B75-Tattersfield_Rowson:Basteria-2010 26/06/2011 14:51 Page 39 Tanzartemon gen. nov., a new land snail genus (Gastropoda, Pulmonata, Streptaxidae) from Morogoro District, Tanzania Peter Tattersfield DepartmentofBiodiversityandSystematicBiology,NationalMuseumofWales,CathaysPark,CardiffCF103NP,UK;[email protected] [correspondingauthor] & Ben Rowson DepartmentofBiodiversityandSystematicBiology,NationalMuseumofWales,CathaysPark,CardiffCF103NP,UK TanzartemonisdescribedasanewgenusintheStreptaxidae.Twospecies,T.seddonaeandT.mkungwensis,aredescribedfrom sub-montaneforesthabitatintheUluguruMountainsandfromtheoutlyingpeakofMountMkungwe,MorogoroDistrict, Tanzania.Thegenusischaracterisedbyitsgloboseordepressedgloboseshellmorphology,thepresenceintheshellapertureof 39 oneormoreparietaldenticlesandofseveralstrong,peglikedenticlesthatsitimmediatelyinsidetheperistomeonthebasal, palatalorcolumellarmargins.Thegenitaliaarecharacterisedbythepresenceofapenialsheaththatenclosesthepenis,butdoes notincorporatealoopofthevasdeferens. ThespecimensdonotresembleanyknowngenusintheEastAfricanfaunabecauseofthecombinationoftheirgloboseshell andstrongaperturaldentition.PossiblerelationshipswithothermorphologicallysimilarstreptaxidsknownfromtropicalWest Africa,particularlyLamelligerAncey,1884,arediscussed,but,basedonshellmorphologyandgenitalanatomyweconcludethat TanzartemonislikelytobeanEastAfricanendemicgenusthatisnotcloselyrelatedtoWestAfricanforms.Weconsideritlikely thatTanzartemonhaseitherevolvedinsitufromanothermorphologicallydissimilarAfricanlineageorfromlineagesfound otherwiseonlyinAsiaorSouthAmerica,and,assuch,itisofbiogeographicalinterest. Keywords:Gastropoda,Streptaxidae,Tanzartemongen.nov.,taxonomy,EasternArcmountains,UluguruMountains,MountMkungwe,forest,Tanzania. Introduction variousothergeneraorsubgenera.Richardson(1988)listed atotalof102speciesinthegeneramostrelevanttothis Small,axially-deviatedstreptaxidswithaperturaldenti- study,i.e.HaploptychiusMöllendorff,1905,Indoartemon tionoccurintropicalWestAfrica,SouthAmerica,andAsia; Forcart,1946,OdontartemonPfeiffer,1856(ofwhichLamel- theyarenotknowninMadagascar.AccordingtoGude liger,Ancey,1884,wasconsideredasynonym),Oophana (1902),thereisnofossilrecordforthegroup,whichhave Ancey,1884,PerrottetiaKobelt,1905,andStreptartemon variouslybeenassignedtoStreptaxisJ.Gray,1837,orto Kobelt,1905.Disagreementoverthetypespeciesofeach Basteria75(1-3):39-50 B75-Tattersfield_Rowson:Basteria-2010 26/06/2011 14:51 Page 40 taxon(alternativesbeingfromdifferentcontinentsinsome height(mm);H,shellheight(mm).Forcollections:BMNH, cases),widevariationinmorphology,andalackof NaturalHistoryMuseum,London,UK;MRAC,Musée knowledgeofevolutionaryrelationshipsamongStreptaxi- Royaldel’AfriqueCentral,Tervuren,BelgiumNMT, daehaveresultedinrapidlychangingclassificationsofthis NationalMuseumofTanzania,DaresSalaam,Tanzania; group(e.g.Pfeiffer,1856;Ancey,1884;Tryon,1885;Thiele, NMW,NationalMuseumofWales,Cardiff,UK;. 