Description of immature stages of Hoplia mexicana Harold and H. squamifera Burmeister (Coleoptera, Melolonthidae, Hopliinae)

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

REVISTA

BRASILEIRA

DE

Entomologia

AJournalonInsectDiversityandEvolution

Systematics,

Morphology

and

Biogeography

Description

of

immature

stages

of

Hoplia

mexicana

Harold

and

H.

squamifera

Burmeister

(Coleoptera,

Melolonthidae,

Hopliinae)

Miguel

Ángel

Morón

,

Andrés

Ramírez-Ponce

_,

_Concepción

_{Ramírez-Salinas}

_,

Hortensia

Carrillo-Ruiz

a_{ReddeBiodiversidadySistemática,InstitutodeEcología,Xalapa,Mexico}

b_{LaboratoriodeBiologíaEvolutiva,CentroTlaxcaladeBiologíadelaConducta,UniversidadAutónomadeTlaxcala,Tlaxcala,Mexico} c_{ElColegiodelaFronteraSur(ECOSUR),SanCristóbaldeLasCasas,Chiapas,Mexico}

d_{LaboratoriodeEntomología,EscueladeBiología,BeneméritaUniversidadAutónomadePuebla,Puebla,Mexico}

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r

t

i

c

l

e

i

n

f

o

Articlehistory: Received10April2015 Accepted9November2015 Availableonline19December2015 AssociateEditor:MarcelaMonné Keywords: Biology Morphology Pupae Taxonomy Whitegrubs

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b

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ThirdstagelarvaeandpupaearedescribedbasedonspecimenscollectedinMexico:Oaxaca(Cerro

Zem-poaltepetl),andChiapas(Amatenango),respectively.Pupalcharactersaredescribedforthefirsttime

forAmericanHopliinae.Habitusimagesandfiguresofdiagnosticcharactersaswellascommentsonthe

differencesbetweentheselarvaeandthoseofHopliacallipygeLeConte,1856andH.equinaLeConte,1880,

theonlyHopliinaelarvaepreviouslyknowninNewWorld,arealsoincluded.

©2015SociedadeBrasileiradeEntomologia.PublishedbyElsevierEditoraLtda.Thisisanopen

accessarticleundertheCCBY-NC-NDlicense(http://creativecommons.org/licenses/by-nc-nd/4.0/).

Introduction

The immature stages of world Hopliinae are poorly known. Partialdescriptionsexistforthethird instarlarvaeoftheNorth American species Hoplia callipyge LeConte, 1856 (= H. oregona LeConte,1856; =H. cazieriBoyer,1940) and H.equina LeConte (Ritcher,1949).Additionally,therearelarvaldescriptionsforsome RussianspeciesofthegenusHopliaIlliger,1803andthecentral EuropeanH.philanthus(Fuessly, 1775)(Medvedev,1952;H ˚urka andBíl ´y,1966).Micó(2001)redescribed thelarva ofH. philan-thusanddescribedthatofH.chlorophanaErichson,1848,basedon specimensfromSpain,bothwithdetailedillustrations.Ansarietal. (2006,2008)detailedthelifecycleofH.philanthusandcommented abouttheimportanceandbiologicalregulationofthisturfpest.

LarvaeofNorthAmericanspeciesofHopliamaybecharacterized asfollows:headwithoutstemmata;labrumsymmetrical; maxil-larystridulatoryareawithrowof9–11shortteethwithanteriorly directedpoints;laciniawithlongitudinalrowof3unci,distal2 uncifusedbasally,subapicaluncusmuchsmaller;epipharynxwith plegmatia,plegmataquiteshort;proplegmatiapresentorabsent; epizygumabsent;haptomerumwithsinglehelus;dexiophobaand

∗ Correspondingauthor.

