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Description of the larva ofArgia percellulata(Odonata: Coenagrionidae) PDF

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InternationalJournalofOdonatology Vol.15,No.1,March2012,45–50 Description of the larva ofArgia percellulata (Odonata: Coenagrionidae) RodolfoNovelo-Gutiérrez*andJoséAntonioGómez-Anaya InstitutodeEcología,A.C.ReddeBiodiversidadySistemática.CarreteraantiguaaCoatepec351,El Haya,Xalapa,Veracruz,Mexico (Received14December2011;finalversionreceived10January2012) ThelarvaofArgiapercellulataisdescribedandfigured.ItfallsintothegroupofArgialarvaewithavery prominentligulaandonepalpalseta,butdiffersfromitsclosestrelativesbyhavingtibiaeusuallywithtwo well-defineddarkrings,posteriormarginofsternite6smooth,andposteriormarginofsternite7smooth mediallyandwithspiniformsetaelaterally.Larvaewerefoundundercobblestonesinopen,wide,shallow streamswherethewaterflowwasslowtomoderate,closetotheshorelinewithinacloudforest.Thelarva iscomparedwithfourotherspeciesapparentlycloselyrelated. Keywords: Odonata;Zygoptera;damselfly;Argiapercellulata;larvaldescription;Mexico Introduction Argia percellulata Calvert is an endemic Mexican species with a restricted distribution thus far known only from the states of Chiapas, Puebla, Oaxaca and Veracruz (González-Soriano & Novelo-Gutiérrez, 2007; Garrison, unpublished data), ranging from 380m at RíoTecpatlán, Zihuateutla,Puebla(Cuevas-Yañez,2007),to1980matthevicinityofSierradeJuárez,Oaxaca (Garrison, pers. com.). The original vegetation at all localities was cloud forest, although in Coatepectheforesthasbeenlargelyreplacedbycoffeeplantationsandpastureland.Larvaewere notrearedtoemergence,butweareconfidentoftheiridentitybecausetheonlyotherlargespecies ofArgiafoundinCoatepecover20yearsofintensivecollectionsisA.lacrimans(Hagen),thelarva ofwhichisquitedifferentinseveralfeatures(Novelo-Gutiérrez,1992).Moreover,infourmature larvaethewingvenationisquitevisiblebytransparency,revealingsixandfivepostquadrangular cells on forewings and hind wings, respectively, as described by Calvert (1902) for adults of A.percellulata.Finally,twolarvaefromCoatepecwerecapturedinthesamespotwhereasmall breedingpopulationofA.percellulataoccurseveryyear. *Correspondingauthor.Email:[email protected] ISSN1388-7890print/ISSN2159-6719online ©2012WorldwideDragonflyAssociation http://dx.doi.org/10.1080/13887890.2012.660412 http://www.tandfonline.com 46 R.Novelo-GutiérrezandJ.A.Gómez-Anaya Figure 1. Dorsal(a)andventral(b)viewsofF0malelarvaofArgiapercellulata(gillsdetached). Methods Larvae were collected with a D-frame aquatic net and preserved in the field in 96% ethanol. Afterwards, they were transferred to vials containing 80% ethanol. Mandibular nomenclature followsWatson(1956).Drawingsweremadewithaidofacameralucidaonastereomicroscope ZEISSStemiSV6.PhotographswereobtainedwithaCANONPowerShotG10digitalcamera mountedinastereomicroscopeZEISSStemi2000-C.Measurements(inmm)weremadewitha calibratedocularmicrometerasfollows:Headwidth,acrosscompoundeyes;totallength,dorsally fromanteriormostmarginoflabrumtoposteriormostmarginofabdominalsegment10;abdomen length,ventrallytoposteriormostmarginofabdominalsegment10;hindfemora,alongmidline ofexternalsurface.S1–S10=abdominalsegments1to10.F0=lastinstarlarva. LarvaofArgiapercellulata(Figures1–12) Specimensexamined SevenF0larvae(fourmales,threefemales):Mexico:Veracruz,Coatepec,RíoHuehueyapan,La Marina (19◦ 27(cid:3) 09(cid:3)(cid:3) N, 96◦ 58(cid:3) 53(cid:3)(cid:3) W; 1196m), 19April 2000, R.Arce leg (one male); same databut1May2002,R.Noveloleg(onefemale);Puebla,Tecpatlán(20◦14(cid:3)28(cid:3)(cid:3)N,97◦51(cid:3)35(cid:3)(cid:3) W;380m),3May2002,K.Cuevasleg(threemales,twofemales).Allspecimensdepositedat ColecciónEntomológicadelInstitutodeEcología,A.C.