1935[1992translation];Bequaert&Clench,1936;Van Thenumberofshellwhorlsarecountedusingthe BenthemJutting,1954;Zilch,1960,Richardson,1988; methoddescribedinKerney&Cameron(1979) Schileyko,2000).Worksprovidingrelevantanatomicaldata arefew.Eachgenus’relationshiptoStreptaxis(whosetype Systematic descriptions speciesistheBrazilianHelixcontusaFérussac,1820)isopen toquestion.Streptaxiscontususislarge,onlymoderately FamilyStreptaxidae deviatedandedentate,whiletheother(sub)generaare foundedonsmaller,dentatespeciesthatrangefrom Tanzartemongen.nov. moderatelytoverystronglydeviated. TheonlyAfricantaxonamongtheseisLamelliger.There Tanzartemonischaracterisedbyitsgloboseordepressed issomedisagreementoverthecircumscriptionofthisgenus globoseshellandthepresenceofstrongaperturaldentition anditsrelationshipwithOdontartemonPfeiffer(nonKobelt, includingatleastoneparietallamellaandseveralstrong 1880).WechoosetoadopttheviewofBequaert&Clench denticlesonthepalatal,columellarorbasalmargins.The (1936),Degner(1934),Thiele(1935)andAdam(1962b)in shellmaybeslightlyaxiallydistortedorundistorted.The whichLamelligerisconsideredavalidgenus,ratherthan genitaliaarecharacterisedbythepresenceofapenialsheath thatofSchileyko(2000),Baker(1928)andZilch(1961)who thatenclosesthepenis,butdoesnotincorporatealoopof 40 consideredLamelligeranobjectivejuniorsynonymof thevasdeferens. OdontartemonPfeiffer.Thisdisputeisnomenclaturalrather Etymology:fromthecountryTanzaniaandtheGreek thansystematic,so,thusrecognised,Lamelligercontainsthe masculinenoun‘artemon’,aprincipalpulleyorforesail, samefourspecies:L.troberti(Petit,1841)(typespecies), usedasacomponentofseveralotherstreptaxidnames. L.distorta(Jonas,inPhilippi,1843),L.maassiDegner,1934, Gender:masculine. and,provisionally,anunnamedspeciesdescribedin Typespecies:T.seddonaespec.nov. Schileyko(2000)asOdontartemon“sp.B”.AlthoughAdam Otherspecies:T.mkungwensisspec.nov. (1962b)concludedthatL.distortawasnotaLamelliger,hedid notassignittoanothergenus,soherewediscussitalong Tanzartemonseddonaespec.nov.(Figs1-4,6,8,11-15) withtheotherthreetaxa.AllfourarestrictlyWestAfrican, occurringinAngola,Guinea,Guinea-Bissau,Liberiaand Typematerial:Holotypeplus2paratypescollectedbyP.Tattersfield, SierraLeone(Bequaert&Clench,1936),withL.distortaalso Z.Habibu,C.MichaelandlocalTanzanianassistantsfromTegetero occurringinGhanaandIvoryCoast(Adam,1962b).The villageinJanuary1996bydirectsearching(Table1).Theholotype remainingaxially-deviatedstreptaxidsofAfricaare (NMW.Z.1996.148.00027)isadeadshell.Paratypes1 assignedtothegenusGonaxisTaylor,1877,andtovarious (NMW.Z.1996.148.00028)and2(NMT)werecollectedalive;paratype2 othergeneraoftenconsideredsubgeneraofit.Allareeither isjuvenile. edentateorhaveasingleparietaltooth(Bequaert&Clench, 1936). Typelocality:UluguruNorthForestReserve,UluguruMountains, Thefollowingabbreviationsareused,forshells:d, MorogoroDistrict,MorogoroRegion,Tanzania.Location:6.94°S aperturediameter(mm);Dmax,shelldiameter-majoraxis 37.69°E.Allspecimenswerecollectedfromleaflitterintall,largely (mm);Dmin,shelldiameter-minoraxis(mm);h,aperture intactsub-montaneforestlyingonsteepandstronglyundulating Basteria75(1-3) B75-Tattersfield_Rowson:Basteria-2010 26/06/2011 14:51 Page 41 topography,ontheeastsideoftheUluguruMountains,aboveTegetero betweenthem.Theribbinggraduallybecomesweakerafter Mission,atanaltitudeofabout1500mabovesealevel.Annualrainfall whorl5¾andthebodywhorl(whorl6ofparatype)is intheareaisestimatedas2900-4000mm/yearandthereisnomarked smoothandglossy. dryseason.Furtherdetailsofthecollectinglocalityareprovidedin Theperistomeisdiscontinuousacrosstheparietalregion Tattersfield(1999)andinLovett&Pócs(1993). althoughtheparietalareaisslightlythickenedandmarked byaridgeintheshellthatdevelopsintoanelongated Shell.