Generic relationships within the tribes Cratocentrini and Phasgonophorini (Hymenoptera: Chalcididae)

39 Pages·1997·11.2 MB·English

by  G A W Wijesekara

J.HYM.RES.
Vol.6(2),1997,pp.297-335
Generic RelationshipsWithin the Tribes Cratocentrini and
Phasgonophorini (Hymenoptera: Chalcididae)
G.A.W.WlJESEKARA
MarylandCenterforSystematicEntomology,DepartmentofEntomology,Universityof
Maryland,CollegePark,Maryland,20740[Currentaddress.DepartmentofEntomology,P.O.
Box11,HorticulttiralResearchandDevelopmentInstitute,Peradeniya,SriLanka]
—
Abstract. According tomost recentclassification, thesubfamilyChalcidinaeisdivided into
fourtribes:Brachymeriini,Chalcidini,Cratocentrini,andPhasgonophorini.Ithasbeensuggested
thatthetribesPhasgonophoriniandCratocentriniaresistergroupsandtogethertheyformthe
sistergrouptoamonophyleticgroupconsistingofBrachymeriiniandChalcidini.Acladisticstudy
wasconductedtotesttherelationshipbetweenCratocentriniandPhasgonophoriniandtoestab-
lishthegenericrelationships,usingallknowntaxaineachtribe.Brnchi/meria,Chalcis,andDirhinus
wereusedasoutgroups.ParsimonyanalysisusingthebranchandboundsearchoptionofPAUP
(Ver. 3.1.1), resulted in 14minimum lengthtrees. Noneofthe treescouldberooted tomake
CratocentriniandPhasgonophorinisistergroups.Reanalysisofdataaftersuccessiveweightingof
charactersresultedinasinglemostparsimonioustreethatisidenticaltooneoftheoriginal14
trees.Thistreewasselectedasthepreferredhypothesis.Theseresultscorroboratetherelation-
shipsfortribesinChalcidinaesuggestedfrommypreviousanalysisofchalcididphylogeny.In
additionthisstudyestablishedthegenericrelationshipswithinCratocentriniandPhasgonophorini
forthefirsttime.TheresultssuggestthatMegachakisandTrigoniirellaarethebasallineagesof
tribestheCratocentriniandPhasgonophorini,respectively.Becausethesetwogenerawerenot
usedtorepresentthetribesinmypreviouscladisticticanalysis,theeffectoftaxonsamplingon
my chalcidid phylogeny was tested byincludingthesetwotaxa intheanalysis.Thisanalysis
showednoeffectoftaxonsamplingonpreviousphylogeneticanalysis.Sincethisanalysisuseda
separatesetofcharactersthanthefamilylevelphylogenyanalysis,thetwodatasetswerecom-
binedandresultingdatamatrixof41taxaand65characterswasanalyzed.Thisanalysisresulted
in 14minimumlengthtreesandsuccessiveweightinggave10minimumlengthtreesonestep
longer than any ofthe 14 minimum length trees from equally weighted data. There issome
disagreementbetweenthephylogenetichypothesisresultingfromthisanalysisforChalcididae
andresultsofmypreviousanalysisoffamilyphylogeny.Thisillustratestheeffectofbiasedtaxon
andcharactersamplingontheresultsofphylogeneticanalysis.
Cratocentrini and Phasgonophorini which isextenciedbeyond theshortsyn-
(Hymenoptera:Chalcididae)aretwomor- tergum(Fig.56).Althoughthemajorityof
phologically distinct groups within the Phasgonophorinialsohavealongovipos-
subfamily Chalcidinae. Host records itor, thesyntergumhasextendedconceal-
(available only for few species) indicate ingtheelongatedovipositorsheaths(Figs,
thatthespeciesofbothtribesareparasites 55, 57). Both groups were extensively
ofwoodboringbeetlelarvae,anunusual studied by Steffan (1950a, 1950b, 1956,
host asssociation in Chalcididae (Boucek 1959,and1973).Atpresenteachtribecon-
1988). Theyareamongthelargestchalci- sistsofeightgenera,alldistributedinthe
dids,varyingfrom4to20mm.inlength, tropicalareasoftheworld.
