Non-indigenousanimal species naturalizedin Iberianinlandwaters
EmiliGarc´ıa-Berthou, DaniBoix,andMiguelClavero
INTRODUCTION
Invasions byhuman-introduced non-indigenousspecies(NIS)are one ofthe mainthreats tobiodiversityandadrivingforceofglobalchange(Vitouseketal.
1997, Macketal.2000,ClaveroandGarc´ıa-Berthou2005).TheIberian Peninsula (IP)isahotspot ofbiodiversity(Me´dailandQue´zel 1999)anda knowledge ofthe invasive speciesinhabiting itisessential forconservation andenvironmentalmanagement.Naturalized vertebrates andplants intheIP have receivedconsiderable attention(seee.g.Vila`etal.2001,Pleguezuelos
2002,Sobrino etal.2002,Lloretetal.2004,Alcaraz etal.2005),but its invasiveinvertebrates areverypoorlyknown.Althoughtherearemanyrecords ofsomeinvertebrate invasivespecies,particularly crustaceans,there arevery fewavailable reviewsofselectedtaxa ofinvertebrateinvaders inthe IP(e.g. Espadaler and Collingwood2001).Theaim ofthis chapter istoreview the animal speciesnaturalized inIberianinland waters, including vertebrates and free-livingand parasitic invertebrates.Asusual, the taxonomy and biogeog- raphy ofvertebrate speciesaremuch better known than forinvertebrates,so our data for invertebrates should be regarded as a preliminary check-list. Similarly,the parasites ofnon-commercialaquatic speciesare poorlystudied and the data inthe IPmostlycomefromstudies ofthe eel,Anguillaanguilla (Linnaeus), thus certainly underestimatingthe range ofintroduced parasites (Blanc1997,2001).Wefeel,however, that itisimportanttoprovidesuch a
.
firstcheck-listbecausemany of theinvertebratesinvolvedarenowadays com- monintheIPandformany ofthem itislargelyunknownevenbybiologists that they are not indigenous to the IP. Increasing the awareness on the introduced statusandcurrent distribution ofthesespeciesisessentialtoreduce theirspreadandimpact.
Wecompiledanimal speciescited(byMarch2006)ascurrently naturalized inIberianinlandwatersfromthescientificliterature andunpublishedSpanish Ph.D.theses ( speciesfromestuaries and saline coastal lagoons but excluded purely marine taxa and terrestrial animal speciesnot strictly linkedtoaquatic ecosystems.Welistinvertebrate and vertebrate speciesintroduced byhumansand currently naturalized, i.e. species thatreproduceandsustain populations inthewildwithout human intervention(seee.g.Richardson etal.2000,Pysˇeketal.2004).Afewuncer- taincasesarelistedinaseparate table.Theintroduced originof parasite invertebrates isparticularly uncertainbut wefollowedBlanc(1997,2001), whohasrecentlyprovidedacomprehensive listofaquatic parasitesintroduced toEurope,together withtheirnativedistribution.
NATURALIZEDANIMALSINIBERIANINLANDWATERS
Theinvertebrate andvertebratespeciesnaturalized inIberianinlandwatersare listedinTables1 and 2, respectively.Afewcases,forwhich itisnot clear whether thespeciesisindigenous totheIPorwhether theyhave established, are listedin Table3. We found 45 invertebrate and 28 vertebrate species certainly naturalized atpresentinIberianinlandwaters.
Amongthe45invertebrates, 12wereparasites(mostlyPlatyhelminthes flatworms), mainly offreshwater fishand introduced to Europe from Asia with common carp (Cyprinuscarpio),goldfish(Carassiusauratus),orJapanese eel(AnguillajaponicaTemminckandSchlegel)(seeBlanc1997,2001); several oftheparasiteshavenowbeenrecordedonfishspeciesindigenousorendemic to the IP(seereferences in Table1). The remaining 33 invertebrates were free-livingspecies,mostly crustaceans (18 species)or molluscs (6 species). Mostofthe 28 vertebrates were fish(23 species),and there was noaquatic birdnaturalized andonlyoneamphibian andonereptilianspecies.
Thecontinent oforiginwassignificantly differentbetween vertebrates and invertebrates (independencetest;x2 ¼37:1,df¼7,P0.0005)becausemost
naturalized vertebrateswerenativetotherestofEurope(43%ofthe28species) orNorth America (29%),originsthat inturn wererare among invertebrates (0and12%,respectively),whichpredominantlycamefromAsia(38%).There wasnosignificantvariation inoriginbetweenfree-livingandparasiticinverte- brates (x2 ¼6:5, df¼5, P ¼0.26) or between crustaceansand molluscs (x2 ¼3:8,df¼5,P¼0.59).
Table 1 Non-indigenous invertebrate speciesnaturalized ininlandwatersofthelberianPeninsula (IP).Whenaspeciesis probablyan oldintroduction,buttheintroductiondateislargelyunknown,aquestion mark isgivenasthefirstrecorddate.Thehabitats fornon- parasite speciesarecodedas:1,streamsandrivers(excludingestuaries);2,lakesandreservoirs;3,pondsandpools;4,ricefields;and5, estuarine orsalinewaters.
