Table Of ContentOdonatohgica36(3):263-273 September1.2007
Some Libellulidaelarvae from theYungas
Forest, Argentina:Macrothemishahneli Ris,
Brechmorhoganubecula(Rambur) and
Dasythemismincki clara Ris
(Anisoptera)
N.Von Ellenrieder
Institute deBioyGeociencias,MuseodeCiencias Naturales deSalta,
Universidad Nacional deSalta,Mendoza2,AR-4400Salta,Argentina
[email protected]
ReceivedMarch 15,2006/Reviewed andAcceptedAugust4,2006
A first description of the larva of M. hahneli isprovided. The larva ofB.
nubecula,previously described based onasinglespecimenofdoubtful identity, is
here redescribed based onbonafide specimensbelongingtothatsp. Thelarva of
D. minckiclaraisfound toagree overallwith that ofD. m. mincki,differingonly on
someminordetailsprobablydue togeographicvariation.
INTRODUCTION
Thelarvaeofthethreelibellulidspecies dealtwithhereinhabitthesameaquat-
icenvironmentsinthe ArgentineYungas. MacrothemisHagen, 1868andBrech-
morhoga Kirby, 1894are primarily neotropical genera;Macrothemiscontains40
species andBrechmorhoga 16,whereastheexclusively SouthAmericanDasythemis
includes onlyfour(GARRISON etal.,2006). Only six larvaeof Macrothemis,
viz. M.aurimaculataDonnelly, 1984;M. celeno(Selys inSagra, 1857);M. inacuta
Calvert, 1898;M. musivaCalvert, 1898;M. pseudimitans Calvert, 1898andM.
tessellatainequiunguis Calvert, 1895,andsevenofBrechmorhoga, viz. B.mendax
(Hagen, 1861); B.pertinax pertinax (Hagen, 1861);B. praecoxpraecox(Hagen,
1861);B. praedatrix (Calvert, 1909);B. rapax rapaxCalvert, 1898; B. travassosi
Santos, 1946and B. vivax Calvert, 1906, havebeen described so far(GARRI-
SON etal„ 2006). SANTOS(1969) provided a description forB.nubeculabased
onone larva fromGuanabara, Brazil whichwas reareduntil emergenceof the
adult; hewashoweverunsure aboutthespecific identityofthatfemalespecimen.
264 N.vonEllenrieder
Figs1-9.Ultimatelarval instarofBrechmorhoganubecula (Ramb.):(1)leftproepimeron,dorsalview;
- (2)generalaspect,dorsal view;- (3)rightantenna,dorsal view;- (4)rightmandible,innerview;-
(5)leftmandible,inner view;- (6)prementum,dorsalview;- (7)labialpalp,dorsolateral view;- (8)
abdominal segments 2-10,lateralview;- (9)abdominal segments8-10, dorsalview.
Somelibellulid larvaefromtheYungasForest,Argentina 265
Sincehis descriptiondoesnotfullyagreewiththeexuviaeofrearedbonafideiBre-
chmorhoga nubecula. aredescription ofthatspecies is presented here.
TwolarvaeofDasythemis Karsch, 1889weredescribedtodate:D. venosa(Bur-
meister, 1839)andD. m.mincki(Karsch, 1890)(CARVALHO etal.,2002). Here
thelarvaof D. minckiclara Ris 1908is compared withthatofthenominatesub-
species ofD. mincki.
BRECHMORHOGA NUBECULA(RAMBUR, 1842)
Figures 1-9
Material.— (depositedinthe MuseodeCiencias Naturales deSalta):Ultimate larvalinstar:
Argentina,Jujuyprovince,El Pantanoso,23°31’17”S,64°35’13”W,609m.a.s.l.,3/4-XI-2005,vonEl-
lenrieder leg.,1exuviae(maleemergedinlaboratory),2larvae;Saltaprovince: ElRey NationalPark,
ArroyoLaSala,24°43’41.2”S,64°40’ 16.7”W,971 m.a.sl,30-XI-2005,vonEllenrieder leg.,2larvae;
Lesser, 1312m.a.s.l.,9-1-2006,24°40’57”S,65°28’39”W,vonEllenriederleg., 1larva.Youngerinstars:
Samelocalities;ElPantanoso: 1larva;Lesser,10-XI-05:1 larva;ArroyoLaSala:5larvae;Saltaprov-
ince; BantuNational Park, rocky streamaffluent toRio Baritii,22°29T6.5”S,64°45’36.4”W, 1490
m.a.s.1.,20-VIII-05:2larvae.
