Rediscovery ofMesagrion leucorrhinum(Zygoptera: Megapodagrionidae): a “formal” description of female and ultimate stadium of larva with notes on habits

2011·0.34 MB·

by  León A. Pérez-Gutiérrez

InternationalJournalofOdonatology
Vol.14,No.1,March2011,91–100
Rediscovery of Mesagrion leucorrhinum (Zygoptera:
Megapodagrionidae): a “formal” description of female
and ultimate stadium of larva with notes on habits
LeónA.Pérez-Gutiérreza andJenileeM.Montes-Fontalvob
aGrupodeinvestigaciónenBiodiversidaddelCaribecolombiano,DepartamentodeBiología,
UniversidaddelAtlántico,km7antiguavíaPuertoColombia,Barranquilla,Colombia;
bPostgradoenEntomología,MuseodelInstitutodeZoologíaAgrícola(MIZA),Facultadde
Agronomía,UniversidadCentraldeVenezuela,Apartado4579,Maracay2101-A,Venezuela
(Received6October2010;finalversionreceived14February2011)
AdultfemaleandultimatestadiumlarvaofMesagrionleucorrhinumareformallydescribedandillustrated
basedonmaterialfromthreelocationsinAntioquia,MetaandCundinamarcaDepartments,Colombia.
Thespeciesissexuallydimorphic.The♀isdistinguishablefromotherrelatedgenerabyapairofnotches
intheprothoracicanteriorlobeandshareswithHeteropodagrionandDimeragrionfemalesayellowish,
scarcelysclerotizedregiondorsallybetweentheposteriormarginofS7andanteriorborderofS8.Thelarva
isverysimilartoHeteropodagrion.Differencesforseparatingthemare:thepro-,meso-andmetathoracic
supracoxal processes are less prominent in Mesagrion, and the length of the terminal filament of the
middlegillisnotablylongerinMesagrion.Thespecimenswerealsocomparedwithotherrelatedgenera.
Observationsonhabitsareadded.
La Hembra y la larva en último estadio de Mesagrion leucorrhinum son formalmente descritas e
ilustradasbasadasenmaterialdeColombiadetreslocalidadesenlosdepartamentosdeAntioquia,Meta
yCundinamarca.Existemarcadodimorfismosexualenlaespecie.La♀esdistinguibledeotrosgéneros
relacionadosporunpardemuescasenellóbuloanteriordelprotórax,dorsalmentepresentaenelborde
posteriordelS7ybasedelS8unazonaescasamenteesclerotizadadecoloramarillo,similaralaquese
observaenhembrasdeHeteropodagrionyDimeragrion.LalarvaesmuysimilaraladeHeteropodagrion.
Diferenciasparasepararlasson:losprocesossupracoxalesenpro-,meso-ymetatóraxsonmenospromi-
nentesenMesagrion,yeltamañodelfilamentoterminalenlabranquiamediaesmarcadamentemáslargo
enMesagrion.Losejemplaresfueroncomparadoscongénerosrelacionados.Adicionalmentesepresentan
datossobrehábitos.
Keywords: Odonata; Damselfly; Megapodagrionidae; Mesagrion; Heteropodagrion; female; larva;
Colombia
Introduction
MesagrionSelys,1885isamonotypicgenus.Itssolespecies,M.leucorrhinum,wasknownonly
fromthemaleuntilrecently.ThefemalewasdiagnosedandillustratedbyGarrisonetal.(2010)
butnotformallydescribed.Thisgenusisendemictothecentral-easternsectoroftheColombian
∗Correspondingauthor.Email:[email protected]
ISSN1388-7890print/ISSN2159-6719online
©2011WorldwideDragonflyAssociation
DOI:10.1080/13887890.2011.570162
http://www.informaworld.com
92 L.A.Pérez-GutiérrezandJ.M.Montes-Fontalvo
Andes.Inadditiontothetypelocality(Cristalina,easternslopesofthecentralCordillera,Antioquia
Department)newfindingshavebeenmadeatotherplaces,namelyAlban,ChirajaraandGuaya-
betal(CundinamarcaDepartment)andVeredaLaVictoria(Villavicencio,MetaDepartment),sig-
nificantlyexpandingtheknowndistributionofthespecies.Ithasbeenconsideredofconservation
interestunderthestandardsofInternationalUnionforConservationofNature(IUCN)(Paulson,
2004)duetoadvancedhabitatdeteriorationandpoorknowledgeaboutitsreproductivebiology
andpopulationecology.Thephylogeneticpositionofthegenusisunknown,butarelationshipwith
Heteropodagrion Selys has been proposed based on adult morphology (Garrison & von Ellen-
rieder,2005).Behaviorally,Mesagrionisoneofthefewrepresentativesinthefamilythatclosesits
wingswhenperched(Donnelly,1992),afeaturesharedintheNewWorldonlywithHeteropoda-
grion,ParaphlebiaSelysinHagen,andThaumatoneuraMcLachlan(Garrisonetal.,2010).
