Squamapion alpinum n. sp. from Austria (Coleoptera, Apionidae) Author Giusto, Carlo text Zootaxa 2018 2018-04-11 4407 3 401 414 journal article 30293 10.11646/zootaxa.4407.3.7 18df27be-d100-4d62-bf13-8442f78aabe7 1175-5326 1216638 1AC82902-42BB-4BF3-AAB0-F5065928063B Squamapion alpinum n. sp. ( FIGS 1–2 , 7–10 , 19–22 , 31–31, 37–38, 43 , 46, 49, 52–53 ) Type locality. VIRGEN: MARIN (OSttIROL, AUStRIA ). Distribution. AUStRIA : OSttIROL, KäRNtEN , LOwER AUStRIA . Host plant. Clinopodium alpinum (L.) KUNtzE. Diagnosis. A Squamapion SPEcIES bELONGING tO tHE S. vicinum -GROUP AS dEfINEd bY WANAt (1997) , dIffERING MAINLY fROM Squamapion vicinum (KIRbY, 1808) IN tHE fOLLOwING cOMbINAtION Of cHARActERS: bOdY—ON AVERAGE— SMALLER, SEXUAL ROStRAL dIMORPHISM LESS EVIdENt, fEMALE ROStRUM tHIcKER ANd SHORtER, PROtHORAX LESS AbRUPtLY cONStRIctEd At APEX ANd LESS cOARSELY PUNctUREd, ELYtRAL HUMERAL cALLI LESS dEVELOPEd, PROtHORAcIc ANd ELYtRAL dORSAL OUtLINES MORE ROUNdEd, LEGS SHORtER ANd dIffERENt SHAPE Of GENItALIA. At fIRSt SIGHt, tHE SMALLESt SPEcIMENS cOULd bE EASILY cONfUSEd wItH tHE bIGGESt SPEcIMENS Of Squamapion atomarium (KIRbY, 1808) , bUt tHEY cAN bE dIStINGUISHEd bY tHE MORE ROUNdEd MESOROStRAL dILAtAtION, MALE ROStRUM fEEbLY tAPERING tOwARdS APEX, MORE cONVEX EYES, bEttER dEVELOPEd HUMERAL cALLI Of ELYtRA, QUItE UNIfORMLY ROUNdEd ELYtRA wItH tHE MAXIMUM wIdtH IN tHE MIddLE (NOt IN tHE bASAL tHIRd ANd tHEN NARROwEd tOwARdS tHE APEX), LONGER LEGS ANd dIffERENt SHAPE Of GENItALIA. FIGURES 1–6. habitus. 1–2: Squamapion alpinum , 1: ♂ holotype, 2: ♀ paratype, both from Virgental: Marin.—3–4: Squamapion vicinum , 3: ♂, from Mödling, 4: ♀, from Stronach.—5–6: Squamapion atomarium , 5: ♂, 6: ♀, both from Hodonín: env. Pánov. Scale bar: 1 mm. Type material. HOLOtYPE ( ♂ ): OSttIROL, AUStRIA / VIRGENtAL / LEG. DR. AL. KOFLER // VIRGEN / 24.8.2000 // MARIN / 1400 M // Apion cf. alpinum dEt. A. KOfLER // FERDINANDEUM / INNSBRUCK / SLG. A. KOFLER / 2011-154 ( TLFN ). PARAtYPES (59 ♂♂, 51 ♀♀): OSttIROL, AUStRIA / VIRGENtAL / LEG. DR. AL. KOFLER // VIRGEN / 24.8.2000 // MARIN / 1400 M // Apion cf. alpinum dEt. A. KOfLER // FERDINANDEUM / INNSBRUCK / SLG. A. KOFLER / 2011-154 (1 ♀ CG; 1 ♂, 2 ♀♀ TLFN); AUSTRIA OST-TIROL / VIRGENTAL MARIN / 1999.08.13 1380m / leg. W. Suppantschitsch (1 ♂, 1 ♀ CG; 4 ♂♂, 4 ♀♀ NMW); AUSTRIA TIROL / Matrei Virgental Marin / 1999.08.14 1380m / leg. W. Suppantschitsch (1 ♂, 1 ♀ CG; 1 ♂, 2 ♀♀ NMW); Austria, Osttirol / Virgental / leg. Dr. Al. Kofler // MARIN / 24.7.96 // AN / Acinos // Apion /? n. sp. ♀ / dEt. A. KOfLER // PROPE vicinum (2 ♀♀ NMW); Virgen: Marin (OSttIROL: / VIRGENtAL) 1.400 M / 31.VII.2016 / C. GIUStO LEG. // ON: Clinopodium alpinum (L.) KUNtzE (1 ♂, 1 ♀ BMNH; 19 ♂♂, 11 ♀♀ CG; 1 ♂, 1 ♀ EC; 1 ♂, 1 ♀ KS; 1 ♂, 1 ♀ MNHN; 1 ♂, 1 ♀ MR; 1 ♂, 1 ♀ MW; 1 ♂, 1 ♀ TAUI); Austria Tirol Matrei / Virgental Burg / 1999.08.13 gestreift / leg. W. Suppantschitsch (1 ♂ CG; 2 ♂♂ NMW); AUSTRIA TIROL / Matrei Virgental Burg / 1999.08.14 1400m / leg. W. Suppantschitsch (1 ♂, 2 ♀♀ NMW); OST-TIROL / LEG. A. Kofler // Obermauern / 28.8.88 // Burg / 1400 m (1 ♂, 2 ♀♀ NMW); Virgen: ObERMAUERN: / BURG (OSttIROL: VIRGENtAL) / 1.400 M 31.VII.2016 / C. GIUStO, M. KAHLEN LEG. // ON: Clinopodium alpinum (L.) Kuntze (1 ♂, 1 ♀ CG); Osttirol, Austria / Virgental / leg. Dr. Al. KOFLER // Obermauern / 10.8.1989 // AN Acinos / alpinus // Apion / n. sp. // FERDINANDEUM / INNSBRUCK / SLG. A. KOFLER / 2011-154 (1 ♂ CG; 2 ♀♀ TLFN); OST-TIROL / leg. A. KOfLER // ObERMAUERN / 27.7.78 // A. vicinum / dEt. DIEcKMANN (3 ♂♂, 2 ♀♀ NMW); OST-TIROL / leg. A. Kofler // Obermauern / 16.7.88 (2 ♂♂ NMW); OST-TIROL / leg. A. Kofler // Obermauern / 18.8.88 (1 ♂ NMW); Osttirol, Austria / Virgental / leg. Dr. Al. Kofler // Obermauern / 10.8.1989 // AN Acinos / alpinus (4 ♂♂ NMW); Osttirol, Austria / Virgental / leg. Dr. Al. Kofler // Obermauern / 10.8.1989 // Apion / n. sp. (3 ♀♀ NMW); Virgen: Mellitz: Rabenstein / (Osttirol: Virgental) 1.350- / 1.380 m 31.VII.2016 / C. GIUStO, M. KAHLEN LEG. // ON: Clinopodium alpinum (L.) Kuntze (4 ♂♂ CG); Matrei / Ostt. 25.6.34 // 494 // A. vicinum Kirby / det. Vorišek, 1969 (1 ♀ NMW); Heiligenblut / leg. H. Franz // Apion / dEt. H. WAGNER / vicinum Kb. (1 ♀ NMW); AUSTRIA Kärnten / Karawank., Hoch- / Obir, 9.8.1983 / leg. Döberl // Apion / atomarium Kb. / dEt. MESSUtAt // Squamapion / sp. nov. / det. M. Wanat 1997 / DEI