barb
us b
arbu
s mac
edon
icus
, Bar
bus b
arbu
s the
ssal
icus
1EN
BAL
32Ba
rbus
pelo
ponn
esiu
s9V
alen
cien
nes,
1842
Barb
us p
elopo
nnes
ius p
elopo
nnes
ius
1(E
NW
E)33
Barb
us p
erga
mon
ensis
Kar
aman
, 197
1N
EW1
OTH
ER34
Barb
us p
resp
ensis
Kar
aman
, 192
4N
O C
HA
NG
E1
NEN
NO
35Ba
rbus
rebe
liK
olle
r, 19
26Ba
rbus
pelo
ponn
esiu
s reb
eli1
ENBA
L
36Ba
rbus
sper
chie
nsis10
Step
hani
dis,
1950
Barb
us c
yclo
lepis
sper
chie
nsis,
Bar
bus c
yclo
lepis
1EN
NO
chol
orem
atic
us37
Barb
us st
rum
icae
Kar
aman
, 195
5Ba
rbus
cyc
lolep
is str
umic
ae1
ENBA
L38
Car
assiu
s aur
atus
11(L
inna
eus,
1758
)N
EW1
ALN
39C
aras
sius c
aras
sius
(Lin
naeu
s, 17
58)
NO
CH
AN
GE
1O
THER
40C
aras
sius g
ibeli
o12(B
loch
, 178
2)C
aras
sius a
urat
us g
ibeli
o1
OTH
ER41
Cho
ndro
stom
a pr
espe
nsis
Kar
aman
, 192
4N
O C
HA
NG
E1
NEN
NO
42C
hond
rosto
ma
vard
aren
seK
aram
an, 1
928
Cho
ndro
stom
a va
rdar
ensis
1EN
BAL
43C
teno
phar
yngo
don
idell
a(V
alen
cien
nes,
1844
)N
O C
HA
NG
E1
ALN
44C
yprin
us c
arpi
o13Li
nnae
us, 1
758
NO
CH
AN
GE
1O
THER
45G
obio
bul
garic
usD
rens
ky, 1
926
Gob
io g
obio
bul
garic
us, G
obio
gob
io b
alca
nicu
s1
ENBA
L46
Gob
io c
f.sk
adar
ensis
14K
aram
an, 1
936
NEW
1EN
BAL
47G
obio
fera
eens
isSt
epha
nidi
s, 19
73G
obio
gob
io fe
raen
sis1
ENN
O48
Hyp
opht
halm
icht
hys m
olitr
ix(V
alen
cien
nes,
1844
)N
O C
HA
NG
E1
ALN
App
endi
x I
(con
tinu
ed)
(con
tinu
ed)
Medit. Mar. Sci., 8/1, 2007, 91-166 157
12
34
56
Nam
eAu
thor
ityPr
evio
us N
omen
clat
ure
Fres
hwat
erEn
dem
ic49
Hyp
opht
halm
icht
hys n
obili
s(R
icha
rdso
n, 1
845)
NO
CH
AN
GE
1A
LN50
Ladi
geso
cypr
is gh
igii
(Gia
nfer
rari,
192
7)La
dige
socy
pris
ghig
ii gh
igii
1EN
AEG
51Le
ucas
pius
deli
neat
us(H
ecke
l, 18
43)
NO
CH
AN
GE
1O
THER
52Lu
ciob
arbu
s alb
anic
us(S
tein
dach
ner,
1870
)Ba
rbus
alb
anic
us1
ENW
E53
Luci
obar
bus g
raec
us(S
tein
dach
ner,
1896
)Ba
rbus
gra
ecus
1EN
CE
54Pa
chyc
hilo
n m
aced
onic
um(S
tein
dach
ner,
1892
)Ru
tilus
mac
edon
icus
1EN
BAL
55Pa
chyc
hilo
n pi
ctum
(Hec
kel &
Kne
r, 18
58)
Pach
ychi
lon
pict
us1
ENBA
L56
Para
bram
is pe
kine
nsis
(Bas
ilews
ky, 1
855)
NO
CH
AN
GE
1A
LN57
Pela
sgus
epi
rotic
us
(Ste
inda
chne
r, 18
96)
Para
phox
inus
epi
rotic
us e
piro
ticus
1EN
WE
58Pe
lasg
us la
coni
cus
(Kot
tela
t & B
arbi
eri,
2004
)N
EW1
ENW
E59
Pela
sgus
mar
atho
nicu
s(V
inci
guer
ra, 1
921)
Pseu
doph
oxin
us st
ymph
alic
us m
arat
honi
cus
1EN
CE
60Pe
lasg
us p
resp
ensis
(Kar
aman
, 192
4)Pa
raph
oxin
us e
piro
ticus
pre
spen
sis1
NEN
NO
61Pe
lasg
us st
ymph
alic
us(V
alen
cien
nes,
1844
)Ps
eudo
phox
inus
stym
phal
icus
stym
phal
icus
1EN
WE
62Pe
lasg
us th
espr
otic
us(S
teph
anid
is, 1
939)
Pseu
doph
oxin
us st
ymph
alic
us th
espr
otic
us1
ENW
E+63
Petro
leuci
scus
bor
ysth
enic
us(K
essle
r, 18
59)
Leuc
iscus
bor
ysth
enic
us1
OTH
ER64
Petro
leuci
scus
smyr
naeu
s(B
oule
nger
, 189
6)N
EW1
OTH
ER65
Phox
inus
cf.
