Trophic roles of scavenger beetles in relation to decomposition stages and seasons

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ww w . r b e n t o m o l o g i a . c o m

REVISTA

BRASILEIRA

DE

Entomologia

AJournalonInsectDiversityandEvolution

Biology,

Ecology

and

Diversity

Trophic

roles

of

scavenger

beetles

in

relation

to

decomposition

stages

and

seasons

Noelia

I. Zanetti

a,b

_,

_Elena

_C.

_Visciarelli

b

_,

_Néstor

_D.

_Centeno

a,∗

a_Laboratorio_de_Entomología_Aplicada_y_Forense,_Departamento_de_Ciencia_y_Técnica,_Universidad_Nacional_de_Quilmes,_Buenos_Aires,_Argentina b_Cátedra_de_{Parasitología}_Clínica,_Departamento_de_Biología,_Bioquímica_y_Farmacia,_Universidad_Nacional_del_Sur,_Buenos_Aires,_Argentina

a

r

t

i

c

l

e

i

n

f

o

Articlehistory:

Received29November2014 Accepted9February2015 Availableonline8April2015 AssociateEditor:RodrigoKruger Keywords: Carrionfauna Ecologicalroles Forensicentomology Insectsuccession

a

b

s

t

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Carcassesrepresentatrophicandreproductiveresourceorshelterforarthropods,whicharea repre-sentativecomponentofthedecompositionprocess.Fourexperiments,oneperseason,wereconducted inasemi-ruralareaofBahíaBlanca,Argentina,tostudythetrophicrolesofcadavericbeetles, evaluat-ingtheabundance,compositionanddominanceduringalldecompositionstagesandseasons.Species ofnecrophagous,necrophilousandomnivoroushabitswerefound.Abundance,compositionand domi-nanceofbeetlesinrelationtotheirtrophicroleschangedaccordingtoseasonsanddecompositionstages. Guildsandpatternsofsuccessionwereestablishedinrelationtothoseperiods.Trophicrolescouldbean indicatorofbeetleassociationswithdecompositionstagesandseasons.

Introduction

Carcasses represent a trophic and reproductive resource or shelterfora varietyofanimalssuchasarthropods,which area representativecomponentofthedecompositionprocess.Leclercq

(1974)proposedanecologicalclassiﬁcationbasedonthetrophic

roles of the cadaveric fauna: necrophages (feed from corpses), necrophiles(predate on or parasitize other arthropods), omni-vores(feedfromcorpsesand predateonotherarthropods)and incidentals (look for refuge or appear at random). The car-rion arthropod community is mainly represented by insects

(Carvalhoet al.,2000).These arecapableof detectingchemical

changes,sotheirappearance inone particularstage of decom-positionisnotlikelytobeaccidental(Maga ˜na,2001;Archerand

Elgar,2003; Dekeirsschieter et al., 2011; VonHoermann et al.,

2011,2012).Seguraet al.(2011) suggestedthat decomposition

stages are a primary factor in determining theinsect commu-nity,becausethespeciesexhibitadaptationsthatallowthemto exploitandusecorpsesunderdifferentconditionsandperiodsof time.

Thedecomposition process and the cadaveric fauna succes-sioncanbeinﬂuencedbycircumstancesthatcadaversgothrough afterdeath and many variables have to be consideredin each situation: climatic conditions, season, geographical region, sun exposure,synanthropy,latitude,substratetype,larvalmigration,

∗ Correspondingauthor.

E-mail:ncenteno@unq.edu.ar(N.D.Centeno).

amongstothers(Greenberg,1971;CattsandGoff,1992;Goff,1992;

Kócarek,2001;Centenoetal.,2002;Forbesetal.,2005;Fariaetal.,

2007;BattánHorensteinandLinhares,2011;Dekeirsschieteretal.,

2011; Matuszewskiet al., 2014).Thus, the studyof these

vari-ablesisimportantfortheestimationofpostmorteminterval(PMI) andotherapplicationsinforensics(Keh,1985;Anderson,2001;

Centenoetal.,2002;Doureletal.,2010;BattánHorenstein and

Linhares,2011; Ortloff et al.,2012; Pers. comm., Zanettiet al.,

2015).

