BASTERIA, 65:17-25, 2001 TheterrestrialMolluscaofEasterIsland(Gastropoda, Pulmonata) Christopher+B. Boyko &James+ R. Cordeiro Division ofInvertebrate Zoology,AmericanMuseumofNatural History, Central ParkWestat79th Street,New York,NewYork, 10024 [email protected] The terrestrial Mollusca ofEaster Islandaresurveyed,based onpublishedrecords, museum material, and recent collections in 1999. Six species areknown withcertaintyfrom Easter Island,five ofwhichhave nearlyworldwide distributions.All areknown orsuspectedtobe human introductions.Allopeasgracile(Hutton)isreportedforthe firsttimefromEaster Island and thepreviousrecord ofDeroceraslaeve(Müller)issuggestedtobemisidentified D.reticula- tum(Müller). Keywords: Gastropoda,Pulmonata,biogeography,introduced species,Pacific,Easter Island. INTRODUCTION ThesmallsouthPacificlandmassof EasterIsland, orRapa Nui,is situatednearly 3800 km fromthe SouthAmericanmainland(Chile) and over2200km from PitcairnIsland, it's nearestneighbour tothe west. Theislandisofvolcanicoriginandcontainsno conti- nentalor sedimentary rocks (Fischer & Love, 1993),only aggregations of shellandcoral sandonandneartheislandsfewbeaches.Theterrestrialinvertebratefaunaissparseand only the insects have beenstudied in any detail(Campos &Pena, 1973). There are few records ofterrestrialmolluscs on EasterIslandin the literatureand nosynthesis ofthe data.Thisis in marked contrast to ourknowledge ofthe marinemolluscs, which was summarizedby Rehder's (1980) monograph and added to by otherrecent papers (e.g. Osorio, 1991;1995). Thepresentstudy, basedonallknownliteraturerecords aswellas a collectionby oneoftheauthors(CBB) in 1999,summarizes theinformationavailableon the terrestrialmolluscsofEaster Island.This reportis confinedto thosetaxa whichare trulyterrestrialanddoesnot includemarinepulmonate taxa(e.g.,Siphonaria, Melampus; seeRehder [1980] foradetaileddiscussion ofthese taxa). The first record ofany terrestrialmollusc from Easter Island was a “Parmarionsp." (Ariophantidae) by Fuentes (1914). Noariophantid slugs have subsequently beenrepor- ted from Easter Island and, in the absence ofvoucher material, this record remains unconfirmed.Wesuspect thatFuentes'(1914)Parmarionwas actually oneoftheintrodu- cedslugspecies subsequently reported. Dall (1908) reported the nowcosmopolitan (ori- ginally circum-mediterranean) slugMilaxgagates(Draparnaud, 1801) from underrocks nearthe shore.Odhner (1922) reported this species as well, and addedtwo more taxa: Limaxarborum Bouchard-Chantereaux, 1838,andhis new genus and species Pacificella variabilisOdhner, 1922.Although not implicitly statedby Odhner, P.variabiliswas des- cribedas the first endemicterrestrialmolluscfromEasterIsland. However, itwas later shown toalso occur on many ofthe Pacificislands to the west (see Cooke & Kondo, BASTERIA, Vol. 65,No. 1-3,2001 18 1961). Theonly other reports ofterrestrialmolluscs from Easter Island were those of Rees(1955), who alludedtothepresence ofHelixaspersaMidler, 1774,andSteele(1957), whomerely compiled alist ofprevious records. Walden(1962) correctedthe identifica- tionofthe EasterIslandLimaxspecies toLimax(Lehmannia) valentiana(Ferussac, 1821), basedondissectionofthe specimens reported byOdhner (1922), butdidnot reportany newmaterial. Kuschel (1963) discussed the relationships of the terrestrial Easter Island faunawith selectotherPacificislandsandcitedfivespecies ofmolluscs. Heonly cited onetaxon by name(Melampuspascus Odhner, 1922),but the Milax, Limax andPacificella, as well as a , secondspecies of Melampus[later shownby Rehder(1980) tobea misidentifiedM. pas- cus] mustlogically have comprised hislist. Klemmer& Zizka (1993) repeated Kuschel's (1963)fivetaxa,including