Lignyodes bischoffi (Blatchley, 1916)

Caldara, Roberto, 2017, Morphological evidence from immature stages further suggests Lignyodina being close to Tychiina (Coleoptera, Curculionidae, Curculioninae, Tychiini), Zootaxa 4320 (3), pp. 426-446 : 429-436

Lignyodes bischoffi (Blatchley, 1916)

Material examined. Larvae : 86 LARvAE Of dIffERENT INSTARS cOLLEcTEd fROM SEEdS Of gREEN ASh ( Fraxinus pennsylvanica ) ON 20.08.2014 (35 LARvAE), 20.09.2014 (40 LARvAE), 0 7.08.2016 (11 LARvAE) ANd 20 LARvAE Of dIffERENT INSTARS cOLLEcTEd fROM SEEdS Of cOMMON ASh ON 20.10.2014 IN ThE CITy PARk Of LubLIN (GPS 51°14'02.1"N 22°33'40.1"E); 89 LARvAE Of dIffERENT INSTARS cOLLEcTEd fROM SEEdS Of gREEN ASh ON 26.09.2014 ALONg ZEMbORzyckA STREET , LubLIN (GPS 51°12'21.5"N 22°33'53.9"E), IN MIddLE-EASTERN POLANd .

Pupae: 10 SPEcIMENS ObTAINEd IN APRIL 2015 fROM SEEdS Of gREEN ASh INfEcTEd by LARvAE, cOLLEcTEd ALONg ZEMbORzyckA STREET ON 26.09.2014 ANd ThEREAfTER STOREd IN bOXES WITh SOIL AT OuTdOOR TEMPERATuRE ANd ObSERvEd uNTIL ThE EMERgENcE Of AduLTS.

Description of mature larvae. Measurements (IN MM). BOdy LENgTh: 5.2–6.5 (MEAN 5.7). ThE WIdEST PLAcE IN ThE bOdy (AbdOMINAL SEgMENTS II–VI) MEASuRES uP TO 1.7. HEAd WIdTh: 0.5–0.7 (MEAN 0.6).

General. BOdy ELONgATEd, SLIghTLy cuRvEd, ROuNdEd IN cROSS SEcTION ( FIgS 7 , 18–19 ).

Colouration. PALE TO dARk bROWN hEAd ( FIgS 7 , 18–19 ). ALL ThORAcIc ANd AbdOMINAL SEgMENTS WhITE TO WhITISh yELLOW ( FIgS 7 , 18–19 ).

Vestiture. SETAE ON bOdy ThIN, ORANgE, dISTINcTLy dIffERENT IN LENgTh (MINuTE TO vERy ShORT OR RELATIvELy LONg).

Head capsule ( FIg. 1 ). HEAd ELONgATEd, fLATTENEd LATERALLy, ENdOcARINAL LINE PRESENT, LESS ThAN hALf ThE LENgTh Of fRONS. FRONTAL SuTuRES ON hEAd dISTINcT, EXTENdEd TO ANTENNAE . ONE STEMMA (ST), IN ThE fORM Of A SMALL dARk PIgMENTEd SPOT WITh cONvEX cORNEA, LOcATEd ON EAch SIdE ANTEROLATERALLy, bEhINd ANTENNA. Des1 ANd des2 LOcATEd IN MIddLE Of ThE cENTRAL PART Of EPIcRANIuM, ShORT des1 NEAR TO ThE MIddLE PART Of EPIcRANIuM, ANd vERy ShORT TO MINuTE des2 NEAR TO SIdE Of EPIcRANIuM; LONg des3 LOcATEd ANTERIORLy ON EPIcRANIuM cLOSE TO fRONTAL SuTuRE OR ALMOST WIThIN IT ; MINuTE des4 LOcATEd IN ThE cENTRAL PART Of ThE EPIcRANIuM NEAR TO ONE SENSILLuM ; LONg des5 LOcATEd ANTEROLATERALLy ( FIg. 1 ). Fs 1 ANd fs3 AbSENT; fs2 MINuTE ANd LOcATEd MEdIALLy; LONg fs4 LOcATEd ANTEROLATERALLy; ANd vERy ShORT TO MINuTE fs5 LOcATEd LATERALLy, cLOSE TO fs4 ( FIg. 1 ). Les 1 AS LONg AS des5, ANd les2 vERy ShORT; ves1–2 ShORT. EPIcRANIAL AREA WITh MORE POSTEPIcRANIAL SETAE ANd SENSILLA (TOgEThER 7).