Materialandmethods
Terminology follows Westfall and May (1996) for body characters, Riek and Kukalová-Peck
(1984)forwingveinnomenclature,andWatson(1956)forlarvalmandibularformula.Alldimen-
sionsaregiveninmillimetersandweremadewithacalibratedocularmicrometer.Illustrations
were made with the help of a camera lucida coupled to aWild M-8 stereoscope and are not to
scale.SpecimensaredepositedatUniversidaddelAtlánticoRegiónCaribe(UARC),Barranquilla,
Atlántico,ColombiaandattheMuseodelInstitutodeZoologíaAgrícolaMIZA,Maracay,Aragua,
Venezuela.Abbreviations:S=abdominalsegment(e.g.S7=abdominalsegment7).
Descriptionoffemale
Materialexamined
Fifteen (cid:2)(cid:2): COLOMBIA: Cundinamarca Department, Guayabetal, altitude 1270 m, 1 (cid:2), 10
December 2007, L.A. Pérez, deposited at UARC, Barranquilla,Atlántico, Colombia; 1 (cid:2) same
locality, but 13 June 2008, L.A. Pérez, deposited at the Museo del Instituto de ZoologíaAgrí-
cola(MIZA),Maracay,Aragua,Venezuela.13(cid:2)(cid:2)samelocality,but7February2009,L.A.Pérez,
depositedatUniversidaddelAtlánticoRegiónCaribe(UARC),Barranquilla,Atlántico,Colombia.
Overall coloration, morphology and wing venation of female are very similar to male (Ris,
1918,Garrison&vonEllenrieder,2005)(Figure1a–c).
Head(Figure2a)
Labrum rounded, yellow with black anterior margin, anteclypeus yellow, postclypeus yellow
with dark stripe along posterior margin (presumably showing varying degree of color intensity
accordingtoindividualmaturity),genaegreyish-brown,antefronsshiningblackandflat,frontal
edge prominent with long hairs on both sides, vertex black, postocular spots absent, occipital
borderbrownwithafewlonghairs.
Thorax
Largely black with distinctive yellow stripes on lateral sclerites. Prothorax largely black
(Figure2b),anteriorlobeyellow,laterallyblackish,withtwoexcisionsonanteriormargin,middle
lobewithreddishdorsolateralspot,propleuronandprocoxaedark,posteriorlobeyellowwithblack
center.Pterothoraxwithlonghairpencilatexternalangleofeachmesostigmalplate.Middorsal
Mesagrionleucorrhinumfemaleandlarva 93
Figure 1. Mesagrionleucorrhinum:(a)female,lateralview,Alban(Cundinamarca),6July2010;(b)immaturefemale,
lateral view, Alban (Cundinamarca), 6 July 2010; (c) male, lateral view, Rio Claro (Antioquia), 4 July 2009. All
PhotosbyLP.