Indigenous
Firstrecord
Habitat/
LacasaMilla´n1999
JuaristiandSalvador2006
Table 1Continued.Indigenous
GROUPSpeciesdistribution / Firstrecord Habitat/
intheIPParasite / ReferencesfortheIP / Otherreferences
PLATYHELMINTHESTREMATODA
Phyllodistomumfolium(Olfers,1816)Ponto–Caspian? / 2004 / parasite / Periba´n˜ezetal.2006
PLATYHELMINTHESCESTODA
BothriocephalusacheilognathiYamaguti Asia / ? / parasite / LacasaMilla´n1992 / Blanc1997,2001
[¼Bothriocephalusopsarichthydis
(Yeh)] NEMATODA
[¼P.jenkinsi(Smith)] MOLLUSCABIVALVIA
Corbiculafluminea(Mu¨ller)Asia,Africa,
andAustralia
19811,3,5Vidal-AbarcaandSua´rez1985,McMahon2000
Escotetal.2003
Dreissenapolymorpha(Pallas)Ponto–Caspian18801,2Vidal-AbarcaandSua´rez1985, Altabaetal.2001
Hall
Ficopomatusenigmaticus(Fauvel)
(¼MercierellaenigmaticaFauvel) CRUSTACEABRANCHIOPODA
IndianOcean19245Rioja1924,
Fischer-Piette1951
Artemiafranciscana(Kellog)North,Central,
andSouthAmerica
1980s5Amatetal.2005
WlassicsiapannonicaDaday / Eurasia / 1990s / 4 / Martinoyetal.2006 / Leonietal.1999CRUSTACEAOSTRACODA
DolerocyprissinensisSars / Asia / 1986 / 3,4 / Fore´setal.1986, / Rossietal.2003
Baltana´setal.1996
Ilyodromusviridulus(Brady) / Australia and / 1996 / 4 / Baltana´setal.1996 / Rossietal.2003
NewZealand
Cyprissp.(¼Cyprissubglobosa
Sowerby) / America,Africa,
andAsia? / 1986 / 2,4 / Fore´setal.1986,
Baltana´setal.1996 / Whatleyetal.2003
Isocyprisbeauchampi(Paris) / Africa / 1976 / 2,4 / Armengol1976, / Rossietal.2003
Baltana´setal.1996
Stenocyprismajor(Baird)Asia19863,4Fore´setal.1986,
Baltana´setal.1996
Strandesiavavrai(Mu¨ller)Africa19834PauloandMoutinho 1983, Baltana´setal.1996
Strandesiavinciguerrae(Masi)Africa19864Fore´setal.1986, Baltana´setal.1996
Tanycyprissp.Asia19883,4Fore´s1988,
Baltana´setal.1996
Rossietal.2003
Rossietal.2003
CRUSTACEACOPEPODA
AcartiatonsaDanaNorthand
SouthAmerica
1990s5Sobral1985,Frischetal.2005
LernaeacyprinaceaLinnaeusAsiaparasite Morenoetal.1986,Gutie´rrez- GalindoandLacasa-Milla´n
2005
Blanc1997,2001
SouthAtlanticcoasts
Table 2Non-indigenous vertebrate species naturalized in inland waters ofthe
Iberian Peninsula (IP). The habitats are coded as: 1, rivers (excluding estuaries);
2,lakesandreservoirs;3,ricefields;and4,estuarine orsalinewaters.
GROUPSpecies
Indigenous distribution
Firstrecord
intheIPHabitat ReferencesfortheIP
PISCES
Abramisbjoerkna(Linnaeus)Europe19951,2Doadrio2002
Abramisbrama(Linnaeus)Europe20042Benejametal.2005
Alburnusalburnus(Linnaeus)Europe19921,2Doadrio2002
Ameiurusmelas(Rafinesque)NorthAmerica19101,2Doadrio2002
CarassiusauratusLinnaeusAsia17th century 1,2,3 Doadrio2002
CobitisbilineataCanestriniEurope20021Doadrio2002
CyprinuscarpioLinnaeusEurasia17th century1,2Doadrio2002
EsoxluciusLinnaeusEurope19491,2Doadrio2002
Fundulusheteroclitus
(Linnaeus)
NorthAmerica19704Doadrio2002
Gambusiaholbrooki(Girard) / NorthAmerica / 1920 / 1,2,3,4 / Doadrio2002Herichthysfacetum(Jenyns) / SouthAmerica / 1985 / 1,2 / Doadrio2002
Huchohucho(Linnaeus) / Europe / 1970 / 1 / Doadrio2002
Lepomisgibbosus(Linnaeus) / NorthAmerica / 1910 / 1,2 / Doadrio2002
Micropterussalmoides / NorthAmerica / 1955 / 1,2 / Doadrio2002
(Lacepe`de)
Oncorhynchusmykiss(Walbaum) NorthAmerica 19th century1,2Doadrio2002
Perca fluviatilisLinnaeusEurope19751,2Doadrio2002
PoeciliareticulataPetersSouthAmerica20001,4Doadrio2002
Pseudorasboraparva
(TemminckandSchlegel)
Asia20011Caiolaand
Sostoa2002
Rutilusrutilus(Linnaeus)Europe19101,2Doadrio2002
Salvelinusfontinalis(Mitchill)NorthAmerica 19th century1,2Doadrio2002
Sanderlucioperca(Linnaeus)Europe19751,2Doadrio2002
Scardiniuserythrophthalmus
(Linnaeus)
Europe19101,2Doadrio2002
SilurusglanisL. / Europe / 1974 / 1,2 / Doadrio2002AMPHIBIAANURA
DiscoglossuspictusOtth / Africa / 1900 / 1 / Pleguezuelos2002
REPTILIACHELONIA
Trachemysscripta(Schoepf) / America / 1985 / 1,2,3,4 / Pleguezuelos2002
MAMMALIA
MustelavisonSchreber / NorthAmerica / 1978 / 1,2,3,4 / Ruiz-Olmoetal.1997,
PalomoandGisbert2002
MyocastorcoypusMolina / SouthAmerica / 1970 / 1 / PalomoandGisbert2002
Ondatrazibethicus(Linnaeus) / NorthAmerica / 2002 / 1 / Elosegi2004
Table 3 Animal species possibly introduced to inland waters of the Iberian
Peninsula (IP), but with uncertain status. Some species are cryptogenic (Carlton
1996),i.e.itisverydifficulttoknow whether they areindigenous orintroduced; the otherspecieshavebeenreportedinthewildbutitis uncertainwhether theyhave establishedpermanentpopulations (naturalized).