−Adultshelldepressedglobular(Figs1-2)with5¾ parietallamellafurtherintotheaperture(seebelow).The whorls(holotype).Slightlydistortedalongcolumellaraxis peristomeisstronglyreflectedtoformabroadlipthat (byabout10°)andapparentlyasymmetricallycompressed extendsaroundallsectionsoftheapertureapartfromnear betweenupperandbasalsurfaces.Thisasymmetryresults thejunctionoftheparietalandpalatalareaswheretheperis- inanovaloutlinewhenviewedfromabove(Figs1-2),with tomeissharpandnotreflected.Theumbilicusissealed, theapertureprojectingfurtherfromthecolumellaraxisthan remainingonlyasashallowdepression.Theapertureis theoppositesideoftheshell.Thebodywhorlandpenulti- crescent-shapedwithsub-equalpalatalandbasalsections matewhorlsarebothwidestneartheapertureandthe andashortercolumellarsection.Thefour-foldapertural majoraxisoftheshellextendsthroughtheaperturalregion dentitionintheadultspecimensconsistsofthreedenticles (Figs1-2).Whenviewedfrombelow,theprecedingwhorls thatsitimmediatelyinsidetheperistomeplusadeeper, areentirelyobscuredbythebodywhorl,withnooverlap single,spiral,elongatedparietallamella.Therearetwo evident. large,rounded,peg-likedenticlesonthebasalsectionand Thebaseoftheshellissmoothandglossywithveryfaint oneonthepalatalmargin,whichallsitjustinsidelip;these andratherirregularspiralstriaethatarevisibleunderhigh denticlesareabout0.3-0.5mmlong.Theinnerendofthe magnificationonly.Theuppersurfaceoftheshellisalso parietallamellastartsapproximatelyoppositethepalatal glossyandwhorls1,2and3aresmooth;extremelyfaint denticleandextendsforalengthofabout1.7mm,withthe 41 spiralscratchesremainonveryfreshshells(paratypes1and outerendfinishingjustoutsidetheplaneoftheaperture. 2).Whorls4-5¾haveregularlyspacedradialribsthatare Thereisafurthershort,linearswellingontheparietal strongestnearthesutureandfadeonthelowerpartsofthe surfacedeeperwithintheaperture.Theonlyknownjuvenile whorl.Theribsonwhorl4arelocallybeadedatintervals (Fig.3)isedentateandwithoutthereflectedlip,andhasan approximatelyequaltothegapsbetweentheribs;these open,symmetricalumbilicus. combinetointroduceaspiralelementintothesculpture. Shellmeasurements:SeeTable1. Thewidthsoftheribsareapproximatelyequaltothegap Bodycolour.−Live-preservedadultspecimenmainly palecream,withapalecream-yellowdigestivegland,anda veryslightorangetingetotheoptictentacleretractor Specimen Accession H Dmax Dmin h d Whorls No. muscles.Thelive-preservedjuvenilespecimenissimilarly Holotype NMW.Z.1996. 5.4 8.6 6.8 2.7 4.6 5¾ coloured,butthedigestiveglandislightbrown. 148.00027 Genitalia(Figs9-14).−Penisnotablylong(9.0mmina Paratype1 NMW.Z.1996. 5.7 8.7 7.1 2.9 4.8 6 withdrawnstateor0.7whorls),tubular,withoutappen- 148.00028 dices.Thepenisissubstantiallyswollenbasallyandslightly Paratype2 NMT 3.4 6.2 5.7 2.7 3.0 4¾ swollenapically,entirelyenclosedinathinsheath.The (juvenile) sheathiscontiguouswiththepenialretractorapicallyand penialwallbasally,beingcrumpledorcreasedaroundthe Table1.Accessionnumbers,shelldimensionsandnumberofwhorlsof apicalfifthofthepenis,andwithasecondsmallretractor Tanzartemonseddonaegen.&spec.nov. musclearisingneartheatrium,attachedtothebodywall. Vasdeferensflattened,free,andrelativelystraight,arising Tattersfield, P. & Rowson,B.-Tanzartemongen.nov.(Streptaxidae)fromTanzania B75-Tattersfield_Rowson:Basteria-2010 26/06/2011 14:51 Page 42 42 Figs1-4.Tanzartemonseddonaegen.