BeetlesareespeciallyusefulinforensiccasesinwhichDiptera havenotbeensampledorwhentheyhavealreadyleftthebody and only beetlescan befound(Centeno et al.,2002).But also, somespeciesofColeopterahavebeenobservedinearlierstages ofdecompositionandtheycanappearindifferentwavesinabody, thus,furtherstudiesonthisorderneedtobeconducted.

Theaimofthisstudywastoanalyzetheabundance,composition anddominanceofcadavericbeetlesperseasonanddecomposition stageinaccordancewiththeirtrophicroles(guilds).

Materialsandmethods

This study is a part of a greater study of decomposition andcadavericsuccession(Pers.comm.,Zanettietal.,2015).The experiments were carried out in a ﬁeld property of the Uni-versidadNacional delSur,locatedat BahíaBlanca(38◦4141S, 62◦1510W),BuenosAiresprovince,Argentina.

Twelvedomesticpigs (SusscrofaL.)weighing15–16kgwere used.Threeofthemwerekilledeachseasonbyastabtotheheart (theusualprocedureofcommercialbutchers)1hbeforeexposure, http://dx.doi.org/10.1016/j.rbe.2015.03.009

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andwerekeptinsideaplasticbagtokeepawayinsectsuntilthe beginningoftheexperiment.TheEthicalCommissionofthe Uni-versidadNacionaldelSurapprovedthisprocedure.Eachcarcass waslaidinsideacagetoprotectitfromscavengervertebrates,and underdirectsunalongtransectsseparated100mfromeachother. Tocomplementthesampling,sixpitfalltrapswereplacedaround eachcageatadistanceof50cmfromthecarcass(experimental traps);twopereachlongsideandonepereachwideside.For con-trolpurposes,othersixpitfalltrapswiththesamespatialpattern wereplacedat15–30mfromthelastcage.

Literature typically refers to decomposition as a process of ﬁvestagesthatarenotnecessarilydiscrete(Bornemissza,1957;

Schoendly and Reid, 1987; Moura et al., 2005) but which can

beused asa guide or referencepoints. We followed the crite-rionestablishedbyCenteno etal.(2002)todeﬁnethestagesof decomposition.Carcasseswerevisiteddailyuntilconcludingthe experiment (severaldays afterthecarcassreachedtheremains stage).Beetles fromon, underand insidethecorpseswere col-lectedmanuallywithentomologicalforceps.Beetlesweretaken fromcadavers(perday)onlyif10%oftheindividuals(this percent-agewasconsideredforeachfamily)wereavailableinordertoavoid anyalterationinthedecompositionand/orsuccessionprocesses duetolossofinsects,andpreservedintoplasticcontainerswith ethanol70%.Somelarvaewererearedforidentiﬁcationofthe spec-imensandcultureswithadultswereestablishedforfutureresearch (Pers.comm.,Zanettietal.,2015).Taxonomicaldeterminationor corroborationofsomespecies/morphospecieswasperformedby somespecialists.Thespecimensweredepositeddry-mountedor in alcohol at Cátedra deZoología de InvertebradosII, Departa-mentodeBiología,BioquímicayFarmacia,UniversidadNacional delSur.

Speciesandmorphospeciesweregroupedinaccordancewith theirtrophicroles.Firstly,weevaluatedthecompositionand abun-danceofeachgroup,calculatingthemeanandSEofeachoneper samplingunit(observationsand/orcaptureoncarcassorcapture insixpitfalltraps)andday,duringeachseasonanddecomposition stage.Secondly,weanalyzedthedominanceofeachgroup.

Results

Beetlesofnecrophagous,necrophilousandomnivoroushabits werefound.Tables1–3showthespecies/morphospeciesthatwere detected.

Table2

Species/morphospeciesofnecrophagoushabits.

Families Species/morphospecies

Dermestidae Dermestesmaculatus(DeGeer,1774) Tenebrionidae Scotobiusclathratus(Guérin-Méneville,1834)

Scotobiusmiliaris(Billberg,1815)

Scotobiusmuricatus(Guérin-Méneville,1834) Scarabaeidae Aphodiusmilitaris(LeConte,1858)

Aphodiuspseudolividus(Balthasar,1941) Ataeniusplatensis(Blanchard,1844)

Canthonornatusbipunctatus(Burmeister,1873) Degallieridiumsp.

Pseudocanthonsp. Onthophagussp.

