Studies on Brazilian Pseudophasmatidae (Phasmatodea) with the description of a new species of Agrostia Redtenbacher and new records for Metriophasma Uvarov and Parastratocles Redtenbacher

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REVISTA

BRASILEIRA

DE

Entomologia

A Journal on Insect Diversity and Evolution

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

Systematics,

Morphology

and

Biogeography

Studies

on

Brazilian

Pseudophasmatidae

(Phasmatodea)

with

the

description

of

a

new

species

of

Agrostia

Redtenbacher

and

new

records

for

Metriophasma

Uvarov

and

Parastratocles

Redtenbacher

Raphael

Aquino

Heleodoro

∗

,

Diego

Matheus

de

Mello

Mendes,

José

Albertino

Rafael

InstitutoNacionaldePesquisasdaAmazônia(INPA),ProgramadePós-Graduac¸ãoemEntomologia,Manaus,AM,Brazil

a

r

t

i

c

l

e

i

n

f

o

Articlehistory:

Received3October2016 Accepted13March2017 Availableonline24March2017 AssociateEditor:DanielaTakiya Keywords: Phasmida Phasmids Stickinsects Taxonomy

a

b

s

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Pseudophasmatidaecurrentlyincludes65speciesrecordedfromBrazil.Inthispaper,weincreasethis numberto68speciesbydescribingAgrostiaﬂavimaculatasp.nov.,basedonmaleandfemale speci-mensfromAcreandAmazonasstates.Furthermore,Parastratoclesmultilineatus(Rehn)andMetriophasma (Metriophasma)diocles(Westwood)arerecordedfortheﬁrsttimefromBrazilbasedonspecimensfrom Rondônia,AcreandAmazonasstates.AkeytospeciesofAgrostiaisalsogiven.

Introduction

Phasmatodeacurrentlyincludesapproximately3000described species (Brock et al., 2016), of which 207 occur in Brazil. AmongBrazilianspecies,65belongtoPseudophasmatidae. Agros-tiaRedtenbacher,1906isoneofthePseudophasmatidaegenera recordedfromBrazilandcurrentlyincludesthreevalidspecies:A. amoenaRedtenbacher,1906describedfromPanamá,A.bipunctata

Redtenbacher,1906describedfromBrazil(Amazonas,Coari)and A.ega(Westwood,1859)describedfromBrazil(Amazonas,Tefé, formerlyknownasEga).BesidesAgrostia,MetriophasmaUvarov, 1940,andParastratoclesRedtenbacher,1906alsooccurinBrazil. Metriophasmaincludes13validspecies,with4speciesrecorded fromBrazil,while Parastratocleshas12valid species,withonly Parastratoclesﬂavipes(Redtenbacher,1904)currentlyknownfrom Brazil.

Inthispaper,wecontributetotheknowledgeofBrazilian Pseu-dophasmatidaeby describing a newspecies of Agrostia and by recordingfortheﬁrsttimeMetriophasma(Metriophasma)diocles (Westwood, 1859) andParastratocles multilineatus(Rehn, 1904)

∗ Correspondingauthor.

E-mail:raphaelnatal36@gmail.com(R.A.Heleodoro).

fromBrazil,aswellasproviding biologicalinformationofthese species.

Materialandmethods

AllstudiedspecimensarepinnedanddepositedatColec¸ãode InvertebradosdoInstitutoNacionaldePesquisasdaAmazônia(INPA). Identiﬁcationofspecimenswasmadewiththefollowingliterature:

Redtenbacher(1906),Uvarov(1940),andZompro(2004). Speci-menswerecomparedwithpicturesofthetypematerialavailable from the Phasmida species ﬁle http://phasmida.speciesﬁle.org/

(Brocketal.,2016).SpecimensofAgrostiawerecollectedatnight, onawhitesheetilluminatedwitha250Wmercuryvaporlamp afterspecimenswereattractedtothelight.Anotherfemale spec-imenwaspreviouslydepositedatINPAcollection.Specimensof MetriophasmaandParastratocleswerefoundinthevegetationby activelysearchingduringnighttime.Specimenswerekilledwith ethylacetate(EtOAc).Photosoflivespecimensweretakenwith aNikonD3000digitalcamera,withan18–55mmlens.Photosof pinnedspecimensweremadewithaLeicaDFC500digitalcamera ﬁttedona LeicaMZ205 stereomicroscopeconnected toa com-puterwithLeicaApplicationSuitesoftware.Thissoftwareincludes anAuto-Montagemode(Syncroscopysoftware)usedtocombine multiplelayersofphotosintoasinglefocusedimage.Labeldata http://dx.doi.org/10.1016/j.rbe.2017.03.003

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isgivenaspresentedonthelabels.Squarebrackets([])indicate complementarydatanotpresentonspecimenslabels.