&spec.nov.1,holotype(NMW.Z.1996.148.00027);2,Paratype1(NMW.Z.1996.148.00028);3,Paratype2,juvenile (NMT);4,do,apexandsculpture. Basteria75(1-3) B75-Tattersfield_Rowson:Basteria-2010 26/06/2011 14:51 Page 43 ratherbasallyfromtheprostate,taperingtowardsthepenis smooth,white,withsmallindistinctvesiclesvisible andenteringthepenialsheathsubapically.Thevasdeferens throughout,andwithafewscatteredthinanchoringmuscle isapparentlycontiguouswiththeapicalpartofpenisand strands.Anteriorductexitingglandsubapicallyandbend- notclearlydemarcated.Thepenialretractorisshort, ingoveranteriorpartofgland;posteriorglandexitingjust attachedtothecolumellarmuscle,andtheenteringpenial posteriortomidpointofgland.Bothductsroundednearest sheathtoattachtoanapicalbendofthevasdeferens.The thegland,becomingflattenedandstretchednearerthebuc- wallsofthepenisarethickerthanthoseofthesheath. calmass,butevenlythickthroughouttheirlength. Interiorsurfaceofpenishasthreedistincttypesofsurface Radula(Fig.8).−Buccalmasslong(11.0mm),slender gradingintooneanotherasfollows.Apicalpartofpenis(or (1.25mm).Radulaofapproximately65steeplyv-shaped vasdeferens):indistinct,reticulateelasticsculptureof rows(angleupto45ocentrally),with23-28teethineach incompletetransverseseptaeandlowscattered,scale-like half-row.Noevidenceofacentraltooth.Lateralteethslen- sub-rectangularpads.Middlepartofpenis:low,indistinct, der,aculeate,flattened,decreasinginsizegraduallylaterally longitudinalpilastersoccasionallyanastomosing.Basalpart withoutasharptransitiontomarginals.Innerlateralswith ofpenis:regular,shallowrhombicdepressions,each longbasalplates,theendofwhichgivestheimpressionofa containingalowcentrallongitudinalpadbearingasingle slightswellingabouttwo-thirdsofthewayalongthelength chitinoushook;sculpturebecominglessregularandgrading ofthetoothwhenteetharedisplacedortransparent. intoindistinct,irregularpilasterstowardsatrium.Hooks Etymology.−thespeciesisnamedinhonourofDrMary small(0.04mmlong),orange-brown,simpleinform,little B.Seddon,inrecognitionofhercontributiontothedevelop- recurved,withsub-circularbases.Totalnumberofhooksin mentofmalacologicalresearchcapacityinTanzania. penis:about82(counted).Albumenglandshort,wedge- shapedandwithauniformstructureofverysmall,indis- Tanzartemonmkungwensisspec.nov.(Figs5,7) tinctvesiclesoracini.Hermaphrodicticductdiverticulum 43 (talon)enlarged,butcompactandconvoluted,nothidden Typematerial:Holotype(NMW.Z.2003.001.00003)plustwoparatypes withinalbumengland.Bursacopulatrix(=gametolyticsacor (NMW.Z.2003.001.00004;NMT)collectedbyB.RowsonandC.Ngereza spermatheca)small,oval,apparentlyempty,andattending bydirectsearching(Table2).Allmaterialwascollectedasdeadshells. albumengland.Bursacopulatrixductveryslenderapically, butsubstantiallywiderbasallyandexitingvaginaonlyjust Typelocality:MkungweCatchmentForestReserve,MorogoroDistrict, belowthepointatwhichthevasdeferensleavesthe Tanzania.MkungweisanisolatedoutlieroftheUluguruMountains, prostate.Oviductalglandflattened,broad,withaciniof whichliesabout20kmtotheeastofthemainUlugurumassif.Location: oviductalglandperhapslargebutalmostindiscernible,the 6.90°E37.91°S.Allspecimenswerecollectedfromsub-montaneforestat surfaceappearinguniform.Noeggsorembryosinoviduct. analtitudeofabout900mabovesealevel.Lovett&Pócs(1993)provide Vaginabroadandmuscular,withratherthickwallsand furtherdetailsabouttheforesthabitatinMkungweCatchmentForest withashortroundedpouchbasally.Interiorsurfaceof andestimaterainfallonthewettereasternandupperslopesoftheRe- vaginaconsistingofelastic,anastomosinglongitudinal serveas1700-2900mm/yearwithmisteffectonthesummitandwithout pilasters,withasinglelobe-likelongitudinalpilasterpartly amarkeddryseason. occludinglumennearexitofbursacopulatrixduct.Interior surfaceofbasalpouchreticulate,consistingofshallowsub- Shell.