Larvae werefoundalongrockysectionsofriversand streams,under stones.
description. - Integument granulose andwithfewsetae.
Head.- Trapezoidal, slightly more than twiceas wide as long, narrowed
posteriorly, withoccipital margin straight, andcephalic lobesbroadly rounded,
not bulging (Fig. 2).Antenna7-segmented, with thirdantennomere thelongest
(Fig. 3). Prementum(Fig. 6) almostas wideas long (ratio width/length0.95-1),
with 7-9lateralsetae on dorsalsurface, arranged as 2+S-2+7; ligula prominent;
articulationprementum/postmentum atlevelofanteriormarginofmesocoxae.
Labialpalp with 6-8 long setae (Fig. 7); movablehook thick, sharply pointed,
as long as palpal setae; distalmargin ofpalp with 8-10crenulations, eachwith
3-4setaeatbase. Mandibles(Figs 4-5)withfollowing formula(sensu WATSON,
1956):L 12340abb’,R 1F234yabb’
Thorax.-Lateralmargins ofneckmembranestronglysclerotized,rounded,
bearinglong setae. Rowofstout, stiffsetaealonganteriormarginofproepister-
numandinferiormarginofpropleuron; lateralmarginofproepimeron straight
(Fig. 1).Wing pads extending to posterior margin ofabdominalsegment 5 or
midlength of6. Legs short(hind leg not surpassing distalmarginof analpyra-
midwhenextendedto sidesofbody); femorawiththreedarkbands, sometimes
diffuse; tibiaewiththreedarkbandsandtarsi withdark apexes(Fig. 2).
Abdomen.- Colorpattern as showninFigure2. Segments 2-9withblunt
mediodorsalprojections; those onsegments 2-6 well developed, on 7-9 almost
imperceptible laterally (Fig. 8). Lateralspines of segments8and 9 shorterthan
half ofmiddorsallength oftheirsegments;posterior margin of sternite 10not
visibleindorsalview; cerciaslong as 0.4-0.5of epiproct (Fig. 9).
Measurements (inmm;averageand standard deviation;range insquarebrackets;N = 6
266 N.vonEllenrieder
unlessindicatedotherwise).- Head mediodorsal length:2.59± 0.23[2.3-2.85](N = 5); head max,
width: 5.92 ± 0.18 [5.6-6](N =5); headwidth/ length:2.3± 0.25 [1.96-2.61](N = 5):prementum
length: 4.35 ± 0.14 [4.2-4.5];prementummax.width: 4.2± 0,14 [4-4,4];prementummax.width/
length:0,97± 0.02[0.95-1],FemurI:2.88 ±0.17[2,7-3,2];II:3.66±0.13 [3.5-3.S](N = 5); III:4.96
±0.19 [4.8-5.3](N=5).Tibia 1:3.45 ±0.19 [3.2-3.7];II:3.98 ±0.15 [3.S-4.2](N=5);III:5.52±0.18
[S.2-5.6](N =5).Innerwingpads:7.14+0,05 [7.1-7.2](N=5);externalwingpads:6.86 ±0.18 [6.6-
7.10](N =5).Abdomen lengthwith appendages:13.84±0,46 [13,2-14,5](N=5); lateralspineonS8
length:0.51 ± 0.08 [0.4-0.6];lateral spineonS9length:0,47 ±0.05 [0.4-0.5];epiproct length:1.66±
0.07[1.6-1.75];cercuslength:0,72± 0,04[0.7-0.8];cercus/epiproctlength:0.43± 0.04[0.4-0.5];para-
proctlength:1.67 ±0.14[1.5-1.8], Totallengthwithappendages:21.68±0.77[20.9-22.9] (N = 5).