Müncheberg / Col—03314 (1 ♀ SDEI); AUSTRIA Kärnten / Karawank., Hoch- / Obir, 5.8.1986 / leg. Döberl (1 ♂ CG; 1 ♂, 3 ♀♀ SDEI); AUSTRIA Kärnten / Karawank., Hoch- / Obir, 28.8.1985 / leg. Döberl (1 ♀ CG); Karavanchen, Cr. / Ne.Obir // Hölzel leg. // Coll. / Klagenfurt / via Kofler (1 ♂ NMW); AUSTRIA KÄRNTEN / Eisenkappel Hochobir / 1999.08.15 1700m / leg. W. Suppantschitsch (1 ♂ NMW); Eisenkappel / 11.8.75 Kä. / E. Ulbrich leg. // Oistra / 1577 m / Apion / vicinum (1 ♀ SDEI); HOLdHAUS / BAdEN // COLL. MUS. VINdOb. (1 ♀ NMW). Etymology. THE NAME—fROM tHE LAtIN AdJEctIVE “ alpinus ”, PERtAINING tO tHE ALPS—REfERS tO tHE dIStRIbUtION AREA, LOcAtEd wItHIN tHE ALPINE cHAIN, INHAbItEd bY tHE NEw SPEcIES. Description of holotype . BOdY INtEGUMENt bLAcK; ANtENNAE tEStAcEOUS wItH dARKENEd tERMINAL SEGMENt Of fUNIcLE ANd dARKER cLUb; fEMORA bLAcK; tIbIA dARK bROwN tO NEARLY bLAcK, ONLY bASES Of PROtIbIAE ARE LIGHtENEd. VEStItURE wHItE tO cREAM-wHItE cOMPOSEd Of PILIfORM ScALES REGULARLY dIStRIbUtEd OVER tHE ENtIRE bOdY, JUSt cONcENtRAtEd AROUNd tHE EYES, PROcOXAE, INtERcOXAL PROcESS Of MESOVENtRItE, MESO- ANd MEtANEPIStERNA ANd MESOEPIMERA; ON ELYtRA, ScALES, AS LONG AS INtERVALS’ bREAdtH, ARE ARRANGEd IN A SINGLE REGULAR ROw ON INtERStRIAE, At bASE—bEtwEEN tHE 2Nd ANd tHE 3Rd ELItRAL StRIAE—SOME ScALES fORM cONfUSEd ROwS; StRIAL ScALES ARE SLIGHtLY fINER ANd dIStINctLY SEPARAtEd fROM EAcH OtHER. ONLY tHE twO bASAL tHIRdS Of PROROStRUM bEAR SHORt ANd fINE fEEbLY VISIbLE PILIfORM ScALES. LENGtH Of SPEcIMEN (bL): 1.85 MM. ROStRUM AS IN FIGS 7–8 ; RELAtIVELY SHORt ANd StOUt ; IN dORSAL VIEw, bASAL HALf Of PROROStRUM SLIGHtLY NARROwING fROM MESOROStRAL dILAtAtION fORwARd, APIcAL HALf cYLINdRIcAL, IN PROfILE dIStINctLY ANd REGULARLY cURVEd, PROROStRUM SLIGHtLY NARROwING APIcAd; SHAGREENEd, IRREGULARLY ANd SHALLOwLY PUNctAtE, MAtt, SHINING ONLY IN tHE APIcAL tHIRd Of PROROStRUM; RL: 0.47 MM, PRRL: 0.36 MM, MtRL: 0.10 MM, PRRAw: 0.11 MM, PRRMw: 0.11 MM, MSRw: 0.15 MM, MtRw: 0.14 MM, bL/RL: 3.94, PL+EL/RL: 3.66, RL/PL: 1.09, RL/MSRw: 3.13, RL/PRRMw: 4.27, MSRw/PRRMw: 1.36, MSRw/PRRAw: 1.36. HEAd cONIcAL ANd tRANSVERSE; HL: 0.25 MM, Hw: 0.35 MM; fRONS bARELY dEPRESSEd, ALMOSt