Conopleura aliena Smriglio, Mariottini & Calascibetta, 1999

[HERE fIRST cONSIdEREd juNIOR SubjEcTIVE SyNONym Of Tritia lima (DILLwyN, 1817): SEE bELOw]

Type locality: Temperate northern Atlantic. CENTRAL TyRRHENIAN SEA (ITALy, MEdITERRANEAN SEA) (FIg. 1D1).

Material examined: Temperate northern Atlantic. CENTRAL TyRRHENIAN SEA (ITALy), 06/1987, 350 m— HOLOTyPE—1 SH—6.8×4 (MZB 12694) (FIg. 1D1, 4A–H, 4W; APPENdIx 1: FIg. 1C). Off ISOLA dELLE FEmmINE (PALERmO, ITALy), 105 m—PARATyPE—1 SH—5×3.5 (SC PRIVATE cOLLEcTION, ONLy PHOTOS ObSERVEd AS IT wAS NOT POSSIbLE TO ExAmINE THE SHELL de visu: AuTHORS’ dATA) (FIg. 1D 2).

Known distribution and bathymetric range: ONLy kNOwN fROm THE cENTRAL MEdITERRANEAN SEA, wHERE TwO SHELLS wERE REPORTEd fROm 105 ANd 350 m dEPTH. Temperate northern Atlantic. ITALy (SmRIgLIO et al. 1999: CENTRAL TyRRHENIAN SEA ANd Off ISOLA dELLE FEmmINE) (FIg. 1D1–2).

Description: SPEcIES wITH A SmALL SHELL, LIgHT, gLObOSE, SLIgHTLy ELONgATE, ANd wITH A TRuNcATE SPIRE, Of ~ 7 mm mAxImum HEIgHT ANd 4.5 wHORLS IN THE HOLOTyPE (FIg. 4A–C). SHELL cOLOuR ALmOST wHITISH-yELLOwISH (FIg. 4A–C). PROTOcONcH Of ~1 wHORL, SmOOTH buT PARTIALLy EROdEd (FIg. 4G–H, 4W). APEx dEPRESSEd (FIg. 4H). TELEOcONcH Of 4.5 cONVEx wHORLS IN bIggEST SPEcImEN kNOwN, INcREASINg IN dIAmETER uP TO LAST wHORL, cONSTITuTINg LITTLE LESS Of TOTAL SHELL HEIgHT ANd SIgNIfIcANTLy ExPANdEd AT SHOuLdER ANd SubSEquENTLy NARROwINg (FIg. 4A–C). SuTuRE dISTINcT, SLIgHTLy INcISEd wITH A SuLcuS, SHOuLdER NOT cORONATE (FIg. 4A–D, 4F). SHOuLdER SuLcuS mAdE Of A SERIES Of VERy SmALL SEPTA SEPARATEd by RAdIATINg LAmELLAE ORIgINATINg fROm ANAL SINuS (FIg. 4D). AxIAL ScuLPTuRE fINE, POSSIbLy cONSTITuTEd by SEVERAL SubSEquENT gROwTH LINES, AxIAL RIbS PRESumAbLy PRESENT (FIg. 4A–C, 4E). SPIRAL ScuLPTuRE IN THE cONVEx PART Of THE wHORL cONSISTINg Of 7–9 fAINT PRImARy cORdS, OfTEN fLExuOuS wHEN cROSSINg gROwTH LINES (FIg. 4A–C). APERTuRE wIdE, OVAL, uNIfORmLy wHITISH-yELLOwISH (FIg. 4A–C). UmbILIcuS OR fALSE umbILIcuS AbSENT (FIg. 4A–C). COLumELLAR EdgE wITH SLIgHTLy ExPANdEd cALLuS ANd EVIdENT cOLumELLAR RIb (FIg. 4A). OuTER LIP THIN wITH AbSENcE Of STROmbOId NOTcH (FIg. 4B). ANAL SINuS dEEP, LARgE, U-SHAPEd, cLOSE TO SuTuRE (FIg. 4A–B). SIPHONAL cANAL THIck, SHORT ANd bROAd (FIg. 4A). ANATOmy, RAduLA, ANImAL cOLOuR ANd OPERcuLum uNkNOwN.