carinablack,mesepisternumwithcompleteyellowantehumeralstripe,ofroughlysamewidthover
itsentirelength,mesinfraepisternumblack,mesocoxablack,metepisternumblackwithcomplete
yellowstripewhichiscurvedtowardsthemetapleuralsuture.Metinfraepisternumblack,meta-
coxaeblack.Metepimeronblackwithyellowspotatmetapleuralsuture;inventralviewmetathorax
completelyyellow.Legswithfemorablackondorsalsurface,ventralsurfacereddish,moreintense
onforelegs,inmaturespecimensyellowish;innerrowofhindfemoraarmedwitheightsetiform
spursapproximatelyofsamesizeandalittlelongerthanthespacebetweenthem.Tibiaeyellow-
ish,thoseofhindlegsarmedwitheightsetiformspurs(innerrow)decreasingdistally,butfirst,
secondandthirdspursofsamesize.Tarsalclawwithinnertoothapproximatelyhalfsizeoftip.
Wings
Wings hyaline, with margins smoky at anal region, venation black, Pt reddish (brownish in
immature specimens), of same shape and size in both wings, inferior margin covering less
94 L.A.Pérez-GutiérrezandJ.M.Montes-Fontalvo
Figure 2. MorphologicaldetailsofMesagrionleucorrhinum.Female(a)head,dorsalview;(b)prothorax,dorsalview;
(c)forewing;(d)hindwing;(e)distalportionofabdomen,lateralview.Male(f)genitalligula,ventralview;(g)genital
ligula,lateralview.
than two cells, proximal margin very oblique. Petiolation reaching level of arculus, no supple-
mentary sectors between RP2 and IR2. Fw (Figure 2c) 27–31mm, usually with two complete
antenodals, occasionally one incomplete supernumerary Ax, 18–22 postnodals, IR2 arising
beyond first Px, RP2 arising between sixth and eighth Px; RP3 arising at or immediately
beyond subnodus, CuP close to level of first primary Ax, CuA arising between second Ax
and nodus. Hw (Figure 2d) 28–31.5mm, two complete antenodal crossveins, 17–20 Px, RP2
arising between fifth and sixth Px, RP3 arising right before subnodus. IR2 arising at first
postnodal.
Mesagrionleucorrhinumfemaleandlarva 95
Abdomen
Inimmaturespecimensabdominalsegmentsscarletredwithdistalyellowringsandnarrowyellow
mid-dorsalline;maturespecimenswithgreyish-brownabdomen.Indorsalview,betweenendof
S7andanteriormarginofS8,adistinctiveyellowish,scarcelysclerotizedregion.Novulvarspine
onS8,basalplatelarge.Posteriorgonapophysesslightlysurpassingtipsofcerci(Figure2e).
Diagnosticremarks
WeobservedthatyoungfemalesofM.leucorrhinumareidenticaltomalesincolorationinlife,
withstrikingredcolorandyellowdistalringsonabdominalsegments,andredforelegs.Thered
abdomen and the yellowish portion between S7 and S8 are very distinctive, and together with
the black and yellow color pattern of labrum and clypeus easily distinguish the species in its
naturalenvironment.ApairofnotchesintheprothoracicanteriorlobedistinguishesfemaleM.
leucorrhinumfromsimilarspecies,suchasH.superbum.
Additionalnotesonmales
Male genital ligula with inner fold vestigial (Figure 2f, g), shaft of genital ligula sclerotized to
middleregion,with6–7setaeoneachside.Apicalsegmentmembranous,tipconvex,notchedand
expanded,inlateralviewwithbilobulatedtrumpet-shapedextensions,closelyapressedtomiddle
regionofligula.
Descriptionoflarva
Materialexamined
1 (cid:3) ultimate stadium exuviae, 1 (cid:3), 4 (cid:2) ultimate stadium larvae, COLOMBIA: Cundinamarca
Department,Guayabetal,altitude1270m,1(cid:2),10December2007,L.A.Pérez.Othermaterial
considered:13specimensofearlystadia,samedata,preservedinethanol.
Larvayellow-ochraceouswithirregular,darkspotsonthoraxandabdomen(Figure3a).
