Description of the third instar larva of Saccharoscaptus laminifer (Dechambre) (Coleoptera: Melolonthidae: Dynastinae)

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REVISTA

BRASILEIRA

DE

Entomologia

AJournalonInsectDiversityandEvolution

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

Systematics,

Morphology

and

Biogeography

Description

of

the

third

instar

larva

of

Saccharoscaptus

laminifer

(Dechambre)

(Coleoptera:

Melolonthidae:

Dynastinae)

Miguel

Ángel

Morón

ReddeBiodiversidadySistemática,InstitutodeEcología,Xalapa,Mexico

a

r

t

i

c

l

e

i

n

f

o

Articlehistory:

Received4April2017 Accepted17July2017 Availableonline29July2017 AssociateEditor:AdrianaMarvaldi Keywords: Morphology Taxonomy Biology Whitegrubs Sugarcane

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ThelarvaofpentodontineS.laminiferisdescribedforﬁrsttimebasedonspecimenscollectedunder rootsofsugarcaneinSantaCruz,Bolivia.Diagnosticstructuresareillustratedandthedifferencesand similaritieswithotherpreviouslydescribedlarvaeofSouthAmericangeneraofPentodontiniareoutlined. AkeytothelarvaeofsomeAmericangeneraofpentodontinesisincluded.

Introduction

IntheAmericas,thetribePentodontiniincludes32generaand

151speciesthatoccurfromsouthernCanadatoArgentina,with

fourgeneraandninespeciesintheWestIndies(Endrödi,1969,

1985;MorónandGrossi,2015;RatcliffeandCave,2015; López-Garcíaetal.,2016),butonlythelarvaeofninegenera(Aphonus

LeConte, Bothynus Hope, Coscinocephalus Prell, Euetheola Bates,

LigyrusBurmeister,NeoryctesArrow,OrizabusFairmaire,

Oxygril-ius Casey, Philoscaptus Bréthes) represented by 15 species are

described(Ritcher,1966;Morón,1976;Lumaret,1991;Morónand

Ratcliffe,1996;Morelli,1997;Ramírez-Salinasetal.,2004,2010; Pereiraetal.,2013),includingscarcedataaboutthebiologyofeach species.

During2012,theauthorreceivedsamplesofmediumsizewhite

grubs collected under roots of sugarcane ﬁelds of Santa Cruz

province,Bolivia.AfterbreedinginthelaboratoryofInstitutode

InvestigacionesAgrícolasElVallecito(IIAV),UniversidadAutónoma

GabrielRenéMoreno,SantaCruz,theselarvaewereidentiﬁedas

PhiloscaptuslaminiferDechambre,1979(Copa-BazanandMorón,

2014).Thisspecieswasknownonlybythemaleholotypecollected

in“Para,Brazil”duringXIXcentury.Recently,thisspeciesbecame

thetypespeciesofSaccharoscaptus(MorónandGrossi,2015).The

aimsofthepresentworkaretodescribethethird-instarlarvaofS.

laminifer,andtoprovideanupdatedkeytothelarvaeofAmerican

Pentodontini.

E-mail:Moronmiguel.moron@inecol.mx

Materialandmethods

The classiﬁcation of families of Scarabaeoidea used in the

presentpaperwasproposedbyEndrödi (1966)and updatedby

Morón(2010)andChermanandMorón(2014).Termsand

charac-tersusedinthedescriptionoflarvaarethoseofRitcher(1966),

Morón (1987),and Morónet al.(2014). Drawingsofdiagnostic

structures weremade usinga Leicastereomicroscope 0.8–5.0×

associatedwithacameralucida,andmeasurementswereobtained

withanocularmicrometer.Photographswereobtainedwithaidof

avideosystemassociatedtoaCelestron44206stereomicroscope.

Voucherspecimensweredepositedatthereferencecollectionof

IIAV,SantaCruz,Bolivia,andColecciónEntomológica Institutode

Ecología,A.C.Xalapa,Mexico(IEXA).