phox
inus
15(L
inna
eus,
1758
)Ph
oxin
us p
hoxin
us1
OTH
ER66
Phox
inus
stry
mon
icus
Kot
tela
t, 20
07Ph
oxin
us p
hoxin
us1
(EN
NO
)67
Pseu
dora
sbor
a pa
rva
(Tem
min
ck &
Sch
lege
l, 18
46)
NO
CH
AN
GE
1A
LN68
Rhod
eus a
mar
us(B
loch
, 178
2)Rh
odeu
s ser
iceu
s am
arus
1O
THER
69Rh
odeu
s mer
idio
nalis
Kar
aman
, 192
4Rh
odeu
s ser
iceu
s am
arus
1EN
BAL
70Ro
man
ogob
io e
limei
us16
(Kat
toul
as, S
teph
anid
is &
G
obio
ura
nosc
opus
elim
eius
1
ENBA
LEc
onom
idis,
197
3)71
Rutil
us p
anos
iBo
guts
kaya
& Il
iado
u, 2
006
Rutil
us yl
ikie
nsis
1EN
WE
72Ru
tilus
pre
spen
sis(K
aram
an, 1
924)
Rutil
us o
hrid
anus
pre
spen
sis1
NEN
NO
73Ru
tilus
rutil
us(L
inna
eus,
1758
)Ru
tilus
rutil
us m
ariza
, Rut
ilus r
utilu
s doj
rane
nsis,
1
OTH
ERRu
tilus
rutil
us v
egor
iticu
s74
Rutil
us sp
.Spe
rchi
os17
In K
&F2
007
NEW
1EN
CE
75Ru
tilus
ylik
iens
isEc
onom
idis,
199
1N
O C
HA
NG
E1
ENC
E
App
endi
x I
(con
tinu
ed)
(con
tinu
ed)
Medit. Mar. Sci., 8/1, 2007, 91-166158
12
34
56
Nam
eAu
thor
ityPr
evio
us N
omen
clat
ure
Fres
hwat
erEn
dem
ic76
Scar
dini
us a
carn
anic
usEc
onom
idis,
199
1N
O C
HA
NG
E1
ENW
E77
Scar
dini
us e
ryth
roph
thal
mus
(Lin
naeu
s, 17
58)
NO
CH
AN
GE
1O
THER
78Sc
ardi
nius
gra
ecus
Step
hani
dis,
1937
NO
CH
AN
GE
1EN
CE
79Sq
ualiu
s cf.
cii18
(Ric
hard
son,
185
6)N
EW1
OTH
ER80
Squa
lius p
elopo
nnen
sis19
(Val
enci
enne
s, 18
44)
Leuc
iscus
cep
halu
s pelo
ponn
ensis
1
ENW
E81
Squa
lius k
eadi
cus
(Ste
phan
idis,
197
1)Le
ucisc
us k
eadi
cus
1EN
WE
82Sq
ualiu
s mor
eotic
us20
(Ste
phan
idis,
197
1)N
EW1
ENW
E83
Squa
lius o
rphe
usK
otte
lat &
Eco
nom
idis,
200
6Le
ucisc
us c
epha
lus m
aced
onic
us1
ENBA
L84
Squa
lius p
amvo
ticus
21(S
teph
anid
is, 1
939)
NEW
1EN
WE
85Sq
ualiu
s pre
spen
sis(F
owle
r, 19
77)
NEW
1N
ENN
O86
Squa
lius s
p. A
oos
In K
&F2
007
Leuc
iscus
cep
halu
s var
dare
nsis
1EN
BAL
87Sq
ualiu
s sp.
Evia
22In
K&
F200
7N
EW1
ENC
E88
Squa
lius s
p. E
vino
s23In
K&
F200
7Le
ucisc
us c
epha
lus a
lbus
, Leu
cisc
us "s
valli
ze"
1EN
WE
89Sq
ualiu
s var
dare
nsis
Kar
aman
, 192
8Le
ucisc
us c
epha
lus v
arda
rens
is1
ENBA
L90
Teles
tes b
eotic
us(S
teph
anid
is, 1
939)
Pseu
doph
oxin
us b
eotic
us1
ENC
E91
Teles
tes p
leuro
bipu
ncta
tus
(Ste
phan
idis,
193
9)Ph
oxin
ellus
pleu
robi
punc
tatu
s1
ENW
E+92
Tinc
a tin
ca(L
inna
eus,
1758
)N
O C
HA
NG
E1
OTH
ER93
Trop
idop
hoxin
ellus
hell
enic
us(S
teph
anid
is, 1
939)
NO
CH
AN
GE
1EN
WE
94Tr
opid
opho
xinell
us sp
artia
ticus
(Sch
mid
t-Rie
s, 19
43)
NO
CH
AN
GE
1EN
WE
95Vi
mba
mela
nops
(Hec
kel,
1837
)N
O C
HA
NG
E1
ENBA
LC
obiti
dae
96C
obiti
s ara
chth
osen
sisEc
onom
idis
& N
alba
nt, 1
997
NO
CH
AN
GE
1EN
WE
97C
obiti
s hell
enic
aEc
onom
idis
& N
alba
nt, 1
997
NO
CH
AN
GE
1EN
WE
98C
obiti
s mer
idio
nalis
Kar
aman
, 192
4N
O C
HA
NG
E1
NEN
NO
99C
obiti
s ohr
idan
aK
aram
an, 1
928
NEW
1EN
BAL
100
Cob
itis p
unct
icul