fLAt, RUGOSELY PUNctAtE ANd StRONGLY MIcROScULPtUREd wItH A SHORt MEdIAN fOVEA; EYES MOdERAtELY LARGE ANd REGULARLY cONVEX, EYL: 0.19 MM; Hw/EYL: 1.84. ANtENNAE ( FIG. 31 ) INSERtEd At bASAL 0.21 Of ROStRUM, SLENdER, wItH fINE, POORLY EREct SEtAE; AL: 0.67 MM, AScL: 0.13 MM, AL/PRRL: 1.86, AScL/MSRw: 0.87. PROtHORAX ( FIG. 1 ) tRANSVERSE, cAMPANULAtE, wItH cURVEd ANd bISINUAtE SIdES, cONStRIctEd bEHINd ANtERIOR MARGIN ANd bEfORE bASE, wItH PROMINENt HINd ANGLES, wIdESt At bASE; dISc MOdERAtELY cONVEX, dENSELY PUNctAtE, PUNctURES cOARSE ANd tHIcK, ROUNd, 22–27 µM IN dIAMEtER, SEPARAtEd bY LESS tHAN ONE tHIRd Of tHEIR dIAMEtER; INtERSPAcES cONVEX, StRONGLY MIcROREtIcULAtE, MAtt; PREScUtELLAR fOVEA JUSt bROAdER tHAN dIScAL PUNctURES ANd NOt LONGER tHAN tHREE PUNctURES; PL: 0.43 MM, Pb: 0.57 MM, Pw: 0.51 MM, PAw 0.41 MM PAcw: 0.41 MM, PL/Pw: 0.84, Pbw/PAw: 1.39, Pw/Hw: 1.46, PL/Pbw: 0.75. ScUtELLUM SUbtRIANGULAR wItH cURVEd SIdES, JUSt LONGER tHAN wIdE, fLAt ON dISc, GLAbROUS ANd StRONGLY MIcROREtIcULAtE, MAtt. ELYtRA ( FIG. 1 ), IN dORSAL VIEw, SUbREctANGULAR IN OUtLINE, wItH MOdERAtELY cURVEd SIdES, wIdESt IN tHE MIddLE ANd wItH HUMERAL cALLI wELL dEVELOPEd; INtERStRIAE 1.7–2.0 tIMES AS wIdE AS StRIAE, fLAt, RUGOSELY MIcROScULPtUREd wItH wEAK tRAcES Of ALMOSt INdIStINct PUNctURES; StRIAE SHARPLY MARGINAtEd, cAtENULAtE, wItH SEPtAE bEtwEEN PUNctURES dIStINctLY dEEPENEd; EL: 1.29 MM, Ebw : 0.75 MM, Ew : 0.83 MM, EL/ Ew : 1.55, EL/PL: 3.00, Ew / Pw : 1.36, Ew / Pbw : 1.46, Ebw / Pw : 1.47, Ebw / Pbw : 1.32 . LEGS MOdERAtELY SLENdER; HINd fEMORA tHIcKENEd; PROtIbIAE fEEbLY INcURVEd OUtwARdS, RELAtIVELY SHORt; tARSI MOdERAtELY ELONGAtE, 1 St SEGMENt 2.1 tIMES AS LONG AS wIdE, 2 Nd ISOdIAMEtRIc, 0.63 tIMES AS LONG AS 1 St, 3Rd ALMOSt ISOdIAMEtRIc, 0.85 tIMES AS LONG AS wIdE, wItH LObES wELL dEVELOPEd, ONYcHIUM 1.2 tIMES AS LONG AS 2 Nd SEGMENt; PRfL: 0.48 MM, PRfw: 0.15 MM, PRfL/ PRfw: 3.20, PRtL: 0.58 MM, PRtw: 0.08 MM, PRtL/PRtw: 7.25, PRtL/PL: 1.35. MEtAtHORAcIc wINGS wELL dEVELOPEd. FIRSt twO VENtRItES dENSELY PUNctAtE, PUNctURES cOARSE ANd tHIcK, ROUNd, 20–25 µM IN dIAMEtER, SEPARAtEd bY LESS tHAN ONE tHIRd Of tHEIR dIAMEtER; INtERSPAcES cONVEX, StRONGLY MIcROREtIcULAtE, MAtt; VENtRItES