Trophic strategy of ant-eating Mexcala elegans (Araneae: Salticidae): looking for evidence of evolution of prey-specialization

2011·5 MB·English

by  Stano Pekár

2011.TheJournalofArachnology39:133-138
Trophic strategy ofant-eating Mexcala elegans (Araneae: Salticidae): looking for evidence of
evolution ofprey-specialization
StanoPekar: DepartmentofBotanyandZoology, FacultyofSciences, Masaryk University, Kotlafska2,611 37 Brno,
Czech Republic. E-mail: [email protected]
CharlesHaddad: DepartmentofZoology«feEntomology,UniversityoftheFreeState,P.O. Box339, Bloemfontein9300,
SouthAfrica
Abstract. WeinvestigatedthetrophicstrategyofMexcalaelegansPeckham&Peckham1903,anant-eatingsalticidspider
fromSouthAfrica,inordertogainbaselineinformationconcerningtheevolutionofpreyspecialization.Westudiedits
naturalprey,preyacceptance,andchoiceusingavarietyofpreyspecies.Initsnaturalhabitat,thespidercapturedonly
ants,mainlyitsmimeticmodelCamponotiiscinctelliis,indicatingthatthespeciesisastenophagousant-eater.However,in
thelaboratory,M.eleganscaptured 12differentinvertebratetaxawithefficiencysimilartothecaptureofants,suggesting
thatitiseuryphagous.Forthecaptureofantsbutnotforotherprey,itusedaspecializedprey-capturebehavior.Inprey-
choiceexperiments,thespidersdidnotpreferantstoflies.Wefoundnoevidenceforneuralandbehavioralconstraints
related to identification and handling ofprey. Our results suggest that M. elegans is aeuryphagous specialist using a
specializedant-eatingcapturestrategyinwhichpreyspecializationhasevolvedasabyproductofriskaversion(“enemy-
freespace”hypothesis).
Keywords: Prey,huntingbehavior,myrmecophagy,mimicry,evolution
Stenophagy,theutilizationofanarrowpreyrange,maybe Spidershavebeenfoundtobemainlyeuryphagous(Nentwig
a product of an innate response due to evolutionary 1987),buttherearequiteafewcasesofstenophagousspecies.
transitions and fitness trade-offs or a proximate response Evidenceforstenophagyismainlyanecdotal.Themostfrequent
duetospecificenvironmentalconditions;i.e.,dominanceofa type of stenophagy observed is myrmecophagy; spiders in
certain prey species. In the former case, such species are several families (e.g., Zodariidae, Gnaphosidae, Theridiidae)
stenophagous specialists because they are not able to catch demonstratespecializationinantpredation(Heller1976;Carico
and utilizealternative prey. Inthelattercase, such predators 1978; Pekar2004). Whilethemajorityofsalticidspidersrarely
are stenophagous generalists since they possess versatile feedsonants(e.g.,Nentwig1986;Guseinov2004),sometropical
adaptations allowing them to capture and process a variety speciesaremyrmecophagous(Cutler1980;Wing1983;Jackson
ofpreyinenvironmentswithdiverseprey(Sherry 1990). &VanOlphen1992;Lietal. 1999;Allan&Elgar2001;Jackson
Evolutionofstenophagousspecialistshasbeenexplainedby & Li 2001). These myrmecophagous species use a specialized
a number of hypotheses (particularly in herbivores). The tactictocaptureants(e.g.,Jackson&VanOlphen1992;Jackson
enemy-free space hypothesis postulates that stenophagy has & Li 2001). However, no salticid species is known to prey
evolved as a byproduct of using host/prey as a refuge or exclusivelyonants.