Remarks: THE dIffERENcES IN SHELL mORPHOmETRy Of C. aliena ANd THOSE Of SPEcIES REPORTEd AS cONgENERIc IN THE cuRRENT LITERATuRE ARE SHOwN IN TAbLE 1. REmARkAbLy, SEVERAL C. aliena SHELL fEATuRES dO NOT mATcH THE TwO OTHER TAxA AScRIbEd TO Conopleura HINdS, 1844 . THESE INcLudE: I) PROTOcONcH wHORLS (~1 vs ~2.25–2.5 IN OTHER Conopleura TAxA) (FIg. 3K–L, 3V –Z, 4G–H); II) TELEOcONcH gENERAL ASPEcT (ObTuSE, SLIgHTLy ELONgATE wITH A TRuNcATE SPIRE vs SLENdER ANd bIcONIcAL wITH AN AcumINATE SPIRE) (FIg. 3A–C, 3F–G, 3M–Q, 4A–C) ANd ScuLPTuRE (fAINT vs STRONg PRImARy cORdS) (FIg. 3A–C, 3F–G, 3I, 3M–Q, 3T, 4A–C); III) fALSE umbILIcuS (AbSENT vs PRESENT) (FIg. 3J, 3U, 4A); IV) APERTuRE gENERAL ASPEcT (wIdE ANd OVAL vs NARROw, SINuOuS, ANd AxIALLy ELONgATEd) (FIg. 3A, 3F, 3M, 3P, 4A); V) mORPHOLOgIcAL fEATuRES Of THE cOLumELLAR EdgE (cOLumELLAR RIb PRESENT vs AbSENT) (FIg. 3J, 3U, 4A) ANd Of THE OuTER LIP (STROmbOId NOTcH PRESENT vs AbSENT) (FIg. 3G, 3Q, 4B). HOwEVER, ON THE OTHER HANd, OTHER fEATuRES fIT THE gENuS Conopleura, THE mOST ImPORTANT ONES bEINg THE PRESENcE Of A SHOuLdER SuLcuS wITH SEPTA SEPARATEd by RAdIATINg LAmELLAE ANd AN ANAL SINuS (FIg. 3H, 3R, 3D), wHIcH HAVE SuggESTEd ITS ORIgINAL ATTRIbuTION TO THE AbOVE-mENTIONEd gENuS.

THE mAjOR dIScREPANcIES bETwEEN C. aliena ANd cONgENERIc SPEcIES ImmEdIATELy RAISEd quESTIONS AbOuT THE SPEcIfIc, gENERIc, ANd fAmILIAR ATTRIbuTION Of THIS SPEcIES. SmRIgLIO et al. (1999) TENTATIVELy AScRIbEd THIS TAxON TO Conopleura IN AbSENcE Of SOfT PARTS, buT ITS gENERIc ATTRIbuTION wAS SubSEquENTLy uTILIzEd IN dATAbASES ANd cHEckLISTS (E.g. GOfAS 2009; TEmPLAdO & VILLANuEVA IN COLL et al. 2010). HOwEVER, THE PRESENcE Of A cOLumELLAR RIb dOES NOT fIT THE SuPERfAmILy CONOIdEA sensu BOucHET et al. (2011) AS A wHOLE. SmRIgLIO et al. (1999) ALSO ORIgINALLy HIgHLIgHTEd SImILARITIES Of C. aliena wITH SOmE mEmbERS Of THE fAmILy NASSARIIDAE IREdALE, 1916 (1835), ANd IN PARTIcuLAR cOmPAREd THE NEwLy dEScRIbEd TAxON wITH Tritia lima (DILLwyN, 1817) (fORmERLy Nassarius lima: SEE GALINdO et al. 2016), A MEdITERRANEAN ENdEmIc INcORREcTLy dEScRIbEd AS A REcENT SPEcIES fROm THE EAST INdIES (CHEmNITz 1795; DILLwyN 1817; CERNOHORSky 1984), wHOSE TyPE mATERIAL wAS fIRST TRAcEd ANd fIguREd HERE (FIg. 4I – J; APPENdIx 1: FIg. 1D). HOwEVER, dIffERENcES wITH RESPEcT TO T. lima LISTEd by SmRIgLIO et al. (1999) INcLudE PROTOcONcH ANd TELEOcONcH fEATuRES, AS wELL AS THE PRESENcE Of THE ANAL SINuS (SEE bELOw fOR ExPLANATION). IN AddITION, T. lima IS cONcEIVEd AS POSSESSINg A SERIES Of dENTIcLES ON THE INTERNAL PART Of THE OuTER LIP.