III–V IMPUNctAtE, StRONGLY MIcROREtIcULAtE, MAtt. TEGMEN ( FIGS 37–38 ) wItH tEGMINAL APOdEME LONGER tHAN fORKEd bASAL PIEcE; tEGMINAL PLAtE wItH MEdIAN cARINAE MOdERAtELY SEPARAtEd, REAcHING ScLEROtISEd PLAtES ANd cURVEd OUtwARdS AbOVE tHEM; ScLEROtISEd PLAtES bEARING 9 (LEft) OR 11 (RIGHt) PORES; APIcAL MEMbRANOUS LObES RUdIMENtARY ANd UNItEd, wItH SOME MIcROcHAEtAE MOStLY At OUtER MARGINS; wHOLE fOLdEd MARGINS Of tEGMINAL PLAtE wItH VERY fINE, dENSE LONfItUdINAL cARINAE. PENIS ALMOSt SYMMEtRIcAL, SLIGHtLY dOwNcURVEd IN PROfILE ( FIGS 19–22 ); tEctUM wItH MELANIzEd MEdIAN cARINA REAcHING NEARLY HALf LENGtH Of tHE LObE; ENdOPHALLUS, IN ORIfIcIAL REGION, wItH twO AGGREGAtIONS Of dENSE, ALMOSt UNIfORM ANd MINUtE SPINES. SPIcULUM GAStRALE AS IN FIG. 43 . Description and variation of male and female paratypes . APARt fROM MORPHOLOGIcAL VARIAbILItY SUMMARIzEd IN TAb. 1, MALE PARAtYPES ARE VERY SIMILAR tO tHE HOLOtYPE . IN AddItION tO EXPEctEd MORPHOLOGIcAL dIffERENcES (SEE TAb. 1), fEMALE dIffERS fROM MALE AS fOLLOw: ANtENNAL fUNIcLE SLIGHtLY dARKER. PROROStRUM wItH fINER ANd SPARSER PILIfORM ScALES VANISHING IN tHE APIcAL HALf . EYES dIStINctLY SMALLER. ANtENNAE AS IN FIG. 32 . ELYtRA ( FIG. 2 ) , IN dORSAL VIEw, wItH cURVEd SIdES. GONOcOXItES AS IN FIG. 46 ; StYLI SUbtRIANGULAR, 1.4–1.5 tIMES AS LONG AS wIdE ANd wItH 7–8 APIcAL SEtAE NOt LONGER tHAN HALf StYLUS LENGtH; cOXItES LONG , dIVERGENt wItH fINE MEdIAN cARINA; fOREtUbE (NOt fIGUREd) cOVEREd wItH MINUtE ANd SIMPLE MIcROtRIcHIA. StERNItE VIII ANd SPERMAtHEcA AS IN FIGS 49 , 52–53 . Bionomy. AS REPORtEd AbOVE, AdULtS Of S. alpinum wERE cOLLEctEd ON Clinopodium alpinum (L.) KUNtzE ( LAMIAcEAE ) ANd, EVEN tHOUGH NO LARVA HAS bEEN fOUNd INSIdE StEMS OR ROOtS, It cAN bE cONSIdEREd StRIctLY MONOPHAGOUS ON tHIS PLANt ( FIGS 59–60 ). WEEVILS wERE SwEPt fROM tHERMOPHILOUS ANd XEROtHERMOPHILOUS SOUtHfAcING MEAdOwS, MOStLY ON SILIcEOUS-LIMEStONE SOILS ( FIGS 57–58 ). PHENOLOGIcAL dAtA INdIcAtE tHAt AdULtS, fROM MIddLE ALtItUdES (1,350–1,700 M), ARE ActIVE fROM MIddLE Of JULY tO LAtE AUGUSt. THE ONLY KNOwN SPEcIMEN fROM LOw ELEVAtION (230 M) IS tHE fEMALE fROM BAdEN bEI WIEN. Remarks. MEASUREMENtS ANd