Results

It isdifﬁcult tooffera setof clearcut diagnosticcharacters

to distinguish the larvae of the tribe Pentodontini from those

ofothertribesof Dynastinae.Butasa preliminaryintroduction

basedonRitcher(1966),Morón(1987),MorónandRatcliffe(1996),

and recent experience of the author, the following diagnostic

combinationoflarvalcharactersofAmericanPentodontiniis

pro-posed:maximumwidthofheadcapsule3–8mm;with0–2anterior

frontalsetae;labrumslightlyasymmetrical,bothlateralmarginsof

epipharynxrounded;apexoflaciniawithtransverserowof3unci

fusedbasally,theintermediateuncussmallerthanlateralunci;left

mandiblewithS4reducedorabsent;lastantennomerewith1–8

http://dx.doi.org/10.1016/j.rbe.2017.07.004

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sensoryspots;eachtarsalclawwith2setae;dorsaofabdominal

segmentsVIIIandIXwithoutshort,spiniformsetae.

KeytothethirdinstarlarvaeofAmericanPentodontini

(ModiﬁedfromPereiraetal.,2013)

Note. Diloboderus is not included in the tribe Pentodontini,

becausebasedonlarvalcharacters,itwastransferredtothetribe

Agaocephalini (Pardo-Locarno and Morón, 2006; Neita-Moreno

etal.,2014;MorónandGrossi,2015).

1.Rasterwithpalidiaandseptula...2

1.Rasterwithoutpalidiaandseptula...3

2.Palidiamonostichous,eachpalidiumconsistingof7–10

com-pressedpaliwhosetipsareslightlyhooked.Palidiaandseptulanot

extendingacrossloweranallip.Lastantennomerewith2dorsal

sensoryspots...Euetheola

2. Palidia polystichous, each palidium consisting of a

patch of 5–7 irregular, longitudinal rows of sharp,

cylin-drical, spine-like pali. Palidia and septula extending across

lower anal lip. Last antennomere with 2-4 dorsal sensory

spots...Ligyrus(Ligyrodes)

3.Innermarginofleftmandible(betweenscissorialandmolar

areas)withasmalltooth(S4)...4

3.Innermarginofleftmandiblebetweenscissorialandmolar

areassmoothorslightlyirregularbutnotdentate...8

4.Lastantennomerewith2dorsalsensoryspots...6

4.Lastantennomerewith5-12dorsalsensoryspots...5

5.Ocelliabsent.Lastantennomerewith5-6dorsalsensoryspots.

Maximumheadcapsulewidth4.7mm...Neoryctes

5.Ocellipresent.Lastantennomerewith12dorsalsensoryspots.

Maximumheadcapsulewidth10.9mm...Bothynus

6. Respiratory plates of abdominal segments I-VIII similar in

size...7

6.RespiratoryplatesofabdominalsegmentsIandVIIIslightly

smaller than those of segments II-VII, which are similar in

size._._._.._._.._._.._._.._._.._._.._._.._._.._._.._._..._._.._._.._._..._._.._._.._._...Ligyrus(s.str.)

7. Color of cranium yellowish-brown, surface with numerous

smallpits.Eachsidewith1prominent,exteriorfrontalseta.Ocelli

vague...Oxygrylius

7.Colorofcraniumreddish-brown,surfacedeeplyandsparsely

punctate. Each side without exteriorfrontal setae.Ocelli well

deﬁned...Coscinocephalus

8.Lastantennomerewith1dorsalsensoryspot.Maxillary

stridu-latoryteethwithanteriorlyprojectingpoints...10

8. Last antennomere with 4-8 dorsal sensory spots.

Maxil-lary stridulatory teeth truncate, without anteriorly projecting

points...9

9.RespiratoryplatesofabdominalsegmentsI–VIIInearlysimilar

insize.Lastantennomerewith4–5dorsalsensoryspots.Frons

withmanysecondaryshortsetae...Philoscaptus

9.RespiratoryplatesofabdominalsegmentsI–IVsimilarinsize,

thoseofabdominalsegmentsV–VIIIprogressivelysmaller

poste-riorly.Lastantennomerewith6–8dorsalsensoryspots.Fronswith

scarcesecondaryminutesetae...Saccharoscaptus

10. Respiratory plate with 13-25 oval micro-holes

across any diameter. Maximum width of head capsule

6.0–6.5mm...Orizabus

10. Respiratoryplate with12–20 elongate,irregularly shaped

micro-holesacrossanydiameter.Maximumwidthofheadcapsule

3.8–5.2mm...Aphonus

Saccharoscaptuslaminifer(Dechambre,1979).Third-instarlarva

(Figs.1–14)

