Dragonfly Predation on Butterflies in a Tropical Dry Forest

Author(s): Alfonso Alonso-Mejia and Monica Marquez

Source: Biotropica, Vol. 26, No. 3 (Sep., 1994), pp. 341-344
Published by: Association for Tropical Biology and Conservation
Stable URL: http://www.jstor.org/stable/2388856
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BIOTROPICA26(3): 341-344 1994

NOTES

DragonflyPredation on Butterfliesin a Tropical Dry Forest

Keywords: butterflies;CostaRica; Guanacaste;Lepthomisvesiculosa;Libellulidae;Odonata;Palo Verde;
predation;Siproetastelenes;unpalatability.

ADULT DRAGONFLIES FEED upon a varietyofprey(Corbet1980), includingoccasionalbutterflies and moths

(Poulton 1906, Daecke 1915, Hobby 1933, Bell & Whitcomb1961, Dunkle 1989, White & Sexton
1989). Most of the publishedinformation suggeststhat butterflies are only a small componentof a
dragonflies'diet.Here,we describea systemwheretheGreatPondhawkDragonfly, Lepthomisvesiculosa
Fabricius(Odonata: Libellulidae),preysheavilyupon a wide varietyof adult butterfly species.We also
observedL. vesiculosarejectingseveralindividualsof Siproetastelenes(Lepidoptera,Nymphalinae)and
discussthepossibleroleof defensivechemicalsfoundin butterflies thatmay deterinvertebrate predators.
The GreatPondhawkDragonfly is a bold, swiftflyerthatfeedspredominantly on largeflying insects
(Dunkle 1989). It is a relativelylargedragonfly (length54-64 mm, abdomen 39-46, hindwings35-
43) thatoccursin marsh-bordered ponds and lakes fromthesoutheastern UnitedStatesand theAntilles,
southto Paraguay(Carle 1982).
Observationsweremade at Palo Verde National Park (PVNP), GuanacasteProvince,Costa Rica.
This area is a mosaicof secondarygrowthforestand abandonedpastures,delimitedby marshesand the
TempisqueRiver.The terrestrial vegetationis characterized as lowlandtropicaldryforest(Gill 1989).
We observedpredationby dragonflies at threesites.At the firstsite,we walked in an open marsh
(300 x 75 m) near the airstripdose to the OrganizationforTropicalStudies(OTS) station.During
mostof the studythe marshhad no standingwater.At the secondsite,a dirtroad, we sampled 2500
x 4 m fromtheAdministration Headquartersof PVNP to thecattletanks(i.e., waterhole). The third
sitewas at the foresttrail"La Venada" (600 x 1 m). We walked thesetransects twicea day, in the
morning(0800-1200 hr) and in the afternoon (1400-1800 hr) from2 June to 1 August 1992. We
founda highabundanceof butterflies, sincemostspecieshad emergedduringtherainyseason(DeVries
1987).
Evidenceforpredationwas both directand indirect.We watchedL. vesiculosaattacking,carrying
off,and feedingon 31 individualsof 19 speciesof butterflies (Table 1). We followedeach of these
dragonfliesuntilit finishedfeedingon the butterfly. We observedthatthe dragonflies avoided feeding
on the butterfly wings,so theyfellto the groundas theyseparatedfromthe thorax.We collectedthe
rightforewing forspeciesidentificationand measurement. We noticedat the base of the wingsa series
of "bite marks" leftby the dragonfly. This typeof damage was consistent, easy to identify,and very
differentfromdamage caused by birds(Wourms & Wasserman1985). Indirectevidenceconsistedof
butterflywingsfoundon thegroundthathad thesamebite-mark patternfoundon thewingsofbutterflies
we observedbeingattacked.We countedbutterflies onlyif we foundtherightforewing(RFW; we did
notindude otherwingsto avoid overestimations); and ifclearlyrecognizabledragonfly damage appeared
on thewings.
During the study,L. vesiculosaattacked241 butterflies (35 species)and 10 diurnalmoths(three
species;Table 1). Theyattackedbutterflies throughout the day (0830 to 1756 hr; N = 26), and killed
individualsthat rangedin size from 11 mm (e.g., Microtia elva) to 45 mm RFW (e.g., Dryadula
phaetusa).The overallmean size of the preyed-uponbutterflies was 30.7 mm (N = 199). Gill (1989)
reporteda totalof 172 butterfly speciesin an area nextto PVNP, of which68 specieswere observed
flyingat the timeof the study(A. Alonso-M.,pers. obs.). Therefore, the Great PondhawkDragonfly
preyedupon 52 percentof the butterfly speciesfoundin the area at thattimeof theyear.
Lepthomis vesiculosacapturedbutterflies in the air by seizingthe dorsalsurfaceof the thorax.They
would typicallybite the head immediately and incapacitatethe prey(N = 31; 19 species).When the
dragonfliessettledon a branch,theyused theirforelegsto manipulatethebutterfly intoa feedingposition
and fed on the thoraxand abdomen.They neverate the wings.In some instances,dragonflies flewfor
shortdistancesif theyweredisturbedor neededto repositionthebutterfly. It took a dragonflyabout 30
min to consumeeach preyitem (mean = 35.5; SD = 5.2; N = 6 observations).They did not attack
perchedbutterflies. In severalinstances(N = 12; fourspecies) butterflies avoided being capturedby