SmRIgLIO et al. (1999) ORIgINALLy SuggESTEd THAT dIffERENcES IN THE PROTOcONcHS Of C. aliena ANd C. striata wERE duE TO THE fAcT THAT PROTOcONcHS Of C. aliena wERE LOST, ANd THAT A dOmEd INTERNAL cALLOuS PLug wAS ONLy VISIbLE—A cHARAcTER THAT THEy HAd NOT ObSERVEd IN T. lima . OuR SEM ObSERVATIONS Of THE C. aliena HOLOTyPE SuggEST THAT THE PROTOcONcH, ALTHOugH bLuNT, IS AcTuALLy PRESENT ANd cOmPRISEd Of ~1 wHORL (FIg. 4G, 4W). THE INfERREd PROTOcONcH Of C. aliena IS SImILAR TO THAT Of T. lima IN TERmS Of THE NumbER Of wHORLS (~1 vs 1.25, RESPEcTIVELy) ANd THEIR SHAREd fAINT PROTOcONcH/TELEOcONcH bOuNdARy ANd TELEOcONcH ScuLPTuRE (FIg. 4V –W) (SEE ALSO NEgRI & CORSELLI 2016). ON THE OTHER HANd, THE PROTOcONcH Of C. aliena IS INdEEd LARgER, AT LEAST IN THE HOLOTyPE (IT wAS NOT POSSIbILE TO ExAmINE THE PARATyPE), wITH RESPEcT TO THAT Of T. lima (FIg. 4V –W), buT THE SAmE HOLdS EVEN mORE wHEN cOmPARINg IT TO THE OTHER TwO Conopleura TAxA. THIS mAy bE ExPLAINEd by THE PRESENcE IN C. aliena Of AN AbNORmALITy THAT wILL SubSEquENTLy RESuLT IN A HyPERmORPHIc SHOuLdER (SEE bELOw fOR dIScuSSIONS, ANd FIg. 6Q–W fOR A SImILAR SITuATION IN THE wHORLS Of Buccinum labyrinthus GmELIN, 1791 vs Buccinum undatum LINNAEuS, 1758).

SmRIgLIO et al. (1999) ALSO HIgHLIgHTEd THE PRESENcE Of STRONg AxIAL RIbS IN T. lima TELEOcONcHS, wHIcH ARE AbSENT IN C. aliena . THIS mORPHOLOgIcAL fEATuRE, HOwEVER, HAS NO OVERALL TAxONOmIc VALuE, AS T. lima mAy OR mAy NOT HAVE AxIAL ScuLPTuRE, wITH INTERmEdIATE fORmS bETwEEN THE TwO ExTREmE cHARAcTER STATES (VARIAbILITy IN T. lima TELEOcONcH fROm C. aliena TyPE LOcALITy: FIg. 4P–T). WHEN SEEN fROm bELOw, THE HOLOTyPE Of C. aliena ALSO SHOwS A POSSIbLE AxIAL ScuLPTuRE Of fAINT RIbS (FIg. 4E). FINALLy, THE PRESENcE/AbSENcE Of dENTIcLES ON THE OuTER LIP dEPENdS ON gROwTH STAgE (PRESENT IN fuLLy gROwN LIPS vs AbSENT IN PARTIALLy gROwN LIPS ANd “juVENILES”), SuggESTINg THAT THIS dIAgNOSTIc cHARAcTER ALSO cANNOT bE cONSIdEREd (FIg. 4I –T).

HOwEVER, IN AgREEmENT wITH SmRIgLIO et al. (1999), AN ANAL SINuS ANd SHOuLdER SuLcuS wITH SmALL SEPTA ARE uSuALLy LAckINg IN T. lima . CAREfuL ObSERVATIONS Of C. aliena SHELLS HIgHLIgHTEd A dIScONTINuOuS LAmELLAR ScuLPTuRE IN THE SHOuLdER SuLcuS (FIg. 4D, 4F–H), EVEN wITH SOmE dIffERENcES bETwEEN HOLOTyPE ANd PARATyPE (THE LAmELLAR ScuLPTuRE IN THE HOLOTyPE STARTS AT 2.5 wHORLS, wHILST IN THE PARATyPE IT STARTS AT 2.25 wHORLS). IN AddITION, SEPTA IN THE SPEcIES Of Conopleura HINdS, 1844 START SOON AfTER THE PROTOcONcH ANd ARE REguLARLy ARRANgEd (FIg. 3H, 3R), THEREfORE SuggESTINg THAT SucH A PEcuLIAR ScuLPTuRE Of THE SHOuLdER SuLcuS REPRESENTS A dEfININg cHARAcTER Of THE gENuS. THOSE IN C. aliena START ONLy AfTER AN AbRuPT bREAk ON THE SHELL SHOuLdER (FIg. 4F–H), THEREfORE SuggESTINg THAT THIS IS NOT A dIAgNOSTIc cHARAcTER, buT A mEcHANIcAL AbNORmALITy.