Head (Fig. 1): Width of head capsule 7.0–7.3mm. Surface

reddish brown, moderately rugo-punctate. Frontal suture and

clypeofrontalsuturedistinct.Frons(Fig.1):with2exteriorfrontal

setaand1posteriorfrontalsetaoneachside;eachanteriorangle

offronswith1seta,andanteriorfrontalsetaeabsent;remaining

cranial surface with 3–5 dorso-epicranial setae, 4–6 epicranial

setae,4–6para-ocellarsetaeoneachside.Ocelliabsent.Clypeus:

Formtrapezoidal,with2lateralsetaeateachside,withoutcentral

setae. Surface of postclypeus reddish-brown, well sclerotized

and rugo-punctuate;surface of preclypeuslightbrown. Labrum

rugo-punctate,slightly asymmetrical,2posteriorsetae,3–4

lat-eralsetaeoneach sideand 2centralsetae.Epipharynx (Fig.2):

formsuboval,widerthanlong,asymmetrical,bothlateraledges

rounded.Haptomeralprocessofepipharynxprominentandentire;

rightchaetoparia with45–50stout setae; leftchaetoparia with

30–35stoutsetae,withoutsensillae;gymnopariawide,withsome

scatteredslender,smallsetae;acropariawith8–10straight,long,

thicksetae;rigthacanthopariawith12–14short,curved,

spine-likesetae;leftacanthopariawith10–11short,curved,spine-like

setae;pedium wide,elongate. Dexiotormanarrowand sinuose;

laeotorma narrow,straight, shorter than dexiotorma; epitorma

curvatetowardcenterofpedium; pternotormarounded.

Dexio-phobaabsent;laephobaclearly developedbetweenhaptolechus

andinnersideoflaeotorma,formedby12–16longslenderﬁmbriae.

Sclerotizedplateofrightnesium,nearlyfusedwithdexiotorma,

triangular,withacuteapex;senseconeonleftnesiumrepresented

bylongitudinal,slightlycurved,well-sclerotizedplate,apexwith

3–4sensillascarcelydeﬁned.Crepiswell-marked.Rightmandible

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1 1 mm 2 mm 1 mm 1 mm 1 mm 0.5 mm 0.5 mm 0.5 mm 5 4 2 3 9 12 11 10 8 8a 6 7

Figs.1–12.ThirdinstarofSaccharoscaptuslaminifer.1,head,frontalview.2,epipharynx.Mandibles,ventralview:3,right.4,left.Lastantennomere:5,dorsal.6,ventral.

7,detail,maxillarystridulatoryteeth,lateralview.8,labium,hypopharynxandrightmaxilla,dorsalview.8a,detail,unciatapexoflacinia,mesialview.9,prothoracic respiratoryplate.10,protarsalclaw.11,mesotarsalclaw.12,metatarsalclaw.

roundedtooth(S3)afterscissorialnotchweaklydeﬁned;scrobe

with3–4 stout setae.Dorsal surface with row of 7–8 slender,

longsetae. Ventralsurface withelongate-oval stridulatoryarea

formedby14–16narrowlyseparatedridges;ventralprocess

well-developed,curved,rounded.Brustiawith2–3stout,longsetae.Calx

large,2basolateralsetae.Molarareawith3wide,convex,ridged

lobes(M1-3)withoutsetae.Leftmandible(Fig.4)Scissorialregion

with3 teeth;teeth 1and 2vaguely separatedfromtooth3 by

scissorialnotch.Scrobewith5–6slender,longsetae.Dorsalsurface

withrowof7–9slender, longsetae;aciawell-developed,

trian-gular, and apicalsetae absent,11–12 basolateral setae.Ventral

surfacewithelongate-ovalstridulatoryareaformedby15–17

nar-rowlyseparatedridges;ventralprocesswell-developed,rounded;

dorsomolarareawithrowof 6 slender,moderately longsetae;

brustiawith4–5stout,shortsetae.Molarareawith2lobes,ﬁrst

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10 mm

1 mm

13 14

Figs.13–14. ThirdinstarofSaccharoscaptuslaminifer.13,lateralviewoflivinglarva.14,raster,ventralview.