341

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342 Alonso-Mejfaand Marquez

TABLE 1. preyeduponbythedragonfly
Butterflies Lepthomisvesiculosaat Palo VerdeNational Park, Guanacaste
Province,CostaRica duringJuneand July(wetseason)of 1992.

Dragonflypredation Forewinglength

Directly Inferredfromdiscardedwings (mm)

Butterfly
species observed Marsh Edge Forest Mean (N) Range
Papilionidae
Papilioninae
Eurytidesepidaus - 4 44.0 (3) 43-45
Pieridae
Coliadinae
Anteosmaerula - 1 38.0 (1)
Aphrissaboisduvalii - 2 34.0 (2)
A. statira 1 1 33.0 (1)
Euremadaira 1 1 4 2 22.2 (5) 15-26
E. dina 1 - 42 9 24.1 (45) 19-26
E. nise - 2 20.5 (2) 18-23
Kricogonia
lyside - 13 2 27.9 (14) 20-31
Phoebis
agarithe - 15 - 34.5 (15) 32-37
P. argante 1 1 3 - 33.8 (4) 32-35
P. sennae 2 1 9 - 31.0 (5) 28-33
Pierinae
Appiasdrusilla - 7 1 32.0 (5) 27-37
Ascia monuste 2 2 17 - 30.8 (17) 29-36
A. josephina - 1 - 40.0 (1)
Nymphalidae
Nymphalinae
Anartiajatrophae - 2 - 27.0 (2) 26-28
Euptoietahegesia 3 8 1 32.8 (9) 31-35
Hamadryasfebrua - 3 36.0 (3) 34-38
H. glauconome 2 1 6 33.3 (6) 31-36
Junoniaevarete - 2 1 28.7 (3) 28-29
Marpesiapetreus - 1 14 37.7 (13) 34-40
Siproetastelenes 4 3 9 41.7 (10) 40-43
Smyrnablomfildia 1 2 35.0 (2) 33-37
Heliconiinae
Agraulisvanillae 2 14 39.5 (8) 38-40
Dryadulaphaetusa - 1 45.0 (1)
Dryasiulia - 2 43.5 (2) 43-44
Heliconiuscharitonius - 1 38.0 (1)
Melitaeinae
Microtiaelva 1 1 1 13.0 (2) 11-15
Charaxinae
Anaea aidea 4 1 7 1 30.0 (8) 29-32
Zaretisellops 1 1 34.0 (1)
Danainae
Danaus eresimus 1 - 1 1
Satyrinae
Taygetisandromeda 1 1 35.0 (1)
Lycaenidae(3 spp) - 2 1
Hesperiidaesp 1 1
Moths (3 spp) 3 1 9 17.3 (7) 11-30
Total 31 15 205 21 30.7 (199) 11-45

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 Notes 343

droppingto the groundand remainingthere.Dragonfliesflewabove the butterflies but neverattacked