Antennae LOcATEd AT ThE ENd Of ThE fRONTAL SuTuRE ON EAch SIdE, MEMbRANOuS ANd SLIghTLy cONvEX bASAL ARTIcLE bEARINg ONE cONIcAL SENSORIuM, RELATIvELy LONg; bASAL MEMbRANOuS ARTIcLE WITh 3 SENSILLA ThAT dIffER IN LENgTh ANd 2 PORES ( FIg. 3 ).

Clypeus ( FIg. 4 ) APPROXIMATELy 3 TIMES AS WIdE AS LONg WITh 2 vERy ShORT TO MINuTE cls, LOcALIzEd POSTEROLATERALLy, ANd 1 SENSILLuM; ANTERIOR MARgIN cONcAvE.

Mouth parts. LAbRuM ( FIg. 4 ) APPROXIMATELy 2 TIMES AS WIdE AS LONg, WITh 3 PAIRS Of PILIfORM lms Of dIffERENT LENgThS; lms2 LONg, SLIghTLy EXcEEdS ANTERIOR MARgIN Of LAbRuM; lms1 ANd lms3 ALMOST EquAL IN LENgTh, bOTh dISTINcTLy ShORTER ThAN lms2; lms1 LOcATEd cLOSE TO ThE MARgIN Of ThE cLyPEuS, lms2 LOcATEd ANTEROMEdIALLy ANd lms3 LOcATEd ANTEROLATERALLy; ANTERIOR MARgIN dOubLE SINuATE. EPIPhARyNX ( FIg. 5 ) WITh 3 fINgER-LIkE als ThAT dIffER IN LENgTh; WITh 3 ShORT ams: ams1 fINgER-LIkE ANd ENLARgEd IN MIddLE, ams2 ANd ams3 PILIfROM ANd RELATIvELy ELONgATEd; WITh 2 RELATIvELy ShORT ANd fINgER-LIkE mes; LAbRAL ROdS (LR) ELONgATEd, APIcAL PART MORE ScLEROTIzEd, ANd bASAL PART SLIghTLy ENLARgEd. MANdIbLES ( FIg. 2 ) dISTINcTLy bROAd, bIfId, WITh TEETh Of uNEquAL hEIghT; SLIghTLy TRuNcATE; bOTh mds RELATIvELy LONg, PILIfORM, ALIgNEd LONgITudINALLy ANd LOcATEd IN dISTINcT hOLES. MAXILLA ( FIg. 6 ) STIPES WITh 1 stps, 2 pfs ANd 1 mbs, WITh ONE SENSILLuM cLOSE TO mbs; stps ANd pfs1–2 RELATIvELy LONg, mbs ShORT; MALA WITh 5 PILIfORM dms; 3 vERy ShORT TO MINuTE vms, ALMOST EquAL IN LENgTh; vms dISTINcTLy ShORTER ThAN dms. MAXILLARy PALPI WITh TWO PALPOMERES; bASAL PALPOMERE WITh 1 ShORT mxps ANd TWO SENSILLA; LENgTh RATIO Of bASAL ANd dISTAL PALPOMERES: 1:0.8; dISTAL PALPOMERE WITh ONE SENSILLuM ANd A gROuP Of cONIcAL, cuTIcuLAR APIcAL PROcESSES. PRAELAbIuM ( FIg. 6 ) OvAL-ShAPEd ANd dISTINcTLy ELONgATEd, WITh 1 RELATIvELy LONg prms; LIguLA WITh SINuATE MARgIN ANd 2 ShORT ligs; PREMENTAL ScLERITE WELL vISIbLE. LAbIAL PALPI WITh TWO PALPOMERES; LENgTh RATIO Of bASAL ANd dISTAL PALPOMERES: 1:0.9; dISTAL PALPOMERE WITh ONE SENSILLuM ANd ShORT, cuTIcuLAR APIcAL PROcESSES; bASAL PALPOMERE WITh 1 dORSAL SENSILLuM. POSTLAbIuM ( FIg. 6 ) WITh 3 pms, ALL pms LOcATEd LATERALLy; pms1 ANd pms3 ShORT, pms2 dISTINcTLy LONgER ThAN OThERS.