subrectangular.Stipes largerthanwide. Galeawithmany stout

setae and 1 well-developed uncus at apex. Lacinia with many

stout setae and apicaltransverse row of 3 unci(middle uncus

shorterthanothers),basallyfused(Fig.8a).Maxillary palpus

4-segmented,segment4aslongassegment2.Stridulatoryareawith

7blunt,truncaterigdesandanteriortruncateprocess(Figs.7–8).

Hypopharynx(Fig.8).Glosawith20–55 slender,long setaeand

14–18short setae.Hypopharyngealsclerome asymmetrical,

con-cavemedially,dentiformprocessproduceddorsallywithrounded

apex; left lateral lobe without setae; right lateral lobe with 4

slender,moderaterlylongsetae.Leftmarginwithrowof10–12

moderatelylongﬁmbriaedirectedtowardcenterofscleromeand

rowof5–6ﬁmbriaonrightmargin.Antenna:with4antennomeres,

1to3nearlywithsamelengtheachtoother;apicalantennomere

twicelongthanantennomere1.Apicalantennomerenearlyoval

indorsalorventralview;dorsalsurfacewith6–8sensoryspots

(Fig.5);ventralsurfacewith4–5sensoryspots(Fig.6).

Thorax(Fig.13):Pronotalscleromewide,irregularlyandweakly

sclerotized, with 4–5 slender, long setae. Prothoracic spiracle

(Fig. 9)0.25mm long,0.20mm wide;respiratory plate reddish

brown,regularlyshapedasaclosed“C”,bullaslightlyprominent,

convex;distancebetweenrespiratorylobesmuchlessthan

diame-terofbulla;platewith25–30microholesacrossdiameteratmiddle,

holeswithirregularedges(400×).Dorsumofprothoraxwith3–4

irregulartransverserows of 24–38long, slendersetae,without

spiniformsetae.Mesoprescutumwithtransverse,irregularrows

of16–22long,slendersetae;mesoscutellumwithtransverserows

of14–18long,slendersetae.Metaprescutumwithirregularrows

of26–34long,slendersetae;metaescutellumwithrowsof18–22

long,slendersetae.Legs:Tarsalclawswithenlargedapicalprocess,

1basoexternalseta,and1internal,preapicalseta(Fig.10).Tarsal

clawonmesothoraciclegslightlylongerthanthoseofprothoracic

leg(Fig.11).Tarsalclawofmetathoraciclegshorterthanclawsof

precedentlegs(Fig.12).Coxa,trochanter,andtibiotarsusofalllegs

withmanyslendersetae.

Abdomen(Fig.13):abdominalspiraclesI–IV2.5mmlongand

2.0mmwide.SpiraclesonsegmentsV–VIequalinsize(2.3mm

longand1.9mmwide);spiracleonsegmentVIIslightly smaller

thanpreceding(2.2mmlongand2.0mmwide)andspiracle on

segmentVIIIsmallerthanallpreceding(1.8mmlongand1.8mm

wide).AbdominalsegmentIonprescutumwithirregular

trans-verserowsof10–14long,slendersetaeand20–26short,spiniform

setae;subscutumwith10–12long,slendersetae,without

spini-formsetae;scutumwithabout38–44short,spiniformsetaeand

18–20long,slendersetae;scutellumwithabout10–14short,

spini-formsetaeand26–34long,slendersetae.Allspiracularareaswith

14–16long,slendersetae.AbdominalsegmentsII–VIonprescutum

withirregularrowsof14long,slendersetaeand28short,

spini-formsetae;subscutumwith14–16longsetae,withoutshortsetae;