E-mail:miguel.moron@inecol.mx(M.Á.Morón).

laeophobaabsent;haptolachuswithoutmicrosensilla;spiraclesof abdominalsegments1–3similarinsize,spiraclesofsegments4–8 muchsmaller;analopeningY-shaped;rasterconsistingof subtri-angulartegesof30–60fairlylonghamatesetaewithcurvedtips; claws offorelegslargeand falcate,claws ofmiddle legsmuch reducedinsize,clawsofhindlegsminuteorabsent(Ritcher,1966). LarvaeofEuropeanspeciesofHopliasharealloftheabovecited characters,except:maxillarystridulatoryareawithrowof12–14 shortteethwithanteriorlydirectedpoints;laciniawith subapi-cal uncus large; epipharynx withoutproplegmatia; spiracles of abdominalsegments4–6diminutive(Micó,2001).

Duringrecentyears,wehadtheopportunitytorearmanyscarab larvaecollectedfromforestsoil,grasslandsandcornfieldsof south-eastern Mexico,and someof them belongtospecies ofHoplia. This material allowed us to describe in the present paper the thirdinstarlarvaeforH.mexicanaHarold,1869andH.squamifera Burmeister,1844,forwhichtheimmaturestageswereunknown. Wealsodescribethepupaeofthesetwospecies,alifestagethat wasunknownforanyNewWorldhopliinescarab.

Materialandmethods

Throughoutthedescriptionsanddiscussions,wefollow termi-nologyemployedbyRitcher(1966)andMorón(1986).Drawings ofdiagnosticstructuresweremadeusingaLeicastereomicroscope http://dx.doi.org/10.1016/j.rbe.2015.11.012

0085-5626/© 2015 Sociedade Brasileira de Entomologia. Published by Elsevier Editora Ltda. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).

2 M.Á.Morónetal./RevistaBrasileiradeEntomologia60(2016)1–7

associated with a camera lucida at 0.8–5.0× magnification. Measurements were taken using an ocular micrometer (Leica microsystems,Germany).Voucherspecimens weredepositedat ColecciónproyectoDiversidadenSistemasdeCultivos,ElColegio delaFronteraSur(ECOSUR),andColecciónEntomológica,Instituto deEcología,A.C.Xalapa,México(IEXA).

Taxonomy

Descriptions

HopliamexicanaHarold,1869

Thirdinstar(Figs.1–11and13).Description.Head(Fig.2). Max-imum width of head capsule: 1.8–2.0mm. Surface of cranium smooth,withscatteredminutepunctures,paleyellowish.Frontal andepicranialsutureshidden.Fronswith2exteriorfrontalsetae, 3–4posteriorfrontalsetae,1anteriorfrontalseta,1anteriorangle seta,3dorsoepicranialsetae,8–9epicranialsetaeoneachsideand 7–9setaebehindeachantennal support.Clypeuswith2central setaeand 4lateralsetaoneach side.Labrumnearly symmetri-cal withrounded lateral marginsirregularly bordered, anterior margin projected forward with irregular border, 6–7 posterior setae,1 central seta and 3–4lateral setaeoneach side. Stem-mataabsent. Epipharynx (Fig. 5)1.0–1.1mm wide, 0.8–0.9mm long,epizygumabsentand zygumasairregular,ovate, reddish yellowplate.Haptomerumweaklyraised,with4heli;each pleg-matiaformedby8–10shortplegmata;proplegmatiaabsent.Each acanthopariawith10–12spinosesetae.Dexiophobaandlaeophoba absent.Dexiotormanarrowed,slightly sinuose.Laeotorma elon-gateandnarrowed,withsclerotizedplateunciform,raised.Sense coneshort.Crepisweakly defined.Leftchaetoparia with28–30 spinose and medium size setae mixed. Right chaetoparia with 30–36short,stoutsetae.Mandibles(Figs.3and4)without ven-tralstridulatoryarea;scissorial areainbothmandibleswithout distalblade.Right mandiblewith1smallpreapicaltooth,inner marginsimplycurved,anddistallobe(M1)ofmolarareascarcely developed.Calxshort.Leftmandiblewith1smallpreapicaltooth,

innermarginsimplycurved,brustiamoderatelysetose.Aciaabsent. Maxillae(Figs.6–9).Malawith1uncusonapexofgalea,3unci sur-roundedby3–4stouthelionapexoflacinia;stridulatoryareawith 8smallteethwithanteriorlydirectedpoints.Hypopharyngeal scle-rome(Fig.10)asymmetricalwithraisedandcurvedprojectionon therightside.Fourthantennomereelongate,withdistalhalf nar-rowed,with1ovate,dorsalsensoryspotonbasalhalfand2ventral sensoryspots.