the"two"Melampusspecies, andlistedPacificella variabilis as an EasterIslandendemic. Themostrecent informationavailableonEasterIslandterrestri- almolluscswasin ashortpaperbyNaranjo-Garcla &Appleton (1999), wherethey added HelixaspersaandDeroceraslaeve(Miiller, 1774) to thefaunallist. In 1999,oneofus (CBB) spentthreeweeks onEasterIsland aspart ofa surveyofthe marine invertebratefauna; there was also opportunity to collect terrestrial fauna on numerous occasions, especially when the weather was too poor for marine work. Localitieswere sampled throughout the island, withthe exception of the Mt.Terevaka areawhich was not visited. True to the nature of an islandwith depauperate biota, the samples yielded very few specimens ofterrestrial molluscs, with only a single species (Helix aspersa) being foundin substantialnumbers.Allofthe terrestrialmolluscswere found nearareas ofhumanhabitation, current or past, andnone were foundin more remote areas ofthe island such as the Poike peninsula. Additionally, only a very few examplesofeachspecies werefounddespite intensivesearches ateachlocation.Limited asthese samples were, they containedtwo cosmopolitan taxa previously unreported, or reportedbutmisidentified,fromEasterIsland,andprobably introducedafter 1917when the last major collectionsofterrestrialmolluscs were made (see Odhner, 1922). These samples, combinedwith Odhner's (1922) specimens, allowed thiscomprehensive sum- maryofthe terrestrialmollusctaxa. Themostimportant conclusionthatcanbe drawn is that Easter Islandappearsto have noendemic terrestrialmolluscanfauna,which is perhaps not surprisinggiven itslocationandextreme isolation. Unless otherwisenoted, allliteraturecitationsreferto recordsofspecimens collected from Easter Island. The specimens collected in 1999are deposited in the American MuseumofNatural History, NewYork (AMNH); all of Odhner's specimens are in the collectionsoftheSwedishMuseumofNaturalHistory (SMNH). In'Materialexamined' thenumberofspecimens isaddedafterthecollectionno. SYSTEMATICACCOUNT FamilyAchatinellidae Pacificella variabilisOdhner, 1922 Pacificella variabilis Odhner,1922: 249-252,pi. 8, figs 15-17;Steele, 1957: 113 (list);Klemmer &Zizka, 1993:24. Tornatellinops variabilis (Odhner); Cooke & Kondo, 1961: 172-175, figs 76a-h,77; Naranjo-Garcia & Appleton, 1999: 120, 121. Boyko &Cordeiro: TerrestrialMolluscaofEasterIsland 19 Material examined. —Sta.707,EasterIsland (RapaNui),2.vii.l917,coll. K.Backstrom(SMNH 1668/5 shells,syntypes). Distribution. — This species is widely distributed on Pacific islands, from Truk (Carolines) to Easter Island (Preece, 1995: 287), and north to the Mariana Islands (Bauman, 1996: 19). Themapgiven by Cooke& Kondo(1961: fig.77) includedatleast 67 islandsfromwhichthe species was known atthattime. Many ofthose islands have distinctive morphotypes (Preece, 1995:287) thatmayrepresent endemic subspecies or even species, so the true range of this taxon (as definedby the type specimens from Easter Island) is unclear. Although originally described from Easter Island (Odhner, 1922),the species mostlikely was transported thereby humans froman unknownsour- ce population elsewhereinthe Pacific. Remarks. —Although the habitatof thisspecies onEasterIsland hasnotbeen repor- ted, itisabundantonbothnativeandintroducedplants onPitcairnIsland(Preece, 1995: 287). Itwaserroneously calledanEasterIslandendemicby Klemmer& Zizka(1993: 24). Itsplacement inPacificella Odhner, 1922,ratherthanTornatellinops Pilsbry, 1915,was con- firmedbybothPreece (1995) andCowieetal. (1995). Nospecimens were foundin 1999. Odhner's (1922) specimens in SMNH are syntypes, as are the "paratypes" in the Bishop Museum, Hawaii,citedby Cooke&Kondo (1961: 172). Family Limacidae Lehmanniavalentiana(Férussac, 1821) Limax arborum Bouchard-Chantereaux;Odhner, 1922: 249;Steele, 1957: 113 (list). Not L. arborum Bouchard-Chantereaux,1838,asynonym ofL. marginatusMiiller,1774. Limax (Lehmannia)valentiana (Ferussac);Walden, 1962:71-92,figs 5-7. LimaxmarginatusMidler;Naranjo-Garcia&Appleton,1999: 120,121. NotL. marginatusMiiller,1774. Material examined. — Sta. 705, Easter Island (Rapa Nui), 2.vii.l917, cod. K. Backstrom (SMNH 28094/7). Distribution.