Trogidae Omorgusbatesi(Harold,1872) Polynoncusgemmingeri(Harold,1872)

Table3

Species/morphospeciesofomnivoroushabits.

Families Species/morphospecies Anthicidae Anthicidaesp.

Cleridae Necrobiaruﬁpes(DeGeer,1775) Nitidulidae Nitidulacarnaria(Schaller,1783)

Abundanceofguilds Necrophiles

Itwasthemostabundantgroup.Morethan38species were observed. In cadavers, in order of frequency and abundance, theywereCarpelimusspp.,Aleocharinaespp.,Philonthus ﬂavolim-batus (Erichson, 1753), Creophilus maxillosus (Linnaeus, 1758) (Staphylinidae); Euspilotus (Hesperosaprinus) pavidus (Erichson, 1834),Phelistersp.,Euspilotus(sensustricto)ornatus(Blanchard, 1843), Euspilotus (sensustricto) lacordairei (Marseul,1855) and Euspilotus (sensu stricto) patagonicus (Blanchard, 1843) (His-teridae); while in traps, Hylitus tentyroides (Lacordaire, 1830) (Tenebrionidae) and the tribes Harpalini, Cyclosomini, Lebiini, Pterostichini,Bembidiini(Carabidae)werethemostrepresentative. InFig.1AandBthemeanabundanceofnecrophilesper decom-positionstageand seasonaccordingtothesamplingmethodis presented.Thisgroupwasmoreabundantinspringandsummer. Duringspring,theabundancewasgreaterinAdvancedDecay,and insummerthereweremoreindividualsinActiveDecayfollowed byAdvancedDecay.Intraps,thegroupshowedasimilarbehavior Table1

Species/morphospeciesofnecrophiloushabits.

Families Species/morphospecies Families Species/morphospecies

Archeocrypticidae Archeocrypticustopali(Kaszab) Histeridae Phelistersp.

Melyridae Astylussp. Xerosaprinus(X)diptychus(Marseul)

Carabidae Pelmatellusegenus(Dejean) Dendrophilinaespp.

Bradycellusviduus(Dejean) Histerinaesp.1 Selenophorus(S)punctulatus(Dejean) Histerinaesp.2

Tetragonoderuschalceus(Chaudoir) Tenebrionidae Blapstinuspunctulatus(Solier) Tetragonoderuslaevigatus(Chaudoir) Gondwanocripticusplatensis(Fairmaire) Carbonelliaplatensis(Berg) Hylitustentyroides(Lacordaire) Argutoridiusoblitus(Dejean) Leptynoderesstrangulata(Fairmaire) Notaphusposticalisplatensis(Negre) Trichotonroigi(FerrerandMoragues) Histeridae Carcinopssp. Scarabaeidae Archophileuruschaconus(Kolbe)

Euspilotussp. Archophileurusfodiens(Kolbe)

Euspilotus(H)caesopygus(Marseul) Staphylinidae Aleocharinaespp. Euspilotus(H)connectens(Paykull) Carpelimusspp.

Euspilotus(ss)lacordairei(Marseul) Creophilusmaxillosus(Linnaeus) Euspilotus(H)niger(Arriagada) Haematodessp.

Euspilotus(ss)ornatus(Blanchard) Haematodesbicolor(Laporte) Euspilotusparenthesis(Schmidt) Philonthusﬂavolimbatus(Erichson) Euspilotus(ss)patagonicus(Blanchard)

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Abundance/c/day

Abundance/st/day

A

B

Winter Spring Summer Autumn

Seasons Seasons F B ACD ADD R 88 80 72 64 56 48 40 32 24 16 8 0 36 34 32 30 28 26 24 22 20 18 16 14 12 10 8 6 4 2 0

Fig.1. Meanabundance(±SE)ofnecrophilousbeetlesperdecompositionstages andseasons.(A)Cadavers(c).(B)Traps(st).F,Fresh;B,Bloated;ACD,ActiveDecay; ADD,AdvancedDecay;R,Remains.

asincadavers,buttheabundanceincreasedfromBloatedstageto AdvancedDecay.

Necrophileswerealsofoundasinmatures.Weobservedlarvae belongingtoHisteridae,Staphylinidae,Tenebrionidae and Cara-bidae.