GROUPSpecies
Possibly indigenous
totheIP
Uncertain
establishmentReferences
MOLLUSCAGASTROPODA
Ferrissiawautieri(Mirolli)Anderson2005 [¼F.clessiniana(Jickeli)] yes no
CRUSTACEADECAPODA
Austropotamobiusitalicus (Faxon)/Austropotamobius pallipes(Lereboullet)
PISCES
yesnoGrandjean etal.2001
AcipenserbaeriBrandtnoyesElviraandAlmodo´var2001
Aphaniusfasciatus(Valenciennes)noyesDoadrio2002
Ctenopharyngodonidella
(Valenciennes)
noyesJ.M.Queral2005,personal communication
Ictaluruspunctatus(Rafinesque)noyesDoadrio2002
Oncorhynchuskisutch(Walbaum)noyesDoadrio2002
Tincatinca(Linnaeus)yesnoDoadrio2002
AMPHIBIA
BufomauritanicusSchlegelnoyesPleguezuelos2002
RanacatesbeianaShawnoyesPleguezuelos2002
Ranaridibunda,Ranakl.
esculenta,Ranalessonae
REPTILIACHELONIA
noyesAranoetal.1995,
Garc´ıa-Par´ısetal.2004
Pelodiscussinensis(Wiegmann) noyesPleguezuelos2002
AVES
Aixgalericulata(Linnaeus)noyesGAE2006
Ansererythropus (Linnaeus)noyesGAE2006
Brantacanadensis(Linnaeus)noyesGAE2006
Oxyurajamaicensis(Gmelin)noyesGAE2006
MAMMALIA
Castor fiberLinnaeusyesyesCen˜aetal.2004
Themain habitat alsodifferedbetween vertebrates and invertebrates (inde- pendencetest;x2 ¼22:8,df¼4,P0.0005),becausetheformerweremostly presentinstreamsandrivers(26ofthe28species werepresentinstreamsand rivers)orlakesandreservoirs,whereas severalinvertebrates wereonlypresent inestuaries/salinewaters(e.g.severaldecapodcrustaceansintroduced intothe Guadalquivir Riverthroughballastwater)orinricefields(namelyostracods).
Themechanism ofintroductionisobviouslyalsodifferentforinvertebrates and vertebrates, because most of the former are accidental introductions (e.g.Asianostracodsinricefields,ballastwater,etc.),whereasmostfishspecies havebeenintroduced intentionally(nowadays illegally). Therefore,naturalized vertebratesandinvertebratesshowedoppositepatterns, withtheformer(mostly fish)intentionallyintroduced from the rest ofEurope or North America to Iberian streams and reservoirs and most invertebrates originating fromAsia andaccidentallyintroduced toestuaries orricefields.
UNCERTAINCASES
Wefoundfourspeciesforwhichitisuncertainwhether thespeciesisindigen- oustotheIPand13speciesthat theymaynothaveestablished(Table3).An interesting caseillustrating boththelackofknowledgeoninvasivespeciesand thepowerofmoderngenetictechniques isthecrayfishoftheAustropotamobius pallipesspeciescomplex.Untilthe1980sthepopulations intheIPweregener- allyregarded asanendemicspeciesorsubspeciesinstrong declineduetothe introductionoftheoomyceteAphanomycesastaciSchikorawithNorthAmerican crayfish(Mart´ınezetal.2003).Grandjean etal.(2000) showedthattwospecies (A.pallipesand Austropotamobiusitalicus)could be distinguished within the speciescomplexandthat Spanishpopulations wereveryclosetosomeItalian populations, sotheymightbeofanthropogenicorigin,asalready proposedby Albrecht(1983), andshouldberegardedasA.italicus.Grandjean etal.(2001) demonstratedadrasticbottleneckinSpanishpopulations butdiscussedseveral potentialmechanisms alternativetothehypothesisofintroductionbyhumans. Withfurther geneticanalyses, Tronteljetal.(2005) supported the anthropo- genicoriginfortheSpanishpopulationsbutdidnotfindunequivocal separation betweenA.pallipesandA.italicus(butseealsoSchulzandGrandjean 2005).