Head
Head slightly wider than meso- and metathorax, antenna curved with seven segments, the two
basalsegmentsdorsallydarkerandwithabundanthairs.Labiumshort,almostsquare(Figure3c),
baseofprementumatrestlocatedbetweenfirstpairofcoxae,indorsalviewwithslightlycurved
rowofverysmalltransverseridgesoneachside.Middlelobeslightlyconvex,withbluntdenticles
(Figure 3d),mediancleftshallow,indorsalviewpalpallobewithoutsetaeorspines,inventral
view with long and thin hairs; three palpal hooks, the medial one longest, basal hook shortest;
mandibleswithincisorandmolarteeth;formula:L1+2345O a (m1,2,3,4,5)bb(cid:3) (1<3<2<
4<5)/R1+2345y a b (1<3<2<4<5)inbothmandiblesa >b (Figure 3e–g);molars
numbervarybetweenfiveandseven(twospecimenswithsevenmolars,onewithsixmolarsand
onewithfivemolars),posteriormarginofcompoundeyewithsinuosity,fronsflat.
Thorax
Prothoraxlaterallyblack,flatsidesformingaprominentkeel,armedanterolaterallywithtwosmall
butdistinctsupracoxalprocesses,ofwhichtheposteriorprocessislarger.Mesothoraxpalewith
96 L.A.Pérez-GutiérrezandJ.M.Montes-Fontalvo
Figure 3. FinalstadiumlarvaofMesagrionleucorrhinum.(a)Habitus;(b)lateralmarginofmeso-andmetathorax;(c)
head,ventralview;(d)labiumandleftlabialpalp,dorsalview;(e)leftmandible,medialview;(f)leftmandible,posterior
(ventral)view;(g)rightmandible,medialview;(h)distalsegmentsoffemaleabdomenshowinggonapophysesandleft
gill,ventralview;(i)malecerci,mediodorsalview;(j)malecerci,lateralview;(k)mediangill,dorsalview.
low, rounded lateral process in the middle region. Metathorax pale with prominent laterodistal
processes (Figure 3b). Diverging wing pads extending to end of S9 with pattern of dark spots
atbase,legsshort,hindfemurextendingbackwardsparalleltoabdomentoposteriormarginof
S7. Femur and tibia laterally thickened with distinguishable carinal margins, armed with hairs
andspines.
Mesagrionleucorrhinumfemaleandlarva 97
Abdomen
Short, S1–8 with prominent lateral keels and long hairs, S9 and S10 constricted and narrow.
Female gonapophyses with rounded tips, surpassing posterior margin of S10, without hairs
or spines (Figure 3h). Male gonapophyses not obvious in last stadium, cercus kidney-shaped
with dorsal margin beset with small setae, paraproct ventrally flat and lobed (Figures 3i, j).
Caudal gills not petiolated, without dorsal keels, tracheation not evident; epiproct with well
developed lateral keels; lateral and middle gills with articulated terminal filament and with
dense hair cover (Figure 3k), terminal filament in middle gill approximately twice the size of
lateralones.
Discussion
ReviewingdescriptionsofotherMegapodagrionidaelarvae,thespeciesmostcloselyresembling
M.leucorrhinumisHeteropodagrionsanguinipes(Tennessen,2010).Characterssharedbetween
thesetwogeneraare:presenceofaslightlycurvedrowofverysmalltransverseridgesoneach
side of prementum, and presence of pro-, meso- and metathoracic processes (more prominent
inHeteropodagrion).SuperficialsimilaritieswiththelarvaofSciotropiscyclanthorum(Rácenis,
1959)describedbyDeMarmels(2004)maybenoted.Themostnoticeabledifferencesbetween
otherrelatedgeneraarepresentedinTable1.