Thorax. PROThORAX dISTINcTLy SMALLER ThAN MESO- ANd METAThORAX. SPIRAcLE bIcAMERAL. PROThORAX ( FIg. 8 ) WITh 4 LONg ANd 4–6 vERy ShORT TO MINuTE prns ON SMALL, WEAkLy PIgMENTEd dORSAL ScLERITE; ThIS ScLERITE SubdIvIdEd INTO TWO TRIANguLAR PLATES MEdIALLy; 1 LONg ANd 1 vERy ShORT TO MINuTE ps. MESOThORAX ( FIg. 8 ) WITh 2 LONg ANd 2 vERy ShORT TO MINuTE pds; 1 vERy ShORT as; 1 vERy ShORT ss; 1 LONg eps; 1 LONg ps. ChAETOTAXy Of METAThORAX ( FIg. 8 ) ALMOST IdENTIcAL TO ThAT Of MESOThORAX. EAch PEdAL AREA Of ThORAcIc SEgMENTS WELL SEPARATEd, WITh 1 LONg ANd 3–4 vERy ShORT TO MINuTE pda.

Abdomen. AbdOMINAL SEgMENTS I–VI Of ALMOST EquAL LENgTh, NEXT AbdOMINAL SEgMENTS dEcREASINg gRAduALLy TO ThE TERMINAL PARTS Of ThE bOdy. AbdOMINAL SEgMENT X REducEd TO fOuR ANAL LObES Of uNEquAL SIzE, ThE dORSAL bEINg dISTINcTLy ThE LARgEST, ThE LATERAL PAIR EquAL IN SIzE, ANd ThE vENTRAL LObE vERy SMALL. ANuS LOcATEd TERMINALLy. SPIRAcLES bIcAMERAL, ThE EIghT AbdOMINAL SPIRAcLES LOcATEd LATERALLy, cLOSE TO ThE ANTERIOR MARgIN Of AbdOMINAL SEgMENTS I–VIII. AbdOMINAL SEgMENTS I–VII ( FIgS 9–10 ) WITh 1 vERy ShORT TO MINuTE prs; 2 LONg ANd 3 vERy ShORT TO MINuTE pds (ORdER: LONg, ShORT, ShORT, LONg, ShORT); 1 vERy ShORT TO MINuTE ss; 1 LONg ANd 1 vERy ShORT TO MINuTE eps; 1 LONg ANd 1 vERy ShORT TO MINuTE ps; 1 MINuTE lsts. AbdOMINAL SEgMENT VIII ( FIg. 10 ) WITh 1 LONg ANd 2 vERy ShORT TO MINuTE pds (ORdER: ShORT, LONg, ShORT); 1 LONg ANd 1 vERy ShORT TO MINuTE eps; 1 LONg ANd 1 vERy ShORT TO MINuTE ps; 1 MINuTE lsts. AbdOMINAL SEgMENT IX ( FIg. 10 ) WITh 2 ds (1 ds LONg, 1 ds MINuTE); 2 ps (1 ps LONg, 1 ps MINuTE); ANd 1 vERy ShORT sts. AbdOMINAL SEgMENT X ( FIg. 10 ) WITh 2 MINuTE SETAE (ts).

Description of pupae. Measurements (IN MM). BOdy LENgTh 3.10–4.50; WIdEST PART 1.30–2.50 (SEE TAbLE 3). Colouration. BOdy WhITE OR LIghT yELLOW. CuTIcLE SMOOTh.