These genetic techniques might also prove useful for tench [Tincatinca (Linnaeus)] in the IP. Tench is indigenous to many parts of Europe but consideredintroduced intoItaly(Bianco1998)and Portugal (Almac¸a1995). Thislatter country shares itslargestriverbasins(Duero,Tajo,and Guadiana rivers) with Spain. There are doubts about its indigenous status in Spain (Doadrio2002).Infact,Go´mezCaruana and D´ıazLuna(1991) consideredit introduced into the IParound the 17th century. There are records oftench stocking bymonks in Spanish and Portuguese ponds several centuries ago (Almac¸a 1995, Garc´ıa-Berthou and Moreno-Amich 2000). As far as we know, no phylogeographic study on tench has been performed, in contrast tomanyotherEuropeancyprinids,although theycouldbemosthelpfulin clarifyingitsnativedistribution.
Asimilar,moresolvedexampleof‘‘cryptogenic’’species(seeCarlton1996) isthe case ofthe freshwater snail Physellaacuta(Draparnaud).Thisspecies was firstdescribedfromEurope(Drapanaud,1805), namely fromthe River
Garonne, near Bordeaux(France).Thisspeciesiswidelydistributed inthe IP andtherestofwesternEurope(Vidal-AbarcaandSua´rez1985)andinhabitsall typesoffreshwaters. Ithasbeengenerally regarded asindigenous tocontin- entalEurope(Haas1929,Germain1930,MacanandCooper1977,Girodetal.
1980,Vidal-AbarcaandSua´rez1985)anditspresenceinNorthAmericawas not reported until the 1990s(Wu etal. 1997).Nowadays, three types of evidenceindicate,however, that P.acutaisindigenous toNorthAmericaand nottoEurope:(i)thelackofrecordsofPhysellashellsfromEuropeansediments olderthan the 18th century (Lozek1964); (ii)recent studies using internal morphology comparisons (Anderson 2003)and reproductive isolation experi- ments(Dillonetal.2002)showingthatatleastonePhysellaspeciesfromNorth America[Physellaheterostropha(Say)]isactuallyP.acuta;and(iii)somehistor- icaldataofthecottontradebetweenFranceandtheUnitedStatesinthe18th century that could explain the arrival ofthis speciestothe RiverGaronne, whereitwasfirstobserved(Anderson2003).
ThecaseofP.acutaillustratestheimportance ofhistoricaldataandthefossil recordastoolsfortheidentificationofoldintroductionsbyman.Fossilrecords have been very helpful to establish the introduced nature ofostracods and suggestthat dispersalbyman ofmany otherinvertebrates isveryoldandhas beengenerallyneglected(McKenzieandMoroni1986,Rossietal.2003).
Theothergroupof speciesinTable3arespeciesthat havebeenreportedin thewildbut itisuncertainwhether they have established. Thereareseveral other NISthat havebeenrecordedinthewild(seee.g.ElviraandAlmodo´var
2001, Pleguezuelos2002)buthavecertainlynotestablishedpermanent populations.
ECOLOGICALIMPACT
TheecologicalimpactofmostoftheseNIS islargelyunknownwithafew exceptions.Theredswampcrayfish,Procambarusclarkii,hasalteredthefunc- tioning and structureofmany aquatic ecosystemsinthe IPreducing macro- phytes and associated species, among other impacts (Geiger et al. 2005, Rodr´ıguezetal.2005, Chapter28).Theeastern mosquitofish(Gambusia holbrooki)hasbeenexperimentallydemonstratedtoaffectendemiccyprinodon- tiform fishes[Aphaniusiberus(Valenciennes) and Valenciahispanica(Valenci- ennes)] byresource and interference competition (Rinco´n etal.2002). The zebramussel(Dreissenapolymorpha)isoneofthebestknown invasivespecies and, althoughitisa veryoldintroductioninto Portugal, only recently has itbeenintroduced toSpainthroughtheEbro River,whereitiswidespread nowadays andmightaffect theendangered giantpearlmussel,Margaritifera auriculariaSpengler (Altaba etal.2001).Thezebramussel isstillnot wide- spread in the IP, but it will probably be fostered by the illegal, poorly controlled introductionand translocationsoffishthat are stillveryfrequent.
The polychaete Ficopomatusenigmaticus(Fauvel) isvery abundantin some Spanishcoastallagoonsandprobablyprofoundlyaffectsitsecosystemfunction- ingbecauseitbuildslargereef-likeaggregates (Schwindt and Iribarne 1998). Manypiscivorousfish havebeenintroduced intotheIPandsomeunique ecosystemssuchasLakeBanyoleshavebeenprofoundlyalteredandarenow- adays completely dominated by NIS (Garc´ıa-Berthou and Moreno-Amich
2000).