RAtIOS, QUANtIfIEd ANd SUMMARIzEd IN TAb. 1, cLEARLY INdIcAtE tHAt MORPHOLOGIcAL VARIAbILItY Of S. alpinum , S. vicinum ANd S. atomarium IS PARtIcULARLY wIdE. ALMOSt ALL RANGES Of VARIAbILItY PARtLY OVERLAPP MAKING tHE PROPER REcOGNItION Of SPEcIES NOt ALwAYS EASY; PRObAbLY, fOR tHIS REASON, IN tHE PASt, EMINENt SPEcIALIStS AS H. WAGNER ANd L. DIEcKMANN MISIdENtIfIEd S. alpinum wItH S. vicinum wHILE OtHER ENtOMOLOGIStS cONfUSEd It wItH S. atomarium (SEE “ TYPE MAtERIAL”). DESPItE tHE PRESENt dEtAILEd StUdY Of MORHOLOGIcAL cHARActERS, RELAtIONSHIPS AMONG tHE tHREE AfOREMENtIONEd SPEcIES REMAIN UNcERtAIN ANd, ON tHE bASIS Of OUR StILL POOR KNOwLEdGE Of tHE SYStEMAtIcS Of tHE GENUS Squamapion , It IS MORE OPPORtUNE tO SPEAK AbOUt SIMILARItY RAtHER tHAN AffINItY. GIVEN tHIS, S. alpinum At fIRSt SIGHt RESEMbLES S. vicinum tHANKS tO ItS RELAtIVELY bIG bOdY SIzE ANd tO tHE INcOSPIcUOUS VEStItURE cOMPOSEd Of wHItE tO cREAM-wHItE PILIfORM ScALES, ON ELYtRAL INtERVALS HARdLY tOUcHING EAcH OtHER IN A SINGLE REGULAR ROw (IN S. atomarium VEStItURE IS dIStINct ANd tHE wHItE tO cREAM cOLOREd ScALES ARE PILIfORM ANd SUbtLY LANcEOLAtE wItH cLEARLY POINtEd tIPS, ON ELYtRAL INtERVALS MOStLY tOUcHING EAcH OtHER OR EVEN SLIGHtLY OVERLAPPING IN A SINGLE ROw) ( FIGS 1–6 ). ON tHE cONtRARY, ItS ROStRUM, ALtHOUGH SHOwING A MORE REdUcEd SEXUAL dIMORPHISM tHAN tHE OtHER twO SPEcIES, IS SIMILAR tO tHAt Of S. atomarium EVEN If, IN tHE LAttER SPEcIES, MESOROStRAL dILAtAtION IS A LIttLE bIt dENtIfORM ANd MALE PROROStRUM IS MORE EVIdENtLY tAPERING tOwARdS tHE APEX ( FIGS 7–18 ). FIGURES 7–18. head and rostrum in dorsal and lateral view. 7–10: Squamapion alpinum , 7–8: ♂ holotype, 9–10: ♀ paratype, both from Virgental: Marin.—11–14: Squamapion vicinum , 11–12: ♂, from Mödling, 13–14: ♀, from Stronach.—15–18: Squamapion atomarium , 15–16: ♂, 17–18: ♀, both from Hodonín: env. Pánov. Scale bar: 0.5 mm. FIGURES 19–30. penis and penal apex in ventral and lateral view. 19–22: Squamapion alpinum , holotype, from Virgental: Marin.—23–26: Squamapion vicinum , from Kraków.—27–30: Squamapion atomarium , from Hodonín: env. Pánov. FIGURES 31–45. antenna, tegmen and spiculum gastrale. 