Taxonomy

Stratocleinae:Stratocleini AgrostiaRedtenbacher,1906

Comments.ThenewspeciesisplacedinAgrostiabasedonthe

followingdiagnosticcharactersprovidedbyRedtenbacher(1906): metathoraxdistinctlyshorterthanmesothorax;tegminaellipsoid;

forefemurcurvedatbasalthird;andforefemurandhindfemur dorsally bicarinated.Additionally, Agrostia is theonly genus in Stratocleiniwiththefollowingcombinationofcharacters:head, proandmesonotumopaque;mesonotumatleasttwotimeslonger than pronotum; tegmina elongated,longer than large, dorsally withconspicuousspotofvariablecoloration;costalareaofalae withspotsconcolor tospots integmina;and alaenot reaching tergum8.

Agrostiaﬂavimaculatasp.nov.

(Figs.1–17)

Figs.1–9.Agrostiaﬂavimaculatasp.nov.holotype♂.(1)Habitus,dorsalview.(2)Idem,lateralview.(3)Idem,ventralview.(4)Headandprothorax,dorsalview.(5)Head, lateralview.(6)Tegmina,dorsalview.(7)Apexofabdomen,dorsalview.(8)Idem,lateralview.(9)Idem,ventralview.

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Diagnosis.Ocelliorange (Figs. 1,4, 10).Tegmina with ellip-soidyellowspotatmedianregion(Figs.1,3,6,10,13,14).Cercus laterallyﬂattened(Figs.7–9;15–17).Malesubgenitalplate longi-tudinallydividedintohemisternitesapically(Fig.9).

Description.Holotype♂ (Figs. 1–9).Head. Dorsally smooth,

lightbrown,withconcavityatvertex;coronalsutureconspicuous, black(Fig.4).Fronswithblackarchedspots(Fig.4).Vertexwithtwo parallelarchedpalestripes(Fig.4).Genalightyellow(Figs.2,5). Clypeus trapezoidal. Labrum U-shaped. Ocelli ellipsoid, orange (Figs. 1,4).Eyes globose,lightbrown, withlongitudinalarched stripe(Fig.5).Antennacoveredbysetae,dorsallyblack,ventrally light brown; ﬁrst ﬂagellomere 2 times longer than pedicellus (Fig.4).Maxillaryandmandibularpalpiyellow,shiny,coveredby setae.

Thorax. Pronotum rectangular, 1.5 times longer than wide, withlongitudinal black and light yellow stripes;latero-distally

with small light yellow spots (Fig. 4). Mesonotum rectangu-lar, slender, elongated, 6 times longer than wide and 3 times longerthan pronotum; withlongitudinal parallel stripes, alter-natingblackandlightyellow;blackstripeswithsmallscattered light yellow spots (Figs. 1 and 2).Metanotum rectangular, 2.5 timesshorterthanmesonotum,shiny,smoothandlight yellow. Prosternumtrapezoidallight yellow,withsmallrounded tuber-cles.Mesosternumrectangular,slender,elongated,6timeslonger thanwide,lightbrownwithseveralscatteredsmalllightyellow tubercles and no visible median longitudinal carina. Metaster-numrectangular,elongated,broad,3timeslongerthanwide,1.5 timesshorterthanmesosternum;withsurfacerough,lightbrown (Fig.3).

Legs.Alllegslaterallycoveredbysetae;withfemoraandtibiae light yellow and with small black spot at apex of femora and baseof tibiae;(Figs.1–3).Midand hindtarsiwiththird toﬁfth

Figs.10–17.Agrostiaﬂavimaculatasp.nov.♀.(10)HabitusofAcrespecimen,dorsalview.(11)Idem,lateralview.(12)Idem,ventralview.(13)Tegminaofspecimenfrom Acre,dorsalview.(14)TegminaofspecimenfromAmazonas,idem.(15)Apexofabdomen,dorsalview.(16)Idem,Lateralview.(17)Idem,ventralview.

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Figs.18–20. Agrostiaamoenasyntype♂.(18)Habitus,dorsalview.(19)Tegmina,lateralview.(20)Apexofabdomen,lateralview.PhotosweretakenbyPaulBrock,copyright NaturhistorischesMuseum,Vienna.