ata
Erk’
akan
, Ata
lay-
Ekm
ekçi
N
EW1
OTH
ER&
Nal
bant
, 199
810
1C
obiti
s pun
ctili
neat
aEc
onom
idis
& N
alba
nt, 1
997
NO
CH
AN
GE
1EN
NO
102
Cob
itis s
teph
anid
isiEc
onom
idis
& N
alba
nt, 1
997
NO
CH
AN
GE
1EN
NO
App
endi
x I
(con
tinu
ed)
(con
tinu
ed)
Medit. Mar. Sci., 8/1, 2007, 91-166 159
12
34
56
Nam
eAu
thor
ityPr
evio
us N
omen
clat
ure
Fres
hwat
erEn
dem
ic10
3C
obiti
s stru
mic
aeK
aram
an, 1
955
NO
CH
AN
GE
1EN
BAL
104
Cob
itis t
richo
nica
Step
hani
dis,
1974
NO
CH
AN
GE
1EN
WE
105
Cob
itis v
arda
rens
isK
aram
an, 1
928
NO
CH
AN
GE
1EN
BAL
106
Saba
nejew
ia b
alca
nica
(Kar
aman
, 192
2)Sa
bane
jewia
aur
ata
balc
anic
a 1
ENBA
LN
emac
heili
dae
107
Barb
atul
a ba
rbat
ula
(Lin
naeu
s, 17
58)
Orth
rias b
arba
tulu
s1
OTH
ER10
8O
xyno
emac
heilu
s bur
esch
i(D
rens
ky, 1
928)
Orth
rias b
rand
ti bu
resc
hi
1EN
BAL
109
Oxy
noem
ache
ilus p
indu
s(E
cono
mid
is, 2
005)
Orth
rias p
indu
s1
ENBA
L11
0O
xyno
emac
heilu
s the
ophi
liiSt
oum
boud
i, K
otte
lat &
Bar
bier
i, 20
06
NEW
1O
THER
Icta
luri
dae
111
Icta
luru
s pun
ctat
us(R
afin
esqu
e, 1
818)
NEW
1A
LNSi
luri
dae
112
Silu
rus a
risto
telis
Gar
man
, 189
0N
O C
HA
NG
E1
ENW
E11
3Si
luru
s gla
nis
Linn
aeus
, 175
8N
O C
HA
NG
E1
OTH
ERC
lari
idae
114
Cla
rias g
arie
pinu
s(B
urch
ell,
1822
)N
EW1
ALN
Esoc
idae
115
Esox
luci
usLi
nnae
us, 1
758
NO
CH
AN
GE
1O
THER
Cor
egon
idae
116
Cor
egon
us c
f.la
vare
tus24
(Lin
naeu
s, 17
58)
Cor
egon
us la
vare
tus
1A
LNSa
lmon
idae
117
Onc
orhy
nchu
s kisu
tch
(Wal
baum
, 179
2)N
O C
HA
NG
E1
ALN
118
Onc
orhy
nchu
s myk
iss(W
alba
um, 1
792)
NO
CH
AN
GE
1A
LN11
9Sa
lmo
dent
ex25
Hec
kel,
1852
Salm
o tru
tta d
ente
x1
ENBA
L12
0Sa
lmo
fario
ides
26(K
aram
an, 1
938)
Salm
o tru
tta m
acro
stigm
a, S
alm
o tru
tta d
ente
x1
ENBA
L12
1Sa
lmo
letni
ca(K
aram
an, 1
924)
NEW
1A
LN12
2Sa
lmo
cf. m
aced
onic
us27
(Kar
aman
, 192
4)Sa
lmo
trutta
mac
edon
icus
1EN
BAL
123
Salm
o pe
lago
nicu
sK
aram
an, 1
938
Salm
o tru
tta p
elago
nicu
s1
ENBA
L12
4Sa
lmo
peris
teric
usK
aram
an, 1
938
Salm
o tru
tta p
erist
eric
us1
NEN
NO
App
endi
x I
(con
tinu
ed)
(con
tinu
ed)
Medit. Mar. Sci., 8/1, 2007, 91-166160
12
34
56
Nam
eAu
thor
ityPr
evio
us N
omen
clat
ure
Fres
hwat
erEn
dem
ic12
5Sa
lmo
sala
rLi
nnae
us, 1
758
NEW
1A
LN12
6Sa
lmo
sp.L
ouro
s28In
K&
F200
7N
EW1
ENW
E12
7Sa
lmo
cf. t
rutta
29Li
nnae
us, 1
758
NEW
1A
LN12
8Sa
lveli
nus f
ontin
alis
(Mitc
hill,
181
4)N
O C
HA
NG
E1
ALN
Mug
ilida
e12
9C
helo
n la
bros
usR
isso,
182
7N
O C
HA
NG
E0
OTH
ER13
0Li
za a
urat
a(R
isso,
181
0)N
O C
HA
NG
E0
OTH
ER13
1Li
za ra
mad
a(R
isso,
182
7)N
O C
HA
NG
E0
OTH
ER13
2Li
za sa
liens
(Riss
o, 1
810)
NO
CH
AN
GE
0O
THER
133
Mug
il ce
phal
usLi
nnae
us, 1
758
NO
CH
AN
GE
0O
THER
Athe
rini
dae
134
Ath
erin
a bo
yeri30
Riss
o, 1
810
NO
CH
AN
GE
0O
THER
Vale
nciid
ae13
5Va
lenci
a let
ourn
euxi
(Sau
vage
, 188
0)N
O C
HA
NG
E(1
)EN