Cratocentrini are distinctive among chal- Steffan (1959)suggested thatCratocen-
cidids in having an elongated ovipositor trini and Phasgonophorini evolved from
298 JournalofHymenopteraResearch
the same "stem". He hypothesized that with abdominal tergite I reduced and
their specialized abdomens were conver- shorterthantergiteII (Megahcolus,Paras-
gently evolved as a result of ovipositing typiura, Stypiura,andStenochalcis).Steffan
intosimilartypesofhosts. Boucek(1988) (1956)impliedthathistribeswerenatural
stated that these tw^o tribes were sister groupsandgavemanysynonymsforspe-
groupsandformedasistercladetoChal- cies of Phasgonophora and Cratocentrus.
cidini+Brachymeriinibuthedidnotpro- Steffan(1973)alsorevisedthegeneraStyp-
videanycharacterevidence. iura (six species) and Parastypiura (three
Mypreviousstudyofhigherlevelphy- species) of the Neotropical region. Since
logeny of Chalcididae (Wijesekara 1997) Steffan's work, two more genera have
indicatedthatCratocentriniandPhasgon- beenaddedtothetribe:KopinataandTri-
ophoriniarenotcloselyrelated.Phasgon- gonurella (Boucek 1988). AtpresentPhas-
ophoriniisthesistergrouptoBrachimer- gonophorini consistsofeightgeneraand
iini,andthesetwotribesplusHaltichelli- 57species.
nae,Dirhininae,andEpitraninaeformthe
sister group to Cratocentrini (Fig. 74). Cratocentrini
However my study of family phylogeny The genera Larradomorpha Stadelmann
usedonlytwoexamplarsofCratocentrini 1792, Marres Walker 1841, and Acantho-
and Phasgonophorini. Both tribes consist chalcis Cameron 1884 were originally
of eight genera, and it therefore seemed placed in Leucospidae, and Cratocentrus
that a study of generic relationships Cameron 1907 was originally placed in
amongallknowngeneraofCratocentrini Haltichellinae. Masi (1944) subsequently
andPhasgonophoriniwouldbeusefulnot referred these genera to Brachymeriinae.
only to establish generic relationships Later,Cratocentriniwasestablishedtoin-
within the tribesbut also to testthe op- cludethesegeneraplusMacrochalcisMasi
posinghypothesesregardingtherelation- 1945 (=AUocentrus Cameron 1911); and
shipbetweenthesetwotribes. MegachalcisCameron1903(Steffan1950b).
TAXONOMICHISTORY TwhheotardidbeedwafsourrenveiwsedgenbeyraS,teSfpfaatnoce(n1t9r5u9s),,
Many taxa included in Cratocentrini Philocentrus,Acrocentrus,andVespomorpha.
and Phasgonophorini were described in Narendran (1984) synonymized Allocen-
thenineteenthcenturyandclassifiedwith- trus with Megachalcis, and Boucek (1992)
indifferenttaxonomicgroups.Masi(1944) synonymized Larradomorpha with Marres.
placedthemallwithinthesubfamilyBra- Atpresentthetribeconsistsofeightgen-
chymeriinae.Bothtribeswereestablished eraand23species.
bySteffan(1950a, 1950b). MATERIALANDMETHODS
—
Phasgonophorini Specimens. Thisstudywascarriedout
Steffan(1950b)establishedthePhasgon- using the collection at the United States
ophorini to include Phasgonophora West- National Museum of Natural History,
wood1832,TrigoiiuraSichel1865,Sti/piurci Washington D. C. Additional specimens
Kirby 1883, Megalocolus Kirby 1883, and were borrowed from the following insti-
Stenochalcis Masi 1929. He also described tutions: The Natural History Museum,
a new genus, ParastypiurnwithinBrachy- London;UniversityofCalicv:t, Kerala,In-
meriinae.Hesuggestedthatthetribecon- dia;South AfricanMuseum, CapeTown,
sisted oftwogroupsofgenera: thePhas- SouthAfrica;PlantProtectionResearchIn-
gonophoragroup,withabdominaltergite1 stitute, Pretoria, South Africa; and Muse-
larger than the tergite II (Phasgonophora um NationaldeHistoireNaturelle, Paris,
and Trigonura), and the Stypiura group. France.