Thedistribution, abundance,andimpactofintroduced parasitesintheIPis largelyunknownbut somespeciessuch asLernaeacyprinaceaarewidespread (Morenoetal.1986,Gutie´rrez-Galindoand Lacasa-Milla´n2005)and several ofthem have now been recorded on endemic fishspecies(seereferences in Table1).TheswimbladdernematodeAnguillicolacrassus,whichwastransferred fromitsindigenous host (the Japanese eel,A.japonica)to the European eel (A.anguilla), can severelyimpairswimbladder function (andthus possibly spawningmigration)andhascausedmortalitiesin bothfarmedandwild populations in the presence ofother stressors (Kirk2003).Similarly to the case ofcrayfish plague, Gozlanetal.(2005) have recently shown that the topmouth gudgeon,Pseudorasboraparva,anAsiaticcyprinidhighlyinvasivein Europeand recently introduced to the IP,carries a pathogen that strongly affectsindigenouscyprinids.
Giventheenormous impactofthefewwell-investigatedinvasivespecies,the considerablenumber ofintroduced species,andthepresenceintheIPofmany endemicspeciesofplants (Me´dailand Que´zel1999),freshwater fish(Doadrio
2002),andamphibians (Pleguezuelosetal.2002),theoverallpotentialimpact ofthesenaturalized speciesisenormous and should beurgently investigated. Theroomformanagementand educational improvement bypublicadminis- trations toprevent further introductionsandtranslocationsandtoreducethe spreadofinvasivespeciesisevenlarger.Wehopethispaperwillcontribute to theimprovedunderstandingandcontrolofinvasivespeciesinEuropeanwaters.
ACKNOWLEDGEMENTS
We thank J.F. Gutie´rrez-Galindo, F. Padro´s, and anonymous reviewers forhelpful comments. Thisstudy was financially supported bythe Spanish Ministry of Science and Technology (REN2003–00477) and the Ministry ofUniversities, Research and Information Society (DURSI),Government of Catalonia(DistinctionAwardforuniversityresearch 2004toEGB).
REFERENCES
Albrecht, H.1983.Besiedlungsgeschichte und urspru¨nglich holoza¨neVerbreitung der europa¨ischenFlußkrebse(Decapoda:Astacidae).Spixiana6,61–77.
Alcaraz,C.,A.Vila-Gispert,andE.Garc´ıa-Berthou. 2005. Profilinginvasivefishspecies: theimportanceofphylogenyandhumanuse.DiversityandDistributions11,289–298. Almac¸a,C.1995.Fishspeciesandvarietiesintroduced intoPortuguese inland waters.
Publicac¸o˜esavulsasdoMuseuBocage,Lisboa,Portugal.
Altaba,C.R.,P.J. Jime´nez,andM.A.Lo´pez.2001.Eltemidomejillo´ncebraempiezaa invadirlosr´ıosespan˜ olesdesdeelcursobajodelr´ıoEbro.Quercus118,50–51.
Amat, F.,F.Hontoria, O.Ruiz,A.J.Green,M.I.Sa´nchez,J.Figuerola,and F.Hortas.
2005.The American brine shrimp as an exotic invasive species in the western
Mediterranean.BiologicalInvasions7,37–47.
Anderson, R.2003.Physella(Costatella)acutaDraparnaudinBritainand Ireland –its taxonomy, origins and relationships to other introduced Physidae. Journal of Conchology38, 7–21.
Anderson,R.2005.Anannotatedlistofthenon-marinemolluscaofBritainandIreland. JournalofConchology38, 607–638.
Aranda, C.,R.Eritja,andD.Roiz.2006.Firstrecordandestablishment ofthemosquito
AedesalbopictusinSpain.MedicalandVeterinaryEntomology20, 150–152.
Arano, B.,G. A.Llorente,M.Garc´ıa-Par´ıs,andP.Herrero.1995.Speciestranslocation menacesIberianwaterfrogs.Conservation Biology9,196–198.
Armengol, J.1976.Crusta´ceosacua´ticosdelCotodeDon˜ana. Oecologiaaquatica 2,
93–97.
Bagun˜a`,J.,E.Salo´,and R.Romero. 1980.Lesplana`riesd’aigu¨esdolcesaCatalunya i lesIllesBalears.I.Clausistema`ticaidistribucio´ geogra`fica.Butllet´ıdelaInstitucio´ Catalana d’Histo`riaNatural 45, 15–30.
Bakke,T.A.,P.D.Harris,andJ.Cable.2002.Hostspecificitydynamics:observationson gyrodactylidmonogeneans.InternationalJournalforParasitology 32, 281–308.
Baltana´s, A., B.Beroiz,and A. Lo´pez.1996.Listafaun´ıstica y bibliogra´ficade los
ostra´codosno-marinos (Crustacea, Ostracoda)delaPen´ınsula Ibe´rica,IslasBaleares eIslasCanarias.Asociacio´nEspan˜oladeLimnolog´ıa,Madrid,Spain,71pp.