themon theground.However,theywould attackbutterflies thatflewagain. Minno and Emmel(1993)
reported L. vesiculosafeedingon thebutterfliesAscia monuste (Pierinae)and Papilio andraemon (Papilioni-
nae) in BiscayneNational Park,Florida.We also observedL. vesiculosapreyingupon otherdragonflies
(e.g., Erythemis plebeyaOdonata: Libellulidae;N = 8).
The numberof observedattacksand the numberof wingsfoundsuggeststhatL. vesiculosapreyed
upon more butterflies along the forestedge (205 individuals;31 species) than in the othertwo sites
(Table 1). This mightbe theresultof a higherbutterfly densityand diversity at the forestedge (Brown
1991; A. Alsonso-M.,pers. obs.). However,we sampled a largerarea at the edge of the forestand
dragonflies mighthave been easierto see as well as wingseasierto find.
The greaternumberof individualstakenof some species(e.g., Euremadina; Table 1) may reflect
theirabundanceratherthan the preferences or selectionby the dragonflies, althoughsome butterflies
mightbe easierto catch.At highdensities,otherinsectsincludingbees (Wright1944), beetles(Soria &
Machado 1982), locusts (Stortenbeker1967), and bollwormmoths (Bell & Whitcomb 1961) are
frequently consumedby dragonflies. Pritchard(1964), however,emphasizedthe importanceof small
prey,even fordragonflies thatfeedon largeprey.
The GreatPondhawkDragonfly rejected5 out of 12 Siproetastelenes,
a nymphaline partiallyrejected
by some bird species(Brower 1984), but acceptableto others(Brower& Brower1964; Chai 1986,
1990). We watchedfourattackson S. stelenes. One was completely eaten,whereas,thedragonflies rejected
theotherthreewhilechewingon thehead. It seemedas iftheydroppedthebutterfly whentheyencountered
(i.e., tasted)the hemolymphof the butterfly. A varietyof sensillahave been describedforthe mouth
partsof dragonflies, some of whichare presumablychemoreceptors (Petryszak1977). In Costa Rica, S.
stelenesfeedson Acanthaceae(DeVries 1987). Some membersof thisfamilyare knownto have iridoid
glycosides, chemicalcompoundsknownto be sequesteredby butterflies and knownto be toxicto some
vertebrate predators(Bowers1991). Furtherstudiesareneededto determine whetherS. stelenesis capable
of sequestering thosecompoundsand the mechanismto deterdragonflies.
Sunny and dry places, like Guanacaste, are generallydeficientin butterflies such as Heliconius
(Heliconiinae),Parides (Papilioninae),and ithomiines(Ithomiinae)knownto be unpalatableto birds
(Chai 1990). However,L. vesiculosadid not rejectotherbutterfly speciesknownto be unpalatableor
partiallyunacceptableto severalspeciesof birds.These includeAscia monuste (Pough & Brower1977;
Chai 1986, 1990) withno rejections out of 19 individualsand Agraulisvanillae (Broweret al. 1963,
Brower1984) withnonein 14. Similarly, Whiteand Sexton(1989) reportedthedragonhunter Hagenius
brevistylus (Odonata, Gomphidae) preyingupon monarchbutterflies (Danaus plexippus,Lepidoptera,
Danainae), an insectlong knownto be unpalatableto birds(Brower1984). Sincedragonflies do not eat
the wings,theymay be less likelyto encounterunpalatablesubstances.Therefore, the chemicaldefense
of some butterfly speciesagainstbirdsis apparently in deterring
ineffective dragonfly predation.
Our findings thatL. vesiculosafeedsextensivelyand somewhatselectively on severalspeciesofbutterflies
raisessome importantquestions.For example,what is the significance of invertebratepredationin the
evolutionofbutterfly Is therea palatability
palatability? spectrum(sensuBrower1984) in butterfly toxicity
and/orin theindividualresponseof dragonflies to thosechemicals?Studiesare neededon theimportance
of invertebrate predationon invertebrates, and its interactionwithdefensivechemicals.
We thankL. Brower,R. Edwards,S. Malcolm,K. Silvius,and M. Zalucki forcommentson early
versionsof themanuscript, and I. Carreraforfieldassistance.This studywas supportedby Pew Charitable
Trustsawards by OTS to AAM and MM, by a Tropical Conservationand DevelopmentProgram
(University of Florida)grant,and by a DGAPA-UNAM (Mexico) fellowshipto AAM.

BELL, R., AND W. H. WHITCOMB. 1961. Erythemissimplicollis predatorof the boliwormmoth.

(Say) a dragonfly
Fla. Entomol.44: 95-97.
BOWERS,M. D. 1991. Iridoid glycosides.In G. A. Rosenthaland M. R. Berenbaum(Eds.). Herbivores,their
interactions withsecondaryplantmetabolites,vol. I, pp. 297-325. AcademicPress,San Diego, California.
BROWER, L. P. 1984. Chemicaldefensein butterflies.In R. I. Vane-Wrightand P. R. Ackery(Eds.). The biology
of butterflies,pp. 109-134. AcademicPress,London,England.
and plantpoisons:a studyin ecologicalchemistry.
, ANDJ. V. Z. BROWER. 1964. Birds,butterflies, Zoologica
49: 137-159.