−Shellgloboseandsymmetricalwithwhorlsexpand- rhombicdepressions. inguniformlywithoutaxialdistortion;whorls7(holotype). Salivarygland(Fig.15).−Single,5.1mmlong,occupying Allspecimensarewornandretainlittleperiostracumbutthe abouthalfawhorlinlength,taperingposteriorly,concave smallfragmentsthatremainareglossyandretainevidenceof andoverlyingtheoesophagusonthecolumellarside,the finespiralmicroscultureonthebaseoftheshell;acoarser,ra- anteriorpartcurvedslightlytowardsthecolumella.Surface dial,ribbedsculptureispresentontheuppersurface. Tattersfield, P. & Rowson,B.-Tanzartemongen.nov.(Streptaxidae)fromTanzania B75-Tattersfield_Rowson:Basteria-2010 26/06/2011 14:51 Page 44 Aperturecrescentshapedwithsub-equalcolumellar,basal andpalatalsectionsthatareratherpoorlydefinedbecause ofthecurvatureoftheperistome.Theshellsarecalcified andtheperistomeappearstobecontinuousacrossthepari- etalzone,whichisthickenedandridged.Theperistomeis stronglyreflectedtoformalipthatextendsaroundall sectionsoftheapertureapartfromacrosstheparietalzone andaroundthejunctionoftheparietalandpalatalareas whereitissharpandnotreflected.Theumbilicusissealed, remainingonlyasashallowdepression. Thesix-foldaperturaldentitionconsistsoffourdenticles thatsitimmediatelyinsidetheperistomealongthebasal andcolumellarmargins,andtwoelongatedparietal lamellae.Thefourbasal/columellarprocessescomprise threestrong(c.0.4-0.5mmlong)denticles–oneeachatthe palatal/basalandcolumellar/basaljunctions,andthethirdin thecentreofthebasalzone.Thefourthdenticleislocatedon thecolumellaandismuchweaker.Theoutermostparietal lamellaextendsbeyondtheplaneoftheperistomeanditsits oppositethedenticleatthebasal/palataljunction;itisabout 2.8mminlengthandextendsaroundthespiralintothe 44 aperture.Thedeeperparietallamellaisshorter(about 0.8mmlong)andpegshaped,anditsitsatrightanglesto thelongerlamella. Shellmeasurements:SeeTable2. Differentiation.−Thestrongaperturaldentitionand generallygloboseshellshapeindicateastrongaffinitywith T.seddonae.However,T.mkungwensismaybedistinguished byitssymmetrical,uniformlyexpandingspireandabsence ofsignificantaxialdistortion,thepresenceofasecondmore deeplysetparietaldenticleandtheoccurrenceofathird, weakdenticleonthecolumellarmarginoftheperistome. Anatomy.−Notknown. Etymology.−fromthetypelocality,MountMkungwe. Fig.5.Tanzartemonmkungwensisgen.&spec.nov.,holotype (NMW.Z.2003.001.00003). Basteria75(1-3) B75-Tattersfield_Rowson:Basteria-2010 26/06/2011 14:51 Page 45 Comparisons Introduction).Lamelligerischaracterisedbyastrongcontin- uousparietallamella,whichrunstovaryingdegreesonthe Shellmorphology.