WE+
Poec
iliid
ae13
6G
ambu
sia h
olbr
ooki
31G
irard
, 185
9G
ambu
sia a
ffini
s(1
)A
LN13
7Po
ecili
a cf
.lat
ipin
na32
Lesu
er, 1
821
NEW
1A
LNC
ypri
nodo
ntid
ae
138
Aph
aniu
s alm
irien
sisK
otte
lat,
Barb
ieri
&
NEW
0EN
WE
Stou
mbo
udi,
2007
139
Aph
aniu
s fas
ciat
us(V
alen
cien
nes,
1821
)N
O C
HA
NG
E0
OTH
ERG
aste
rost
eida
e14
0G
aste
roste
us g
ymno
urus
Cuv
ier,
1829
Gas
tero
steus
acu
leatu
s0
OTH
ER14
1Pu
ngiti
us h
ellen
icus
Step
hani
dis,
1971
NO
CH
AN
GE
1EN
CE
142
Pung
itius
pla
tygas
ter
(Kes
sler,
1859
)N
O C
HA
NG
E(1
)O
THER
Mor
onid
ae14
3D
icen
trarc
hus l
abra
x(L
inna
eus,
1758
)N
O C
HA
NG
E0
OTH
ERSy
ngna
thid
ae33
144
Syng
nath
us a
baste
rR
isso,
182
7N
O C
HA
NG
E0
OTH
ER
App
endi
x I
(con
tinu
ed)
(con
tinu
ed)
Medit. Mar. Sci., 8/1, 2007, 91-166 161
12
34
56
Nam
eAu
thor
ityPr
evio
us N
omen
clat
ure
Fres
hwat
erEn
dem
icC
entr
arch
idae
145
Lepo
mis
gibb
osus
(Lin
naeu
s, 17
58)
NO
CH
AN
GE
(1)
ALN
146
Mic
ropt
erus
salm
oide
s(L
a C
epèd
e, 1
802)
NEW
(1)
ALN
Perc
idae
147
Perc
a flu
viat
ilis
Linn
aeus
, 175
8N
O C
HA
NG
E1
OTH
ER14
8Sa
nder
luci
oper
ca
(Lin
naeu
s, 17
58)
Stizo
stedi
on lu
ciop
erca
1O
THER
149
Zing
el ba
lcan
icus
34(K
aram
an, 1
936)
Zing
el str
eber
bal
cani
cus
1EN
BAL
Cic
hlid
ae15
0O
reoc
hrom
is ni
lotic
us(L
inna
eus,
1758
)N
EW(1
)A
LNBl
enni
idae
151
Sala
ria e
cono
mid
isiK
otte
lat,
2004
NEW
1EN
WE
152
Sala
ria fl
uvia
tilis
(Ass
o, 1
801)
NO
CH
AN
GE
(0)
OTH
ERG
obiid
ae15
3Ec
onom
idic
hthy
s pyg
mae
us(H
olly
, 192
9)N
O C
HA
NG
E(1
)EN
WE+
154
Econ
omid
icht
hys t
richo
nis
Econ
omid
is &
Mill
er, 1
990
NO
CH
AN
GE
1EN
WE
155
Knip
owits
chia
cau
casic
a35(B
erg,
191
6)N
O C
HA
NG
E(1
)O
THER
156
Knip
owits
chia
goe
rner
iA
hnel
t, 19
91N
EW(1
)EN
WE
157
Knip
owits
chia
mill
eri
(Ahn
elt &
Bia
nco
1990
)N
O C
HA
NG
E(1
)EN
WE
158
Knip
owits
chia
thes
sala
(Vin
cigu
erra
, 192
1)N
O C
HA
NG
E(1
)EN
NO
159
Prot
eror
hinu
s sem
illun
aris
(Hec
kel,
1837
)Pr
oter
orhi
nus m
arm
orat
us1
OTH
ER16
0Zo
steris
esso
r oph
ioce
phal
us(P
alla
s, 18
14)
NO
CH
AN
GE
0O
THER
Pleu
rone
ctid
ae16
1Pl
atic
hthy
s fles
us(L
inna
eus,
1758
)N
O C
HA
NG
E0
OTH
ER
App
endi
x I
(con
tinu
ed)
LE
GE
ND
Col
umn
1:Sp
ecie
s an
nota
tion.
All
spec
ies
with
val
id n
ame
as w
ell a
s un
desc
ribe
d op
erat
iona
l tax
a na
mes
are
num
bere
d an
d co
nsid
ered
as
sepa
rate
taxo
nom
ic u
nits
. C
olum
n 2:
Scie
ntifi
c N
ame.
Val
id s
cien
tific
nam
e or
ope
ratio
nal t
axon
omic
uni
t nam
es a
re g
iven
. C
olum
n 3:
Aut
hori
ty.S
peci
es a
utho
r an
d da
te a
re g
iven
. Und
escr
ibed
taxa
are
giv
en o
nly
if pr
ovid
ed b
y K
OT
TE
LA
T &
FR
EY
HO
F (2
007a
) (h
ereb
y re
ferr
ed a
s K
&F
2007
) w
ithon
e ex
cept
ion
(Eud
onto
myz
on s
p. L
ouro
s, an
ope
ratio
nal n
ame
prov
ided
by
this
stu
dy).
Medit. Mar. Sci., 8/1, 2007, 91-166162
Col
umn
4:Pr
evio
us N
omen
clat
ure.