This content downloaded from 205.133.226.104 on Mon, 14 Dec 2015 20:13:09 UTC
All use subject to JSTOR Terms and Conditions
344 Alonso-Mejfaand Marquez

AND C. T. COLLINS. 1963. Experimental studiesof the mimicry.7. Relativepalatabilityand

miillerianmimicryamong neotropicalbutterflies of the subfamilyHeliconiinae.Zoologica 48: 65-84.
BROWN, K. S. JR. 1991. Conservation of neotropicalenvironments: insectsas indicators.In M. Collinsand J. A.
Thomas (Eds.). The conservationof insectsand theirhabitats,pp. 350-401. Academic Press,London,
England.
CARLE, F. 1982. A contribution to the knowledgeof the Odonata. UnpublishedPh.D. Dissertation,Virginia
Polytechnic Instituteand StateUniversity, Virginia.
CHAI, P. 1986. Field observations and feedingexperimentson the responseof rufous-tailed jacamars(Galbula
ruficauda)to free-flying butterflies
in a tropicalrainforest.Biol. J. Linn. Soc. 29: 161-189.
1990. Relationshipsbetweenvisual characteristics of rainforestbutterfliesand responsesof a specialized
insectivorous bird. In M. Wicksten(Ed.). Adaptivecolorationin invertebrates, 106 pp. 31-60. Texas A&M
UniversitySea GrantCollege,Galveston,Texas.
CORBET, P. S. 1980. Biologyof Odonata. Annu. Rev. Entomol.25: 189-217.
DAECKE, E. 1915. Note on Hageniusbrevistylus chewinga femalePapilio turnus.Entomol.News 26: 335.
DEVRIES, P. 1987. The butterflies of Costa Rica. PrincetonUniversity Press,Princeton,New Jersey.
DUNKLE, S. W. 1989. Dragonfliesof the Florida Peninsula,Bermuda,and the Bahamas. ScientificPublishers,
Gainesville,Florida.
GiLL,D. E. 1989. A naturalist'sguide to the OTS Palo Verde Field Station,El Refugiode Fauna Silvestre'Dr.
RafaelLucas RodriguezCaballero',GuanacasteProvince,Costa Rica. San Jose,Costa Rica. Organizationfor
TropicalStudies.
HOBBY, B. M. 1933. The preyof Britishdragonflies. Trans.Entomol.Soc. South Eng. 8: 65-76.
MINNO, M. C., AND T. C. EMMEL. 1993. Butterflies of the Florida Keys. A Mariposa PressEdition,Gainesville,
Florida.
PETRYSZAK, A. 1977. The sense organsof the mouth parts in Libellula depressaL. and L. quadrimaculataL.
(Odonata). Acta Biol. Cracov. Ser. Zool. 20: 87-100.
POUGH, F. H., AND L. P. BROWER. 1977. Predationby birdson greatsouthernwhitebutterflies as a functionof
palatability,sex, and habitat.Am. Midl. Nat. 98: 50-58.
POULTON, E. B. 1906. Predaceousinsectsand theirprey.Trans.R. Entomol.Soc. Lond. 1906: 399-401.
PRITCHARD, G. 1964. The preyof adult dragonflies in westernAlberta.Can. Entomol.96: 821-825.
SORIA,S. J., ANDA. B. M. MACHADO.1982. GynacanthabifidaRambur (Odonata, Aeshnidae),a new enemyof
Xyleborus spp (Coleoptera,Scolytidae),a pest of cocoa in Bahia, Brazil.Rev. Theobroma 12: 257-259.
STORTENBEKER, C. W. 1967. Observationson thepopulationdynamicsoftheRed Locust,Nomadacrisseptemfasciata
(Serville),in its outbreakareas. Agric.Res. Rep. (Wageningen)No. 694.
WHITE, S. D., AND 0. J. SEXTON. 1989. The monarchbutterfly (Lepidoptera:Danaidae) as preyforthe dragonfly
Hageniusbrevistylus (Odonata: Gomphidae). Entomol.News 100: 129-132.
WOURMs,M. K., AND F. E. WASSERMAN. 1985. Bird predationon Lepidopteraand the reliability of beak-marks
in determining predationpressure.J. Lepid. Soc. 39: 239-261.
WRIGHT, M. 1944. Some randomobservations on dragonfly habitswithnoteson theirpredaceousness on bees.J.
Tenn. Acad. Sci. 19: 295-301.
Alfonso Alonso-Mejia and Monica Marquez
Departmentof Zoology
Universityof Florida
Gainesville,Florida32611

BIOTROPICA26(3): 344-348 1994

Roost Interactionsbetweenthe CommonVampireBat

(Desmodus rotundus)and Two FrugivorousBats
(Phyllostomusdiscolorand Sturniralilium)
in Guanacaste, Costa Rica
Key words: Costa Rica; Desmodus; Phyllostomus; roostingbehavior; Sturnira; vampire control.

PRIOR TO THE INTRODUCTION of domestic animals to Latin America, vampire bat (Desmodus rotundus)
populations were probably small due to the limited numbers of large wild mammals as a source of blood
(Turner 1975, Greenhall 1988, Lord 1988). The introductionof domestic livestockprovided the common
vampire with an abundant food supply and must have enabled populations to increase dramatically. D.

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