Residence and territorial characteristics of Libellulidae species in a neotropical assemblage (Anisoptera)

Odonatologica 37(3):213-220 September 1,2008

Residence and territorial characteristics of

Libellulidae species in a neotropical assemblage

(Anisoptera)

D.C. Resende¹* and P. DeMarco+Jr.²

ReceivedFebruary 25,2007 / Revised andAcceptedDecember30,2007

INTRODUCTION

During

territorial

_{defence, aggressive}

attacks among

heterospecific

odonate malesarecommon

(HASSAN,

1978; MAY, 1980)

and

_competition

_and aggres-sive interactions couldcause some

species

to

change

theirhabitat selection

_(MAY,

1980;RUSSELL etal., 1998). These

_aggressive

_{interactions may}cause a sepa-ration of niches among

species.

Thus the

_{co-occurring species,}

Ischnura

_elegans

(Vander L.), Coenagrion pulchellum (Vander L.), Enallagma cyathigerum (Charp.)

and

_{Erythromma najas (Hans.),}

for

_example,

have

_enough

differentiationin their

oviposition

sites

_(JOHANNSSON,

_{1978). Sympetrum flaveolum (L.)}

malesuse

1 LaboratoriodeEcologiaeSolos,Curso de Ciencias Biologicas,

CentroUniversitario do Leste de MinasGerais,R.BarbaraHeliodora,725BomRetiro, BR-35160-215, Ipatinga, MG, Brazil

dcresende@ig.com.br

2

Laboratorio deEcologiaTeorica e_{Sintese, Departamento}deBiologia Geral, Universidade Federal de_{Goias, BR-74001-970, Goiania, Goias,}Brazil

pdemarco@icb.ufg.br

*

Correspondingauthor

Duringterritorial behaviour, aggressiveattacksamongheterospecificodon. 3 3 are commonand _maycause aseparationofniches, basedonthepreferredsites for territorial defence.Here,territorial behaviour and the characteristics of territories in <JdofErythrodiplaxmedia, Micrathyriacatenataand M.hesperisaredescribed and their territorial_{fidelity, capturing}andmarkingof <3 3arediscussed. In all_spp. stud-ied,there was aclear distinction among the microhabitats defendedasterritories.In bothMicrathyriaspp., 3 3 seemtodefend territorieswith definedresources.In E. media,the defendedresources areless evident. Its 3 3arehighlyaggressiveand show highterritorial fidelity but, apparently, theylose theterritoryif_{they stayaway}from waterforatleast oneday.

lower

_perches

than those of S.

_{sanguineum (Mull.)}

but both

_{species perch}

lower when their

_neighbour

isamale of the other

_{species (REHFELDT}

&

_HADRYS,

1988).

Differences in territorial

_quality

between odonate

_species

are little evident. In some

species,

males defended territories selected

_by

the female basedon the

quantity

of

_oviposition

resourceavailable

_(TSU

BAK1&ONO, 1986; ALCOCK,

1987a;

ALCOCK,

1987b;

KOENIG, 1991; MEEK & HERMAN, 1991; BAT-TIN,

1993),

whereasin other

_species,

territorial defenceseemstobe_morerelated tothe chance of males

_detecting

females

_arriving

atthe

_{pond (ALCOCK,}

_1987a,

1987b;

VAN BUSKIRK,

1986;

BATTIN,

1993;

DE MARCO & RESENDE,

2004)

and in this case, the size of the

territory

candetermine

reproductive

success ofsome

species,

since it increases the chance of male

_visualizing

females

_(VAN

BUSKIRK,

1986).

The

_perches

orsubstrates forthe male

_perching

ina

territory

canalso be diverse(PARR, 1983;OSBORN & SAMWAYS, 1996),and

_change

mainly

asafunction of their

availability.

The

_objective

of this

_study

_was todetermine the criteria for selection of ter-ritories

_by

malesof

Erythrodiplax

mediaBorror,

Micrathyria

catenata

Calvert,

and M.

_hesperis

Ris,

by observing

their territorial

_behaviour,

territorial

_fidelity,

and the characteristics of the

_respective

territories.

METHODS

STUDY AREA - The work wascarried outduring February-Aprilin the Centro deEstudos de Florestas Naturaisof Universidade Federal deViposa,Minas Gerais(20°45’S, 42°5TW).The

re-gionalclimate is Cwa(Kdppen classification), moderatelyhumid sub-tropicalwithahidric deficit betweenMayand_September(GOLFARI, 1975).The_averageannual rainfall is 1450_mm,relative air humidityca_{80% and the temperature}varyingfromamaximum of26,1°Ctoaminimum of 14,0°C

(VALVERDE, 1958).