Necrophages

Therewere found13 species. Dermestesmaculatus(DeGeer, 1774)(Dermestidae)wasthemostabundantinallseasons.This species was followed, in order of frequency and abundance in cadavers, by Polynoncus gemmingeri (Harold, 1872) (Trogidae); Scotobius muricatus (Guérin-Méneville, 1834) (Tenebrionidae); Onthophagus sp. and Canthon ornatus bipunctatus (Burmeister, 1873)(Scarabaeidae);while intraps,Scotobiusmiliaris(Billberg, 1815) (Tenebrionidae);Aphodius pseudolividus(Balthasar, 1941) andAtaeniusplatensis(Blanchard,1844)(Scarabaeidae)werethe mostrepresentative.Fig.2AandBshowsthemeanabundanceof necrophagesalongthedecompositionprocessduringeachseason andconsideringthesamplingmethod.Thisgroupwasmore abun-dantinwinterandthisseasonwascharacterizedbyanincreasein thenumberofindividualsfromBloatedstagetoAdvancedDecay.In spring,necrophagesshowedasimilarpatternasinwinter,butwith thedifferencethattheabundancedidnotdecreaseinRemains.In traps,AdvancedDecayandRemainswerethemostabundantstages duringwinterandspring.Insteadinsummer,specimensweremore abundantinActiveDecay.

Inadditiontothenecrophagousadults,wecollectedlarvaeof DermestesmaculatusandScarabaeidae.

Omnivores

Three species were observed in cadavers: Nitidula carnaria (Schaller,1783)(Nitidulidae)andNecrobiaruﬁpes(DeGeer,1775)

Abundance/c/day

Abundance/st/day

A

B

Seasons

Seasons F B ACD ADD R 38 36 34 32 30 28 26 24 22 20 18 16 14 12 10 8 6 4 2 0 2.0 1.9 1.8 1.7 1.6 1.5 1.4 1.3 1.2 1.1 1.0 0.9 0.8 0.7 0.6 0.5 0.3 0.2 0.0 0.1

Fig.2.Meanabundance(±SE)ofnecrophagousbeetlesperdecompositionstages andseasons.(A)Cadavers(c).(B)Traps(st).F,Fresh;B,Bloated;ACD,ActiveDecay; ADD,AdvancedDecay;R,Remains.

(Cleridae)weremorerepresentativeinabundanceandfrequency, while in traps was Anthicidae sp. In Fig. 3A and B the mean abundanceperdecompositionstageandseasonaccordingtothe samplingmethodis illustrated.Thisgroup wasmore abundant insummerandduringthisseason,thenumberofspecimenswas greaterinAdvancedDecayandActiveDecay.Duringspring,there weremorespecimensinthelasttwostagesandinsummer,the AdvancedDecayshowedanotableincrease.

Aswiththeotherguilds,wefoundinmatures.Inthiscase,larvae belongedtoNitidulacarnariaandNecrobiaruﬁpes.

Dominanceofguilds

Thedominanceofbeetleguildswasanalyzedandtheresults are shown in Fig.4.In winter, necrophages was thedominant guild. Theirrelative frequency washigh duringall the decom-position process, increasingfrom the Bloated stage (52%) until Remains(90%).Atthesametime, theproportionof necrophiles decreased and omnivores appeared. During spring, necrophiles constitutedthemostrepresentativeguild.Thedominanceofthis group decreased from the Bloated stage (95%) until Remains (46%). In summer,proportions weresimilar betweenthe three groups.TheBloatedstagewasrepresentedbynecrophiles(55%) followedbynecrophages(45%).InActiveDecay,necrophiles con-tinueddominate(57%), whileatthesametime thepresenceof necrophages decreased notably (15%) and appeared omnivores (28%).InAdvancedDecay,omnivoresbecamedominant(45%)and werefollowedbynecrophileswhosepresencedecreased(39%).In Remains,thegreaterrelativefrequencyofthepopulationwas rep-resentedbyomnivores,withanincreaseintheirvalues(62%),while necrophiles decreased notably (7%) and necrophages increased (31%).Duringautumn,necrophilesshowedagreaterproportion; theothergroupswerealmostinequal proportions.Therelative

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Abundance/c/day

Abundance/st/day

A

B

Seasons Seasons F B ACD ADD R 34 32 30 28 26 24 22 20 18 16 14 12 10 8 6 4 2 0 1.2 1.1 1 0.9 0.8 0.7 0.6 0.5 0.4 0.3 0.2 0.1 0.0