Thorax. Respiratory plates light yellow, scarcely curved, “C” shaped(Figs.1and13)0.12–0.14mmlong,0.10–0.11mmwide, bullasmall,slightlyraised,rounded,distancebetweenlobeslonger thandorso-ventraldiameterofbulla;microscopicholesof respi-ratory plate elongate-oval in outline and arranged in irregular transverserows.Lateralscleromeofpronotumnotdefined. Pro-prescutumwith20–22longsetaeregularlydistributed;proscutum with8–9slenderlongsetae;mesoprescutumwithtransverserow of10–12longsetae;mesoscutumwith12–16setae; mesoscutel-lumwithrowof 8–10slenderlongsetae; metaprescutumwith transverserowof12–16mixedshortandlongsetae;metascutum with10–12setae;metascutellumwithtransverserowof10–12 mixedshortandlongsetae(Figs.1and13).Protarsalclawslong, narrowed,sharplypointed,with1prebasalinnersetaand1lateral external seta, moderatelylonger than mesotarsalclaws; meso-tarsalclawslong,sharplypointed,withpre-basalandlateralsetae; metatarsalclawshort,rounded,with2setae,muchshorterthan mesotarsalclaws.Apicalmetatarsomereshortened,rounded,with ventralsideclearlyconvex.

Abdomen.Respiratoryplateslightyellow,slightlycurved,“C” shaped;onsegmentsI–IV withsimilardiameter,0.12–0.13mm long,0.10mmwide;platesonsegmentsV–VIIareslightlysmaller, 0.10–0.11mm length, 0.09mm width. Plate on segment VIII is smallerthanpreceding.DorsaofabdominalsegmentsI–VIeach withdensevestitureofyellowstout,shortsetae;dorsaofsegments VII–IXwithtransverserowsof20–28slender,shortsetae.Venter ofabdominalsegmentsI–VIIIwithtransverserowsof14–18 slen-der,longsetae;venterofsegmentIXwithtransverserowsof12–16 mixedshortandlongsetae.

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10

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Figs.6–12.Hopliamexicana.6–11:thirdinstarlarva.Maxillae:dorsalview:6,left.7,right.Mesialview:8,left.9,right.10,labiumandhypopharynx.11,raster.12,MALE pupa,lateralview.

Raster(Fig.11)witheachpalidiaformedby9–11pali, conver-genttowardbasalanddistalextremes,septulawide,oval;tegilla with20–26short,stoutsetae;campuswith14–18slender,short setae;barbulamuchmoredenselysetose.Dorsalanallipwithdense vestitureofshortsetae.Ventralanallipwithscatteredshortsetae towardsidesanddistalbordernarrowlynotchedatmiddle.Anal slit“Y”shaped.Approximatedorsalbodylength16–17mm.

Description: body length 7.5–7.8mm. Widest width

4.4–4.5mm. Head. Surface glabrous, strongly deflexed; frons convexwithtworoundedprominences;clypeuswidelyconcave onthecenter;labrum,mandibles,maxillaeandpalpsdiscernible; antennalthecabrieflyexpanded,stoutwithapexrounded;eyes small (Fig. 14). Thorax: surface glabrous. Pronotum convex, surface slightly irregular, anterior angles prominent, posterior anglesrounded.Meso-andmetanotumwell-differentiated.Elytral