defense (Brower 1958). The neural constraints hypothesis We investigated the prey capture behavior of a salticid
(Jermyetal. 1990)suggestsaninabilitytorecognizecuesfrom spider Mexcala elegans Peckham & Peckham 1903 in South
other than preferred prey. The physiological trade-off Africa. Mexcalaelegansappearstobean inaccurate Batesian
hypothesis (Singer 2001) is relevant when the predator is mimicofa few ground-livingant species. It is adistinctively
constrained in utilization of other than its preferred food. polymorphicspider, with threecolorvariations: 1)a metallic
And, the optimal-foraging hypothesis (Singer 2008) predicts silver-gray bodywith black triangularabdominal markingin
lowerefficacyinthecaptureofalternativeprey. late instar immature and adult specimens, resembling silver-
Revealingthetrophicstrategyofaspeciesrequiresmultiple grayground-dwellingants(Fig. lA),presumablyCamponotiis
approaches. Analysis of natural prey alone cannot provide cinctellus that are common on the ground surface and low
completeevidenceforatrophicstrategy.Suchdataneedtobe foliagein northeastern South Africa; 2) a metallicsilver-gray
supplemented by extensive laboratory prey acceptance and body adorned by two pairs oflarge yellow abdominal spots
choiceexperiments. This is because the natural preyanalysis (Fig. IB)inadultspecimensresemblinglargeground-dwelling
reveals only the realized trophic niche that measures actual winglessfemalemutillidwasps;and3)ametallicblueprosoma
diet use and results from the effect of both intrinsic and andbrightmetallicgreenabdomeninearlyinstarimmatures,
extrinsic variables. In contrast, laboratory experiments can possiblyinaccurateantmimics.
reveal the fundamental trophic niche that is determined by OtherspeciesofthegenusMexcalafeedontheirantmodels
intrinsic variables only (Bolnick et al. 2003). Furthermore, (Curtis 1988). Therefore, we predicted that M. elegans also
trade-offs (behavioral, morphological, or physiological) that hunts its model ants, thus supporting the enemy-free space
constrainpreyutilizationinstenophagousspecialistscanonly hypothesis. In order to reveal any trade-offs, neural or
bedeterminedexperimentally.Thegatheredevidencecanthen behavioral, thatwould lead to support alternativeevolution-
beusedtodrawconclusionsonthetrophicstrategy. ary hypotheses, we performed both field and laboratory
133
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134 THEJOURNALOFARACHNOLOGY
\
A
—
Figure 1. Mexcalaeleganscapturingantsinthefield.A.FemaleofthegraycolorvariationcapturingCaniponotuscinctellus\B.Femaleof '
thespottedvariationcapturingCaniponotussp.2.
surveys. Afterexaminingnaturalpreycaptureinthefield,we and 12 h laterinitiated anewtrialwithadifferent prey. Ifa
tested the ability ofthis species to catch and eat alternative prey was accepted, we initiated the next trial 24 h later. For
prey in the laboratory, and also whether it prefers ants to each trial, we recorded whether the prey was attacked and
alternativeprey. subsequentlyconsumed. Intrialswithantortermiteprey,we j
METHODS also recorded the latency to attack (i.e., time between the
Field survey.—We investigated the natural prey of M. slpaitdeenrcyotroiepnatraatliyosnist(io.we.a,rtdimtehebeptrweeyenanthdetahtetacakttaacnkd)garanbdbitnhge
elegans during field trips to Ndumo Game Reserve, South thepreyin thechelicerae).