Volume6,Number2,1997 299
—
Taxa Used mid Character Selection. All (reversals)ofcharactersoverparallelevo-
valid genera of Cratocentrini and Phas- lution of characters (convergence), was
gonophorini were included in this study usedtooptimizethecharacterstates.Six-
(Appendix1).Inaddition,representatives teeningrouptaxaandthreeoutgrouptaxa
ofothermajorcladesofChalcididaeiden- werecodedfor40characters.Thecharac-
tifiedinmyearlierfamilylevelstudy(Wi- ter optimization was studied using
jesekara 1997) were also included. These MacClade version 3.0 (Maddison and
taxaservedasoutgroupsforassessingre- Maddison1992). —
alantdioPnhsahsipgsonwoipthhoirnintih.etArirbeesprCersaetnotcaetnitvreinoif meSatsaubirleistyofAntarleyesisst.abilTihtyerfeoraprheylodigfefneerteinct
BrachymeriiniwasincludedbecauseCra- hypotheses. These include the branch
tocentrini + Phasgonophorini have been lengths (Bremer 1994), Bremer support
included in Brachymeriinae and Brachy- (decayindex),andbootstrapvalues.Boot-
meriinigroupedwithPhasgonophoriniin strapping (Felsenstein 1985) and Bremer
my previousanalysis.Sincemyprevious support(Farrisetal.1994)provideabetter
analysis of family phylogeny indicated measure oftree stability than thebranch
that Brachymeriini plusPhasgonophorini length(Bremer1994).
formthesistergrouptoacladeconsisting Thedegreeofcharactersupportforvar-
ofHaltichellinae, DirhininaeandEpitran- ious nodes of the phylogenetic tree was
inae, I also included Dirhimis as an out- evaluated using bootstrap analysis (Fel-
group to represent this sister clade. The senstein 1985) and rescaled branch sup-
tree wasrooted by including a represen- port (decay) index (Bremer 1988, 1994).
tative of Chalcidini (Chalcis), which the AutoDecay version 3.0 (Eriksson 1995)
previousanalysissuggestedtobethemost wasusedtocalculatethedecayindices.—
basallineageinthefamily. Abbreviations Used in Figures 1-65.
Comparative morphology was studied DAS:dorsalaxillarysurface,FRD:foram-
for as many species as possible for each inal depression, FRN: frenum, HC: hind
genus(SeelistofCratocentriniandPhas- coxa,HF:hindfemur,HS:horizontallydi-
gonophorinispeciesstudied.Appendix1). rectedspur,HT:hindtibia,HTS:hindtib-
Characters that varied among thegenera ial spur, LBR: labrum, LS: lateral sulcus,
wereselected fortheanalysis.Characters LM: lamella, MAN: mandible, MEP: me-
thatprovedtobesynapomorphiesforthe sosternal process, MES: mesepisternum,
relevanttribesinmypreviousstudywere MNS: mesonotal spiracle, MS: malar
also included. The generic autapomor- sulcus, MV: marginal vein, OCE: ocellus,
phieswerenotinclud—edintheanalysis. OS: outerspur, PET: petiole, PMV: post-
Character Analysis. Cladistic analysis marginalvein,PRE:prepectus,PRM:pos-
wasperformedusing"PhylogeneticAnal- terior pronotal margin, PRN: pronotum,
ysis Using Parsimony" (PAUP), version PRO: propodeum, SCA: supraclypeal
3.1.1 (Swofford 1993). The branch and area,SCR:scrobe,SMV:submarginalvein,
bound search option, which guarantees STV: stigmal vein, TD: tarsal depression,
finding all shortest possible trees, was TEG: tegula, TOR: torulus, TR: trocanter,
used. All the multistate characters were TV: tergiteV, TVIl: tergiteVII, VER:ver-
treated asunordered (non-additive;Fitch tex.
1971). Although there are many criteria
thatcanbeusedtoordercharactersIpre- RESULTSANDDISCUSSION
fernottoassumetheorderingofthestates CharactersSelected
in multistate characters prior to cladistic
analysis (Wijesekara 1997). ACCTRAN Forty morphological characters were
optimization,whichfavorssecondaryloss scoredforatotalof19taxa(Table1).Eight
300 JournalofHymenopteraResearch
Table1. DatamatrixforPhasgonophoriniandCratocentrini.