Subs
peci
es r
epla
cem
ent n
ames
and
not
atio
n on
cha
nges
with
res
pect
to E
CO
NO
MID
IS (
1991
) ar
e as
follo
ws:
NO
CH
AN
GE
: No
rece
nt n
ame
chan
ges
(i.e
. spe
cies
nam
es r
etai
ned
as g
iven
by
EC
ON
OM
IDIS
, 199
1); N
EW
: New
add
ition
s to
the
Gre
ek f
resh
wat
er f
ish
chec
klis
t, no
t in
clud
ed in
EC
ON
OM
IDIS
(199
1) u
nder
the
curr
ent t
axon
nam
e.
Col
umn
5:"F
resh
wat
er re
side
nce"
. Fis
h kn
own
to b
e re
stri
cted
to fr
eshw
ater
s in
Gre
ece
are
dist
ingu
ishe
d fr
om sp
ecie
s spe
ndin
g a
part
of t
heir
live
s in
the
sea
or in
wat
ers w
ith h
igh
salin
ity. T
ypic
ally
fres
hwat
er s
peci
es th
at u
sual
ly d
o no
t rea
dily
ent
er s
eaw
ater
con
ditio
ns a
re d
enot
ed w
ith 1
; Eur
yhal
ine
spec
ies,
brac
kish
wat
er o
r m
arin
e sp
ecie
s th
atar
e to
lera
nt o
f or s
pend
par
t of t
heir
live
s in
fres
hwat
er c
ondi
tions
are
den
oted
with
0. W
here
unc
erta
inty
exi
sts o
n th
e be
havi
our o
f the
se fi
sh in
Gre
ece
a pr
ovis
iona
l des
-ig
natio
n is
giv
en in
par
enth
eses
. C
olum
n 6:
End
emic
ity:l
evel
of e
ndem
ic s
tatu
s at
the
natio
nal s
cale
, i.e
. res
tric
ted
dist
ribu
tiona
l ran
ge o
r en
dem
ic s
tatu
s re
lativ
e to
the
terr
itori
al b
ound
arie
s of
Gre
ece.
EN
WE
: End
emic
to W
este
rn G
reec
e an
d/or
the
Pelo
ponn
ese.
E
NW
E+
: "N
ear-
ende
mic
" to
Wes
tern
Gre
ece;
i.e.
res
tric
ted
to G
reec
e, b
ut a
sm
all f
ract
ion
of g
loba
l pop
ulat
ion
exte
nds
into
the
sout
hern
par
t of A
lban
ia.
EN
CE
: E
ndem
ic to
Cen
tral
Eas
tern
Gre
ece,
i.e.
the
Att
iko-
Beo
tia r
egio
n (s
outh
of P
agas
itico
s G
ulf,
incl
udin
g Ft
hiot
is, B
eotia
, Att
iki a
nd E
uboe
a).
EN
NO
: End
emic
to N
orth
ern
Gre
ece;
incl
udin
g T
hess
aly,
but
not
the
Paga
sitic
os G
ulf (
addi
tiona
lly tw
o sp
ecie
s th
at r
each
Att
iko-
Beo
tia r
egio
n ar
e in
clud
ed).
NE
NN
O: "
Nea
r-en
dem
ic" t
o N
orth
ern
Gre
ece;
glo
bal p
opul
atio
ns r
estr
icte
d to
Gre
ece’
s tr
ansb
ound
ary
lake
s (i
.e. P
resp
a or
Doi
rani
).E
NB
AL
: End
emic
to th
e B
alka
ns (
i.e. s
outh
of t
he D
anub
e R
.).E
NA
EG
: End
emic
to th
e A
egea
n Is
land
s.O
TH
ER
: Spe
cies
with
oth
er d
istrib
utio
n pa
ttern
s; th
is in
clud
es w
ides
prea
d sp
ecie
s, an
d sp
ecie
s hav
ing
a lo
caliz
ed p
opul
atio
n on
ly in
Asia
Min
or a
nd th
e ad
jace
nt G
reek
isla
nds.
AL
N: A
lien
spec
ies;
ther
e is
no
subs
tant
iate
d ev
iden
ce th
at th
e sp
ecie
s is
nat
ive
with
in th
e G
reek
terr
itory
. W
here
dou
bts
exis
t a p
rovi
sion
al s
tatu
s is
giv
en in
par
enth
eses
.
CRITICAL COMMENTS ON CHECK-LIST
1 Eudontomyzon hellenicus refers only tothe populations in the Strymon R. Anunidentified Eudontomyzon species hasalso been recorded from the Almopeostributary of the Aliakmon R. but itremains undescribed (ECONOMIDIS& B N RESCU, 1991).
2 Eudontomyzon sp. Louros is not a validtaxa described for the Greek freshwaterichthyofauna (not referred to by anoperational name by K&F 2007). TheEudontomyzon in the Louros basin isgiven this provisional name here sinceits distribution in Greece is extremelylocalized and this taxon has been knownto differ morphologically from E. hel-lenicus for several years now(ECONOMIDIS, 1995). K&F 2007remark that the population from Louros"either belongs to E. stankokaramani orrepresents a distinct, unnamed species".
3 Acipenser stellatus is considered as extir-pated in the Aegean basins (K&F 2007),documentation of its past presence isevident in older publications(PAPACONSTANTINOU, 1988). Re-cent occurrences of the species in theEvros R. seem to be from stocking orescapes from Bulgaria (APOSTOLOU,pers. comm.; KOUTRAKIS &ECONOMIDIS, 2006).