−NeitherspeciesofTanzartemonisidenti- parietalsurfaceintotheshell.InL.maassi(paratypein calwithanyAsianorSouthAmericanspeciesorgenus MRAC97344-97346,fromTammasadou,GuineeFrancaise) knowntous,sowerestrictourshellcomparisonstoAfrican thelamellaextendsforaboutonequarterofawhorlfrom taxa.AnatomicalinformationonStreptaxidaeismore theaperture.Thiscontinuousparietallamellamaybeho- fragmentaryandhomoplasyinshellshapecommon,sowe mologouswiththeshortlamellaandslightareaofdeeper makeanatomicalcomparisonswithbothAfricanandnon- thickeningpresentinT.seddonae,andthedoubleparietal Africantaxa. dentitioninT.mkungwensis.However,incontrastto ThedistortedshellofT.seddonaesuggestsanaffinity T.seddonae,theaperturesofjuvenileL.maassi(MRAC)are withthestreptaxidgenusGonaxisTaylor,1877,whichis furnishedwithaparietallamella. widespreadintheEastAfricanfauna.However,allthe TherearealsosimilaritieswithLamelligerinrespectof speciesattributedbyVerdcourt(2006)toGonaxisandits thebasalandpalataldenticles.Lamelligertroberti(MRAC subgeneraintheEastAfricanfaunaareedentateorhaveat 97349-97350det.DegnerfromBolahun,Liberia)hastwo mostasingleweakparietaltooth,sothepresenceofthe denticles,whicharealmostfusedinsomespecimens,atthe strongaperturaldentitioninTanzartemonservestodifferen- baseoftheaperturalliponthecolumellaoronthebasal tiateitfromGonaxis.Likewise,noneoftheotherspeciescur- sectionoftheaperture(theapertureisroundedsothese rentlyassignedtoGonaxisfromelsewhereinAfrica(e.g.by sectionsarenotwelldifferentiated).InL.maassi(specimens Bequaert&Clench,1936;Adam,1962a,b)havedentition asabove)therearetwosimilardenticleswhichlieonthe beyondasingleparietaltooth.AllAfricanstreptaxidswith columellarsection,andthereisalsoafurther(sometimes undistorted,globoseordepressedgloboseshells(e.g. elongated)denticlelyingdeeperwithintheapertureonthe TayloriaBourguignat,1889)areedentate,withtheexception palatalsurface;thisdenticleisassociatedwithadepression 45 oftheEastAfricanJuventigulellaTattersfield,1998.However, atthesamepositionontheoutsideoftheshell.Sometimes shellsofJuventigulellaareotherwisequiteunlikethatof thereisafurthersmalldenticleatthesameposition,slightly Tanzartemon,beingminute,stronglysculpturedandwith closertotheaperture.TheumbilicusofL.maassiisclosed, differentdentition(Tattersfield,1998). butitisachinkinL.trobertiandopeninthethirdspecies Inbeingbothstronglydentateanddistorted,Tanzartemon L.distortus(materialinBMNH).Fromthefiguresand bearsaresemblanceonlytotheWestAfricanLamelliger(in- descriptiongivenbySchileyko(2000),Odontartemon“sp.B” cludingL.distortaandOdontartemon“sp.B”ofSchileyko;see closelyresemblesL.maassi. Theshellsofalltheabovespeciesaremoredistorted alongthecolumellaraxisthanTanzartemon,totheextentthat Specimen Accession H Dmax Dmin h d Whorls No. theprecedingwhorlprojectsbeyondthebodywhorlwhen Holotype NMW.Z.2003. 6.0 7.8 7.1 2.4 4.0 7 viewedfrombelow.InTanzartemon,theprecedingwhorlis 001.00003 entirelyobscuredbythebodywhorl.Intermsofshell Paratype1 NMW.Z.2003. 6.1 8.4 7.5 2.3 4.1 7 surfacesculpture,theshellbaseandapicalwhorlsofboth 001.00004 maassiandtrobertiaresmoothbuttheotherwhorlsare Paratype2 NMT 6.0 8.3 7.3 2.3 4.3 7 ribbed(weaklyintroberti). Genitalanatomy.−Dataongenitalanatomywerecon- sultedforthefollowingspeciesandsources(generafollow Table2.Accessionnumbers,shelldimensionsandnumberofwhorlsof Richardson,1988,exceptwherenoted):Discartemonstenosto- Tanzartemonmkungwensisgen.&spec.nov. musvanBenthemJutting,1954(Malaysia;Berry,1965);Hap- loptychiussinensis(Gould,1856)(Vietnam;Schileyko,2000); Tattersfield, P. & Rowson,B.-Tanzartemongen.nov.