and posterior wing thecae closely appressed, curved ventrally around thebody;elytral thecaewithirregular depressionsand large, prominent, humeral tubercles (Figs. 12 and 15); thecae of the wings slightly longerthan elytral thecae. Protibiae with apical and preapical tubercles clearly developed. Meso and metatibiaewithapicaltubercle.Abdomen:segmentsI–VIclearly wider and shorter than the distal segments VII–VIII, without dioneiform organs, but segments II–IV with pairs of promi-nent tubercles and segment V withincreased posterior border (Figs.12and 15–16).Pleurallobesrounded. SpiracleIelongate, withfineperitremeandcoveredbywingthecae;spiraclesII–IV ovate and high, prominent, withnarrow sclerotized peritreme; spiraclesV–VIIIclosed.SuturesbetweensegmentsVIIIandXnot complete,partiallyfused.Lastsegmentslightlypruinose,without urogomphi.

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15

16

14

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17

19

20

21

18

Figs.17–21.ThirdinstarlarvaofHopliasquamifera.17,lateralhabitus.18,head,frontalview.Mandibles,ventralview:19,left.20,right.21,epipharynx.

Last abdominal segment of male withlarge dorsal and lat-eralroundedprominences,andgenitalampullaventrallyexposed (Figs.12and14–16);femalewithoutsuchprominencesattheapex ofabdomen.

Specimensexamined.Fourthirdinstarlarvae,fourpupaeand twoexuviae of thirdinstar larvaerearedto adults,collectedat Mexico:Oaxaca: SantaMaríaTlahuitoltepec municipality,Cerro Zempoaltepetl,2400m,8-VI-2010,A.Ramírezcol.(IEXA).

Comments.ThethirdinstarlarvaeofH.mexicanaresemblethose ofH.callipygeandH.equina,butH.mexicanahas4helion hap-tomerum,andpalidiawithwell-definedseptula.Hopliacallipyge andH.equinaeachhave1helusonthehaptomerumandonly sub-triangulartegesoflonghamatesetaewithcurvedtips.Thedense setiferousvestitureonthedorsumofabdominalsegmentsI–VIand thesidesofthelast segment,aidin distinguishingH.mexicana fromtheknownlarvaeof otherMexicanmelolonthine scarabs. Projectionsonthehumeralareaoftheelytraandthemiddleof theabdominalsegmentsofthepupaeof bothsexesmayactas auxiliarysupportsduringpupaldevelopment.These projections possiblyaidinregulatinghumiditynearthebodywall.Thistypeof structureisalsoobservedinpupaeofsomeScarabaeinae(Edmonds andHalffter,1978),andCeratocanthinae(MorónandArce,2003). Biology. Hoplia mexicana is known only from the northern mountainsoftheMexicanstateofOaxaca,atlocalitieswith ele-vationsbetween2300and2500m.Larvaehavebeencollectedin richorganicsoil of oakand coniferous forest.Adults arerarely attractedtoelectriclightsduringsummer,andtheirhostplantsare unknown.

HopliasquamiferaBurmeister,1844

Thirdinstar(Figs.17–24).Description. Head(Fig.18).Maximum widthofheadcapsule:2.1–2.4mm.Surfaceofcraniumsmooth, withscatteredminutepunctures,paleyellowish.Frontaland epi-cranial sutures hidden. Frons with 2 exterior frontal setae, 1 posteriorfrontalseta,anteriorfrontalsetaabsent,1anteriorangle