Fig.3.Meanabundance(±SE)ofomnivorousbeetlesperdecompositionstagesand seasons.(A)Cadavers(c).(B)Traps(st).F,Fresh;B,Bloated;ACD,ActiveDecay;ADD, AdvancedDecay;R,Remains.

frequencyofnecrophileswasmaximumintheBloatedstage(82%) andthendecreasedinActiveDecay(36%),intheotherstagesthere wasnovariation.Theproportionofnecrophagesandomnivores wasmoreorlessconstantbetweenstages;withthedifferencethat

Necrophiles Spring Autumn Summer Winter Omnivores Necrophages

Fig.4.Dominanceofbeetleguildsobservedincadaversperdecompositionstages andseasons,representedinaternarydiagram.Eachlineofthesidesofthetriangleis equaltoa5%ofthetotal.Theblackpointcorrespondstotheequicomposition(33.3% foreachguild).Thegreaterpointineachlineindicatestheinitialdecomposition stageperseason.

therelativefrequencyofomnivoreswasgreaterthannecrophages andtheyscarcelyappearedintheBloatedstage.

InFig.5AandDarepresenttheresultsofdominanceofbeetles capturedintraps.Necrophileswerethepredominantguildduring thefourseasonsandthroughthedecompositionprocess.During winter,necrophilesdominatedduringalltheprocess.Whilethe proportionofthisgroupdecreased,necrophagesand omnivores became notable,particularlytoward Remains.In theother sea-sons,necrophilesalsopredominated,buttheirimportancedidnot decreasetowardtheendoftheprocess.

100% 80% 60% 40% 20% 0% 100% 80% 60% 40% 20% 0% 100% 80% 60% 40% 20% 0% 100% 80% 60% 40% 20% 0% F B ACD ADD R Decomposition stages F B ACD ADD R Decomposition stages B ACD ADD R Decomposition stages B ACD ADD R Decomposition stages Necrophiles Omnivores

A

B

D

C

Necrophages

Fig.5.Dominanceofbeetlesguildscapturedintrapsperdecompositionstageandseasons.(A)Winter.(B).Spring.(C).Summer.(D).Autumn.F,Fresh;B,Bloated;ACD,Active Decay;ADD,AdvancedDecay;R,Remains.

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Discussion

Incadavers,speciﬁcdominancechangedaccordingtoseasons and decomposition stages. We inferred that during winter to theextentthatdecomposition processprogressed, necrophages became dominant and omnivores appeared, while the propor-tionofnecrophilesdecreased.Inspring,whenthedominanceof necrophilesstartedtodiminish,theothergroupsappeared,having necrophagesagreaterrepresentationthanomnivores.During sum-mer,thedecreaseofnecrophilesseemstocontributetoagreater relativefrequencyofomnivores.Thissuccessioncouldbearesult oftemporalsegregationasaformofreducingoverlapping, depre-dationorcompetitionforfoodresources(Zanetti,2013).

Weobservedsomerelationsbetweenthelengthofeachstage ofdecompositionandthepresenceofspecies.Theserelationswere mostlyobservedinwinter:Carpelimusspp.andAleocharinaespp. weremorefrequentinActiveandAdvanceddecaythanBloated stage;E.(Hesperosaprinus)pavidusandN.carnariaweremore fre-quentinAdvancedDecayandRemainsthanintheothersstages; Phelistersp.andE.(sensustricto)ornatusweremorefrequentin AdvancedDecaythan inRemains; andE.(sensu stricto) patago-nicusonlyappearedinAdvancedDecay.Somerelationwasalso observedinspringwithS.muricatus,whichwasmorefrequentin RemainsthaninBloatedstage;andA.pseudolividuswasonlyfound inRemains.AtaeniusplatensiswasonlyfoundinAdvancedDecayin winterandinRemainsduringsummer(Pers.comm.,Zanettietal.). Consideringtheseﬁndings,thelengthofeachdecompositionstage notalwaysisadeterminingfactorperse,insteadseveralfactors couldactincombination.

These results contraposed with those reported by Arnaldos

etal.(2005)whomentionedthatomnivoresappearedatthesame

timethatnecrophiles,maintainingthroughdecompositionstages.