TheclassificationbasedongillmorphologybyKalkman,Choong,OrrandSchütte(2010)and
redefinedbyTenessen(2010)isherereviewed.ReexamininglarvaeofSciotropis,Heteropoda-
grionandDimeragrionandcomparingthemwithdescriptionsofallneotropicalspeciesknown
to date, it becomes evident that the terminal filament of the middle gill can have two patterns:
onegroup,consistingofDimeragrion,Heteragrion,OxistigmaandPhilogenia,hasunarticulated
terminalfilaments;asecondgroup,whichincludesHeteropodagrion,Hypolestes,Paraphlebia,
Mesagrion,ThaumatoneuraandSciotropis,hasarticulatedterminalfilaments,eachwithacon-
striction at its base. Sciotropis, exceptionally, lacks a terminal filament on the middle gill, a
conditionitshareswithsomegeneraoftheoldworld,namelyCaledopteryx (Lieftinck,1976).
Finally,athirdgroupconsistingofTeinopodagrionandprobablyMegapodagrionlacksterminal
filamentonallgills.
InhisdescriptionofthalarvaofThaumatoneura,Calvert(1915)referstogillregenerationasa
reasonforthevariabilityinthesizeofthefilaments.HoweverthatisnotthecaseforMesagrion
andHeteropodagrion.Afterexaminationof19and15specimensrespectively,wefoundthatpro-
portionoffilamentlengthisabsolutelystable,evenindifferentstadia.Thisobservationconfirms
theinformativevaluetothischaracterforseparatinglarvaeofHeteropodagrionandMesagrion,
withtheterminalfilamentsinHeteropodagrionshorterthaninMesagrion.Tocorroboratethiswe
measuredandcomparedheadwidthwiththetotallengthofterminalfilaments,asthesestructures
haveproportionalgrowth(Velasquezetal.,2009);seeTable2.
AfterverifyingthemandibularformulainHeteropodagrionsanguinipeslarvae,wecompared
resultswithMesagrionleucorrhinumandconcludedthatbothgenerasharethesamenumberof
incisors,whilethenumberofmolarsvaryingindividuallybetween5and7.Hence,thischaracter
isuninformativeforseparatingthetwogenera.
Other genera, such as Argiolestes (Lieftinck, 1956), share morphological aspects of the
labial palps and prementum with Mesagrion (three palpal hooks, the middle one longest) and
absence of premental setae. However, differences in details of gill morphology (saccoid in
Mesagrion and fan-like in Argiolestes) (Kalkman et al., 2010) suggest no close relation. The
subquadrateshapeofprementumanditsshortlengtharesimilartoCaledopteryxsarasinilarvae
(Lieftinck,1976).
9
8
Table 1. MorphologicalfeaturesinMegapodagrionidslarvaeknowntodate,comparisonsanddifferencesbetweenTennessen’sandKalkman’sclassificationsystemsandtheircontribution. L.A
.
P
Tennessen,2010 Kalkmanetal.2010 Thiscontribution ére
z
Long Palpal Gills Thoracic -G
u
Taxa References Group1 Group2 Baloon Fan legged teeth filaments processes tié
r
Dimeragrionpercubitale Demarmels,1999 ∗ ∗ 3 NotArticulate Absent rez
Heteragrionalbifrons Novelo-Gutierrez,1987 ∗ ∗ 3 NotArticulate Absent an
d
Heteragrionalienum Novelo-Gutierrez,1987 ∗ ∗ 3 NotArticulate Absent J
Heteragrionaurantiacum Santos,1968 ∗ ∗ 3 NotArticulate Absent .M
.