—This nearly cosmopolitan ground-dwelling (non-arboreal) slug is nati- vetothe IberianPeninsula,but hasbeenwidely spread byhumans(Kerney, 1999: 160). Walden(1962: 88-90) cited this species as occurring on allcontinents except Asiaand Antarctica. InNorthAmerica, it islargely confinedto greenhouses (Chichester & Getz, 1973:36). Remarks.—Thisspecies hasbeenreported fromEasterIslandunderthenameL. arbo- rumor itsseniorsynonymL. marginatus(see Pilsbry, 1948:529). Walden(1962) dissected oneoftheaboveexaminedspecimens, recognized thismisidentification, andnoted that many recordsofL. marginatus from tropical andsubtropical countrieswere actually L. valentiana.Nospecimens were foundonEaster Islandin1999. Deroceras reticulatum(Müller, 1774) (fig. 1) ?Deroceras laeve (Midler);Naranjo-Garcia&Appleton,1999:121. ?NotD. laeve(Midler,1774). 20 BASTERIA, Vol. 65,No. 1-3,2001 Fig. 1. Deroceras reticulatum(Müller, 1774) from AnaTe Pahu, Easter Island (RapaNui), largest specimen, 2.3 cmlength(AMNH300602). Material examined. —Under leaflitter indepressionofcollapsedlava tubenearcavemouth,AnaTe Pahu,Easter Island (RapaNui),21.viii.1999,coll.C. B.Boyko (AMNH300602/2). Distribution.—Thisspecies isnativeto Europe andis foundin mostofNorthAmerica andmany temperateand subtropical countries. It hasbeen introducedand spread by commerce throughout every temperateand subtropical region with European settle- ment(Burch, 1962:83; Robinson, 1999:table 1). Remarks. — This species was the eighth most common mollusc intercepted by the United States Division of Plant ProtectionandQuarantine in import shipments from 1993-1998(Robinson, 1999:table 3). It isoften found confinedto towns,farmlands,or gardens andlives in open places, pastures, arable fields, roadsides,landfills, and urban gardens (Chichester & Getz, 1973;Kerney, 1999);itwasfoundonEasterIslandin anarea showing clearsigns ofpast humanhabitationand some cultivation.Muchoftheleaflit- terwas banana(Musaspec.), whichonly growsin such humidandwind-shelteredareas onEasterIsland(Flenley, 1993: 13). An undeterminedspecies ofDeroceras, close to, orperhaps identicalwith, D. reticula- tum,was foundonPitcairnIslandin 1991and was also suggested to beofrecent intro- duction(Preece, 1995:302). Both this taxon and D. laeve have been reported from Hawaii as introducedtaxa (Cowie, 1997:22). Typically, ofthesetwo slugs, D.reticulatumistheone foundin invasive situations (Robinson, 1999: table 1), but the reverse appears to be true in Hawaii. However, Cowie(1997: 22) notedthatthe two species names mayhavebeen usedinter- Boyko &Cordeiro: TerrestrialMolluscaofEasterIsland 21 changeably whenreferring to Hawaiianspecimens, making identificationsinthe litera- ture questionable. Thepresent specimens from Easter Island are clearly D.reticulatum because they possess awhite-borderedbreathing pore onthe mantle more anteriorly locatedthan thatofD.laeve. Dissectionofone specimen from EasterIsland confirmed thisidentification,basedonthemorphology ofthemale genitalia, especially the charac- teristic shape andthickness ofthe distal portion ofthe penis (see Kerney &Cameron, 1979). Because the commonly introducedDeroceras foundworldwideis D. reticulatum, andbecauseourspecimens arealso thisspecies, westronglysuspect thatthesolerecord of D. laevefrom Easter Island (Naranjo-Garcia & Appleton, 1999: 120) was based ona misidentification.Attempts toborrow thematerialofNaranjo-Garcia &Appleton (1999) from the Universidad Nacional Autonoma de Mexico for direct examination were unsuccessful, dueto the inaccessibility ofthe collectionwhich was being moved at the timeofourrequest. However, Naranjo-Garcia (pers. commun. 