Duetoalocaldam, Nympheacf.amplaand Eleocharis are_veryabundant in the coastalregion.At the shoreline therearesticks and _grassesthatareusedasperches bysomedragonfly species.

BEHAVIOUR AND TERRITORIAL CHARACTERISTICS - _Duringthestudy period,all activities relativetoterritorial defence,matingritual andovipositionwererecorded. Theemphasis

wasondescriptionsof thebehaviour,courtshipritual before mating,male defence behaviourduring ovipositionand ontheovipositionsites.

To characterize the territories defended by E. media,M. catenata and M. hesperis, wewalked around the shoreline of thepondandrecorded,for eachindividual,the type and heightof itsperch, the distance from theperchtotheshoreline,thevegetationsurroundingtheperchand the distance betweenperchesof theconspecificterritorial males. To determineinterspecificdifferences in the ter-ritorial characteristics,weperformed analysesofChi-squarefor_categoricalvariablesandanalyses of variance (ParametricANOVA and Kruskal Wallis)forquantitativevariables.

TERRITORIALFIDELITY— Forthe sake of theenquireinto theuseof the territoriesbymales of the three_species, 12_adjacentsections of fivemeterslengthweremarked in the littoralzoneof the

pond.Maleswerecapturedand markedonthewingswith non-toxicpaint.The date ofcaptureand thesection inwhich themalewasencounteredwererecorded.Duringthesubsequent days,between

10:00 and 14:00. the segmentswerechecked,all marked individuals andanynew onesmarked. In theanalysesof male territorial fidelity,males withatleastfour_recaptureswereusedfor E. media

Territorial characteristics inneotropicalLibellulidae 215

and those with three_recapturesfor Mcatenataand M.hesperis.This difference in criteriawasdueto the lower frequencyofrecapturedindividuals of the lattertwospecies.Aperiodof five consecutive dayswasused todetermine whether the individuals absent from the dam foratleast onedaycould lose their defended territories. Itwasassumed thatamalehadchangeditsterritorywhen itappeared in adifferentsection,notcontiguoustotheoriginally defended segment. Therelationshipbetween territorial changeand the absence of maleswastestedbyaChi2

analysis.

RESULTS

BEHAVIOUR AND TERRITORIAL CHARACTERISTICS

Erythrodiplax

mediaseemstobemore

aggressive

than the

_{Micrathyria species.}

The femaleswereabundantatthe

_pond

and

_couples

in

_copula

wereoftenseen, butno

courtship

ritualwas everobserved. After

_copulation,

the females remained forsomeseconds

_perched

onbranchesnearthe male

_{territory, generally guarded}

by

the male.

_{Finally, they oviposited}

nearthe shoreline of the

_pond,

in locations with

_{submerged vegetation. Oviposition}

_wasnotrestrictedtothe

respective

male

territories, although

the males

_guarded

the females

_during

the entire

_oviposition

period, attacking

othermales that

_approached.

Micrathyria

catenataisnotavery

aggressive species, though

the

_largest

of the three

species

studied.Maleswere

observed,

butnofemales and

matings

were ever seen.

M.

_hesperis

isasmall and likewisenotvery

aggressive species.

A

_single

mat-ing

wasobserved and therewasneithera

courtship

ritualnorapausebetween

mating

and

_oviposition.

The female

_oviposited

in the male’s

_territory

and it_was

guarded by

the male

_flying

above._However,weobserved another female

ovipos-iting unguarded, perched

on aleafof

_Nymphea

and

submerging

herabdomen.

Thetwo

_{Micrathyria species perched}

more

frequently

on

aquatic macrophytes

(73.6%;

n=

53),

when

compared

with E. media

males,

which

perched

on branch-es,grassesor even onthe soil

_(78.9%;

n= _19;Chi2 =_15.76;df= _1;

p<0.001; Fig.

1)

in additionto

_{using macrophytes.}

_{Comparing only}

_{Micrathyria species,}

M. catenata

perched mainly

on

Eleocharis,

whereas M.

hesperis perched

more

com-monly

on the budor onthe leaf of

Nymphea (74%

and

48%, respectively;

Chi2

= _15.34;df= _1;

p<0,01; Figs

lb,

_1c).

The

_{perch’s height}

_{was also different among the three}

_{species (ANOVA;}

F =

12,85;

df=

_{2; p<0,01).}

M.catenata

perched higher

than M.

hesperis (Fig. 2; Tukey

test;

p<0,01).