Characters
1 1111111112 2222222223 3333333334
1234567890 1234567890 1234567890 1234567890
Chalcis 0011011000 0000000000 0001000100 2000010000
Brachymeria 0021013000 0000102000 0010102001 0010010000
Dirhinus 0001110000 0101101000 1010110100 0000010000
Acanthochalcis 1221112001 1101010001 2101020112 1110002020
Megachakis 1221112000 1121010101 2100020112 1110012000
Cratocentrus 1221112020 1102010011 2111020112 1110002020
Marres 1221112000 1102010110 2101010112 1110002030
Vesponwrpha 1221112000 1102010010 2101010112 1110002040
Spatocentrus 1221112001 1102010110 2111010112 1110012010
Philocentrus 1221112001 1102010010 2111020112 11100??0?0
Acrocentrus 1221112020 1102010011 2111010112 1110012010
Phasgonophora 2000000011 1111102010 3200100000 OHIO11000
Trigoiiura 2000000001 1111102010 3200100000 OHIO11000
TrigoinireUa 2100013000 1101102000 3210100000 0010010000
Stypiura 20100OHIO 1102102010 3200101000 0020111101
Megalocolus 20000OHIO 1101102010 3200101000 0010211101
Stenochttlcis 2100011110 1101102000 3200101000 0020213100
Kopinnta 2100001100 1101102010 3200101000 0010111001
Parnfti/piiirn 2000001111 1101102010 3200101000 0011011001
ofthecharacterswerealsousedinthepre- 2. Supraclypealarea.
viousanalysisoffamilyphylogeny.Those 0. Without a modified bridge with
charactersareindicatedbyanasterisk(*) same sculpture as rest of the face
followingthecharacternumber.Thechar- (Fig.3).
actersaredescribedbelow and measures 1. Modifiedtoformabridge,i.e.,with
offitforeachcharacteraregivenintable differentsculpturethan restofthe
2. face(Fig.4).
CharacteristicsoftheHead: 2. Tgoirnuloifltohceatceldypaetustheanadntetrhieorsumparra--
1. S0.izLeaorfget,heaslbarboraumd.asbaseofmandible In Phlayspegaolnoaprheoariisnriedthueceadnt(eFnign.al2).toruli
1. (HFailgf. 1a)s. long as base of mandible Tarheelaorceaatebdeatwwaeeynftrhoemctlhyepecallypmeaalrgmianrgainnd.
2. (LFeisgs.2t)h.anhalfasbroad asbase of antennal toruli is sometimes modified
conBdoitthimoantnroidfbieabsnleheax(vpFeiogs.teh3d)e.antydpiccoanlticghuaolucsidliad- tfmoworesmetinsnpgtehcaeiberssictdrhgoeberaeolfisbdaiafsfeselriaegnnhttdsicncudlliypcptaeuturiseo.nbeIo-nf
brum.Phasgonophorinihasasmalllabrum this bridge-like structure, but the area is
ewrhelraebarsumC,rationcteenrtmreidniiahtaesianresliazteivbeelytwlearegn- mthoerefacoerhlaesvsincgonstainmueoussculwpittuhret.heInressotmoef
Brachymeriini and Phasgonophorini. Al- other groups this is distinctly differenti-
though the structure of the labrum has ated. Cratocentrini differs in having the
been used previously forphylogeneticin- toruli located at the clypeal margin and
ferencewithinChalcidoidea(Darling1988), hence having very reduced supraclypeal
thesizeofthelabrumhasnot. area.
Volume6,Number2,1997 301
3. Malarregion. funiclesegmentslenderandlongandthe
201... WAAitscahurloicunutasapprrseeussleecnnutts((oFFriiggc..a6r5))i..na(Fig.3). Sogtrteohsuestrivsaeenlgdymeswhnoitrdstednaiinsnttigenncnttaloywasarhreodrcsthearrtahacetnedraippsertxoi.-c
Inmostchalcididsthemalarregionhas ofmost Cratocentrini, Brachymeriini and
adistinctsulcusrunningfromtheventral Dirhininae.
marginoftheeyetobaseofthemandible.
ThesulcusisabsentfrommostPhasgon- 7. Antennalscrobe.
ophorini, present as a sulcus in Styphira 0. Morethan2xaslongasbroadnear
and Chalcis, and indicated as a carina in toruliandparallelsided(Fig.7).
Brachymeriini.InCratocentriniitistrace- 1. Lessthan2x aslongasbroadnear
ableandcarina-likeasinBrachymeriini. toruliandalmostpear-shapedwith
abluntlateralmargin(Fig.4).