Africa inJune-Julyand November-December2004-2009(11 Intheprey-choiceexperiment,performedaftertheacceptance
trips in total) that formed part of a larger arachnid experiment with a paired design, we released two non-native
biodiversitysurveyinthereserve. Wecollected64M. elegans preyitemsofsimilarsize(relativeprey/spidersize:0.4-1)atthe ,
spiders in a variety ofhabitats: Acacia nigrescens woodland sametimeintothedishoccupiedbyaspider. Spiders(n = 15) ]
(1.6% of total), A. xanthophloea forest (7.8%), broadleaf werestarvedfortwodayspriortoeachtrial. Weusedanant,
woodland (25%), floodplains (25%), Ficus sycomonis forest Tetramorium caespitiim (Myrmicinae), and a fly. Drosophila
(3.1%),andsubtropicalbush(37.5%). Individualspiderswere melanogaster(Drosophilidae),ortwoantspecies, T. caespitum |
followed for up to 10 minutes to see whether they would andLasiiisniger(Formicinae).Thesetwoalternativetreatments
capture ants and to note the prey capture behavior and wererepeated foreachindividualonarandombasis. Inthese
interactions with different ant species. Ifthey had a prey in paired trials, we recorded which ofthe two prey insects was
their chelicerae, the spiders were collected and preserved in attackedandwhichonewasconsumed.Atleastoneoftheprey
ethanol and brought to laboratory where their sex and the insectswasattackedandconsumedineachtrial.Allexperiments
prey was identified to species level. We measured the size of wereperformedbetween09:00and 16:00h. '
adultmales(// = 15)andfemales(n = 15)and 15antworkers Data analysis. We analyzed data using various methods
of each species captured in the field using an ocular withinR(RCoreDevelopmentTeam2009).Forthefielddata,
micrometerwithinabinocu—larstereomicroscope. weusedANOVAtocomparepreysizeamongimmature,adult
Laboratory experiments. For intensive studies of prey male and adult female spiders. Because there were repeated ’
capture and prey choice, we brought 15 live juvenile M. measuresofthesameindividualsinbothexperiments,weused
elegans (body size 3.5-5.3mm) collected at Ndumo Game Generalized Estimating Equations (GEE) as an alternativeto ’
Reserve to the home laboratory. We housed spiders individ- Generalized Linear Models. This method allows implementa- j
uallyinPetridishes(diam.4.5cm)withafilterpaperattached tionofanassociation(correlation)structurethatcorrectsfortoo 1
to the bottom. A small piece ofcotton moistened at 2-day small standard errors of parameter estimates and inferences I
intervals served as a water resource. Using these spiders, we favoring acceptance of the alternative hypothesis (Hardin &
performed twodifferentexperiments. Hilbe2003).WeusedGEEwithbinomialerrorstructure(GEE- 1|
Intheacceptanceexperiment,weusedacompleterepeated b) to compare capture frequency of the prey acceptance i
measures design, offering each spider (n = 15) each of 17 experiment,sincethe responsevariableswererelativefrequen-
potential prey species in random order (Table2). The prey cies.WeusedGEEwithGammaerrorsandloglink(GEE-g)to ,
were not native to the spider, as the experiments were comparelatenciesamongselectedpreyspecies,astheresponse .
performedinEurope,butweusedonlypreyfromordersthat variablewastime,and variancewasexpected toincreasewith (
alsooccurinSouthAfrica. Therelativebodysizeoftheprey themean.Weusedaproportiontesttocomparethefrequency :
(1.6-8.0mm)tospiderbodylength(3.3-5.3mm)was0.3-2.4. ofattackandconsumptionseparatelyforselectedpreyspecies.
Wreespoonbdsetroveadperaeychittermialwictohnitnin1u5oumsilny,.wIef sstpoipdepresd tdhied trnioatl WMceNeamnaalryzteesdtdtuheetoprpeayi-rcehdoitcrieals.experiments data with the ||
i
PEKAR&HADDAD—TROPHICSTRATEGYOFMEXCALA ELEGANS 135
—
Table 1. Natural prey ofjuvenile, male, and female Me.xcala
elegans specimens determined during field observations in Ndumo
GameReservefrom2004to2009.Thesizeisanaveragetotalbody
lengthofworkersattackedbyspiders.