The

_{perch’s height}

for E. media males_was

_{intermediate,}

not

differ-ing significantly

from that in either of thetwo

Micrathyria species (Fig.

2;

Tukey

test; p=

_0,13).

Furthermore,the

_vegetation

around the territories of the three

_species

wasdifferent

(Chi

2=_50,90;df=_8;

_p<0,01).

E. media defended territories with Eleocharis and grasses

(37%

and

42%, respectively;

Chi

2=_19,89;df=4;

_p<0,01),

whereas M.

cat-enatadefended

mostly

those with Eleocharis_(76%;Chi2=

28,09;df=

and M.

hesperis

those with

_Nymphea

(94%;Chi:

= _18,1;df= _1;

_p<0,01).

E. media males defendedterritories

_along

the shorelineand

_perched

_on

branch-es,grasses,

directly

onthe soil oron Eleocharisatanaverage

height

of 14cm.

Thetwo

_{Micrathyria species perched}

furtherover the

pond

than E.

media,

par-ticularly

in thecaseof M.

hes-peris (Chi

2 = _18,83; df = _1;

p<0,01)

for

_comparison

with M. catenata

(Kruskal-Wal-lis ANOVA;Chi2=_21,62;df

= _1;

p<0,01).

In M. catena-ta, thecentre of the territo-ries

averaged

onemeterfrom theshoreline;males

_perched

higher

thanthe other

species,

primarily

on Eleocharis. The territories of M.

_hesperis

ex-tendedeven furtherout over thewater than those of M. catenata

(Kruskal-Wallis

ANOVA; Chi2 =

5,37; df=

1;_p=0,021;

_{Fig. 3)}

and

_they

mainly perched

onthe leaves orflowers of

_Nymphea.

How-ever, the distance of M.

hes-peris territory

from the shore-line variedand therewasno

homogeneity

of variance in the data

_(Fig.

3; Bartlett Test; Chi2 = _27,128;df =2;

p<0,001).

There was some

differ-encein the distance between

perches

used

_by

males of the three

_{species (F}

=

_11,06;

df

=_2;

_{p<0,01; Fig.}

_4).The dis-tance between the

perches

used

_by

E. media males

_(x

=229

_cm)

_wassimilartothe

distance between these in M.

hesperis

(.v =338cm;

Tukey

test; p=_{0,87). However,} M. catenatamales

perched

near-Fig.I. Relative frequencyof the typeofperchesinViqosa,

MG. Brazil,usedbymales of:(a)E. M.hesperis. M. catena-media , —_(b) ta, - and(c)

Territorial characteristics in_neotropicalLibellulidae 217

ertheir

neighbours (x

= 103

cm)

thanthe othertwo spe-cies

_(Tukey

_test;

_p<0,001).

TERRITORIALFIDELITY

The

_displacement

of males of the three

_species

between sections of the

_pond

was sim-ilar

(F

=

2,24;

df = _2;

p =

0,13; Fig. 5). However,

the

re-capturerate wasvery

low,

es-pecially,

for the

_Micrathyria

species.

Of the 151 marked E.

media males

_only

35_were

_recaptured

atleast four

_times,

whereas of the 114 marked M. catenataand 24 M.

hesperis

males,

only

sevenandfour,

respectively,

were

re-captured

atleastthree times. Thus

_only

the E. media datawereconsidered fur-ther. The

_frequency

distribution of

_displacement

in this

_species

showed thatmost males defendedterritories in thesame

place, moving

about 10to20 meters, with

only

afew individuals

moving

morethan 35meters

(Fig. 6). Furthermore,

in 77% of the occasions when E.

me-dia males

_stayed

away from the

_pond

foratleastone

day

they

lost their section of

ter-ritorial

defence,

whereas this occurred in

_only

34% of the cases when males remained

atthe

_{pond (Chi}

2=

_9,19;

_n=

38;df =_1;

p<0,05).

DISCUSSION

The

_partition

of microhab-itat among the

species

of a

community

is_{a common}

phe-nomenonandcan occur due to

_{interspecific competition.}

Females of

many

Micrathyria species oviposit

on

aquatic plants,

suchas

Nymphea

(PAULSON,

1969),

henceitishardto_{say whether} themicrohabitats selected

by Micrathyria

speciesasdescribedinthis

_study

_area

consequence of

competition

with E. mediaor acharacteristic of the genus. Libel-lulidae

_{mostly oviposit directly}

onthe surface of thewater or onthe surface of wet

_{floating objects, including plant}

material—

exophytic oviposition

—

(COR-E.media, Fig.2. Mean heightofperchesused for malesof

M.hesperisinViijosa, MG,Brazil. Bars in-dicate the standard deviation.