Figs.21–23. Agrostiabipunctatalectotype♂(21)Habitus,dorsalview.(22)Tegmina,lateralview.(23)Apexofabdomen,lateralview.PhotosweretakenbyPaulBrock, copyrightNaturhistorischesMuseum,Vienna.

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Figs.24–26.Agrostiaegaholotype♀.(24)Habitus,dorsalview.(25)Tegmina,lateralview.(26)Apexofabdomen,lateralview.PhotosweretakenbyPaulBrock,copyright NaturalHistoryMuseum,London.

tarsomeresgraduallyturninglightbrown.Areaapicalisinalltibiae conspicuous.

Wings(Fig.6).Tegminaellipsoidal,elongated,3timeslonger thanwide,brown,withsmallblackspotatshoulderhump; longi-tudinalandtransverseveinsyellow;withellipsoidyellowspotat medianregion.Alaeapproximately5timeslongerthantegmina, withcostalareadarkbrown,analareahyaline;longitudinalveins of costal area light yellow, transversal veins of anal area light brown.

Abdomen.Elongated, slender. Abdominal terga dorsally dark brownfrombaseoftergum1 tobasalthird oftergum6.Terga 1–8rectangular.Tergum7rectangular,inlateralview4.3times longerthanhigh;indorsalview1.3timeslongerthantergum8, withdeeplongitudinalmediansulcus.Tergum8rectangularwith deeplongitudinalmediansulcus,inlateralview3.4timeslonger thanhighand1.3timeslongerthantergum9(Figs.7and8). Ter-gum9rectangular,1.2timeslongerthanwide,withdeepmedian longitudinalsulcus;trapezoidalinlateralview,withventral mar-ginconvex,posterior margin emarginated,2 times longerthan tergum10(Figs.7and8).Tergum10 dorsallyﬂattened, anteri-orlyemarginated;lateralmarginsinuous,broadeningfrombaseto apex;posteriormarginconvex;laterallycoveredbysetae(Fig.7). Abdominalsternadarkbrown,withsmall-scatteredyellowspots (Fig.3)withmedianlongitudinalsulcusfromsternum2to8.Sterna 2–7elongated,longerthanwide(Fig.3).Subgenitalplatecovered withsetae,ellipsoidal,longitudinally dividedintohemisternites apically,being1.5timeslongerthantergum9(Fig.9).Paraprocts ellipsoidal,elongated,notcoveredbysubgenitalplate.Cercuslight brown, covered with setae, moderately incurved, laterally ﬂat-tened,withpointedapex(Figs.7–9).

Measurements (mm). Holotype: body length 62.5;

antenna>55.0;dorsalheadlength3.8pronotum2.7;mesonotum 6.4;forefemur10.5;foretibia11.6;midfemur6.4;midtibia5.7; hindfemur10.4;hindtibia10.5;tegminalength7.3;alaelength 31.9.

Paratype: body length 56.3; dorsal head length 3.1; antenna>55.0; pronotum 2.5; mesonotum 6.1; fore femur 10.4;foretibia11.6;midfemur6.1;midtibia5.5;hindfemur10.0; hindtibia10.1;tegminalength6.3;alaelength34.4.

Female(Figs.10–17).Similartomale,butlongerandrobust.

Followingdifferenceslistedbelow.

Head. Slightly quadrate in dorsal view (Fig. 10). Gena dark brown (Fig. 11). Yellow spot at tegmina longer and broader (Figs.13and14).

Abdomen.Tergum7inlateralview2.8timeslongerthanhigh (Fig. 16).Tergum 8 in lateral view1.6 times longerthan high (Fig.16).Tergum 9trapezoidal(Fig.15).Tergum10 coveredby setae,indorsalviewoblong,inlateralviewwithbasalandlateral marginsstraight,posteriormarginemarginated(Figs.15and16). Subgenitalplatelaterallycompressed,graduallybroadeningfrom anteriormargintowardapex,withposteriormarginconcave,being 1.2timeslongerthantergum9(Figs.11,16,17).Cercustriangular, nearlystraight.

Female measurements (mm). Acre specimen: body length

87.0; dorsal head length 5.2; antenna>40.0; pronotum 4.6; mesonotum9.0;forefemur12.0;foretibia13.3;midfemur10.0; midtibia9.7;hindfemur15.4;hindtibia15.2;tegminalength12.0; alaelength51.6.