(IEXA). Diagnosis Amoderatelyrobustlarva(Figure1a),groundcolorbrownwithlightareas,legswithwidedark bandsseparatedbynarrowpaleareas.Caudallamellaemostlygrayorgrayish-violetinbasalhalf, remainderpalewithanindefinite,transversegrayband. Description Head. Widerthanlong,posteriormarginwidelyconcave,dorsalcolorpatternasinFigure1a. Cephaliclobesroundedwithposterolateralmarginsbearingstoutspiniformsetae.Labrumbrown, DescriptionofthelarvaofArgiapercellulata 47 Figures 2–6. DetailsoflarvalmorphologyofA.percellulata.(2)Leftantenna,dorsalview.(3)Mandibles:(a)rightand (b)leftmandibles,ventrointernalview;(c)rightand(d)leftmandibles,externalview.(4)Hypopharynx,ventralview. (5)Ventralviewofleftgaleolacinia.(6)Labium:(a)prementum,dorsalview;(b)endhooksandmovablehookofleft palp,frontalview;(c)rightpalp,dorsalview. setose,setaewhitish;clypeustrapezoidal,brown,withapairofparallel,narrowwhitelinesalong middlethird.Antennaeseven-segmented(Figure2),scarcelylongerthanhead,scapeshortand pale, remainder of antennomeres brown; proportional length of antennomeres (from basal to apical): 0.4, 0.55, 1.0, 0.6, 0.45, 0.25, 0.15. Mandibles (Figure 3) with molar teeth but lacking molarcrest,withfollowingformula:L1’12340ab/R1’1234ya0,bothmandibleswith aventraltransverse,rowofsetaerunningfromexternalmarginofmandibletolevelofinternal border of tooth 4.Ventral pad of hypopharynx subquadrangular with a tuft of 6–7 long, robust setae located subapically to each anterolateral corner, and a transverse row of smaller setae on anterior margin (Figure 4). Maxilla: galeolacinia (Figure 5) with six teeth, three dorsal teeth approximately of the same size, three ventral teeth of differing sizes, apical one the longest, a rowofstiffsetaeprecedingbothventralanddorsalteeth.Labiumyellowishbrown.Prementum- postmentumarticulationreachingposteriorborderofprocoxae.Prementum0.80timesaswide aslong(Figure6a),witharowof17–20spiniformsetaealongdistalhalfoflateralmarginsand agroupof15–21basidorsalspiniformsetaeofdifferingsize;ligulaveryprominent(2.95times wideratitsbasethanlong,measureddorsally),withcloselyset,minuteclaviformsetaeondistal margin.Prementalpalpwithtwoendhooksshorterthanmovablehook,theventral(medial)one longest(Figure6b),internalmarginofpalpfinelyserrate,externalmarginofsmallerendhook with a row of minute, sharp denticles; palp with one long setae basal to movable hook, and an irregularrowof24–26short,stoutspiniformsetaealongdorsalmargin(Figure6c);arowofthree, stoutspiniformsetaeatbaseofpalparticulation. 48 R.Novelo-GutiérrezandJ.A.Gómez-Anaya Figures 7–12. DetailsoflarvalmorphologyofA.percellulata.(7)Sternites7–10showingsetaeandmalegonapophyses. (8) Male gonapophyses, left lateral view; (9) Sternites 8–10 showing setae and female gonapophyses. (10) Female gonapophyses,leftlateralview.(11)Caudallamellae:(a)medianlamella,leftlateralview;(b)laterallamella,external view.