4 Huso huso is now considered as an alienspecies in Greece since all populationsare from stocking and/or escapees fromfish farms (KOUTRAKIS &ECONOMIDIS, 2006). Unfortunatelywe cannot confirm the possible exis-tence of natural populations of this stur-geon (or even its transient occurrence)in the northern Aegean in the past;although the species may possibly have
Medit. Mar. Sci., 8/1, 2007, 91-166 163
existed up until the 19th or early 20thcenturies. A few unconfirmed records inthe Aegean are reported in earlieraccounts (ONDRIAS, 1971;ECONOMIDIS, 1973;PAPACONSTANTINOU, 1988) butthese have later been regarded asextremely doubtful (KOUTRAKIS &ECONOMIDIS, 2006).
5 The existence of Alburnus scoranza isundocumented in Greece. A speciesregarded as Alburnus alburnus wasdetected within the Greek part of theAoos (ECONOMOU et al., 2007) and istentatively given the provisional opera-tional name A. cf. scoranza here, sincethe only Alburnus species in the immedi-ate vicinity and within the SouthernAdriatic biogeographic region is A. sco-ranza (K&F 2007).
6 Alburnus sp. Volvi is an unnamed taxonvery similar to Alburnus alburnus. Speci-mens only from L. Volvi and L. Kerkini(Strymon basin) were collected in a firstprovisional description (FREYHOF &KOTTELAT, 2007a).
7 Alburnus thessalicus belongs to theAlburnus alburnus complex with fourtaxa in Greece (A. thessalicus, A. mace-donicus, A. sp. Volvi, A. alburnus) (K&F2007). These taxa share many character-istics and may be difficult to identify inthe field.
8 Barbus balcanicus populations in thelower Axios were formerly considered asboth Barbus peloponnesius petenyi andBarbus cyclolepis; difficulty of identifica-tion of these fish on morphologicalgrounds is noted in K&F 2007, but thespecies can be distinguished usingmolecular markers (KOTLI’K et al.,2002).
9 Barbus peloponnesius is not recorded inAlbania following K&F 2007, but there
is recent evidence of its existence in theextreme southern part of the country(MARKOVA et al., 2007).
10 Barbus sperchiensis has isolated popula-tions in the Sperchios valley and inNorthern Euboea (formerly consideredBarbus cyclolepis sperchiensis), the Pag-asiticos Gulf’s Cholorema basin (for-merly considered B. cyclolepis cholore-maticus) and in Thessaly (formerly con-sidered Barbus cyclolepis strumicae)(ECONOMIDS & BOGUTSKAYA,2003). This unusual distribution strad-dles a biogeographic boundary andincludes insular populations on EuboeaIsland.
11 Carassius auratus is frequently confusedin the literature with Carassius gibelio,which was formerly treated as "a wildferal form of C. auratus" (K&F 2007).The taxonomic entity of Carassius aura-tus (the goldfish) is not present as anindependent taxonomic unit inECONOMIDIS (1991) although it ispresent in BOBORI & ECONOMIDIS(2006).
12 The status of Carassius gibelio withrespect to its being native in Greece isstill unresolved and controversial; weadopt its status "as probably native" onlyin the Strymon and the Evros followingECONOMIDIS (1991). There is con-siderable doubt as to this species nativestatus in southeast Europe, althoughpopulations have long been naturalized.
13 The status of Cyprinus carpio withrespect to its being native to Greece isunclear; natural populations have prob-ably been established and are natural-ized for centuries. We adopt thisspecies status as "native" in Thessaly,Macedonia, and Thrace according toECONOMIDIS (1991).
14 The existence of Gobio skadarensis is
Medit. Mar. Sci., 8/1, 2007, 91-166164
undocumented in Greece. The speciesfound inhabiting the Aoos(ECONOMOU et al., 2007) is tenta-tively given the operational name Gobiocf. skadarensis in this list since the onlyknown Gobio sp. in the immediatevicinity and in the South Adriatic bio-geographic region is Gobio skadarensis,previously referred to as Gobio gobioalbanicus ( ANDA et al., 2005).
15 Phoxinus phoxinus was until recently theonly species in this genus in Greece.K&F 2007 present a new Phoxinusspecies in the Strymon (Phoxinus stry-monicus), but no classification is pro-vided for the Phoxinus populations ofAliakmon, Loudias, Nestos, Filiourisand Evros; although they state that"Phoxinus populations in the Loudiasand Filiouris drainages are possiblyconspecific". The aforementionedauthors do not give Phoxinus phoxinusas inhabiting any area in the southernBalkan river basins or in Greece. Pend-ing an accurate description of theunclassified populations we propose thefish not in the type locality river (Stry-mon) be provisionally termed Phoxinuscf. phoxinus.
16 Romanogobio elimeius may possiblyinclude a very similar-looking rheophilicgudgeon called Romanogobio kessleribanarescui (ECONOMIDIS, 1991);K&F 2007 were "unable to dististinguishR. banarescui and R. stankovi from R.elimeius on the basis of the available lit-erature and material and therefore ten-tatively treat them as synonyms".
17 Rutilus sp. Sperchios was observed inthe lower Sperchios in the late ‘90s butwas never described. Only two small-sized specimens were collected in 1997(K&F 2007); this taxon needs immedi-ate study.