Benejam, L.,J.Carol, C.Alcaraz, and E.Garc´ıa-Berthou. 2005.First record ofthe
common bream (Abramisbrama)introduced totheIberian Peninsula. Limnetica24,
273–274.
Bianco,P.G.1998.Freshwater fishtransfersinItaly:history,localmodificationoffish composition,andapredictiononthefutureofnativepopulations. Pages165–197in I. J.Cowx,editor.Stockingandintroductions offishes.FishingNewsBooks,Blackwell, Oxford,UK.
Blanc,G.1997.L’introductiondesagentspathoge`nes danslese´cosyste`mesaquatiques:
aspects the´oriques et re´alite´s. Bulletin Franc¸ais de la Peˆche et de la Pisciculture
344/345,489–513.
Blanc,G.2001.Introductionofpathogens inEuropean aquatic ecosystems: attempt ofevaluation andrealities.Pages37–56 inA.UriarteandB.Basurco,editors.Environ- mental impactassessment ofMediterranean aquaculturefarms.CIHEAM-IAMZ, Zaragoza,Spain.
Brinkhurst, R.O.andB.G.M.Jamieson.1971.AquaticOligochaetaoftheworld.Oliver andBoyd,Edinburgh,UK,860 pp.
Brown,D.S.,M.A.A.Gracio,andC.Meier-BrookC.1998.TheAsianfreshwater snail Gyrauluschinensis(Dunker,1848)(Planorbidae)inWestAfrica andEurope.Journalof AfricanZoology112,203–213.
Caiola, N.andA.deSostoa.2002.FirstrecordoftheAsiaticcyprinidPseudorasboraparva
intheIberianPeninsula. JournalofFishBiology61, 1058–1060.
Carlton, J. T. 1996. Biological invasions and cryptogenic species. Ecology 77,
1653–1655.
Cen˜a,J.C.,I.Alfaro,A.Cen˜a,U.Itoitz,G.Berasategui, and I.Bidegain.2004.Castor europeoenNavarra yLaRioja.Galemys16, 91–98.
Clavero,M.andE.Garc´ıa-Berthou.2005.Invasivespecies arealeadingcauseofanimal extinctions.TrendsinEcologyandEvolution20, 110.
Cuesta, J.A., J.E.Garc´ıa-Raso, and J.I. Gonza´lez-Gordillo.1991.Primera cita de Rhithropanopeusharrisii(Gould,1814)(Crustacea, Decapoda,Brachyura,Xanthidae) enlaPen´ınsula Ibe´rica.Bolet´ındelInstituto Espan˜oldeOceanograf´ıa7,149–153.
Cuesta, J.A.,E.Gonza´lez-Ortego´n,P.Drake,and A.Rodr´ıguez.2004.Firstrecord of
PalaemonmacrodactylusRathbun,1902 (Decapoda, Caridea, Palaemonidae) from
Europeanwaters. Crustaceana77, 377–380.
Cuesta,J.A.,E.Gonza´lez-Ortego´n,A.Rodr´ıguez,F. Baldo´,C.Vilas,andP.Drake.2006. Thedecapodcrustaceancommunity oftheGuadalquivirEstuary(SWSpain):seasonal andinter-year changes incommunity structure.Hydrobiologia557,85–95.
Cuesta,J.A.,L. Serrano, M.R.Bravo,andJ.Toja.1996.Fournewcrustaceansinthe
Guadalquivirriverestuary(SWSpain),includinganintroducedspecies.Limnetica12,
41–45.
Dana, E.D.and S.Viva. 2006.Stenopelmusrufinasus Gyllenhal 1836(Coleoptera: Erirhinidae)naturalized inSpain.TheColeopteristsBulletin60, 41–42.
Dillon,R.T.,A.R.Wethington,J.M.Rhett, and T.P.Smith. 2002.Populations of
the European freshwater pulmonate Physa acuta are not reproductively isolated from American Physa heterostropha or Physa integra. Invertebrate Biology 121,
226–234.
Doadrio, I., editor. 2002.Atlas y libro rojo de los peces continentales de Espan˜a. MinisteriodeMedioAmbiente,Madrid,Spain,347 pp.
Draparnaud,J.P.R.1805.Histoirenaturelle desmollusquesterrestresetfluviatilesdela
France.Paris,164 pp.
Elosegi,M.M.2004.Observacio´ndeuna rata almizclera,Ondatrazibethicus(Linnaeus,
1766)enEzkurra(Navarra). Galemys16, 63–64.
Elvira,B.and A.Almodo´var. 2001.Freshwater fishintroductionsinSpain:factsand figuresatthebeginningofthe21stcentury. JournalofFishBiology59, 323–331. Eritja,R.,R.Escosa,J.Lucientes,E.Marque`s, R.Molina,D.Roiz,and S.Ruiz.2005.
Worldwide invasion of vector mosquitoes: present European distribution and challengesforSpain.BiologicalInvasions7,87–97.
Escot,C.,A.Basanta,F.Cobo,andM.A.Gonzalez.2003.SobrelapresenciadeMytilopsis leucophaeata (Conrad, 1831)(Bivalvia,Dreissenacea, Dreissenidae) en elr´ıo Guadalquivir (surdelaPeninsula Ibe´rica).Graellsia59, 91–94.