(Streptaxidae)fromTanzania B75-Tattersfield_Rowson:Basteria-2010 26/06/2011 14:51 Page 46 Figs6-8.Tanzartemongen.nov.6,T.seddonaespec.nov.,paratype1(NMW.Z.1996.148.00028),aperture;7,T.mkungwensisspec.nov.,holotype(NMW.Z. 2003.001.00003),aperture;8,T.seddonaespec.nov.,paratype1(NMW.Z.1996.148.00028),half-rowsfrommedianpartofradula. Hypselartemoncontusulus(Férussac,1827)(Brazil;Barbosaet recommendingthesplittingofgenera(Schileyko,2000; al.,2002);Lamelligermaassi(Degner,1934)(Guinea;Degner, Rowson,2010);thisisevidentlyanimportantdifference.Ina 1934);Lamelligertroberti(Petit,1841)(Guinea/Liberia; largenumberofAfricanandothergenera,thepenialsheath Degner,1934);Odontartemon(i.e.,Lamelliger)“sp.B”(Sierra enclosesaloopofthevasdeferens(seeSchileyko,2000)but 46 Leone;Schileyko,2000);OophanadiaphanopepleaVan thisisnotthecaseinT.seddonae.Secondly,penialhooksin BenthemJutting,1954(Malaysia;Berry,1963);Oophana T.seddonaeoccurstrictlyinthebasalpartofthepenis,unlike obtusa(Stoliczka,1871)(Burma/Myanmar;Blanford& thethreeLamelligerspecieswherehooksarescattered Godwin-Austen,1908);Perrottetiagudei(Fulton,1915) throughout.Thirdly,thepenialretractorjoinsthecolumellar (Vietnam;Schileyko,2000[treatedinStreptaxisbyRichard- muscleinT.seddonae,asitdoesinthemajorityofstrep- son,1988]);Streptartemonglaber(Pfeiffer,1849)(Guyana; taxids.InthisitrecallsL.trobertiandL.maassi,butnotL.sp. Schileyko,2000);andStreptartemonglabernormalis B.,whereitissaidtojointhebodywall(Schileyko,2000). (Jousseaume,1889)(Venezuela;Baker,1925;1926).Dataon Theremaininganatomicallyknowntaxacitedabovehavea other,moredissimilar,streptaxidgeneraarealsogivenby penialsheathcoveringsomeorallofthepenis(theexcep- Degner,1934andSchileyko,2000inparticular.Inaddition, oneofus(BR)hasdissectedmanyadditionalStreptaxidae fromAfrica,Asia,SouthAmericaandtheIndianOcean Figs9-15.Tanzartemonseddonaegen.&spec.nov.,paratype1 (Rowson,2010). (NMW.Z.1996.148.00028).9,genitalia,dorsalview;10,penisandvagina, Therearethreemaindifferencesbetweenthegenitaliaof ventralview;11,insideofvagina;12,insideofpenisandsheath;13, Tanzartemonandthethreeanatomicallyknownspeciesof hooksandrhombicpadsfromatrialpartofpenis;14,hooksfromatrial Lamelliger.Firstly,T.seddonaehasalongpenialsheath,unlike partofpenis;15,salivarygland.Scalebars1mm(9-13,15);0.1mm(14). thethreeanatomicallyknownspeciesofLamelliger,inwhich Abbreviations:ag,albumengland;bc,bursacopulatrix;bd,bursaduct; asheathisabsent(Degner,1934;Schileyko,2000).A fo,freeoviduct;hd,hermaphroditicglanddiverticulum;og,oviductal variously-developedpenialsheathoccursinalltheother gland;pn,penis;pr,penialretractor;ps,penialsheath;sd,salivary streptaxidscitedabove,aswellasmanyothers,thesheath glandduct;vd,vasdeferens;vg,vagina. havingbeenusedasacharacterindefiningsubfamiliesand Basteria75(1-3) B75-Tattersfield_Rowson:Basteria-2010 26/06/2011 14:51 Page 47 47 Tattersfield, P. & Rowson,B.-Tanzartemongen.nov.(Streptaxidae)fromTanzania B75-Tattersfield_Rowson:Basteria-2010 26/06/2011 14:51 Page 48 tionbeingOo.obtusa,whosedescriptioninBlanford& evolvedinsitufromanothermorphologicallydissimilar Godwin-Austen(1908)isnotdetailedenoughtobesure). Africanlineageorfromlineagesfoundotherwiseonlyin ExceptinS.glabernormalis,thepenialretractorjoinsthe AsiaorSouthAmerica.TheEasternArcMountainforests,of columellarmuscle.Allhavepenialhooksthroughoutthe whichthetypelocalitiesofbothTanzartemonspeciesforma interior,exceptforOo.obtusa(againnodata)andfor