11April2001) informed us thatthe specimens they dissected (number unspecified) "fully agreewith the shape drawn of the female(aphallic) form in page 147 of Kerney & Cameron [1979, for D. laeve].” Intheabsenceofacomparison ofadultmale penis morphology, weare hesitant to agreewiththeir conclusions as totheidentity ofthose specimens. Only furthercol- lecting, anddissectionof moremale genitalia, willanswerthe question ofwhetherthere are two species ofDeroceraspresentonEasterIsland. Family Milacidae Milaxgagates(Draparnaud, 1801) LimaxgagatesDraparnaud;Dall,1908:435. Milaxgagates(Draparnaud); Odhner, 1922: 249;Steele,1957: 113 (list); Naranjo-Garcia&Appleton, 1999:120, 121. Material examined. — Sta. 705, Easter Island (Rapa Nui), 2.vii.l917, coll. K. Backstrom (SMNH 28091/3). Distribution. — This is a cosmopolitan species now, native to Europe and the Mediterraneanbasinthathasbeenintroducedintonumerous countriesonallcontinents exceptAntarctica(Robinson, 1999:table 1).Robinson(1999: table1)listedthisspecies as absentfrom easternAsiaandthe oceanicislands ofthe Pacific,butithasbeenfoundin shipmentsofplants fromChinaand Singapore (Dundee, 1974: 16) andoccurs onEaster Island(Dall, 1908;herein). Remarks. —LikeallterrestrialmolluscsonEasterIsland,except Helixaspersa, thisspe- ciesis apparently notvery common as no specimens were foundin1999. Family Subulinidae Allopeas gracile(Hutton, 1834) (fig. 2) Material examined.—Under leaflitter indepressionofacollapsedlavatubenearacavemouth,Ana Te Pahu,Easter Island(RapaNui),21.viii.1999,coll. C.B.Boyko(AMNH300603/9). Distribution.— This species is probably the most widely ranging of all land snails (Pilsbry, 1946: 177-178). Of presumed tropical American origin, it has been introduced intogreenhouses inthesoutheasternUnited States(Burch, 1962: 127),Mexico (Branson 22 BASTERIA, Vol. 65,No. 1-3, 2001 Fig. 2.Allopeasgracile(Hutton,1834)fromAnaTe Pahu,Easter Island (RapaNui), largest specimen, 1.0cm shelllength(AMNH300603). & McCoy, 1965: 10),andthroughout thetropics ofbothhemispheres (Pilsbry, 1946: 177 178;Robinson, 1999:table1). Itis hereinrecordedfrom EasterIsland forthe firsttime. Remarks.—AllopeasgracilewasfoundonEasterIsland inassociationwithpast human habitationandcultivation, inthe samehabitatasDerocerasreticulatum.Although onlycol- lectedas shellmaterialin 1999,itlikely persists onEasterIsland asthe habitatis favou- rableforthespecies. This species was recorded as a probable recent introduction to Pitcairn Island by Preece (1995: 295). Family Helicidae Helix aspersa Müller, 1774(fig. 3) Helixaspersa Miiller;Rees, 1955:93; Naranjo-Garcia& Appleton,1999: 121 Material examined. — On various ornamental plants, vicinity ofHotelTopara'a, HangaRoa,Easter Island(RapaNui),20.viii.1999,coll. C.B.Boyko(AMNH300601/5). Boyko &Cordeiro: TerrestrialMolluscaofEasterIsland 23 Fig. 3.Helixaspersa (Müller,1774).A,photographedinsitu ongroundsofthe Hotel Topara’a,Easter Island (RapaNui); B,specimenscollected from same site; largest specimen,3.1 cmshelllength(AMNH300601). Distribution.