SPECIES PUPA

HW (P) BL (P) BW (P) MED MIN. MAX. MED MIN. MAX. MED MIN. MAX. L. bischoffi ³ 0.65 0.55 0.75 3.50 3.10 4.00 2.20 1.55 2.50 L. bischoffi ♀ 0.77 0.60 0.80 3.90 3.30 4.50 2.20 1.30 2.50 Morphology ( FIgS 11–13 ). BOdy MOdERATELy ELONgATE ANd RAThER SLENdER. ROSTRuM LONg (APPROXIMATELy 6 TIMES AS LONg AS WIdE). ANTENNAE RELATIvELy LONg ANd SLENdER; cLubS cOvEREd by cONIcAL PROTubERANcES. PRONOTuM 2.0 TIMES AS WIdE AS LONg. AbdOMINAL SEgMENTS I–VII Of EquAL LENgTh, AbdOMINAL SEgMENT VIII NARROW, AbdOMINAL SEgMENT IX dISTINcTLy SMALLER ThAN OThERS. UROgOMPhI cONIcAL, RELATIvELy ELONgATEd, WITh dARk PIgMENTEd APIcAL PART, SLIghTLy cuRvEd TO INSIdE.

IdENTIfIcATION Of ThE SEX Of PuPAE IS vERy dIffIcuLT IN ThE STudIEd SPEcIES. ThE dIffERENcES IN SIzE Of bOdy bETWEEN MALES ANd fEMALES WERE RAThER POORLy ObSERvEd. ThE ShAPE Of ThE gONOThEcAE WAS TyPIcAL Of ThE PuPAE Of WEEvILS (bILObEd IN fEMALES ANd uNdIvIdEd IN MALES) buT WAS ALSO WEAkLy vISIbLE IN SOME SPEcIMENS.

Chaetotaxy ( FIgS 11–13 ). SETAE dISTINcT, vERy LONg, yELLOW TO LIghT bROWN; LOcATEd ON cONIcAL PROTubERANcES ON PRONOTuM, hEAd ANd ROSTRuM; STRAIghT, hAIR-LIkE, SLIghTLy cuRvEd. HEAd cAPSuLE WITh 1 vs, 1 os, 1 sos ANd A PAIR Of vERy ShORT, POORLy ObSERvEd pas. ROSTRuM WITh 1 rs LOcATEd MEdIALLy. PRONOTuM WITh 2 as, 2 ls, 2 ds, ANd 3 pls. DORSAL PARTS Of MESOThORAX WITh 2 SETAE PLAcEd MEdIALLy. ChAETOTAXy Of METAThORAX IdENTIcAL TO ThAT Of MESOThORAX. EAch APEX Of fEMORA WITh 2 fes Of EquAL LENgTh. DORSAL PARTS Of EAch AbdOMINAL SEgMENTS I–VIII WITh 2 vERy LONg SETAE. ALL AbdOMINAL SETAE LOcATEd ON SMALL cONIcAL PROTubERANcES; vENTRAL PARTS Of AbdOMINAL SEgMENTS I–VIII WIThOuT SETAE. AbdOMINAL SEgMENTS I–VIII EAch WITh A PAIR Of MIcROSETAE PLAcEd LATERALLy. AbdOMINAL SEgMENT IX WITh A PAIR Of dORSAL SETAE, ANd NEXT PAIR LOcATEd ON uROgOMPhI.

Biological observations. FEEdINg ANd dEvELOPMENT Of L. bischoffi WAS ObSERvEd ON Fraxinus pennsylvanica (gREEN ASh OR REd ASh) ( FIg. 16 ) ANd F. excelsior (cOMMON ASh) ( OLEAcEAE ). ThERE WAS A hugE NuMbER Of F. pennsylvanica INhAbITEd by L. bischoffi ANd ONLy ONE EXAMPLE Of F. excelsior IN ThE AREA Of STudy. ThE WEEvILS ARE cOMMON IN ANThROPOgENIc bIOTOPES (PARkS ANd ALLEyS) ANd ESPEcIALLy INhAbIT TREES WELL EXPOSEd TO ThE SuN ANd SuRROuNdEd by gRASSy AREAS ( FIgS 14–15 ). WE REALIzEd ThAT gREEN ASh IS ThE MOST PREfERREd TREE AS hOST fOR L. bischoffi LARvAE. OvIPOSITION WAS ObSERvEd fROM ThE MIddLE Of SEPTEMbER uNTIL ThE bEgINNINg Of OcTObER ( FIgS 15, 17 ). MORE ThAN 80% Of ThE SEEdS Of F. pennsylvanica WERE INfEcTEd by LARvAE Of L. bischoffi . ThE LEvEL Of INfESTATION Of F. excelsior WAS dISTINcTLy LOWER ANd REAchEd 10%. ONLy ONE Egg PER SEEd WAS fOuNd. IN ThE bEgINNINg Of NOvEMbER, MATuRE LARvAE LEfT ThE SEEdS ( FIgS 18–19 ) ANd ENTEREd ThE SOIL. SOME Of ThEM LEfT ThE SEEdS AfTER ThEy fELL TO ThE gROuNd. ThE LARvAE OvERWINTEREd IN ThE SOIL (cA 5 cM dEEP) WIThOuT buILdINg A SPEcIAL chAMbER. NO LARvAE OvERWINTERINg INSIdE ThE SEEdS WERE ObSERvEd. PuPATION PROcEEdEd AT ThE ENd Of APRIL. AduLTS WERE fOuNd IN ThE SOIL 10 dAyS LATER, buT ThEy dId NOT LEAvE ThE SOIL IMMEdIATELy AfTER METAMORPhOSIS. ThE LARvAE REAREd AT A cONSTANT 20°C, dId NOT START PuPATION EARLIER ThAN ThOSE REAREd AT OuTdOOR TEMPERATuRE. ALMOST 60% Of LARvAE dIEd AS A RESuLT Of INfEcTION by fuNgI. ALL LARvAE ThAT dId NOT PuPATE dIEd.