seta,3–4dorsoepicranial,14–18epicranialsetaeoneachsideand 6–8setaebehindeach antennalsupport.Clypeuswith2central setaeand4lateralsetaeoneachside.Labrumnearlysymmetrical withroundedlateralmarginsirregularlybordered,anteriormargin projectedforwardwithirregularborder,basaltransversekeelwith 5posteriorsetae,1anterior-centralsetaand5–6lateralsetaeon eachside.Stemmataabsent.Epipharynx(Fig.21)1.6–1.9mmwide, 1.0–0.2mmlong.Epizygumabsentandzygumasirregular,ovate, reddishyellowplate.Haptomerumweaklyraised,with4heli;each plegmatiaformedby9–10shortplegmata;proplegmatiaabsent. Eachacanthopariawith9–10spinosesetae.Dexiophobaand laeo-phobaabsent.Dexiotormanarrowed,slightlycurved.Laeotorma elongate and narrowedwith sclerotized plateunciform, raised. Sensecone short. Crepisweakly defined. Left chaetoparia with 30–34spinoseand mediumsizesetaemixed.Rightchaetoparia with28–30short,stoutsetae.Mandibles(Figs.19and20)without ventralstridulatoryarea;scissorialareainbothmandibles with-outdistalblade.Rightmandiblewith1smallrounded,preapical tooth,innermarginsimplycurved,anddistallobe(M1)ofmolar areascarcelydeveloped.Calxshort.Leftmandiblewithout preapi-caltooth,innermarginsimplycurved,brustiamoderatelysetose. Aciaabsent.Maxillae(Fig.23).Malawith1uncusonapexofgalea, 3uncisurroundedby3–4stouthelionapexoflacinia; stridula-toryareawith10–11smallteethwithanteriorlydirectedpoints. Hypopharyngealsclerome(Fig.22)asymmetricalwithcurved pro-jectionontherightside.Fourthantennomereelongate,withdistal halfnarrowed,with1ovate,dorsalsensoryspotonbasalhalfand 2ventralsensoryspots.

Thorax. Respiratory plates light yellow, scarcely curved, “C” shaped (Fig.17)0.11–0.15mm long, 0.10–0.12mm wide, bulla small,slightlyraised,rounded,distancebetweenlobeslongerthan dorso-ventraldiameterofbulla;microscopicholesofrespiratory plateelongate-ovalinoutlineandarrangedinirregulartransverse rows. Lateral scleromeof pronotum not defined.Proprescutum with14–18longsetaeregularlydistributed;proscutumwith5–6

22

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Figs.22–25.Hopliasquamifera.22–24:thirdinstar.22,labiumandhypopharynx.23,leftmaxilla,dorsalview.24,raster.25,femalepupa,lateralview.

slenderlong setae;mesoprescutumwithtransverserowof 8–9 longsetae;mesoscutumwith10–12setae;mesoscutellum with rowof6–8slenderlongsetae;metaprescutumwithtransverserow of9–12mixedshortandlongsetae;metascutumwith8–10setae; metascutellumwithtransverserowof6–8mixedshortandlong setae(Fig.17).Protarsal clawslong,narrowed,sharplypointed, with1 pre-basalinner setaand 1 lateralexternal seta, moder-atelylongerthanmesotarsalclaws;mesotarsalclawslong,sharply pointed,withpre-basalandlateralsetae;metatarsalclawshort, rounded,with2setae,muchshorterthanmesotarsalclaws. Api-calmetatarsomereshortened,rounded,withventralsideclearly convex.

Abdomen.Respiratoryplateslightyellow,slightlycurved,“C” shaped;onsegmentsI–IV withsimilardiameter,0.13–0.14mm long,0.11mmwide; plateson segmentsV–VII slightly smaller, 0.11–0.12mmlong,0.10mmwide.Respiratoryplateonsegment VIIIsmallerthanplateonprecedingsegment.Dorsaof abdomi-nalsegmentsI–VIeachwithdensevestitureofyellowstout,short setae;dorsaof segmentsVII–IX withtransverserows of18–20 slender, short setae.Venter of abdominalsegments I–VIII with transverserowsof10–12slender,longsetae;venterofsegmentIX withtransverserowsof10–12mixedshortandlongsetae.Raster (Fig.24)witheachpalidiaformedby10–12pali,convergenttoward basalanddistalextremes,septulawide,oval;tegillawith24–30 short,stoutsetae;campuswith14–16slender,shortsetae; bar-buladenselysetose.Dorsalanallipwithdensevestitureofshort setae.

Ventralanallipwithscatteredshortsetaeatsidesanddistal bor-dernarrowlynotchedatmiddle.Analslit“Y”shaped.Approximate dorsalbodylength16–18mm.