—Nowacosmopolitan species, native toEurope andthe Mediterranean basin, it has been introduced into numerous countries on all continents except Antarctica, including many Pacificislands (Robinson, 1999:table 1). Remarks. —The occurrenceof this species onEaster Island is not surprising, given that it is perhaps the most widely and most often distributedintroduced pulmonate. Indeed, H. aspersa was the most common mollusc intercepted by the United States Division of Plant Protectionand Quarantine in imports during 1993-1998(Robinson, 1999:table 3). Itis typically foundin gardens, grasslands, woods and dunes(Kerney, 1999:205). On Easter Island, it was found only in the vicinity ofthe solesettlementof Hanga Roa. Although only afewspecimens were collected, itwas observed tobe very abundanton ornamental plants around houses. As with introductionsofthis species elsewhere, itmayhavebeen deliberatelybrought to EasterIslandforuseasfood. Rees (1955: 93) was thefirstauthorto citeH.aspersaas occurringonEasterIsland,but gave noreference for thisinformationand wehave beenunableto track the source of hisrecord. Itispossible thatpart ofRees's (1955: 93) statementaboutdistributionwasa generalized remarkonsnailbiogeography, rather thanalist oflocalitiesfromwhichH. aspersawas known. Support forthis hypothesis comes from thefact thatalthough Rees (1955) listed Pitcairn Islandamong those islands reached by "the common snail," H. aspersa is nototherwiserecordedfromthatisland (Preece, 1995). Likewise, Rees(1955) mentionedHawaiiin the questionable paragraph, butH.aspersa was not well documen- ted from there until a year later (Kondo, 1956: 141); its brief earlier mention in a Hawaiianentomologicaljournal [Anonymus, 1953]no doubtescaped the noticeofmala- cologists. Naranjo-Garcia & Appleton (1999: 121) provided the only other recordofthis species from EasterIsland.ThepresentrecordofH.aspersais thereforeonly the second well-documentedonefrom EasterIsland. 24 BASTERIA, Vol. 65,No. 1-3,2001 ACKNOWLEDGEMENTS Funding toCBB for the InvertebrateSurvey ofEaster Islandwas provided by the U. S. National Park Service (USNPS), Gateway National Recreation Area, Division of NaturalResources,as partofaScienceMuseumofLong Island/Explorers Clubfive-year research expedition to explore the impacts ofEl Ninoevents onWorldHeritage Sites. John Tanacredi(USNPS) is thankedfor making this Survey possible. PaulaMikkelsen (AMNH) and an anonymous individualkindlyreviewed the manuscript andsuggested valuableimprovements. All photographs bySteve Thurston(AMNH) excepting fig. 3B, whichis by CBB. REFERENCES ANONYMUS, 1953.HelixaspersaMuller. —Proceedings ofthe Hawaiian EntomologicalSociety 15:13. BAUMAN, S., 1996. Diversity and decline of land snails on Rota, Mariana Islands. — American MalacologicalBulletin 12: 13-27. BOUCHARD-CHANTEREAUX, M.,1838. Cataloguedesmollusquesterrestres etfluviatiles observesjus- qu'acejoural'etatvivant dans le departmentdu Pas-de-Calais LeRoy-Mabile, Boulogne:1-94. BRANSON, B.A.,& C.J. MCCOY, JR., 1965. Gastropodaofthe 1962 University ofColorado Museum ExpeditioninMexico. —University ofColorado StudiesSeries inBiology 13: 1-16. BURCH,J.B., 1962.Howtoknowthe easternland snails: 1-218.Dubuque,Iowa. CAMPOS,L.,&L.E.PENA, 1973. Losinsectos deIsla dePasqua.—RevueChilena deEntomologia7:217- 229. CHICHESTER, L.F.,&L.L. GETZ, 1973. Theterrestrial slugsofnortheastern NorthAmerica.—Sterkiana 51: 11-42. COOKE, C.M.,Jr.,ScY.KONDO, 1961[I960].Revision ofTornatellinidae andAchatinellidae (Gastropoda, Pulmonata).—Bernice P BishopMuseumBulletin 221: 1-303. COWIE, R.H., 1997. Catalog and bibliographyofthe nonindigenousnonmarine snails and slugs ofthe Hawaiian islands.