YOUNGERINSTARS. — Differfromultimateinstarasfollows:mediodorsalpro-
jections well developed onabdominalsegments 1-9; fewer numberof setae on
labialpalp andprementum; fewernumberof crenulationson distalmargin of
prementum;colorpatternwithabdominalsegments 1-4 dark,andfewernumber
of darkbandsonlegs (2or 1, dependingonsize).
DIAGNOSIS. - Thelarva ofB. nubeculadiffersby the straight lateralmargin
oftheproepimeron from B.rapax(witha digitiformprocess inB.rapax;NOVE-
LO-GUTIERREZ, 1995b),by thelateralspines onabdominalsegments 8-9of
equal lengthfromB.pertinax (that on8abouttwiceas long astheoneon9 in B.
pertinax; NOVELO-GUTIERREZ,1995a), and shorterthanmiddorsallength
ofsegment9 fromB. mendax(as long as itin B. mendax; NOVELO-GUTIER-
REZ, 1995b), bythemediodorsalprojectionsonabdominalsegments2-5 convex
fromB.praecox(withstraight dorsalmargins,parallel tobody axisinB.praecox;
NOVELO-GUTIERREZ,1995b),on segments6-9bluntandlowinlateralview
from B. praedatrix (well developed andacute in B. praedatrix;FLECK, 2004),
andbythenumberofpremental setaeandmandibularformulafromB. travassosi
(7-9 premental setae inB. nubecula, 11 inB. travassosi;mandibleswiththreemo-
lars in B. nubecula,lackingmolarsin B. travassosi; SANTOS& COSTA, 1999).
It canbe distinguished fromthat ofB. vivax, the only sympatric species of the
genusinthe Argentine Yungas, by thedorsal projection on abdominalsegment
3 clearly shorterand narrower than that on4(Fig. 8,whichis almostequal to
thaton4in B. vivax),andby thenumberofpremental setae (7-9inB. nubecula,
and 12-13in B. vivax; NOVELO-GUTIERREZ, 1995b).
MACROTHEMISHAHNELI RIS, 1913
Figures 10-18
Material.— (depositedinthe MuseodeCiencias Naturales deSalta): Ultimatelarval instar:
Argentina,Jujuyprovince, El Pantanoso,23°31T7”S,64°35’13”W,609m.a.s.1.,3/4-XI-2005,vonEl-
lenrieder leg.,1 larva;Salta province:Lesser, 1312 m.a.s.1.,09-1-2006,24°40’57”S,65°28’39”W,von
Ellenriederleg.,4exuviae(lmale,3femalesemergedinlaboratory),1 larva;samebut 11/13-X-2005,
2larvae. Youngerinstars:samelocality:Lesser, 13-X-2005, 2larvae. Jujuyprovince: LaAngostura,
24°27’01.4”S,65°22’43.7”W,1511 m.a.s.1.,8-XI1-2005,col. vonEllenrieder: 1 larva;ArroyoenRta
6aPalmaSola,23°52’12”S,64°22’4”W,534ra.a.s.1.,5-XI-2005,col. vonEllenrieder: 1larva. Salta
province: Baritu National Park, rocky streamaffluent to Rio Baritii,22°29T6.5”S64°45’36.4”W,
Some libellulid larvaefromtheYungasForest,Argentina 267
1490m.a.s.1.,20-VIII-05:2larvae;ElReyNational Park,streamonwaytoPozoVerde,24°44’28.8”S,
64°40’35.8”W,1029 m.a.s.l.,20-VIII-05:1larva.
Larvae werefoundatriverand streammargins,inareasofabundantaquaticvegetation.
DESCRIPTION.- Integument coveredwithsmallspiniform setae.