Ants Spiderpredators
Subfamily/species Size[mm] Juveniles Males Females Total
Formicinae
Anoplolepis
custodiens(Smith) 5.9 0 1 4 5
Camponotus
cinctellus
(Gerstacker) 7.2 6 12 6 24
Camponotussp.2
(maculatusgroup) 8.6 2 3 3 8
Polyrhachissp. 8.6 0 4 5 9
Myrmicinae
Crematogastersp. 3.5 2 0 1 3
Myrmicaria
natalensis(Smith) 6.3 0 1 3 4
Tetramorium Figure 2.—Comparisonofthepreysize(mean ±SE)capturedby
quadrispinosum juveniles,malesandfemalesinthefield.
Emery 3.5 3 0 0 3
Ponerinae attacked, they were less likely to consume Triholium larvae
Pachycondyla and Pardosa spiders (Proportion tests, X" > 5.5, P < 0.02).
tarsata(Fabricius) 16.5 0 0 4 4 Spidersattackedpreythatwereonaverage 1.03oftheirbody
Streblognathus length (Q25 = 0.64, Q75 = 2.2, n = 255). In the choice
PseupRdeooebmteeyrrrstmisiocninae 11.6 0 0 2 2 eslypxipadeserrifslmieeanstt(tsaM,ccksNepdiedmeaarnsrdattcetsoatnsc,skueXdm-e/adn=dL0ac,soiPnuss=uamnIetdnsa=anst1sf5r)a.esqSufiermneitqlluayernltya,-s
Teatrmahpiognueara(Emery) 6.8 2 0 0 2 TetMer.axmcoarliaumeleagnatnss(uMsceNdedmifafreretnesttsp,reXd^a/to>ry0.b4e,hPav>ior0.t5o).catch
Total 15 21 28 64 different prey taxa. Although spiders ignored woodlice and
beetles, they stalked aphids, crickets, bugs, and Theridion
spiders but did not attack them. Spiders grabbed small
springtails, leafhoppers, moths, and flies with their forelegs
— RESULTS and moved them to their chelicerae. In contrast, they
Field survey. In the field, M. elegans captured and repeatedlyattacked termiteshead-on, and thengrabbed hold
consumedtenspeciesofantsfromfoursubfamilies(Table 1). ofthe insect’s thorax. To catch ants, the spider approached
Weobservednopreyotherthanantsbeingcaptured.Among from the rear, maintaining a distance ofthree to four body
ants,themostfrequentpreywasCamponotuscinctellus.Adult lengths from an ant, all the while moving the front legs and
male(bodysize5.3-8.3mm)andfemale(6.1-8.9mm, Fig. 1) abdomen up and down. The spider attacked quickly from
M. eleganscaptured significantly larger ant species {Campo- behind, biting the ant on the abdomen. The spider then
notus, Polyrhachis, Anoplolepis and Myrmicaria) than the retreated and followed its ailing prey with raised forelegs
juveniles, which generally preyed on smaller ants such as (Fig. 3A), maintainingadistanceofabouttwo bodylengths.
Crematogaster, Tetramorhim, and Tetraponera (ANOVA, Once the ant slowed down, the spider grabbed the ant’s
F2,6o = 4.5, P= 0.013, Fig.—2). antenna with its chelicerae (Fig. 3B), and after a minute, it
Laboratory experiments. Although the prey acceptance moveditsholdtothethorax.
experimentshowedthatthespiderswerecapableofattacking Among the four ant and one termite species used in the
diverse prey, and the prey choice experiment showed no trials,thespidersshowedsignificantlydifferentlatencyintheir
preference between prey types, the spiders did respond attacks (GEE-g, = 9.6, P = 0.047, Fig. 4A). Spiders
differently to varying prey types. In the acceptance experi- attacked Lasiusand Messor ants with a significantly shorter
ment, spiders responded differently to the 17 potential prey latency than Formica ants (contrasts, P < 0.02). There was
species. Thefrequencyofattacksdifferedamongthe 17prey also a significantly different paralysis latency among these
species (GEE-b, = 194, P < 0.0001). Spiders did not prey ants (GEE-g, X^4 = 49.4, P < 0.0001, Fig. 4B). Large
attack crickets, beetles, Theridion spiders, or woodlice and FormicaandMessorantshadasignificantlylongerlatencyto
springtailsandbeetlelarvaewereonlyattackedbyhalfofthe paralysis than small Lasiusand Tetramorium ants (contrasts,
spiders. Other prey species such as ants, Pardosa spiders, P < 0.03). Termites of the same size as small ants were
termites, flies, and moths were always attacked (Table2). paralyzed more quickly than all ant species (contrasts, P <
Althoughspidersconsumed themajorityofpreyspeciesthey 0.0001).