—BishopMuseum Occasional Papers50: 1-66. —, N.L.EVENHUIS,&C.C. CHRISTENSEN, 1995.Catalogofthe nativeland and freshwatermolluscs oftheHawaiian islands: 1-248. Leiden. DALL,W.H., 1908. Reports onthe dredgingoperations offthe west coast ofCentral America to the Galapagos,tothe west coastofMexico,and intheGulfofCalifornia,inchargeofAlexander Agassiz, carried onbythe U.S. Fish Commission steamer"Albatross,"during1891,Lieut. Commander Z.L. Tanner,U. S. N., commanding.XXXVII.Reports onthe scientific results ofthe expeditiontothe eastern tropical Pacific, in chargeofAlexander Agassiz, by the U. S. Fish Commission steamer "Albatross," from October, 1904, to March, 1905, Lieut. Commander L. M. Garrett,U. S. N. com- manding. XIV. The Mollusca and the Brachiopoda. — Bulletin ofthe Museum ofComparative Zoology43:205-487, 19pis. DUNDEE, D.S., 1974.Catalogofthe introduced mollusks ofeasternNorthAmerica (northofMexico). — Sterkiana55: 1-37. FISCHER, S.R.,&C.M.LOVE, 1993.Rapanui: thegeologicalparameters.In:S.R.FISCHER, ed., Easter island studies. Contributions to the history ofRapanui in memory ofWilliam T. Mulloy. Oxbow Monograph32: 1-6. FLENLEY,J.R., 1993. Thepresent flora ofEaster Island and itsorigins. In: S.R. FISCHER, ed., Easter island studies. Contributions to the history ofRapanuiin memory ofWilliamT. Mulloy. Oxbow Monograph32:7-15. Boyko &Cordeiro: TerrestrialMolluscaofEasterIsland 25 FUENTES,F., 1914.Contribucion al estudiode lafaunadela Isla dePascua. —Boletin delMuseo Nacional deChile 7:285-319. KERNEY,M.P,1999.Atlas ofthelandand freshwatermolluscs ofBritainandIreland: 1-264.Essex, United Kingdom. , &R.A.D.CAMERON, 1979.Afield guidetotheland snails ofBritain and north-westEurope: 1-288. London,England. KLEMMER,K., 8cG.ZIZKA, 1993.The terrestrial fauna ofEaster Island. In: S.R.FISCHER, ed, Easter island studies. Contributions to the history ofRapanui in memory ofWilliam T. Mulloy. Oxbow Monograph32:24-26. KONDO, Y., 1956.First HelixaspersainHawaii.—TheNautilus 69: 141-142. KUSCHEL, G., 1963. Compositionand relationshipofthe terrestrial faunas ofEaster,Juan Fernandez, Desventuradas,andGalapagosIslands. —Occasional PapersoftheCaliforniaAcademyofSciences44: 79-95. NARANJO-GARCfA, E.,8c C.C.APPLETON, 1999.Dos nuevosregistros demoluscos terrestresintroduci- dos enla IsladePascua,Chile. —Biociencias 6: 119-124. ODHNER,N.H., 1922. Mollusca fromJuanFernandez andEaster Island. —The Natural History ofJuan Fernandez and Easter Island.III.Zoology 11(22):219-254. OSORIO, C., 1991. Charonia tritonis(Linne1758)enIsladePascua (Mollusca:Gastropoda:Cymatiidae). — RevistadeBiologiaMarina,Valparaiso26:75-80. OSORIO, C., 1995.Dos nuevosregistros deIsognomiidae(MolluscaBivalvia)para IsladePascua,Chile. — RevistadeBiologiaMarina,Valparaiso30: 199-205. PILSBRY, H.A., 1946. Land mollusca of North America (north of Mexico). Vol. II. Part I: 1-520. Philadelphia,PA. 1948.Land mollusca ofNorthAmerica (northofMexico).Vol. II.Part II:521-1113. Philadelphia,PA. , PREECE,R.C., 1995.Systematicreview ofthe landsnailsofthe Pitcairn Islands.—BiologicalJournalofthe Linnean Society56: 273-307. REES,W.J., 1955.Escapedescargots. —TheNautilus 68:90-94. REHDER, H.A., 1980. The marine mollusks ofEaster Island (Isla de Pascua) and Sala y Gdmez. — Smithsonian Contributions toZoology289: 1-167. ROBINSON, D.G., 1999.Alieninvasions:the effectsofthe globaleconomyonnon-marinegastropodintro- ductionsintotheUnited States. —Malacologia41:413-438. STEELE,P.H., 1957. Easter Island shells.—The Nautilus 70: 111-113. TURGEON, D.D.,J.F.QUINN,JR.,A.E. BOGAN, E.V. COAN, F.G. HOCHBERG, W.G.LYONS, RM. MIKKELSEN,R.J. NEVES, C.F.E. ROPER, G.ROSENBERG, B. ROTH, A. SCHELTEMA, F.G. THOMPSON,M.VECCHIONE, 8cJ.D.WILLIAMS,1998.Common andscientificnamesofaquatic invertebratesfrom the United States andCanada: mollusks,2nd edition. American Fisheries Society, SpecialPublication 26: 1-526. WALDfiN, H.W., 1962. On the variation, nomenclature,distributionandtaxonomical position ofLimax (Lehmannia)valentianus Férussac (Gastropoda,Pulmonata).—ArkivförZoologi,2nd series 15:71-96.