31–32, 37–38, 43: Squamapion alpinum , 31, 37–38, 43: ♂ holotype, 32: ♀ paratype, both from Virgental: Marin.—33–34, 39–40, 44: Squamapion vicinum , 33, 39–40, 44: ♂, from Kraków, 34: ♀, from Prien am Chiemsee.—35–36, 41–42, 45: Squamapion atomarium , 35, 41–42, 45: ♂, 36: ♀, both from Hodonín: env. Pánov. FIGURES 46–56. gonocoxite, sternite VIII and spermatheca. 46, 49, 52–53: Squamapion alpinum , paratype, from Virgental: Marin.—47, 50, 54: Squamapion vicinum , from Prien am Chiemsee.—48, 51, 55–56: Squamapion atomarium , from Hodonín: env. Pánov. PROtHORAX ANd HUMERAL cALLI Of S. alpinum ARE MORE SIMILAR tO tHOSE Of S. atomarium tHAN tHOSE Of S. vicinum ; tHE LAttER SPEcIES IS cHARActERIzEd bY A StRONGER APIcAL PROtHORAcIc cONStRIctION (ESPEcIALLY IN fEMALE), cOARSER PROtHORAcIc PUNctURAtION ANd ELYtRAL HUMERAL cALLI PARtIcULARLY wELL PROMINENt ( FIGS 3, 4 ). JUSt tHE PRESENcE Of wELLdEVELOPEd HUMERAL cALLI GIVES tO ELYtRA Of S. vicinum A "REctANGULAR OUtLINE" wItH wEAKLY ROUNdEd SIdES ANd MAXIMUM wIdtH POSEd IN tHE MIddLE. ALSO S. atomarium SHOwS ELYtRAL SIdES wEAKLY ROUNdEd, bUt, IN tHIS cASE, tHE MAXIMUM wIdtH IS POSEd IN tHE APIcAL tHIRd ( FIGS 5, 6 ). IN S. alpinum , MALE ELYtRAL OUtLINE, wItH tHE MAXIMUM wIdtH POSEd IN tHE MIddLE ( FIG. 1 ), IS SIMILAR tO tHAt Of tHE MALE Of S. vicinum wHILSt tHAt Of fEMALE, cHARActERISEd bY REGULARLY ROUNdEd SIdES ANd MAXIMUM wIdtH POSEd bEHINd tHE MIddLE, IS dIffERENt fROM ALL tHE REMAINING SPEcIES ( FIG. 2 ). PROtIbIAE Of S. alpinum ARE StRAIGHt OR wEAKLY OUtcURVEd, tHUS INtERMEdIAtE IN tHIS RESPEct bEtwEEN S. vicinum (tIbIAE OUtcURVEd) ANd S. atomarium (tIbIAE StRAIGHt) ( FIGS 1–6 ). ALSO tHE EXAMINAtION Of MALE GENItALIA dOES NOt AId tO cLARIfY RELAtIONSHIPS AMONG tHE tHREE SPEcIES: wHILE S. alpinum ANd S. vicinum SHARE A SIMILAR SHAPE Of PENIS ( FIGS 19–26 ), tEGMEN Of S. atomarium , wItH ItS fOLdEd MEMbRANOUS APIcAL LObES, RESEMbLES tHAt Of S. alpinum ( FIGS 37, 38, 41, 42 ). MORPHOLOGY Of fEMALE GENItALIA INStEAd ( FIGS 46–56 ), IN PARtIcULAR tHAt Of SPERMAtHEcA (bOdY SHAPE ANd cURVAtURE, SHAPE Of NOdULUS, POSItION Of GLANd INSERtION ANd LENGHt Of tHE dUct) ALLOwS fOR A SUbStANtIAL REcOGNItION Of tHREE QUItE dIffERENt PAttERNS ( FIGS 52– 56 ).