M._catenataand

Fig.3. Distance ofperchestothepond marginin malesof E. media.M. catenataand M. hesperis,in_{Vi?osa, MG,}Brazil. Positive_values,in thepond; —_{negative values,away from the}

BET, 1999).

The

_oviposition

on

plant

materialcan

possi-bly

result in

₍₁₎

_an increase in the

_parental

care survival as the eggs would be less

ex-posed

to

_predators

such as fish and other

_aquatic

organ-isms,

or

(2)

adecreasein har-assmentof the

_ovipositing

fe-male

_by

males

_trying

to copu-late.

Males with _a

_high

_degree

of residence should be

dis-placed

less often than males that

_continually

lose territo-rial

_disputes

_{(PARR, 1983;}HARVEY &CORBET, 1985; CONVEY,

1989).

The timeof absence ofaterritorialmalefromwater_canbe relatedtotheenergy res-torationorthe

_period

of

_foraging

(FRIED&MAY, 1983;ANHOLT

_etal.,

1991; MARDEN &ROLLINS, 1994;DE MARCO, 1998).Thiscancauseconfusion in the residence ofa

territory (GRIBBIN

&

_{THOMPSON, 1991), resulting}

in_a greater

displacement

of resident males. DUNHAM

₍₁₉₉₄₎

has

_already

empha-sized that the

_departure

for

_foraging

_canmake it difficultto maintain

_good

ter-ritories

_{and, consequently,}

malescanlose

_{mating opportunities. Nevertheless,}

if territorial defence demandsa

high energetic

cost

_(FRIED

&MAY,

1983), larger

males,

with

_{larger energetic}

reserves, are

supposed

to show a

higher

reproduc-tivesuccess, since

they

stay fora

longer

time in the territories without

_foraging

(HARVEY

&CORBET,

1985).

The distance between in-dividuals of _a

_species

_canbe

directly

relatedtothe

_degree

of

_{aggressiveness}

of territo-rial males

_(VAN

BUSKIRK,

1986).

Differences in the re-source abundancecan also affect male distribution

(MEEK

&

_HERMAN,

1991).

At our

locality

Nym-phea

covers a

large

area of the

_dam,

therefore M. hes-peris malesarein the

position

toestablish territories rather distant from each

_other,

per-Fig.4.Distance betweenperchesusedbymalesof E.media, M.catenataand M._hesperisin Viposa, MG,Brazil.Bars in-dicate the standard deviation.

E.media, M.catenata

Fig.5. Meanmobilityof males of and

M. hesperisinViyosa, MG,Brazil.Bars indicatethe stand-arddeviation.

Territorial characteristics inneotropicalLibellulidae 219

haps

minimising

therewith the

_{intraspecific}

competi-tion.

_{Nevertheless,}

no

differ-encein theresource

availabil-ity

was evident in E. media and M.catenata atthe

_pond,

the_greaterdistance between

perches

in E. media territo-rial maleswas

probably

due

to the fact that this

_species

performs

more

aggressive

in-teractions than does M. cat-enata.

The

_spatial

_arrangementof

species

into habitat units is_a

_complex

result of bionomic

_{characteristics,}

includ-ing oviposition

resources and

perch

choice, and interactions with other

_species

in the

_community.

The bionomicfeaturesare

probably

themost

important

force

driving

the choice of habitat used

_by

a

species

ata

locality, despite

the fact that the

_{co-occurring species}

_can

_affect,

atleast within certain

_limits,

the choice of microhabitat.

ACKNOWLEDGEMENTS

We thank ANDERSON LATINI for field assistanceand TANIASANTOS,MARILIA GAIA and MARIANA MELO for valuable revisions andcomments.Thisstudyreceived financial_support from the Brazilian ConselhoNacionalde_Pesquisas(CNPq).

REFERENCES

ALCOCK, X,1987a. Malereproductivetactics in the libellulid dragonflyPaltothemislineatipes: tem-poralpartitioningof territories. Behaviour 103:157-173.

ALCOCK, J.,1987b. The effects ofexperimental manipulationofresourcesonthe behavior oftwo

ca-lopterygiddamselflies that exhibit resource-defensepolygyny. Can. J.Zool. 65: 2475-2482. ANHOLT, B.R.,J.H. MARDEN& D.M.JENKINS,1991. Patternsofmassgainand sexual

dimor-phismin adultdragonflies (Insecta: Odonata).Can. J. Zool. 69; 1156-1163.