Pupa (Figs. 25 and 26).Description: body length 7.8–8.1mm. Widest width 4.5–4.6mm. Head. Surface glabrous, strongly deflexed;fronsconvexwithtwo roundedprominences;clypeus

Fig.26. MalepupaofHopliasquamifera,withthirdinstarexuviaeonitsabdominal apex.

widelyconcaveonthecenter;labrum,mandibles,maxillaeand palpsdiscernible;antennalthecabrieflyexpanded,stoutwithapex rounded;eyessmall(Fig.25).Thorax:surfaceglabrous.Pronotum convex,surfaceslightlyirregular,anterioranglesbrieflyprominent, posterior angleswidely rounded. Meso- and metanotum well-differentiated.Elytralandposteriorwingthecaecloselyappressed, curved ventrally around body; elytral thecae with irregular

6 M.Á.Morónetal./RevistaBrasileiradeEntomologia60(2016)1–7

Table1

MaincharactersinthepupaeofsomeAmericangeneraofScarabaeoidea.D.O.,dioneiformorgans;H.P.,humeralprojections;P.P.,pronotalprojections;T.P.,tergalprojections; U,urogomphi.

Subfamilies Genera P.P. H.P. D.O. T.P. U References

Melolonthinae PhyllophagaHarris,1827 No No No No Yes Morón(1986)

PolyphyllaHarris,1841 No No No No Yes MorónandNogueira(2015)

ClavipalpusLaporte,Castelnau,1832 No No No No Yes Neita-Morenoetal.(2012)

DemodemaBlanchard,1850 No No No No Yes MorónandSalvadori(2006)

IsonychusMannerheim,1829 No No No No Yes Neita-Morenoetal.(2012)

MacrodactylusDejean,1821 No No No No Yes Ramírez-Salinasetal.(2011)

ManopusLaporteCastelnau,1840 No No No No No Neita-Morenoetal.(2012)

Hopliinae Rutelinae

HopliaIlliger,1803 No Yes No Yes No Presentpublication AnomalaSamouelle,1819 No No Yes No No Micóetal.(2003)

MacraspisMacLeay,1819 No No Yes No No MorónandPaucar-Cabrera(2003)

ChlorotaBurmeister,1844 No No Yes No No JamesonandMorón(2001)

ChrysophoraLeP.&Serville,1828 No No Yes No No Pardo-LocarnoandMorón(2007)

LeucothyreusMacLeay,1819 No No Yes No No Pardo-Locarnoetal.(2006b)

Dynastinae LigyrusBurmeister,1847 No No Yes No No Ramírez-Salinasetal.(2004)

AncognathaErichson,1847 No No Yes No No Ramírez-Salinasetal.(2004)

CoelosisHope,1837 No No Yes No No Pardo-Locarnoetal.(2006a)

GolofaHope,1837 No No Yes No No Morón(1995b)

DynastesKirby,1825 No No Yes No No Morón(1987)

AegopsisBurmeister,1847 No No Yes No No Neita-Morenoetal.(2014)

Cetoniinae EuphoriaBurmeister,1842 No No No No No Ramírez-Salinasetal.(2001)

HoplopygaJ.Thomson,1880 No No No No No MorónandArce(2002)

GymnetisMacLeay,1819 No No No No No MorónandArce(2002)

NeoscelisSchoch,1897 No No No No No Nogueiraetal.(2004)

Trichiinae IncaLePeletier&Serville,1828 No No Yes No No Morón(1983)

ArchedinusMorón&Krikken,1990Krikken,1990 No No Yes No No Morón(1995a)

Orphninae AegidiumWestwood,1846 No No No Yes Yes Morón(1991)

Ceratocanthinae CeratocanthusWhite,1842 Yes Yes No No Yes MorónandArce(2003)

Aphodiinae TermitodiusWasmann,1894 No Yes No Yes Yes Reyes-CastilloandMartínez(1979)

Scarabaeinae PhanaeusMacLeay,1819 Yes No No Yes No Edmonds(1967)