Iannacone(2003)described thatnecrophages weremore

abun-dantandwithagreaternumberofspecies,andwerefollowedby necrophiles.Instead,theoppositehappenedintheactualstudy. Thesedifferencescouldbeattributedtoavarietyofcauses,suchas geographicalregion,climaticconditions,species,typeandsizeof thecarcass,locationofthecadaver,studyarea,methodologyused, amongstothers.

Thestrongassociationofdermestids(necrophages)with win-tercouldberelatedtoalesserpresenceofdipteranlarvaeandtoa longerdecompositionprocess,whichcouldprovidemorecadaveric tissuesduringmoretime,notonlyforfeedingbutalsofor reprodu-cing(Pers.comm.,Zanettietal.,2015).Inthisway,moredipteran larvaecouldconsumeagreatercadavericbiomass,reducingthe foodavailabilityfordermestidsorothernecrophages,meanwhile suchlarvaecouldfavorthepresenceofnecrophilessuchasrove beetles,clownbeetlesorcarabids,whicharepredators ofthem andotherinsectsindifferentstagesofdevelopment(Pers.comm., Zanettietal.,2015),andsomeareparasitesofdipteranpuparia

(Ashe,2002;Miseetal.,2010).Theappearanceofnecrophiles

indi-catestheexistenceofnecrophagousfaunaalreadyinstalledinthe cadaver(CattsandGoff,1992)andthefrequencyofnecrophiles isdirectlyrelatedwiththepresenceoftheprey.Sincethepreys of necrophiles are usually dipteran larvae (Fichter, 1949), it is expectedthat thefrequency of necrophilesdecreases withthe biomassloss and thereduction of thedipteranlarvae. Because omnivores can act as predators, it can be expected that they coexistina lesserdegreewithnecrophiles,but frequentlywith necrophagesbecausetheycanfeedonthemandlikethem,theycan useacarcassasafoodresource.Cleridsdominatedwhenthe fre-quencyofroveandclownbeetlesdecreased(Pers.comm.,Zanetti etal.,2015).Withrespecttocertainauthors,nitidulidsarrivein laterstagesofdecompositionsharingwithdermestids(Payneand

King,1969;OlayaMásmela,2001;ÖzdemirandSert,2009).Thus,

itcouldhappenthatboth,theabsenceorlowfrequencyofaguild,

andthestageofdecomposition,couldinﬂuencetheabundanceand dominanceofcertaingroup.

Coprophagous beetles(scarab beetles) wereobserved inthe studybuttheywereconsideredaspartofthenecrophagousgroup duetothefacultativecapacityofthesebeetlessuchasfeedingfrom tissues.

Conclusion

Differences in the abundance and dominance of beetles in accordancewiththeirtrophicroles(guilds)werefoundper sea-sonand decomposition stages.Thereis anassociation between theseperiodsand thefeedinghabitsofbeetles,asasuccession. The necrophiles (principally rove beetles) dominated between early stages, sharing this with necrophages (particularly der-mestids) in winter. The necrophages were the dominant guild duringwinterprincipallyin thelasttwo stages.Theomnivores showedamajordominanceinsummerduringAdvancedDecayand Remains.

Justasthearthropodsuccessionisusuallyusedintheestimation ofPMI(generallyamaximumPMI),theresultsreportedinthiswork underlinethatknowingorestablishingwhatguildwaspresentor absentandwhenthesameprevailed,canbeofrelevancein eval-uatingimportantalterationsin successionwhichcouldindicate abnormalconditionsinthedecompositionofthecadaver, particu-larlyintheregionwhereourstudywasconducted.Also,knowing howtheguildsbehave,forexample,theappearanceofnecrophiles indicatestheexistenceofnecrophagousfaunaalreadyinstalledin cadavers,shouldbeconsideredwhenexaminingthecrimescene andcollectingentomologicalevidence,aswellastoavoiderrors andintheestimationofPMI.

Conﬂictsofinterest

Theauthorsdeclarenoconﬂictsofinterest.

Acknowledgments

This work was supported by the Universidad Nacional de Quilmes.NoeliaInésZanettiisfellowofCONICET.Authorsalsowish tothanktoRicardoCaminafromUniversidadNacionaldelSurfor helpingwiththestatisticalanalysis,and toGustavoFlores, Fer-nandoAballay,CesarMilton,ArmandoCicchinoandMarianaChani Posse,forhelpinginthetaxonomicaldeterminationor corrobora-tionofsomespecimens.

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