BATTIN, T.X, 1993. Theodonate _{mating system, communication,}and sexual selection: areview. Boll. Zool. 60: 353-360.

CONVEY, P,1989. Influencesonthe choice betweenterritorial and satellite behaviour in male Libel-lula_{quadrimaculata}Linn.(Odonata; Libellulidae).Behaviour109: 125-141.

CORBET, P.S., 1999.Dragonflies:behavior and_{ecology of}Odonata. ComstockPubl. Assoc,, Ith-aca,NY.

DEMARCO, P., Jr,1998. The Amazonian Campina dragonflyassemblage:patternsin microhabitat useand behavior inaforaginghabitat. Odonatologica27; 239-248.

DEMARCO, P,Jr & D.C.RESENDE,2004. Cues for_territorychoice intwotropical dragonflies. Neotrop.Ent. 33: 397-401.

E. me-dia

Fig.6.Frequencydistribution ofmobilityof marked males in_Vigosa-MG,Brazil.

DUNHAM,M„ 1994. The effect ofphysicalcharacters onforaginginPachydiplax longipennis (Bur-meister) (Anisoptera: Libellulidae). Odonatologica23: 55-62.

FRIED,C.S. & M L.MAY,1983.Energyexpenditureand foodintake of territorial malePachydiplax longipennis (Odonata: Libellulidae).Ecol. Ent.8: 283-292.

GOLFARI, L., 1975. Zoneamenloecoldgicodo Estadode MinasGeraispararefloreslamenlo.CPRFC, Belo Horizonte,MG.

GRIBBIN,S.D. &D.J. THOMPSON, 1991.The effects of size andresidencyonterritorial disputes and short-termmatingsuccessin thedamselfly Pyrrhosoma nymphula (Sulzer) (Zygoptera: Coenagrionidae).Anim.Behav.41:689-695.

HARVEY,I.F.&RS. CORBET, 1985. Territorial behaviour of larvae enhances matingsuccessof maledragonflies.Anim.Behav. 33: 561-565.

HASSAN, A.T.,1978._Reproductivebehaviour of Acisoma_panorpoidesinflatum_{Selys (Anisoptera:} Libellulidae). Odonatologica7: 237-245.

JOHANNSSON, O.E.,1978. Co-existence of larvalZygopteracommontothe NorfolkBroads(U.K.), 1. Temporalandspatial separation. Oecologia32; 303-321.

KOENIG,W.D.,1991.Levels of female choice in the white-tailed skimmer Plathemislydia (Odonata, Libellulidae).Behaviour119: 193-224.

MARDEN,J.H. & R.A.ROLLINS,1994. Assessment of energyreservesbydamselflies engagedin aerialcontestsformatingterritories.Anim. Behav.48: 1023-1030.

MAY, M.L.,1980.Temporalactivity patternsofMicrathyriain Central America(Anisoptera: Libel-lulidae). Odonatologica9: 57-74.

MEEK,S.B.&T.B.HERMAN,1991. The influenceofovipositionresources onthedispersionand behavior ofcalopterygiddamselflies. Can.J. Zool. 69: 835-839.

OSBORN, R.& M.J.SAMWAYS, 1996. Determinants of adult dragonfly assemblagepatternsat newpondsin South Africa.Odonatologica25: 49-58.

PARR, M.J.,1983. Ananalysisofterritorialityin libellulid dragonflies (Anisoptera: Libellulidae). Odonatologica12:39-57.

PAULSON,D.R., 1969. Ovipositioninthetropical dragonflygenusMicrathyria (Odonata: Libel-lulidae).Tombo12:12-16.

REHFELDT. G.E. &H.HADRYS,1988.Interspecific competitioninsympatric Sympetrum

sangui-neura(Muller)and S. flaveolum(L.) (Anisoptera: Libellulidae). Odonatologica17:213-225.

RUSSELL, R.W.,M.L,MAY,K.L. SOLTESZ &IW.FITZPATRICK,1998. Massive swarm migra-tions ofdragonflies (Odonata)ineasternNorth America.Am. Midi. Nat. 140: 325-342. TSUBAKI,Y. & T.ONO, 1986._Competitionfor territorialsites andalternative_matingtactics in the

dragonfly, Nannophyapygmaea Rambur(Odonata: Libellulidae).Behaviour 97: 234-252. VALVERDE, O.,1958. EstudoregionaldaZonada Matade Minas Gerais.Revla bras. Geogr.20;

3-79.

VANBUSKIRK, J.,1986. Establishment andorganizationof territories in thedragonfly Sympetrum rubicundulum (Odonata: Libellulidae).Anim. Behav.34: 1781-1790.