FROM ThE SEEdS Of cOMMON ASh INhAbITEd by ThE LARvAE Of L. bischoffi , WE cOLLEcTEd SEvEN SPEcIMENS Of AduLT Psilocera crassispina (ThOMSON, 1878) ( HyMENOPTERA : ChALcIdOIdEA: PTEROMALIdAE ), WhIch hAvE bEEN NOTEd TO bE hyPERPARASITOIdS ON PARASITES Of IMMATuRE STAgES Of WEEvILS ( DINIz & MORAIS 1996), WhEREAS ONE AduLT Of Collyria SP. ( HyMENOPTERA : IchNEuMONIdAE ) WAS fOuNd IN ThE SEEdS Of gREEN ASh. ThE gENuS Collyria SchIødTE, 1839 INcLudES PARASITE SPEcIES ThAT MOSTLy ATTAck WhEAT STEM SAWfLIES, SyMPhyTA ( HyMENOPTERA ), uSuALLy Of ThE gENuS Cephus LATREILLE, 1802. ThE IMMATuRE STAgES Of PARASITES cOLLEcTEd duRINg OuR STudy REMAIN uNIdENTIfIEd.

Remarks. ThIS IS A NATIvE NORTh AMERIcAN WEEvIL ANd AN INvASIvE SPEcIES IN EuROPE. IT WAS REPORTEd fOR ThE fIRST TIME IN SLOvAkIA IN 1960 uNdER ThE NEW NAME Lignyodes slovacicus DIEckMANN, 1970 (DIEckMANN 1970) ANd SyNONyMISEd AfTERWARdS WITh L. bischoffi (DIEckMANN 1974). AfTER SLOvAkIA, L. bischoffi WAS SubSEquENTLy REPORTEd IN AuSTRIA, HuNgARy, MOLdAvIA ANd SOuTh UkRAINE ( LuchT 1987; POdLuSSANy 1996; POIRAS 1991, 1998) ANd IN POLANd ( GOSIk et al. 2001). POIRAS (1991) REPORTEd ThAT MORE ThAN 50% Of SEEdS WERE INfEcTEd by ThE LARvAE Of L. bischoffi . POIRAS (1991) REPORTEd ThAT APPROXIMATELy 5% Of ThE LARvAE dId NOT PuPATE AfTER WINTER ANd REMAINEd IN ThE SOIL TO OvERWINTER AgAIN. IN cONTRAST, WE ObSERvEd ThAT ALL LARvAE ThAT dId NOT PuPATE dIEd. DEvELOPMENT Of L. bischoffi WAS ObSERvEd IN NORTh AMERIcA ON SEEdS Of Fraxinus americana L. ( BARgER & DAvIdSON 1967), ANd ThE SPEcIES ALSO fEd ON F. nigra L. ANd F. pennsylvanica ( CLARk 1980). IN EuROPE, Fraxinus excelsior , L., F. ornus L., F. lanceolata BORkh. ANd F. oxycarpa WILL. hAvE bEEN REPORTEd TO bE hOST PLANTS Of L. bischoffi (DIEckMANN 1974; POIRAS 1991). ThE LIfE cycLE ANd dEvELOPMENTAL bEhAvIOuR Of L. bischoffi ObSERvEd IN POLANd WERE bASIcALLy SIMILAR TO ThOSE REPORTEd IN NORTh AMERIcA ( CLARk 1980) ANd MOLdAvIA ( POIRAS 1991), EXcEPT fOR ThE INfORMATION REgARdINg LARvAE fEEdINg ONLy ON F. pennsylvanica OR F. excelsior ANd OvERWINTERINg INSIdE SEEdS.