18 Squalius cii was first proposed as a ten-tative name for the Squalius of Lesbos(STOUMBOUDI et al., 2006) althoughK&F 2007 refers to this population asSqualius cf. cii. Unidentified Leucisinaecalled "Leuciscus cephalus" (BIANCO,1990) also exist on Samos.
19 The identity of the Squalius in the Ion-ian river basins is controversial andunresolved. Formerly these chubs weregiven as Leuciscus cephalus but it is cer-tain that the chubs west of the Pindosare different species. In the distribution-al compilation we refer to nearly all"Ionian chubs" by the provisional opera-tional name Squalius cf. peloponnensisalthough we do give the specific threetaxa proposed by K&F 2007 in the spe-cific recorded basins. Despite consider-able amount of genetic work(IMSIRIDOU et al., 1997, 1998, 2000;DOADRIO & CARMONA, 1998,2003; SANJUR et al., 2003;ZARDOYA et al., 1999; DURAND etal., 1999b, 2000), further taxonomic re-search involving populations from north-ern and south-western Peloponnese aswell as from the Epirus rivers is neededto resolve the problem of Squaliusspecies distinction in the Ionian basins.
20 Squalius moreoticus described as Leu-ciscus cephalus moreoticus bySTEPHANIDIS (1939) was originallyconsidered as being confined to LakeStymphalia. K&F 2007 give a wider dis-tribution for this taxon (i.e. includingVouraikos R. in the northern Pelopon-nese). We tentatively limit the S.moreoticus distribution to the afore-mentioned lake but its current status isundefined, as the originally describedtaxon may be extirpated or even extinct.The lake completely dried-out in theearly 1990s and the present population
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hosted in the lake may have been intro-duced from another basin – as purport-ed by local fisherman.
21 Squalius pamvoticus was described bySTEPHANIDIS (1939) as Leuciscuscambeda var. pamvoticus presumablyconfined to Lake Pamvotis. K&F 2007give a wider distribution for this taxon,including nearly all the rivers of Epirus(except Aoos). Care is needed in theclassification of populations in therivers of Epirus; and pending furthertaxonomic evidence we treat the Epirusriver taxa as Squalius cf. peloponnensis(STEPHANIDIS, 1939 consideredthese populations as Leuciscus cephaluspeloponnensis). The taxonomy and dis-tribution of Squalius pamvoticus needsimmediate attention.
22 Squalius sp. Evia was previously consid-ered as a distinct form of Leuciscuscephalus vardarensis. This taxon name isprovisionally given for specimens fromthe Manikiotikos R. on Euboea island.The classification of Squalius popula-tions in the streams of northern Euboearemains unresolved (although we tenta-tively place the fish from the Kireas R.under this taxon name in the distribu-tional accounts). Evia should be spelledEuboea, the anglicised classical rendi-tion often used in the scientific bibliog-raphy.
23 Squalius sp. Evinos was proposed as atentative taxonomic unit by K&F 2007for Squalius populations inhabitingMornos, Evinos and Acheloos. Thistaxon is morphologically very similar tosome populations of Squalius pelopon-nensis.
24 Coregonus cf. lavaretus requires identifi-cation due to recent taxonomic changesin K&F 2007.
25 Salmo dentex is mentioned as inhabiting
Aoos (and possibility Alfios) inK&F2007. This species’ existence inGreece needs confirmation.
26 Salmo farioides is now formally consid-ered as the dominant trout of westernGreece, a different taxon from popula-tions east of the Pindos. The populationof the Alfios (Peloponnese) are isolatedand in need of conservation-relevanttaxonomic confirmation (K&F2007).
27 Salmo macedonicus is not recorded as aspecies inhabiting the Greek territory inK&F 2007. These authors do not givedistributional information or the identi-ty of trout in the Strymon, Nestos orEvros; however, in previous works thetaxon S. trutta macedonicus has beendescribed in these areas (KARAMAN,1927). We use the tentative name of S.cf. macedonicus to refer to the fishinhabiting the aforementioned basins.Confirmation of this taxon’s presenceand its distribution in Greece is needed.
28 Salmo sp. Louros is one of the most sur-prising new "taxa" described from theLouros R. DELLING (2003) refers tothis fish as Salmo louroensis a name notaccepted by K&F 2007.
29 Varieties of farmed Salmo trutta fromEuropean hatcheries have been intro-duced into Greek waters particularlynear fish-farming units.
30 The systematic status of Atherina boyeripopulations has been recently ques-tioned. Molecular data (KLOSSA-KILIA et al., 2007) reveal deep geneticdivergence between marine popula-tions of Atherina boyeri and those livingin lagoons and lakes, possibly indicatingthe existence of cryptic or siblingspecies.
31 Gambusia holbrooki has replaced for-mer claims of Gambusia affinis inGreece. K&F 2007 state that there is no
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confirmed record of G. affinis inEurope.
32 Poecilia cf. latipinna has tentativelybeen identified as the alien aquariumescapee which is abundant in Vouliag-meni L., Attiki (KOUTSIKOS pers.comm.); formerly this species wasdescribed as Poecilia sphenops(CHINTIROGLOU et al., 1996).
33 The marine family Syngnanthidae ispoorly documented in freshwaters inGreece. BOBORI & ECONOMIDIS(2006) provide five species of this fami-ly in their freshwater fish fauna list.Pending new information that confirmsthat these species actually reside forlong periods in freshwaters we retainonly Sygnathus abaster in this list, as didECONOMIDIS (1991).