TO EVALuATE THE HyPOTHESIS THAT THE LAmELLAR ScuLPTuRE Of C. aliena REPRESENTS AN AbNORmAL gROwTH, wE cONducTEd A LITERATuRE ANd mATERIAL SEARcH ON AbERRATIONS IN gASTROPOd SHELLS. ON THE ONE HANd, AN ANAL SINuS ANd SHOuLdER SuLcuS wITH SEPTA ARE dIAgNOSTIc cHARAcTERS Of SOmE gENERA (E.g. Conopleura). ON THE OTHER, THESE fEATuRES REPRESENT mORPHOLOgIcAL AbNORmALITIES, OfTEN ASSOcIATEd wITH A HyPERmORPHIc SHOuLdER, IN A wIdE RANgE Of gASTROPOd fAmILIES ANd SPEcIES, INcLudINg CASSIDAE LATREILLE, 1825 (FIg. 5A–D), MURICIDAE RAfINESquE, 1815 (FIg. 5E–I), MITRIDAE SwAINSON, 1831 (FIg. 5K–Q, 6A–L), CONIDAE FLEmINg, 1822 ( Kenyonia pulcherrima: SEE TIPPETT & TuckER 1995; FIg. 6M–P; APPENdIx 1: FIg. 1E), ANd BUCCINIDAE RAfINESquE, 1815 ( Buccinum labyrinthus: SEE PASTORINO & KANTOR 1998; FIg. 6Q–W; APPENdIx 1: FIg. 1F) AmONg OTHERS (SEE ALSO dIScuSSIONS bELOw). ON THE cONTRARy, THE cOLumELLAR RIb Of C. aliena dOES NOT SEEm TO bE AbNORmAL ANd HAS A REguLAR SHAPE, SuggESTINg THAT IT mAy bE A RELIAbLE dIAgNOSTIc cHARAcTER fOR ASSESSINg ITS cORREcT TAxONOmIc IdENTITy. IN PARTIcuLAR, THE cOLumELLAR RIb dELImITINg AdAPIcALLy THE SIPHONAL cANAL IN Conopleura aliena IS STRIkINgLy SImILAR TO THAT APPEARINg IN THE fAmILy NASSARIDAE (FIg. 4). CONVERSELy, IN OTHER SPEcIES Of Conopleura, THE cOLumELLAR EdgE IS bORdEREd by A SORT Of RIm THAT RuNS PARALLEL TO THE mAIN AxIS Of THE SHELL (FIg. 3). WE wERE NOT AbLE TO fINd AbNORmALITIES REPORTEd IN THE cOLumELLAR RIb AS APPEARINg IN C. aliena .

THEREfORE, bASEd ON THE mORPHOLOgIcAL ObSERVATIONS HEREby dIScuSSEd (PROTOcONcH fEATuRES, TELEOcONcH ScuLPTuRE, ANd gENERAL mORPHOLOgIcAL cHARAcTERS), “ Conopleura ” aliena IS HERE ExcLudEd fROm Conopleura ANd mOVEd TO Tritia RISSO, 1826, ANd fuRTHERmORE cONcEIVEd AS dEScRIbEd ON THE bASIS Of TwO AbNORmAL SHELLS Of THE dEEP wATER fORm Of T. lima . AccORdINg TO THE INTERNATIONAL COmmISSION ON ZOOLOgIcAL NOmENcLATuRE, excluded from the provisions of the Code are names proposed for teratological specimens as such (ICZN 2012: ART 1.3.2). BEcAuSE “ Conopleura ” aliena wAS dEScRIbEd AS A PROPER REcENT SPEcIES, IT HAS TO bE cONSIdEREd A juNIOR SubjEcTIVE SyNONym (ICZN 2012: gLOSSARy) Of T. lima . HOwEVER, ONLy dATINg TEcHNIquES (E.g. RAdIOcARbON) mAy TELL uS If ITS dEScRIPTION wAS bASEd uPON fOSSIL REmAINS OR REcENT SHELLS.