(12)(a–d)Malecerci:(a)lateral,(b)dorsolateral,(c)dorsal,(d)ventral;(e)femalecerci,dorsalview.Ce=cercus. Thorax. Pronotaldiskmostlylightbrownwithsomescattered,longsetaeandspiniformsetae on anterodorsal surfaces, lateral margins dark brown, strongly convex, and covered with long setae;propleuradarkbrown.Synthoraxmostlybrownwithawide,palebandonanteriorhalfof mesepimera(Figure1a).Wingsheathsbrown,anteriorpairscarcelysurpassingposteriormargin ofS4,posteriorpairreachingbasalhalfofS5.Legsshort(i.e.,tipofmetatibiaescarcelysurpassing S10whenfullyextended),pro-andmesotibiaelongerthanrespectivefemora;metafemoraand metatibiaesubequal.Femorawithtwolarge,wide,darkbandsconnectedtoeachotherbyathin lineatmiddle,anincompletebasaldarkringwidelyconnectedwithsubbasallargedarkband,the paleinterveningspacesreduced(Figure1a);tibiaewithtwodark,well-definedringsatbasaland middleareas(sometimeswithathird,well-defined,subapicaldarkring),narrowerthanintervening palespaces;dorsalandventralbordersoffemora,externalsurfaceofprofemora,andinternaland external borders of tibiae with robust spiniform setae, these spiniform setae intermingled with long,stiffsetaeondorsalbordersoffemoraandexternalbordersoftibiae;apicointernalborder oftibiaewithsomelarge,robust,reddish,spine-likesetaeintermingledwithabundantlongsetae andtridentatesetae;tarsiwithabundant,yellow,stiffsetaeonventralsurface,andlong,delicate, whitishsetaeondorsum. Abdomen. Wideratbase,reachingmaximumwidthatS3,andthengraduallynarrowingcaudally. Tergiteswithcomplexcolorpattern,butmostlydarkwithapalespotalongmidline,triangular DescriptionofthelarvaofArgiapercellulata 49 ontergites5–9,subquadrangularontergite10,andsubmedian,longitudinalstripetoeachsideof midlineontergites2–9;tergite1pale(Figure1a).LateralmarginsofS1–6withdarkblotchon apicalhalf(Figure1b);lateralmarginsofS1–5slightlyconvex,straightonS6–10;tergites1–4 smooth,5–10withspiniformsetae,scatteredon5andincreasinglyabundantcaudally;atuftof longwhitesetaealongmidlineontergites2–10.Sternites1–6andmiddlethirdof7smooth,lateral thirdsofsternites7–10withspiniformsetae,middlethirdofsternites8–10withclaviformsetae exceptonsmooth,smallcentralareaofsternite10(Figure7).Malegonapophyses(Figures7,8) long,parallel,spatulateandwithtipspointedbutnotacute,reachingposteriormarginofsternite10 (Figure7);ventralborderwith29–31stout,dark,claviformsetae(Figure8).Femalegonapophyses (Figures9,10)exceedingposteriormarginofS10by0.50lengthofS10;lateralvalves,inventral view (Figure 9), with tips pointed, ventral border with 29–30 stout, claviform setae on basal 0.75; in lateral view (Figure 10) with 2–3 spiniform setae on external surface; median valves smooth.Caudallamellae(Figure11)withmoderatelyacuminatetips.Laterallamella(paraproct) (Figure11b)stronglyinflatedonbasal0.55onexternalside,apical0.45laminar,2.5timeslonger thanitswidestpart;alaterobasal,triangular,amber-colored,chitinousplateatbasal0.25oflength oflamellabearingstout,long,spiniformsetae;dorsalandventralborderssmooth,fringedwith delicatewhitesetaeonapical0.40and0.50respectively;mostoflamellagrayorgrayishviolet onbasal0.55,apical0.45whitewithawide,transverse,grayband(Figure11b).Centrallamella (epiproct) (Figure 11a) inflated at basal 0.40 on both sides, almost two times longer than its widestpart;lateral,triangular,amber-colored,chitinousplateextendingtobasal0.25oflengthof lamellaandbearingstoutspiniformsetaeintermingledwithstiffsetae;dorsalandventralborders asdescribedforlaterallamellaandcolorpatternasinFigure11a.Malecercishort(Figure12a–d), globoseindorsalandventralviews(Figure12c,d),roundlypointed.Femalecercitriangularin dorsalview(Figure12e). Measurements (mm). Total length (without caudal lamellae) 15.5–16.3 (n=6); maximum width of head 4.5–4.6 (n=6); hind femur 4.5–4.7 (n=12); abdomen 9.0–10.1 (n=6); paraprocts5.8–6.5(n=4);epiproct4.2–4.8(n=3). Remarks