Figures

FIGURE 7. Lignyodes bischoffi MATURE lARVA, HABITUS, lATERAl VIEW. SCAlE BAR: 1 MM.

FIGURES 14 – 19. 14 — ADUlTS OF Lignyodes bischoffi IN COPUlA ON SEED OF GREEN ASH; 15 — FEMAlE OF Lignyodes bischoffi BORING HOlE FOR OVIPOSITION; 16 — lEAFS OF GREEN ASH WITH TRACkS OF FEEDING OF L. bischoffi; 17 — SEED OF GREEN ASH WITH SPOOR OF OVIPOSITION; 18 – 19 — SEEDS OF GREEN ASH WITH lARVAE OF L. bischoffi.

FIGURES 1 – 6. Lignyodes bischoffi MATURE lARVA: 1 – HEAD, DORSAl VIEW (des – DORSAl EPICRANIAl S., fs – FRONTAl EPICRANIAl S., les – lATERAl EPICRANIAl S., AT – ANTENNA); 2 – RIGHT MANDIBlE (mds – MANDIBlE DORSAl S.); 3 – ANTENNA; 4 – lABRUM AND ClYPEUS (lms – lABRAl S., cls – ClYPEAl S.); 5 – EPIPHARYNX (ams – ANTEROMEDIAl S., als – ANTERIOlATERAl S., mes – MEDIAN S., lR – lABRAl RODS). 6 – MAXIllOlABIAl COMPlEX, VENTRAl VIEW: RIGHT MAXIllA (dms – DORSAl MAlAR S., vms – VENTRAl MAlAR S., mpxs – MAXIllARY PAlPS S., mbs – BASIOVENTRAl S., pfs – PAlPIFERAl S., stps – STIPITAl S.), PREMENTUM AND POSTMENTUM, VENTRAl VIEW (prms – PREMENTAl S., pms – POSTMENTAl S., ligs – lIGUlAR S.). SCAlES BARS: 0.5 MM (1), 0.2 MM (2, 4 – 6), AND 0.1 MM (3).

FIGURES 8 – 10. Lignyodes bischoffi MATURE lARVA, HABITUS, 8 — lATERAl VIEW OF THORACIC SEGMENTS, 9 — lATERAl VIEW OF ABDOMINAl SEGMENT I, 10 — lATERAl VIEW OF ABDOMINAl SEGMENTS VII – X (prns — PRONOTAl S., prs — PRODORSAl S., pds — POSTDORSAl S., as — AlAR S., ss — SPIRACUlAR S., eps — EPIPlEURAl S., ps — PlEURAl S., pda — PEDAl S., lsts — lATEROSTERNAl S., eus — EUSTERNAl S., ds — DORSAl S., sts — STERNAl S., TH 1 – 3 — NUMBER OF THORACIC SEGMENTS, AB 1 – 10 — NUMBER OF ABDOMINAl SEG.). SCAlE BAR: 1 MM.

FIGURES 11 – 13. Lignyodes bischoffi PUPA HABITUS: 11 — VENTRAl VIEW; 12 — DORSAl VIEW; 13 — lATERAl VIEW (SETAE: as — APICAl S., ds — DISCAl S., ls — lATERAl S., os — ORBITAl S., pls — POSTEROlATERAl S., pas — POSTANTENNAl S., rs — ROSTRAl S., sos — SUPERORBITAl S., vs — VERTICAl S.; UR—UROGOMPHI). SCAlE BAR: 1 MM.