EurysternusDalman,1824 No No No Yes No López-GuerreroandMorón(1994)

longitudinal striae and large, prominent, semiconical humeral tubercles(Figs. 25 and 26); thecaeofthe wingsslightly longer than elytral thecae. Protibiae with apical and preapical tuber-clesclearlydeveloped.Mesoandmetatibiaewithapicaltubercle. Abdomen:segmentsI–VIclearlywiderandshorterthanthedistal segmentsVII–VIII,withoutdioneiformorgans,butsegmentsII–IV withpairsofprominenttuberclesandsegmentVwiththicker pos-teriorborder(Figs. 25and26).Pleurallobesrounded.SpiracleI elongate,withfineperitremepartiallycovered bywingthecae; spiraclesII–IVovateandhigh,prominent,withnarrowsclerotized peritreme;spiraclesV–VIIIclosed.SuturesbetweensegmentsVIII andXincomplete,partiallyfused.Lastsegmentslightlypruinose, withouturogomphi.Lastabdominalsegmentofmalewithdorsal andlateralroundedprominences,and genitalampullaventrally exposed(Fig.26);femalewithoutsuchabdominalprominences (Fig.25).

Specimens examined. Seven third instar larvae, collected at Mexico:Chiapas,AmatenangodelVallemunicipality,ElMadronal, 2400m,20-I-1997,cornfield,C.Ramírezcol.(IEXA).Twopupae and2exuviaeofthirdinstarlarvaerearedtoadults,withsame data,except24-X-2005(IEXA).Twothirdinstarlarvae:Chiapas, AmatenangodelValle,28-X-1996,cornfield,C.Ramírez;4third instarlarvaewithsamedata,except4-XI-1996;5thirdinstarlarvae 24-IX-2001(IEXA,ECOSUR).

Comments.ThelarvaeofHopliasquamiferaresemblethoseofH. mexicana,butlackantero-frontalsetae,have9–10wideplegmata oneachplegmatia,andhavewideseptula,whereasHoplia mex-icanahave2antero-frontalsetae,7–8narrowplegmataoneach plegmatiaandnarrowerseptula.

PupaeofbothHopliamexicana,andH.squamiferalack urogom-phianddioneiformorgans.Thisfactisveryinterestingbecause, asfarasweknow,nootherscarabgrouppossessessucha combi-nationofcharacters.Forinstance,withtheexceptionofManopus biguttatusLaporte,1840(Neita-Morenoetal.,2012),knownpupae

of Melolonthinae have urogomphi and lack dioneiform organs (Table1).Ontheotherhand,knownpupaeofRutelinaeand Dynas-tinae,lack urogomphi but, in contrast,have dioneiform organs (Morón,1993).Withthecombinationofthesecharacters,andthe uniquepairedsupportingorgansinthehumeriandtergites,the pupae of Hopliinae are very distinct from othersubfamilies of Melolonthidae.Descriptionofthepupaeofmanyothergeneraof Melolonthidaesurelywill offermorecharacters tobetested in futurephylogeneticanalysesofScarabaeoidea.

Biology.HopliasquamiferahasbeenrecordedfromtheMexican mountainsofnorthernChiapasandOaxacastates,southern Vera-cruzstate,andinwesternGuatemalaatlocalitieswithelevations between1200and2100m.Larvaehavebeencollectedinsoilofcorn fieldsinnorthernChiapas,Mexico.Adultsarefrequentlyattracted toelectriclightsduringspringandsummer,buttheirhostplants areunknown.

Conflictsofinterest

Theauthorsdeclarenoconflictsofinterest.

Acknowledgements

The aidof Adriana E. Castro (ECOSUR) and Susana Guzmán (InstitutodeBiología,UNAM,Mexico,D.F.)duringfieldand tech-nicalworkishererecognized.Thesuggestionsoftwoanonymous reviewerswereveryusefulinimprovingthefinalversionofthe manuscript.WethankBruceGill(Ottawa,Canada)forhisfriendly assistancewiththeEnglishlanguage.Thispaperwassupportedin partbytheresearchline“ColeópterosLamelicorniosdeAmérica Latina”InstitutodeEcología,A.C.Xalapa,Mexico(#2004010011).

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