Espadaler,X.and C.A.Collingwood.2001.Transferred ants intheIberian Peninsula
(Hymenoptera, Formicidae).NouvelleRevued’Entomologie17, 257–263.
Ferreira, M.T. 1985.Occurrence ofthe freshwater medusa Craspedacustasowerbyi Lank.1880 (Hidrozoa:Olindiidae) inPortuguese reservoirs.BoletimdaSociedade PortuguesadeCieˆnciasNaturais 22, 41–46.
Fischer-Piette,E.1951.Stationsdel’anne´lidetubicoleMercierellaenigmaticaFauvelsurla coˆteNordd’Espagne.BulletinduLaboratorieMaritimedeDinard34, 7–9.
Fore´s,E.1988.Elsostra`codesdelsarrossars delDeltadel’Ebre:sistema`tica, ecologia
i distribucio´ geogra`fica. Butllet´ı
15–30.
de la Institucio´ Catalana d’Histo`ria Natural 45,
Fore´s, E.,M.Mene´ndez, and F.A. Com´ın. 1986.Contribucio´n al conocimiento de
Crusta´ceosyRot´ıferosdelDeltadelEbro. Miscella`niaZoolo`gica10, 105–111.
Frisch,D.,H.Rodr´ıguez-Pe´rez,and A.J.Green.2005.Invasion ofartificialponds in Don˜ana Natural Park, southwest Spain, byan exotic estuarine copepod. Aquatic Conservation-Marine andFreshwater Ecosystems15, 483–492.
GAE(Grupo de Aves Exo´ticas de SEO/BirdLife).2006.Aves invasoras en Espan˜a: listadeespeciesenlascategor´ıas CyE. idPrograma¼17.
Gallastegi,I.,A.Rallo,andM.F.Mulcahy. 2002.Areport ofAnguillicolacrassusfrom
Spain.BulletinoftheEuropeanAssociationofFishPathologists22, 283–284.
Garc´ıa-Berthou, E.and R. Moreno-Amich. 2000. Introductionofexotic fishinto a
Mediterraneanlakeovera90-year period.Archivfu¨rHydrobiologie149,271–284. Garc´ıa-Par´ıs, M.,A.Montori, and P.Herrero. 2004.Amphibia, Lissamphibia. Pages
640 inM.A.Ramosetal.,editors.Fauna Ibe´rica,24. MuseoNacional deCiencias
Naturales, Madrid,Spain.
Gelder,S.R.1999.Zoogeographyofbranchiobdellidans(Annelida)andtemnocephali- dans(Platyhelminthes)ectosymbioticonfreshwater crustaceans,andtheirreactions tooneanother invitro.Hydrobiologia406,21–31.
Geiger,W.,P.Alcorlo,A.Baltana´s,and C.Montes. 2005.Impact ofan introduced
Crustacean onthe trophic websofMediterraneanwetlands. BiologicalInvasions 7,
49–73.
Germain, L. 1930.Mollusquesterrestres etfluviatiles.InFaune deFrance21/22. Lechevalier,Paris,France,897 pp.
Girod,A.,I.Bianchi, and M.Mariani. 1980.Gasteropodi,1(Gastropoda: Pulmonata, Prosobranchia:Neritidae,Viviparidae,Bithyniidae,Valvatidae).InS.Ruffo, editor. Guide per il riconoscimento delle specie animali delle acque interne italiane, 7. ConsiglioNazionaledelleRicherche,Verona,Italy,86pp.
Gollasch,S.andK.Riemann-Zu¨rneck.1996.Transoceanic dispersalofbenthic macro- fauna:Haliplanellalineata(Verrill,1898)(Anthozoa,Actinaria)foundonaship’shullin ashipyarddockinHamburgHarbour,Germany.Helgola¨nderMeeresuntersuchungen
50,253–258.
Go´mez Caruana,F.and J.L.D´ıazLuna. 1991.Gu´ıa de peces continentales de la
Pen´ınsula Ibe´rica.Penthalon,Madrid,Spain.
Go´mez-Juaristi,M.andA.Salvador.2006.Anguila–Anguillaanguilla.InL.M.Carrascal and A.Salvador, editors. Enciclopediavirtual delosvertebrados espan˜ oles.Museo NacionaldeCienciasNaturales, Madrid,Spain.
Gozlan,R.E.,S.St Hilaire,S.W.Feist,P.Martin,andM.L.Kent.2005.Diseasethreat to
Europeanfish.Nature (London)435,1046.
Grandjean, F.,D.J.Harris, C.Souty-Grosset, and K.A.Crandall. 2000. Systematics ofthe European endangered crayfish species,Austropotamobiuspallipes(Decapoda: Astacidae).JournalofCrustacean Biology20, 522–529.
Grandjean, F.,N.Gouin,C. Souty-Grosset,andJ.Die´guez-Uribeondo.2001.Drastic bottlenecksintheendangered crayfishspeciesAustropotamobius pallipesinSpainand implicationsforitscolonizationhistory.Heredity86, 431–438.