4. Face. 2. Morethan2xaslongasbroadnear
Mo01.s.tCDiocshntacilnaccvitdeliydosrcohfnalvavteex(Fai(ngF.iagl8.)m.7o)s.tconcave 3. tMfolorarunlgeie-talhniakdnep2leaxtaerar-sasllhomanapgregdaiswnbir(toFhiagds.hna9)er.ap,r
facebecauseoftheverylargescrobalde- toruli and pear-shaped with a
epyreessmiaorngine.xtIenndmionsgtfCrhaolmciedyienaematrhgeisncrot-o sharp, smooth lateral margin (Fig.
bes are smaller and the face is flat or 8).
slightly convex. In Phasgonophorini the Antennalscrobesaregenerallywellde-
faceisdistinctlyconvexbecauseitbulges fined in Chalcididae. In Cratocentrini,
forv^ardfromthevertexandeyemargin. Phasgonophorini, andBrachymeriinithey
5*.0.LoAcabtoivoentofhethleevaelntoefnnlaolwetorrueliy.emar- iafsrroepnrsd.oebeTaphbeelrystcdhruoaebnetsionafrooetrhlweoarnrgdcehraglriconiwdCitrdhast.oofcTehtnih-se
1. gBienlo(wFigt.he4).level oflowereye mar- ttrointihebecclayupseealthmeartgoriunlibaerleowloctahteedlevcellosoefr
oetptmofehyarnoerCutnrgrlhaiaimielntnaio.rlglctiogooehncwrinaauentvl(trriFeeiiwdneghlt.iyiohjelc9euc)aes.omtttnaeoasrdrbiugfeslijetilunewso.nltwtosMlcbpyoateetshclheeitodaewsvPlaehotbhawhaotesevvhrgeeeoleneattvoynhhe-e-eel tmmIsgtolnuhaarielIrrugatngglhlrhitiiooslnnowsyoareemraoriepefnnseetpdtaolyPehorechtecit-ahaehmssetsseahgecrasordcgtpcnoahirebovenaodapi.bblthfaoayorlvpcoraepngicvseiani,avvttiiriyhln,taetagynthiceelsiriestoaailw(nsaneFfhttriloetogerrn.ctienttea4yahern)elde.-.
6. Antenna. In Phasgonophorini the scrobe cavity is
0. Slenderand longwith funicleseg- not margined sharply but in Brachymeria
mentslongerthanbroad(Fig.60). and Cratocentrini itissharply margined.
1. Stout and short with only firstfu- The condition in Brachymeria isdistinctly
nicle segment longer than broad different from that of Cratocentrini. The
(Fig.61). sharp margin of Brachymeria is smooth
Slenderandlongantennaarecharacter- andshinyand resemblestheconditionin
isticofmostPhasgonophorini.Thefunic- Phasgonophorini. The scrobal margin of
ular segments are elongated and of the Cratocentrini is more flange-like and the
same width although slightly decreasing toruli are located relatively furtherapart
in length towards the apex. By contrast, thaninBrachymeriiniorPhasgonophorini
species with stout antenna have the first (Fig.9).
302 JournalofHymenopteraResearch
i
sFpi.g,s.fr1o-n8t.vi1e-3w.oLfofwaecer.f5r-o6n.tMvaileawrosiulfcaucse..15..CChliiinllccitiissspp..26..CBrrataocchcyiimtcnrimssspp..37.-8/'./m.F<a,cv;ei."'"7;.'''Pih""is^gI'oluootpo-ho4n.iStt\^/u(l'iaii(trai.i
8.Brnchymerinsp.
Volume6,Number2,1997 303
Table2. Characterdiagnostics.
Char-
acter
304 JournalofHymenjopteraResearch
Volume6,Number2,1997 305
1. Pronotal width less than 2.5x the 1. Reducedtoaverythinorknob-like
medianlength(Fig. 15). sclerite(Fig.20).