Head. — Trapezoidal, about twiceas wide as long, narrowedposteriorly,
withoccipital margin slightlyconcave, andcephalic lobesbroadly rounded, not
bulging, covered withminutespiniform setae and afew stout long setae (Figs
11-12). Antenna7-segmented, withthirdantennomerethe longest. Prementum
(Fig. 16)almost as wideas long (ratio width/length 0.92-1), with 8-9lateralse-
tae on dorsalsurface; ligula prominent,finely crenulatedalong margin;articula-
tion prementum/postmentum atlevelofanteriormarginofmesocoxae. Labial
palpwith6-8longsetae(usually 7);movablehookthick, sharplypointed, aslong
as palpal setae;distal marginofpalp with8-10obsoletecrenulations, withvery
shallownotches between them(Fig. 15). Mandibles(Figs 13-14)with following
formula(sensu WATSON, 1956): L 12340 abb’,R 123+4y ab
Thorax.-Lateralmargins ofneck membranestronglysclerotized, rounded;
rowofstout,stiffsetae alonganteriormarginofproepisternum, inferiormargin
of propleuron, anddorso-lateral margins ofpronotal disk(Figs 11-12). Prono-
taldisk withdark inverted ‘V-shaped’ spotandacentraltuberclebearingspines
(Figs 11-12). Wing pads extending to midlength of abdominalsegment 5 to 6.
Legs long (hind leg surpassing distal marginofanalpyramid whenextendedto
sidesofbody); femorawitharow oflongstiffsetae alongdorsalmargin; femora
withthreeandtibiaewithtwo darkbands, sometimesincomplete ordiffuse,and
tarsi withdark apexes(Fig. 11). Metacoxawithtriangularventrolateralprocess
(Fig. 10).
Abdomen.- Colorpatternas shownin Figure 11, Segments 3-9withwell
developed spine likemediodorsalprojections; thatonsegment3vertical,remain-
derprogressively decliningposteriorly(Fig. 17).Lateralspinesofsegments8and
9shorterthan halfof middorsallength of theirsegments; posterior marginof
sternite 10not visibleindorsalview; cercias long as 0.55-0.65ofepiproct (Fig.
18).
Measureme nts (inmm; averageand standard deviation;rangein squarebrackets;N =8
unlessindicatedotherwise).- Head mediodorsallength:2.15±0.22[2-2.6](N=7);headmax.width:
4.26±0.18 [4-4.55](N=7); headwidth/length: 1.99± 0.17 [1.75-2.15](N=7);prementumlength:
3.13 ±0.1[3.05-3.35];prementummax.width: 3.01 ±0.07[2.95-3.1];prementummax.width/length:
0.96 ± 0.03 [0.92-1],FemurI;2,44 ± 0.13 [2.25-2.65];II:3.29± 0.21[3-3.6];III:4.19 ±0.14[4-4.3],
Tibia I:2.81 ±0.17[2.55-3.1];II:3.54± 0.13 [3.35-3.75];III:5,08± 0.23 [4.75-5.4],Innerwingpads:
5.1± 0.14[4.9-5.25](N =7);externalwingpads:4.93± 0.08 [4.8-5](N=7).Abdomen lengthwith
appendages:8.56 ±0.42[8.15-9.5];lateralspineonS8length:0.19± 0.04[0.15-0.25];lateralspineon
S9length:0.24± 0.05[0.2-0.35];epiproctlength:0.89±0.05 [0.85-1];cercuslength:0.54 ±0.02[0.5-
0.55];cercus/epiproctlength:0.61 ± 0.03 [0.55-0.65];paraproct length:0.78 ± 0.04[0.68-0.8].Total
lengthwith appendages: 14.41 ±0.51 [13.4-14.96](N=7).
YOUNGERINSTARS.- Differfrom ultimateinstaras follows; fewernumberof
268 N.vonEllenrieder
Figs10-18.Ultimatelarval instarofMacrothemis hahneli Ris: (10)left metacoxa,ventrolateralview;
(11) generalaspect,dorsalview; (12)headandprothorax,lateralview;-(13)rightmandible,inner
view;- (14)leftmandible,inner view;- (15)labialpalp,dorsolateral view;-(16)prementum,dorsal
view;- (17)abdominal segments2-10,lateralview;- (18) abdominal segments8-10,dorsal view.
Somelibellulid larvaefromtheYungasForest,Argentina 269
setaeonlabialpalp andprementum;fewernumberofcrenulationson distalmar-
ginofprementum.