Gu¨nther, H.2004.Trichocorixaverticalisverticalis(Fieber),einenearktische Ruderwanze inEuropa.Mitteilungen desInternationalenEntomologischen Verereins29, 45–49. Gutie´rrez-Galindo, J. F. and M. I. Lacasa-Milla´n. 1999. Monogenea para´sitos de
Cyprinidaeenelr´ıoLlobregat(NEdeEspan˜a)(BarbushaasiPetersen, 1925). Revista
AquaTIC7.
Gutie´rrez-Galindo,J.F.andM.I.Lacasa-Milla´n.2005.Population dynamicsofLernaea cyprinacea(Crustacea: Copepoda)onfourcyprinidspecies.DiseaseofAquaticOrgan- isms67, 111–114.
Gutie´rrez-Yurrita, P.J.,J.M.Martinez,M.Ilheu, M.A.Bravo-Utrera, J.M.Bernardo, and C.Montes. 1999.The status ofcrayfish populations in Spain and Portugal. Pages161–192inF.Gherardi and D.Holdich,editors.CrayfishinEuropeasalien species:Howtomake the bestofabadsituation? A.A.Balkema,Rotterdam, The Netherlands.
Haas,F.1929.Fauna malacolo´gicaterrestre ydeagua dulcedeCatalun˜a.Treballsdel
MuseudeCie`nciesNaturals 13, 1–491.
Habsburgo-Lorena, A.S.1978. Presentsituation ofexoticspeciesofcrayfishintroduced intoSpanishcontinentalwaters. Freshwater Crayfish4,175–184.
Hutchinson,G.E.1931.On the occurrence ofTrichocorixa Kirkaldy (Corixidae, Hemiptera-Heteroptera) in salt water and its zoo-geographical significance. The AmericanNaturalist 65, 573–574.
Kirk, R. S. 2003. The impact of Anguillicolacrassus on European eels. Fisheries
ManagementandEcology10, 385–394.
Lacasa Milla´n, M.I.1992.Introduccio´n alaparasitofaunadepecescipr´ınidos dela cuenca media del r´ıo Llobregat. Ph.D. dissertation, Universidad Auto`noma de Bacelona,Barcelona,Spain.
Laine,A.O.,J.Mattila, and A.Lehikoinen. 2006.Firstrecord ofthe brackish water dreissenidbivalveMytilopsisleucophaeatainthenorthernBalticSea.AquaticInvasions
1,38–41.
Leoni,B.,M.Cotta-Ramusino, and F.G.Margaritora. 1999. Seasonal succession of
CladoceransinaricefieldinItaly.Hydrobiologia391,241–247.
Lloret,F.,F.Medail,G.Brundu,andP.E.Hulme.2004.Localandregionalabundanceof exoticplantspecies onMediterraneanislands:arespecies traitsimportant? Global EcologyandBiogeography13, 37–45.
Lozek,V.1964.Quarta¨rmollusken derTschechoslowakie. Rozpravyu´stredn´ıho u´stavu geologicke´ho31, 1–374.
Macan, T.T.and R.D.Cooper.1977.Akeytothe Britishfresh-and brackish-water
gastropods.Freshwater BiologicalAssociation,Ambleside,UK,46pp.
Mack,R.N.,D.Simberloff,W.M.Lonsdale,H.Evans,M.Clout,andF.A.Bazzaz.2000. Bioticinvasions: causes, epidemiology,globalconsequences, and control. Ecological Applications10, 689–710.
Maillo, P.A.,M.A.Vich,H.Salvado,A.Marques,andM.P.Gracia.2005.Parasites of Anguillaanguilla(L.)from three coastal lagoons ofthe RiverEbrodelta (Western Mediterranean).ActaParasitologica 50, 156–160.
Margalef,R.1974.Ecolog´ıa.Omega,Barcelona,Spain,951 pp.
Martinez,R.,E.Rico,andE.Alonso.2003.CharacterisationofAustropotamobiusitalicus (Faxon, 1914) in a central Spain area. Bulletin Franc¸ais de la Peˆche et de la Pisciculture370–371,43–56.
Martinoy, M.,D.Boix,J.Sala,S.Gasco´n,J.Gifre,A.Argerich, R.Barrera, S.Brucet, A. Badosa, R. Lo´pez-Flores,M.Me´ndez, J.M.Utge´, and X.D.Quintana.2006. Crustacean andaquatic insectassemblagesintheMediterraneancoastalecosystems ofEmporda`Wetlands(NEIberianPeninsula). Limnetica25, 53–70.
McKenzie,K.G.andA.Moroni.1986.Manasanagent ofcrustaceanpassivedispersal via useful plants – exemplifiedby ostracoda ospitiesteri ofthe Italian ricefields ecosystem –and implications arising therefrom. Journal ofCrustacean Biology6,
181–198.
McMahon,R.F.2000.Invasivecharacteristicsof thefreshwater bivalveCorbicula fluminea.Pages315–343inR.ClaudiandJ.H.Leach,editors.Nonindidenous Fresh- water Organisms:Vectors,BiologyandImpacts.LewisPublishers.BocaRaton,FL.