Amazonasspecimen:bodylength75.5;dorsalheadlength4.8; antenna>40.0;pronotum4.4;mesonotum8.6;forefemur12.0;

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Figs.27–31. Parastratoclesmultilineatus♀fromRondônia.(27)Habitusoflivespecimen,dorsolateralview.(28)Habitusofdeadspecimen,dorsalview.(29)Apexofabdomen, dorsalview.(30)Idem,lateralview.(31)Idem,ventralview.

foretibia13.2;midfemur9.7;midtibia9.4;hindfemur15.1;hind tibia15.0;tegminalength10.3;alaelength49.6.

Variations. Femalespecimens fromAmazonas have a larger

yellowspot,withmore deﬁnededges incomparison tofemale specimensfromAcre(Figs.13and14).

Etymology.FromLatinadjectiveﬂavus=yellowandnoun

mac-ula=spot,referringtotheyellowspotonthetegmina.

GeographicalRecords.Brazil:Amazonas,Acre.

Typematerial.Holotype♂(INPA).BRASIL,AC[Acre],Bujari,

FES[FlorestaExtrativistaSustentável]Antimary.25–27.viii.2016. 09◦2002S-68◦1915W, Arm [armadilha] luz, AA Agudelo, FF Xavier,DMMMendes,JARafael[cols.].

Paratypes. Same data as holotype (1♂, ♀ INPA); Brasil, Amazonas, Manaus, Reserva Ducke, 02◦5548S 59◦5830.6W. 04.v.2014.Coletamanual.G.Gomes,col.(1♀INPA).

KeytospeciesofAgrostia

1.Tegminawithyelloworwhiteellipsoidalspot(Figs.6,13,19,22)...2 1.Tegminawithoutellipsoidalspot,insteadwithgreenirregular-shapedspot(Figs.24–26).Brazil(Amazonas)...A.ega(Westwood,1859) 2.Spotontegminayellow,startingatmedianthirdandtouchingtheantero-lateraltegminahump(Figs.6,13,19)...3 2_._Spot_on_tegmina_white,_starting_at_apical_third_and_not_touching_the_{antero-lateral}_tegmina_hump₍_Figs._21–23_)._Brazil

(Amazonas)...A.bipunctataRedtenbacher,1906

3.Spotontegminarangingfrommedianthirdtohalflengthofapicalthirdofthetegmina.Cercicylindricalventrallycurved(Figs.18–20).

Panamá...A.amoenaRedtenbacher,1906

3_._Spot_on_tegmina_not_surpassing_the_median_third_of_the_tegmina._Cerci_laterally_ﬂattened₍_Figs._1–17_)._Brazil_(Acre_and_Amazonas)_._._.._._.._._.._._._A._{ﬂavimaculata}_sp._nov.

Parastratoclesmultilineatus(Rehn,1904) (Figs.27–31)

Comments.Thisspeciesisrecognizedbytheyellowspoton

theanalareaof thealae(Figs. 27 and28).Colorationonliving insectscanvarysigniﬁcantlyfromspecimensknownfromCosta Rica.Theoriginalcolorationisnearlycompletelylostafterdrying process,turningitmostlydarkyellowandblack(Fig.28). Charac-tersonfemaleterminaliaareonlyavailablefromRehn(1904)and

Redtenbacher(1906),withlimitedinformationandwithout illus-trations.Duetothis,wecharacterizethefemaleterminaliabased onspecimenfromRondônia,Brazil.

Description ♀ terminalia. Terga 8–10dorsally withmedian

longitudinalblack stripe(Figs.28 and 29).Terga8–9 rectangu-larindorsalview,trapezoidalinlateralview,allmarginsstraight (Figs.29and30).Tergum10withanteriormarginstraight,slightly

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Figs.32–33.Metriophasma(Metriophasma)diocles♀fromAmazonas.(32)Habitusindorsolateralview.(33)Defensivestance.

sinuouslaterallyandposteriormarginemarginated;inlateralview withanterior,dorsalandposteriormarginsstraight,ventral mar-ginsinuous(Figs.29and30).Subgenitalplatedenselysetose,with medianlongitudinaldarkbrowncarina;swordshaped,narrowing neartheposteriormargin;apexoblong,being1.4timeslongerthan tergum9(Fig.31).Cercusconical,coveredbysetae,withoblong apex.