DIAGNOSIS. ThelarvaofM.hahnelicanbedistinguished fromthoseofM.
aurimaculataand M.inequiunguis bytheabsenceofa dorsalprotuberance onab-
dominalsegment2(present in thesetwo species), andfromthoseofM. musiva,
M. pseudimitans, M. celenoand M. inacutaby the presenceof7-8 palpal setae
(4-6 intheseotherspecies)and obsoletecrenulationson distalmarginoflabial
palp (deepcrenulationsinthesespecies, RAMIREZ& NOVELO-GUTIERREZ,
1999).Itfurtherdiffersfrom M.pseudimitans andM. inacutaby thetriangular
ventrolateralmetacoxalprocess (absent inM.pseudimitans, digitiforminM. ina-
cuta;NOVELO-GUTIERREZ &RAMIREZ, 1998), from M.pseudimitans by
thethickmovable hookoflabialpalp(thicker than 3timesthewidthofpreced-
ing setae, narrowerthantwicewidthofpreceding setae inM. pseudimitans) and
fromM. inacutabythedorsalprotuberance onsegment3 distinclty shorterthan
that on4(of aboutthe same height inM. inacuta) andthe presenceof only 2
molarsonmandibles(three inM. inacuta;NOVELO-GUTIERREZ & RAMI-
REZ, 1998).In the Argentine Yungas M. hahneliis sympatric withM. inacuta
andM. imitans. M. inacutahas, however, a differentlarval habitat, breeding in
themuddy shoresoflenticenvironmentssuch as pondsand lagoons. Thelarva
of M. imitans, adultsofwhich are foundat thesamestreams andrivers where
M. hahnelibreeds, is stillunknown.
DASYTHEMISMINCKICLARA RIS, 1908
Figures 19-26
Material.— (depositedinthe MuseodeCiencias Naturales deSalta): Ultimatelarval instar:
Argentina,Saltaprovince:Lesser, 1312m.a.s.1.,12-XI-2005,24°40’57”S,65°28’39”W,Garrisonleg.,
1exuviae (1femaleemergedinlaboratory).
Larvawasfoundburrowinginmudataswampyareanearacreek.
DESCRIPTION.- Integument uniformlyreddishbrown, coveredwithabundant
long setae.
Head. - Rectangular, aboutoneandahalftimesaswideaslong,withoccipital
margin approximately straight, andcephalic lobesbroadly rounded, not bulging
(Fig. 19);eyes protrudinglaterally anddorsally(Fig. 23). Antenna7-segmented,
with thirdantennomere thelongest (Fig. 20). Prementum(Figs 23-24) slightly
longer thanwide(ratio width/length 0.86), with3longexternalpremental setae
andaninternalcurvedrow of 13-15very shortsetae (Fig.24); ligula prominent,
finely crenulatedalong margin;articulationprementum/ postmentumatlevelof
posterior marginof procoxae.Labialpalp with4long setae (Fig. 25); movable
hookthick, sharply pointed, as long as palpal setae;distal marginofpalpwith
10-11crenulations, withthreetofourshortsetae onthebaseofeachoneofthem.
Mandibles(Figs 21-22) withfollowingformula(sensu WATSON, 1956): L 1234
270 N.vonEllenrieder
Figs 19-26.Ultimate larval instar ofDasythemismincki claraRis: (19)generalaspect, dorsal view;
- (20)rightantenna, dorsalview;- (21)rightmandible,inner view; (22)left mandible,inner view;
- (23)head,ventralview;- (24)prementum, dorsal view;- (25)labialpalp,dorsolateral view;- (26)
abdominal segments 9-10,dorsalview.
Somelibellulid larvaefromtheYungasForest,Argentina 271
yabd, R 123+4xyabed- thereare largesclerotizedareassurrounding accessory
teethdonleftmandibleandcd onright mandible.
Thorax.-Wing padsextending toposteriormarginofabdominalsegment5.
Abdomen.- Lacking lateralspines;posterior marginof sternite10notvis-
ibleindorsalview; cercishorterthanathirdofepiproct (Fig. 26).