Both PhasgonophoriniandCratocentri- The chalcidid prepectus is a semi-an-
ni have a relatively broad pronotum. In nular, transverse sclerite. Delvare (1992)
Brachymeriini and Chalcidini the prono- foundthemedianprocessoftheprepectus
tumisdistinctlyshortanterio-posteriorly. to be phylogenetically informative to re-
13. Pronotalsurface. swoiltvheinCChraaltcoicdeinntirinrielaantidonPshhaispsg.onHopohwoervienri
01.. RFlaaitse(dFig.lat1e5r).al to the median line tthheemlaetdeiraalnaprrmoceofsstdheoepsrenpoetcvtaursy.neInasrtetahde
asipvPrpsur^arhrliieoTaoattsanhsnihne2oe.anogdoottgutotaotR(lhepnluiFaanealnoitmairgceepnbssa.rhshoyeau(imsl1tordFloo6hifrhar)dmmgataaa.i.ioeasisclnrfrsierlabiied1pgol6cdeuhwi)oasci.mnfathomcoripraefeA(onCvFasdnentihsetigreh.aab.aeatsluoeSnrcthmf1moih7poeefot)edm-aw.fdhuilecenridihptraapgkhroenbprreewooyoolsninuodirtsosnopoaneeritrsotfsstiumrulaehooeaemmdlerf-t asaCfssmspttitciprruernaclaziaergerdueteretauplregaolteioichciatcClfsianeettshesnoaurtatfhnslatetroairhcdduwisbnaiouitnvstddmsiceaateitrhexprnaeiestdpovsineduhoemesdepptspotproaweoerCmoowrixdpahneriaftthanotstteohplitrsehoirlceancsinnamixalntodltte/tfaliteh-ComayhnCnelrsh.eelihtimasakalCmtTetlommclrhehchnaileaeaasioerdttsrodtgppoieoitllaicidrapdhtaalneaelaottn.eiodeeeerst.jfu--.uaruTllcdrcitniiiheainIhcIkkfedlicne-nee-
TgbehunemeyprsaareoafreloCcrvaaettreoydcenadtiwsratiiynnicft(rFoiinmg.Mteh1g7e)a.cmheaTldhcieiassne. tPihcahselgloinniopthhoeripnlait,eBirsadcihsytmienrcitienxit,erannadllyH.al-
line closer tothe posteriormargin ofthe 16*. Mesothoracicspiracle.
pronotum,betweenthelateralmarginand 0. Covered by postero-lateral margin
medianline. ofpronotum(Fig. 19).
1. Exposed(Fig.20).
14. Posteriormarginofthepronotum. InmostHymenopterathepronotallobe
0. Broadlyconcave(Fig.10). coversthemesothoracicspiraclecomplete-
Th21e.. sMMheeaddpiiaaellollfyytrthreoiupanondsgtueeldrair(oFr(iFgim.ga.1r5g1)8.i)n.ofthe lctyoocv(eenGrtierbdisnoibnyh1at9vh8ee5)p.tohsCetheamrleicosirodtlihadotaereraacleixcmcaesrppgtiirCnarcalo-ef
pronotumofchalcididsvariesfrombroad- thepronotum.
mlyonclonycaovbesetroveemdarcgoinndaittei.onThiesmaosbtrocaodml-y 17*. Relativesizeandshapeofthetegula.
concave posterior margin. This character 0. Linear, scond axillary sclerite cov-
is probably correlated to the degree to ered(Fig.20).
which the head can be directed back- 1. Both ventrally and posteriorly ex-
wards. In mostPhasgonophorini thecur- panded,ovalshape,secondaxillary
vatureissoacutethatitappearsmedially scleriteofthewingcompletelycov-
triangular and in most Cratocentrini the ered.
p15o)s.terior margin is more rounded (Fig. 2. Vlaernytrascllleyriteexpoafndtehed,wisnegconedxpoasxield-
(Fig. 19)
15*. Externally visible region of the pre- In Chalcidini and Cratocentrini the
pectus tegula is not modified, it is pear shaped
0. Relatively large and elongated, andextendsfrom theanterio-lateralmar-
plate-hke(Fig.19). gin of pronotum to posterio-lateral mar-
306 JournalofHymenopteraResearch
2F0i.gs.Me1s7o-t24h.ora1x7-l1a8t.eraPlrovnieowt,um19.(aIr'rloMwfs^oiwnpdhiocartaei^rualiaislecid.a2r0e.asC)n.ito1c7c.iiTtrniigft-vusipn.i2s1p-.22I.H.AIxVimlsl^a'e^caunodplwsmcust»e/lc»lfurtm..1291-.
PhasgoiwiilwrasiiUahi,22.Mcnachalcissp.,23.Plmi^^oiwplumish/oi/ii,lateralviewofthetlwrax(arrowindicate
thesculpture).24.Chalcissp.,mesepisternum.