scutumwith30–32long, slendersetae and24–28 short,

spini-formsetae;scutellumwith16–18long,slendersetaeand50–54

short,spiniformsetae.AbdominalsegmentVIIwith1transverse

rowof22–26slender,longsetaeand40–48short,spiniformsetae

mixed.AbdominalsegmentVIIIwithirregulartransverserowsof

18–20slender,longsetae,withoutspiniformsetae.Abdominal

seg-mentIXwith2irregularrowsof10–12slender,longsetae,without

spiniformsetae.AbdominalsegmentXwithapproximately40–48

moderate tolong, slendersetae.Pleurallobeswith18–24long,

slendersetae.Raster:withoutpalidia(Fig.14)campuswith6

slen-der,longsetae;tegesformedwith64–72shortsetae,barbulawith

14–18long,slendersetae.Analslitwidelycurve,transverse.Lower

anallipwith80–94stout,shortandmediumsizesetaeondisk,

and76–84stout,longsetaealongdistalborder.Upperanallipwith

90–110stout,shortand mediumsizesetae.Approximatedorsal

bodylength38–52mm.

Remarks.ThefollowingcharacterswillseparatethelarvaeofS.

laminiferfromthoseofallotherknownAmericanDynastinae:

ante-riorfrontalsetaeabsent;externalfrontalsetaepresent;posterior

frontalsetaepresent;clypeus withoutcentralsetae;last

anten-nomerewith6–8dorsalsensoryspots;ocelliabsent;respiratory

platesofabdominalsegmentsV–VIIIprogressivelysmaller;raster

withoutseptulaor palidia;lower anallip withnumerous stout

setae,includinglongsetaethatformafringealongthedistalborder.

Studiedspecimens.Tenthird-instar larvaeaspartofalarge

sampleoflarvaerearedtoadultstage,collectedinBolivia,Santa

Cruz,Warnes,LaJupia,308m, 10-VII-2011,F.CopaBazán(IIAV,

IEXA).

Distribution.ThisspeciesisfoundinsouthernBraziland

east-ernBolivia(MorónandGrossi,2015).

Biology. S. laminiferis a yearly cycle species. Larvae live in

rich organicmattersoils,and duringfourmonths mayfeedon

the roots of sugarcane cultures. In Santa Cruz, Bolivia,

ovipo-sition occurs between December and March,ﬁrst instar larvae

hatchinFebruary–March,secondinstarlarvaearegrowingfrom

March to May, and third instar larvae are active from May

to July; the complete larval development lasts 130–158 days.

Inside a cell build withsoil particles compacted with its own

fecal remains, prepupa requires 5–8 days and pupa another

21–31days.Whentheadultisformeditremainsinsidethecell

for 3–4 months. The male and female emergence and mating

occursbetweenNovemberandDecember(Copa-BazanandMorón,

2014).

Probably,theverylargenumbersoflarvaeobservedinthe

sugar-caneﬁeldsofSantaCruz,Boliviaduring2011(150–200larvae/m2₎

representtheunusualgrowthofpopulationsofwhitegrubsderived

fromsomecombinationoffacts,suchas,scarcelocalrains,frequent

ﬁresingrasslandsandsugarcaneﬁelds,useofcompostas

fertil-izer,deforestationofsurroundinglandsandabuseofinsecticides

appliedtosoilduringmanyyears(Copa-BazanandMorón,2014).

Themostinterestingobservationisthatthisspeciesknownfornear

acenturyfromtheholotype,unexpectedlyappearsinthousandsin

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Conﬂictsofinterest

Theauthordeclaresnoconﬂictsofinterest.

Acknowledgements

FernandoCopaBazan(IIAV,SantaCruz,Bolivia)collectedand

breedthestudiedspecimens.JorgeFrana(INTA,Rafaela,Argentina)

transported the samples. Paschoal Grossi and one anonymous

reviewer provided helpful comments to a previous version of

thismanuscript.Thispaperisacontributiontotheresearchline

“SistemáticayBiologíadeColeópterosLamelicorniosdeAmérica

Latina”supportedbyInstitutodeEcología,A.C.Xalapa,Mexico

(account20030-10011).

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