Measurements (inmm;N= 1).- Headmediodorsal length:5.1;headmax.width;33.5;head
width/length;0.69; prementumlength:4.4;prementum max.width: 3.8;prementum max.width/
length:0.86;antennatotallength:2.2;antennomereI:0.2;II:0.25;III:0.5; IV:0.2;V:0.3;VI:0.45;
VI:0.3.FemurI: 3;II;3.5;III:4.2.Tibia I:3.3;II:3.5;III:4.7.Innerwingpads:5.7; external wing
pads:5.Abdomen lengthwithoutappendages:10.1;abdomenmaximumwidth:5.3;epiproctlength:
1.3;cercuslength:0.35;cercus/epiproctlength:0.27;paraproct length;1.25, Totallengthwith ap-
pendages: 19.4.
DIAGNOSIS. — Thelarvaof D. mincki clarastudied herediffers fromthatof
D. m. minckionly by the numberof crenulationsalong distalmarginof labial
palp (11, versus 12 in D. m. mincki), and by the less definedaccessory molars
ofthe mandibles(consisting on heavily sclerotizedareas rather thanindividual
teethas shownforD. m.minckibyCARVALHOetah, 2002). They agreeoverall
onstructuralcharactersandmeasurements. BothdifferformD. venosa,theonly
otherknow larva fromthegenus, bythe absenceoflateralspinesonabdominal
segments8-9.
DISCUSSION
SANTOS’(1969) descriptionofB.nubeculastatesthatthecolorpatternofthat
larvais uniformly pale reddishbrown, andthatthe tibiaehaveanindistinctapi-
caldarkband.Accordingto thematerialstudiedhere, thecolorpatternincludes
several darkspots onbody anddistinct bands ontibiae(Fig. 2). Themeasure-
ments heprovided forwing pads, head length, epiproct length, andlength and
widthofprementumdo notagreewiththeonesfoundhereeither.Eitherhehad
adifferentspecies ofBrechmorhoga in hand, orhis description was inaccurate,
andsince hedidnot include anillustrationor description ofthe diagnostically
importantabdominalmediodorsalprojections itisnot possible todeterminethe
identityofthatspecimen without examining it.
NOVELO-GUTIERREZ &RAMIREZ(1998)provided thefirstdiagnosesfor
thegeneraBrechmorhoga andMacrothemisbasedonlarvalcharacters. However,
they laterfound(RAMIREZ&NOVELO-GUTIERREZ, 1999),thatseveralof
thecharactersthought diagnosticforBrechmorhoga alsooccurredinsomespecies
ofMacrothemislarvae, suchas dorsalprojections onabdominalsegments2-9(in
M. aurimaculataand M. inequiunguis);; dorsal projections reduced andbluntly
tipped onposterior segments(in M. aurimaculata); andbody lacking long, deli-
cate setae (in M. aurimaculataand M. inequiunguis).
The larva of M. hahnelidiffers furtherfrom the generic larval diagnosis of
Macrothemis(NOVELO-GUTIERREZ & RAMIREZ, 1998; RAMIREZ &
272 N.vonEllenrieder
NOVELO-GUTIERREZ, 1999) by its abdominalsegment 10 and anal pyra-
midnot hiddenlaterally by thelateralspinesofsegment 9,posterior marginof
stemite 10 not visible in dorsal view,and ratiocerci/epiproct of0.55-0.65(all
mentionedas generic characters forBrechmorhoga; RAMIREZ & NOVELO-
-GUTIERREZ, 1999).
Thefactthatmore charactersets previously thought diagnosticforthesetwo
generaare foundto overlap as morelarvaebecome known seemsto indicate, as
doesalsotheanalysis ofadultcharacters(GARRISON & VON ELLENRIED-
ER, 2006), thatthesetwo generaare probably artificial, andthataphylogenetic
analysis encompassing alldescribedspeciesandbothadultandlarvalcharacters
willbe necessaryinordertoresolve theirrelationships andassess theirstatus.
ThefewdifferencesshownbythelarvaofDasythems minckiclarafromthatof
thenominalsubspecies areminorandare likelytheexpected resultof geographic
variability.
ACKNOWLEDGEMENTS
Ithank Drs RODOLFONOVELO-GUTIERREZand ROSSERW.GARRISON forthecriti-
calreadingofthemanuscript.Thisstudy wassupportedbytheConsejoNacional deInvestigaciones
Cientificas yTecnicasdeArgentina(CONICET)and NGSgrantNo. 8016/06.
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