136 THEJOURNALOFARACHNOLOGY
i
Table 2.—Listofpreyusedinlaboratoryexperiment.Thesizeof DISCUSSION
pPreeryceinstaagneaovfercaognesutomteadlbisodofytlheonsgetht.ha«t=wer1e5atrtitaalcskefdor.eachspecies. fieWldeobfsoeurnvdataiocnosntsruagsgteisntgtarosptheincospthraagtoeugysihnabMi.t,elbeugtanlsa.boOruar-
Size % toryexperimentsconverselyindicateaeuryphagoushabit. In
Order/species [mm] Attacked Consumed thefield,M.eleganscapturedonlyants.Thisisconsistentwith
observations oftwo otherspecies ofthisgenus, M. namibica ;
Araneae Wesolowska 2009 and M. ntfa Peckham & Peckham 1902 t
Thericiionsp. 3.0 0 0 from Namibia, that feed on Camponotusfulvopilosus (Curtis \
Pardusasp. 2.5 100 10 1988). In the laboratory, however, M. eleganscaught awide |
Isopoda varietyofprey. So, thefundamental trophicnicheincludesa j
1
PorcellioscaherLatreille 3.5 0 0 wide assortment ofprey, whereas the realized niche includes
i
Collembola onlyants.
SinellaciirvisetaBrook 1.6 45.5 100 Mexcala elegans recognized and captured prey other than
ants as efficiently, or even more efficiently, than ants. Thus
Isoptera neural and behavioral trade-offs resulting in an inability to
Reticiditennessp. 4.7 100 100 recognizecuesfromotherpreyandtocatchnon-antpreywere
Ensifera not present. Thisis incontrasttostenophagousant-eatersof
Acheladomesticus(Linnaeus) 3.5 0 0 thegenusZodarion, forexample,whichareunabletosubdue
Heteroptera preyotherthanants(Pekar2004;Pekar&Toft2009).YetM.
Lygiispratensis(Linnaeus) 6.0 0 0 elegansusedcompletelydifferentbehaviortocatchantsthan
other prey, so this species has clearly evolved a specialized
Sternorhyncha capturestrategythatseemstobeveryeffectiveandsafeforant
AphisfahaeScopoli 1.7 9.1 0 capture,aswehavenotwitnessedasinglesuccessfulreversed
Auchenorhyncha attackbyananttowardthespidersinlaboratoryexperiments
Eupteryxsp. 3.5 81.8 100 (0%,n = 60,pooledacrosstheacceptancetrialswithants). '
Lepidoptera Mexcala elegans used a ‘bite-and-release’ tactic to catch ‘
Plodiainterpunctella(Hubner) 6.5 81.8 100 ants. This specific tactic is also used by other ant-eating '
Hymenoptera salticids, namely Naphrys pulex (Hentz 1846), Aelurillus
muganiciis Dunin 1984, and Tutelina similis (Banks 1895) *
LFaosrinuuscanipgrearte(nLsiinsnaReeutsz)ius 63..35 110000 110000 (Wing 1983;Lietal. 1996;Huseynovetal.2005).Thisspecial -
Messornmtieus(Nylander) 6.0 91.7 100 tactic includes a short leap with a quick bite, followed by
Tetramoriiimcaespiliini release and retreat. Interestingly, a similar tactic is used by
(Linnaeus) 3.5 91.7 100 other non-salticid, ant-eating spiders, such as gnaphosids,
zodariids,andthomisids(Heller 1976;Lubin 1983;Oliveira&
Coleoptera
PThriylblaoorltvirauenliacsaps.tainmeaigimo(Herbst) 83..30 500 00 a(SletEagtdzsawic(amkJaradecsi1kt9seh8toe5nra;l.&hPe1ea9Vkd7aa4-rn)o,nO2;o0lr0pi4f.h)er.e.,onmIbn1itt9he9eas2ll;rebacJeraat;scweikse.s,ee.o,nntohhneetetasahpdle.idaae1bn9rd9sd8o)um,tsehubonaorltaolhxry .t
Diptera tacticsmakingitimpossiblefortheanttodefenditself.