34 To our knowledge Zingel balcanicus hasnot yet been collected within the Greekterritory but records are from very nearthe border in FYR Macedonia. Thisspecies is difficult to collect and it isassumed that it is almost definitelypresent in Greek waters, probably as faras Axioupolis, as stated by K&F2007.The species is also included within thefreshwater fish faunal list of BOBORI& ECONOMIDIS (2006).
35 The Knipowitschia population in thelower Evinos R. was formally referredto as Knipowitschia panizzae (AHNELT& BIANCO, 1991) but this is uncon-firmed; perhaps an unidentified taxon(or taxa) of Knipowitschia exists in sev-eral areas of western Greece (MILLERet al., 2004b).
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Appendix 2Preliminary supplementary checklist of fish species recorded in brackish
and transitional waters of Greek hydrographic basins.
No Species Authority References
Clupeidae1 Sardina pilchardus (Walbaum, 1792) 3, 6
Engraulidae2 Engraulis encrasicolus (Linnaeus, 1758) 6
Gadidae3 Gaidropsarus mediterraneus (Linnaeus, 1758) 4
Belonidae4 Belone belone (Linnaeus, 1761) 3, 6
Atherinidae 5 Atherina hepsetus Linnaeus, 1758 2, 3, 4, 7
Syngnathidae6 Hippocampus guttulatus Cuvier, 1829 37 Nerophis ophidion (Linnaeus, 1758) 78 Syngnathus acus Linnaeus, 1758 4, 79 Syngnathus taenionotus Canestrini, 1871 710 Syngnathus typhle Linnaeus, 1758 3, 711 Syngnathus variegatus Pallas, 1814 3
Scorpaenidae12 Scorpaena scrofa Linnaeus, 1758 6
Triglidae13 Lepidotrigla cavillone (Lacepède, 1801) 314 Chelidonichthys lastoviza (Bonnaterre, 1788) 6
Serranidae15 Epinephelus aeneus (Geoffroy Saint-Hilaire, 1817) 4
Carangidae16 Trachurus mediterraneus (Steindachner, 1868) 6
Sparidae17 Boops boops (Linnaeus, 1758) 618 Diplodus annularis (Linnaeus, 1758) 3, 4, 619 Diplodus puntazzo (Cetti, 1777) 420 Diplodus sargus sargus (Linnaeus, 1758) 3, 4, 621 Diplodus vulgaris (Geoffroy Saint-Hilaire, 1817) 3, 422 Lithognathus mormyrus (Linnaeus, 1758) 3, 4, 623 Pagellus bogaraveo (Brünnich, 1768) 624 Sarpa salpa Linnaeus, 1758 3, 425 Sparus aurata Linnaeus, 1758 3, 426 Spondyliosoma cantharus (Linnaeus, 1758) 4
Moronidae27 Dicentrarchus punctatus (Bloch, 1792) 2, 7
Sciaenidae28 Argyrosomus regius (Asso, 1801) 529 Umbrina cirrosa (Linnaeus, 1758) 6
Mullidae30 Mullus barbatus barbatus Linnaeus, 1758 3, 431 Mullus surmuletus Linnaeus, 1758 4, 6
(continued)
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Appendix 2 (continued)
No Species Authority References
Mugillidae32 Chelon haematocheilus (Temminck & Schlegel, 1845) 733 Oedalechilus labeo (Cuvier, 1829) 2, 4
Clinidae 34 Clinitrachus argentatus (Risso, 1827) 3
Blennidae35 Aidablennius sphynx (Valenciennes, 1836) 336 Parablennius gattorugine (Linnaeus, 1758) 337 Parablennius incognitus (Bath, 1968) 338 Parablennius sanguinolentus (Pallas, 1814) 3, 439 Parablennius tentacularis (Brünnich, 1768) 340 Salaria pavo (Risso, 1810) 3, 6, 7
Gobiidae41 Gobius cobitis Pallas, 1814 342 Gobius cruentatus Gmelin, 1789 343 Gobius geniporus Valenciences, 1837 444 Gobius niger Linnaeus, 1758 345 Gobius paganellus Linnaeus, 1758 346 Pomatoschistus marmoratus (Risso, 1810) 3, 4, 647 Pomatoschistus minutus (Pallas, 1770) 1
Scombridae48 Scomber scombrus Linnaeus, 1758 6
Callionymidae49 Callionymus maculatus Rafinesque, 1810 350 Callionymus risso Lesueur, 1814 3
Scophthalmidae51 Psetta maxima (Linnaeus, 1758) 352 Scophthalmus rhombus (Linnaeus, 1758) 3
Soleidae53 Synapturichthys kleinii (Risso, 1827) 354 Pegusa lascaris (Risso, 1810) 355 Solea solea (Linnaeus, 1758) 3, 4, 6
Note: This supplementary list provides a preliminary compilation of species recorded in transitional waters but not con-firmed to reside in freshwaters. Freshwater and other euryhaline species are commonly present in transitional waters butare not included here since they are in the list of Freshwater Fish (Appendix I). This list is provisional since it is based sole-ly on seven bibliographic references and the species documentation has not been screened or verified by additional fieldresearch. Nomenclature follows fishbase (FROESE & PAULY, 2007).
References: 1 = PAPACONSTANTINOU, 19882 = ECONOMIDIS, 19913 = KOUTRAKIS et al., 20004 = NIKOLAIDOU et al., 20055 = KASPIRIS, 2000.6 = KOUTRAKIS et al., 20057 = BOBORI & ECONOMIDIS, 2006