Biological information. The specimen was found in an

ombrophilousTerraFirmeAmazonianForest,onbushesof approx-imately1.6mhigh.Whenstressed,theinsectopeneditswings, makingtheaposematic colorationmore visible.Furthermore, it alsosprayedastronglystinky(resemblinganherbaceoussmell) substance.

Geographical records. Costa Rica; Brazil [new record]:

Rondônia.

Materialexamined.BRASIL,RO[Rondônia],CandeiasJamari,

UHE Samuel, 17.viii.2016, 8◦5657S–63◦1060W, manual [col-lecting],DMMMendes,FFXavier,AAAgudelo,JARafael[leg.](1♀, INPA).

Xerosomatinae:Prexaspini

Metriophasma(Metriophasma)diocles(Westwood,1859) (Figs.32–36)

Comments. This species is easily distinguished by having

blue spots near base of the alae (Fig. 33). We recorded two coloration patterns for this species: one with body mainly

light yellow (Figs. 32, 33, 36) and another mainly dark brown (Figs.34and35).

Biological records. D. M. M.Mendes tookphotos fromthe

lighterphenotype(Figs.32and33)in2013,atanombrophilous TerraFirmeAmazonianForestinAmazonasState(AM,Brazil),but didnotcollectthespecimen.Thisyear,thesameauthorfound dif-ferentspecimensofsamespeciesinAcreState(AC,Brazil)with lighteranddarkerphenotypes(Figs.34–36)inanombrophilous TerraFirmeAmazonianForestwithbamboos,andcollectedtwo femalesandonemalespecimen.

Both populations (AC/AM) were found on bushes, approxi-matelybetween1.4and2.0mhigh.Whenresting,theseinsects concealtheirpresencewithacamouﬂagebehaviorsimilarto mem-bersofPrisopodidae,whichconsistsofjoiningalllegsneartoits bodytodecreasebodywidth(Fig.36).Whenstressed,thisspecies performedanagonisticdisplaybyopeningandclosingtheirwings repeatedlyshowingtheblueandblackspotsonwings,possibly confusingapotentialpredator.

Geographicalrecords. Colombia;Costa Rica;Panama;Brazil

[newrecord]:Acre,Amazonas.

Examinedmaterial.BRASIL,AM[Amazonas],Manaus,ZF-2,km

34[Specimennotcollected,examinedonlybyphoto].

BRASIL, AC [Acre], Bujari, FES [Floresta Extrativista Susten-tável] Antimary. 25–27.viii.2016. 09◦2002S-68◦1915W, Arm [armadilha]luz,AAAgudelo,FFXavier,DMMMendes,JARafael [cols.](2♀,1♂INPA).

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Figs.34–36.Metriophasma(Metriophasma)dioclesfromAcre.(34)♂Habitus,dorsalview.(35)Livedarkbrownphenotypefemaleinlateralview.(36)Livelightyellow phenotypefemaleindorsalview,displayingcamouﬂagebehavior.

Conﬂictsofinterest

Theauthorsdeclarenoconﬂictsofinterest.

Acknowledgments

ToAlbertodaSilvaNeto,EmilyOlive,andJoãoRafaelOliveira for their support in the laboratory. To Conselho Nacional de DesenvolvimentoCientíficoeTecnológico(CNPq)fortheresearch productivity fellowship to J.A.R. (grant 300.305/2007–9) and for financial resources from Chamada MCTI/CNPq/FNDCT Ação Transversal – Redes Regionais de Pesquisa em Biodiversidade e Biotecnologia N◦ 79/2013 (grant 407.627/2013-8), Fundação deAmparoàPesquisa doEstado do Amazonas(FAPEAM, grant

016/2006); and Programa de Apoio a núcleos de Excelência (PRONEX,proc.1437/2007).

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VerlagW.Engelmann,Leipzig,Germany.

Rehn,J.A.G.,1904.StudiesintheOrthopterousFamilyPhasmidae.Proc.Acad.Nat. Sci.Phila.56,38–107.

Uvarov,B.P.,1940.TwentyfournewgenericnamesinOrthoptera.Ann.Mag.Nat. Hist.11(6),112–117.

Westwood,J.O.,1859.CatalogueofOrthopterousInsectsintheCollectionofthe BritishMuseum.Part1,Phasmidae.London,BritishMuseum.

Zompro,O.,2004.RevisionofthegeneraoftheAreolatae,includingthestatusof TimemaandAgathemera(Insecta,Phasmatodea).Abhandlungendes Naturwis-senschaftlichenVereinsHamburg37,1–327.