Drosophilamelanogaster As the most frequent natural prey of M. elegans were
Meigen 2.0 100 100 Camponotus ants (subfamily Formicinae), we expected that
—
Figure 3. PredatorybehaviorofM.eleganswhencapturingants.A.Spiderstalksattackedantwithraisedforelegs.B.Spidergrabsantennae 1
ofantinchelicerae.
PEKAR&HADDAD—TROPHICSTRATEGYOFMEXCALA ELEGANS 137
—
Figure 4. Comparisonoftheattacklatency(A)andparalysis latency(B)forfourant {Eormica, Lcishts, Xlessor, Telnimorium)andone
termitespecies. Barsindicatemeans,whiskersindicate95%confidenceintervalsofeachmean.
related ants {Formica and Lasins) would be attacked and may provide M. eleganswith higher protection from enemies.
paralyzedmorequicklythanothers.Thespidersattackedfour Thusitappearstofavortheenemy-freespacehypothesis.
ant species used in the acceptance trials at significantly We conclude that the evidence gained on the trophic
different latencies. Slow-moving species {Messor and Lasius) strategy of M. elegans suggests that it is a euryphagous
wereattackedmorerapidlythanfast-movingFormica. Larger specialist, because it has the versatility to catch a variety of
ant species had longer paralysis latencies than small ant prey but uses a specialized prey capture tactic on ants.
species, regardless oftheir taxonomic relatedness, suggesting Observedstenophagyinthefieldhaspresumablyresultedasa
that the venom of M. elegans is not specific for certain byproduct of adaptive dynamics related to risk aversion
subfamilies of ants, as was found in ant-eating Zodarion (avoidingofenemies).
(Pekaretal. 2008). ACKNOWLEDGMENTS
Inthefield,Mexcalaelegansfrequentlycapturesantswitha
greater body length than itself; the largest, Pachycondyla Hamish Robertson (Iziko South African Museum, Cape
tarsata, is double the spider’s body length. Similarly, in Town)isthankedforassistancewiththeidentificationofsome
laboratoryexperiments,thespiderscapturedpreyuptotwice ants; two reviewers and L. Higgins are thanked for useful
their own length, consistent with observations of other comments. This study was supported by Project
mthyeArmbmsseeelcnvocepesha(eog.fgo.u,nseSuosrypaeilrder1as9n4d3t;habPteehkcaaavtricoh2r0a0pl4r)e.tyramdeu-cohffslardgoeersthnaont ntSoph.oarnt0ks0e2do1f6f2otr2h4ep1e6CrmzoieftcshthteRoepMuuinbndlieisrctt.raykEezoeffmivEedledulwcooartkKioZnat,NtYhWoeiuldtNlhdifueamnoids
preclude the presence of physiological trade-offs. We have Game Reserve and to export material for laboratory
not studied the effect ofprey type on fitness aspects such as experiments (permit numbers 1924/2004, 54/2005, 1010/2006,
survival or reproduction. Thus we cannot exclude the 2496/2006,4198/2008and2612/2009).
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IsraelJournalofZoology45:341-359. Manuscriptreceived9September2010, revised2March2011.