LarvaeofArgiapercellulatawerefoundundercobblestonesclosetoshorelineinexposed,wide (e.g.15m),andshallowstreamswherethewaterflowwasslowtomoderate,withinacloudforest. Discussion ArgiapercellulatafallsintothegroupofArgialarvaewithveryprominentligulaandonepalpalseta (Novelo-Gutiérrez,1992)andshowsagreatresemblancetoA.barrettiCalvert(Novelo-Gutiérrez, 2008), A. harknessi Calvert (Novelo-Gutiérrez, 1992), A. insipida Hagen in Selys (Geijskes, 1943), and A. joergenseni Ris (von Ellenrieder, 2007), all sharing the following features: antennalscapepale(exceptinA.insipida;cf.Geijskes,1943,plate3,figureb);ligulastrongly prominent; premental palp with one seta; femora with dark bands wider than intervening pale spaces; sternite 8 with abundant claviform setae on the middle; male gonapophyses spatulate, bluntly pointed, beset with claviform setae on ventral border; female gonapophyses bluntly pointed, lateral valvae with abundant claviform setae on ventral border; male cerci globose; caudal lamellae inflated basally, with a short acuminate tip, ventral and dorsal borders free of stoutsetae.Ofallthesespecies,A.percellulatahasthegreatestnumberofbasidorsalspiniform setaeontheprementum(15–21)andisthelargestinstature. 50 R.Novelo-GutiérrezandJ.A.Gómez-Anaya ThelarvaofA.percellulatadiffersfromtheabove-mentionedspecies[insquarebrackets]as follows: (1) tibiae usually with two dark rings well-defined [one ring, joergenseni; three rings, barretti, harknessi, insipida]; (2) posterior margin of sternite 6 smooth [smooth medially, with spiniform setae at sides in barretti and joergenseni; unknown for insipida]; (3) posterior mar- gin of sternite 7 smooth medially, with spiniform setae at sides [mostly smooth with scattered spiniformsetaeatsides,harknessi;withclaviformsetaeonmiddleandspiniformsetaeatsides, barretti;unknownforinsipida](inthisfeature,similartojoergenseni);(4)malegonapophyses,in ventralview,withexternalmarginstraightinbasalhalf[thismarginstronglysigmoidinbarretti, harknessi,joergenseniandinsipida]. Acknowledgements ThanksareduetoDrs.KenTennessen,MichaelMayandRosserGarrisonfortheircriticismwhichimprovedthecontent andEnglishsyntaxofthefinalversion.WealsothankKarinaCuevas-Yañez,M.Sc.,forthedonationofsomelarvaeofA. percellulata.FinancialsupportwasprovidedbyaCONACYTgrant101542-F. References Calvert,P.P.(1902).Odonata.InF.D.Godman&O.Salvin(Eds.),BiologiaCentrali-Americana:InsectaNeuroptera (pp.74–75).London:R.H.PorterandDulau. Cuevas-Yañez,K.(2007).Losodonatos(Insecta:Odonata)delaHidroeléctricadePatla(ElPozo)ydelRíoTecpatlán, Zihuateutla,Puebla,México.Dugesiana,14,83–91. Geijskes,D.C.(1943).NotesontheOdonataofSurinam.IV.Nineneworlittleknownzygopterousnymphsfromthe inlandwaters.AnnalsoftheEntomologicalSocietyofAmerica,36,165–184. González-Soriano,E.,&Novelo-Gutiérrez,R.(2007).OdonataofMexicorevisited.InB.K.Tyagi(Ed.),Odonata:Biology ofdragonflies(pp.105–136).Jodhpur,India:ScientificPublishers. Novelo-Gutiérrez,R.(1992).BiosystematicsofthelarvaeofthegenusArgiainMexico(Zygoptera:Coenagrionidae). Odonatologica,21,39–71. Novelo-Gutiérrez,R.(2008).DescriptionofthelastinstarlarvaofArgiabarrettiCalvert(Zygoptera:Coenagrionidae). Odonatologica,37,367–373. vonEllenrieder,N.(2007).ThelarvaofArgiajoergenseniRis(Zygoptera:Coenagrionidae).Odonatologica,36,89–94. Watson,M.C.(1956).Theutilizationofmandibulararmatureintaxonomicstudiesofanisopterousnymphs.Transactions oftheAmericanEntomologicalSociety,81,155–205.

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