Heteragrionbariai Demarmels,2004 ∗ ∗ 3 NotArticulate Absent M
Heteragrionbreweri Demarmels,2004 ∗ ∗ 3 NotArticulate Absent on
Heteragrionchlorotaeniatum Demarmels,2004 ∗ ∗ 3 NotArticulate Absent tes
Heteragrionconsors Costa&Santos,1999 ∗ ∗ 3 NotArticulate Absent -Fo
Heteragrionerythrogastrum Ramirez,1992 ∗ ∗ 3 NotArticulate Absent nta
Heteragrionmitratummitratum Demarmels,2004 ∗ ∗ 3 NotArticulate Absent lvo
Heteragriontricelulare Novelo-Gutierrez,1987 ∗ ∗ 3 NotArticulate Absent
Heteropodagrionsanguinipes Tenessen,2010 ∗ ∗ 3 Articulate Present
Hypolestesclara Needham,1941;Westfall,1996 ∗ ∗ 3 Articulate Present
Hypolestestrinitatis Alayo-Soto,1985 ∗ ∗ 3 Articulate Absent
Mesagrionleucorrhinum Thisstudy ∗ ∗ 3 Articulate Present
Oxistigmacaerulans Demarmels,1987 ∗ ∗ 3 NotArticulate Absent
Oxistigmapetiolatum Geijskes,1943 ∗ ∗ 3 NotArticulate Absent
Paraphlebiazoe Novelo-Gutierrez,2008 ∗ ∗ 3 Articulate Absent
Philogeniacarrillica Ramirez&Novelo-Gutierrez,1994 ∗ ∗ 3 NotArticulate Absent
Philogeniacassandra Demarmels,1982 ∗ ∗ 3 NotArticulate Absent
Philogeniamangosisa Bybee&Tenessen,2008 ∗ ∗ 3 NotArticulate Absent
Philogeniapeacocki Ramirez&Novelo-Gutierrez,1994 ∗ ∗ 3 NotArticulate Absent
Philogeniaterraba Ramirez&Novelo-Gutierrez,1994 ∗ ∗ 3 NotArticulate Absent
Sciotropiscyclanthorum Demarmels,2004 ∗ ∗ 3 Articulate,middlegill Present/
withoutfilament Prominent
Teinopodagrioncaquetanum Pérez-Gutierrez,2007 ∗ ∗ 3 Withoutfilament Absent
Teinopodagriondecipiens VonEllenrieder,2006 ∗ ∗ 3 Withoutfilament Absent
Teinopodagrionmeridionale VonEllenrieder,2006 ∗ ∗ 3 Withoutfilament Absent
Teinopodagrionoscillans Demarmels,2001 ∗ ∗ 3 Withoutfilament Absent
Teinopodagrionvallenatum Pérez-Gutierrez,2007 ∗ ∗ 3 Withoutfilament Absent
Thaumatoneurainopinata Needham1911;Calvert,1915 ∗ ∗ 2 Articulate Present
Mesagrionleucorrhinumfemaleandlarva 99
Table 2. Measures of head width and length of terminal filament in middle gill of larvae of
HeteropodagrionsuperbumandMesagrionleucorrhinum.
Lengthofterminal
Species Headwidth filamentinmiddlegill HW/FL* Stadium
H.superbum 3.41 0.86 3..97 Ultimate
3.37 0.93 3.62 Ultimate
3.41 0.89 3.83 Ultimate
3.41 0.93 3.67 Ultimate
3.37 1.03 3.27 Ultimate
1.72 0.52 3.31 Early
3.31 0.86 3.84 Ultimate
3.31 0.86 3.84 Ultimate
2.31 0.72 3.21 Early
M.leucorrhinum 3.24 1.27 2.55 Ultimate
3.17 1.27 2.50 Ultimate
3.21 1.24 2.59 Ultimate
3.24 1.31 2.47 Ultimate
Note:*Ratioofheadwidthtomiddlefilamentlength.
Habitatnotes
AtlocalitieswhereMesagrionleucorrhinumoccurs,acloserelationshipwithsmallwaterfallsof
foreststreamsbetween500and1300mwasobserved.
In their habitat individuals can be numerous, without evidencing competition behavior for
space.Flightissmooth,femalesandmalesareactivewithoutdisturbingoraggressivebehavior.
Itisstillnecessarytoclarifythedetailsofreproductivebiology,butwepresumethatoviposition
takes place on the rock surfaces washed by waterfalls (splash zone). Larvae are camouflaged
through their color which is similar to the rock colors, moss and detritus on which they were
found.Whendisturbed,theymoverapidlywithlateralwavemovements,likePlecopteralarvae.
Acknowledgements
WewishtoexpressourwarmthankstoProf.DrJürgDeMarmelsforthecriticalreadingandsuggestionsforimproving
thismanuscript,toNataliavonEllenriederandananonymousrefereeforthecriticalrevisionofaformerdraft.
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