part,arenowfamousfortheirhighlevelsofendemism, D.stenostomus,whichlackshooksandhas“cornifiedridges” includingbothrecent,specioseradiationsandancientrelicts andasingle“stylet”apically(Berry,1965).Althoughthere withdistantaffinities(Burgessetal.,2007).Intheabsenceof areshellsimilaritiesbetweenthisspeciesandT.seddonae,the geneticdata,themorphologicaldistinctnessthatleadsusto penisofD.stenostomusisquitedifferent.Eachoftheremain- proposeanewgenussuggestsTanzartemonisanexampleof ingspecieshasotherapparentlymoreminorunique thelatter. features(seeSchileyko,2000),asdoesT.seddonae,butAsian, SouthAmericanandAfricanspeciesallsharethesameplan. Acknowledgements Insummary,anatomicalfeaturesdolittletosuggestan especiallycloserelationshipbetweenT.seddonaeandany WethankDrMarySeddonforassistanceduringfieldwork onegenus,whileacloserelationshipwiththeAfrican andforcriticaldiscussionaboutthestatusofthesetaxa. Lamelligerappearsparticularlyunlikely. FieldworkwassupportedbytheUKDarwinInitiativeand Thesamepatternissuggestedbyradularcharacters. bygrantstoPTfromTheBritishEcologicalSociety,The RadulardataareavailablefromH.contusulus,L.maassi, LinneanSocietyofLondonandThePeoplesTrustfor L.troberti,Oo.obtusa,andS.glabernormalis,plusother EndangeredSpecies.ChristineNgerezaandotherstaffof streptaxidsintheliteratureandrecentlyexaminedbyBR. theNationalMuseumsofTanzaniagaveinvaluable ThetwoLamelligerspecieshaveanunusualradulawitha suppportandtheTanzaniaCommissionforScienceand 48 centraltoothandrelativelyfewenlargedinnerlateralsthat Technologygrantedpermissiontoundertakefieldworkand rapidlychangeinsizetowardsthemarginalteeth.Asimilar researchinTanzania.TheTanzaniaDepartmentofForestry patternisshownbytheSouthAmericanS.glabernormalis.In andBeekeeping,CatchmentForestry,andDistrictForestry contrast,theshape,number,gradualchangeinsize,and Officersprovidedsupportandgavepermissiontoworkin apparentabsenceofacentraltoothinT.seddonaeiscloserto theForestReserves.TegeteroMissionandthepeopleof theotheraxiallydeviatedtaxastudiedbyThiele(1933) Tegeterovillageprovidedlocalsupportandassistance andDegner(1934)thantoeitheroftheLamelligerspecies.As duringfieldworkinUluguruNorthForestReserve.Wealso withgenitalanatomy,thetypeofradulaseeninT.seddonae thanktwoanonymousreviewersfortheircommentsand appearstobearathergeneralone,occurringintheBurmese suggestionsonapreviousdraftofthemanuscript. Oo.obtusaandtheSouthAmericanH.contusulus,aswellas manyAfricanspeciesofGonaxis(s.s.ands.l.).Again,aclose relationshipwithLamelligerisnotsupported. References NothingcloselyresemblingLamelliger,oranyofthe othergeneralistedabove,haspreviouslybeenrecorded Adam,W.,1962a.Etudessurlesmollusquesdel’AfriqueCentraleetdes fromcentral,EastorsouthernAfrica,Madagascarorthe régionsvoisines.3.QuelquesespècespeuconnuesdeGonaxis.− IndianOceanislands(tworecordsfor“Rodriguez”basedon Bulletindel’InstitutRoyaldesSciencesNaturellesdeBelgique38 theCumingcollectionhavingbeendismissedaserroneous (17):1-10,pl.1. [Germain,1921;Griffiths&Florens,2006]).Herewepropose Adam,W.,1962b.Etudessurlesmollusquesdel’AfriqueCentraleetdes thatshellandanatomicalfeaturessuggestTanzartemonisan régionsvoisines.4.ContributionàlaconnaissancedugenreLamel- EastAfricanendemicgenusnotcloselyrelatedtoLamelliger. ligerAncey,1884.−Bulletindel’InstitutRoyaldesSciencesNa- Assuchitisofsomebiogeographicalinterest,havingeither turellesdeBelgique38(21):1-11. Basteria75(1-3)