Table Of ContentOdonatologica37(3):193-202 SeptemberI,2008
Ectoparasitic water mitelarvae
of the genusArrenurus
on thedamselfly Coenagrionpuella(Linnaeus)
(Zygoptera: Coenagrionidae)
R.A.Bake¹r,P.J.Mill¹ and A.Zawal2
1SchoolofBiologicalSciences,UniversityofLeeds,Leeds LS29JT, United Kingdom
[email protected]/[email protected]
2DepartmentofInvertebrate Zoologyand Limnology,University ofSzczecin,
Waska 13,PO-71-415 Szczecin, Poland
[email protected]
ReceivedDecember29, 2007/Reviewed andAcceptedFebruary4,2008
Parasitic larvalmites ofthegenus Arrenurushave been found onC.puella. A.bi-
cuspidator,A. cuspidatorand A.maculatormakeupover80%ofthetotalmitesiden-
tified.Theotherspp. foundwereA.bruzelii,A.clavigerand membersoftheA.affinis
complex.Mitesarefoundmainlybetweenthe secondandthird pairsoflegsand be-
hind thethirdpair. A.bicuspidatorand A.cuspidatorsharethese siteswithnumbers
spreadroughlyequallyonboth sites. A.maculalor isfoundalmostexclusively behind
thethirdpair oflegsand onthe first abdominal segment ofthehost.Smaller num-
bersarefound ontheabdominal segments whereA. clavigeristhedominantspecies.
The larvalmites showapreferencefor $hosts.Sizedifferencesbetween theArrenu-
rus spp. areconsidered.
INTRODUCTION
Thebiologyofmitesonodonateshasbeenstudiedby severalworkersincluding
CASSAGNE-MEJEAN (1966), LANCIANI (1970), MITCHELL(1959, 1967,
1968),ROLFF(2000) andZAWAL(2004a,2004b). STECHMANN(1977)pro-
videddescriptions andakey for theidentification of someof theEuropean lar-
valspecies ofArrenurus.ThesubjecthasalsobeenreviewedbyCORBET(1999),
DAVIDS (1997), SMITH (1988) and SMITH & OLIVER (1976, 1986). Until
recently, however,littleattentionhasbeenpaidto thedistributionandsiteselec-
tionofthe variousspecies of larvalArrenurusontheirzygopteranhosts inEu-
rope.ZAWAL (2006a, 2006b,2006c, 2006d, 2007) recently described a number
194 R.A.Baker, P.J.Mill &A.Zawal
ofspecies fromthis genus,basedontheirlarvalmorphologyandthishasprovid-
edsufficientinformationto identify thespecies accurately. BAKER etal.(2007)
published somepreliminary observationsonhostpreferencesandselection sites
of larvalArrenurus species on several odonatehost species. The present paper
examines oneofthese hosts inmore detail,namely C. puella, anditsassociated
parasitic mites.
MATERIAL ANDMETHODS
Thedamselflies werecollected in 2006 duringthepeak periodof adultactivity in areasaround
Szczecin,innorthwestPoland. Damselflyidentification andthesexofindividuals wereverifiedlater.
Material waspooledfromseveral sitesin the area.Additional information fromearlier collections
(BAKER etat,2007)hasbeen used intheanalysisofchoice ofmaleand femalehosts.Damselflies
withattached miteswerepreservedin70%ethylalcohol. Miteswerecountedand theirhostselection
site noted,then removed and mounted individuallyon slidesin Hoyers medium,each slide repre-
sentingonehostspecimen.Countinglarvaewhile attached totheirhostsgivesonlyaroughnumber
and anaccurate count ofmitespresentcanonlybe obtained byremovingand mountingeachone.
Inevitablysomemites werelostinthetransferfromhost toslide,otherlarvaebecame detachedinthe
hostfixativeand theirselection siteistherefore unknown and othersweredamaged,makingidenti-
fication difficult orimpossible.
RESULTS
THEPARASITESFOUND
596larvaehavebeenidentified,usingZAWAL’s(2006a, 2006b, 2006c, 2006d,
2007) descriptions and allof these havebeen measured.Five Arrenurusspecies
havebeenrecordedfrom C.puella: A. bicuspidator, A.bruzelii, A. claviger, A. cus-
pidator and A. maculator, along withmembersofthe A. affinis/neumani/vietsi-
complex. Threespecies, A. bicuspidator (14.3%), A. cuspidator (35.9%), and A.
maculator(32.5%) makeupover 80%ofthe totalnumberidentified.Percentage
figuresfortheotherspecies are: A.claviger 9.4%,A. affinis-complex 6%,A.bruzelii
1.8%.Inadditiontherearesomelarvaethathavebeenidentifiedto Arrenurussp.
only.
PREVALENCEANDINTENSITY OFINFESTATION
306C.puellawereexaminedforparasites andsexed.Of these, 27.5%(84)were
infectedand 222uninfectedand thetotalnumberofmitesfoundwas 738(Fig.
1).The miteload on infectedindividualsvaried from 1-62, withamean of8.8.
Only 22 oftheinfected hosts hadmiteson theirabdomen.Alsothe numberof
mites from this part of the body, compared with the totalnumberfound, was
comparatively low(14%).
ArrenuruslarvaeonComagrionpuella 195
DISTRIBUTIONOFTHE MOSTCOMMONARRENURUS SPECIESONTHE HOST
Thepreferred areaofattachment, forthemajorityof the Arrenurusparasites
identified, is the posterior half ofthe thorax. A. bicuspidator (n = 86) and A.
cuspidator (n = 214) appear to selectsimilar sites for attachment,occurring al-
mostinvariably betweenlegs two andthreeorbehindthethirdpair oflegs. InA.
cuspidator, 95%occurred
between the second and
third pairoflegs andbe-
hindthe third pair, with
numbers spread approx-
imately equally between
these two sites. A. macu-
lator on the other hand
isfoundalmostexclusive-
ly behindthe third pair
of legs with small num-
bers on thefirstabdomi-
nalsegment. Outofato-
tal of 197 A. maculator
identified, 92% occurred
behind legs three on the
thorax (Tab. I).WhenA.
cuspidator and A. macu-
latoroccur aloneontheir
host,thepatternappears
essentially the same, al-
though numbers are too
low to draw any mean-
ingful conclusions. A.
cuspidator, A. bicuspida-
torand A. maculatorap-
pear therefore to prefer
the posterior half of the
thoraxas theirmainarea
ofattachment. Fig. 1.Numbersofmite (allspecies)onall Coenagrionpuellain-
dividuals
The situationis quite
differentforA. claviger(n =56)where89%were foundontheabdomen, mainly
onsegments3 to6,and sometimesinthe inter-segmental region between these
segments. They occurmostcommonly onsegment4.ThusA clavigerappearsto
beanabdominalparasite intermsofitsselectionsite onthehost(Tab. I).
196 R.A.Baker,P.J. Mill&A. Zawal
Table I
Location offourspecies ofArrenurus onadult Coenagrionpuella
AArrrreennuurruuss TThhoorraaxx TT//AA AAbbddoommeenn TToottaall
11//22 22//33 33 11 22 33 44 55 66 77 88 99
bbiiccuussppiiddaattoorr 44 4499 3300 11 22 8866
ccuussppiiddaattoorr 77 9988 110066 33 221144
mmaaccuullaattoorr 22 118822 1133 119977
ccllaavviiggeerr 44 11 11 I1 11 1100 2244 66 77 11 5566
CHOICEBASEDON SEXOFHOST
Using parasitized materialfromearliercollections(BAKER etal., 2007), not
previously analysed forsexpreference,togetherwiththepresentcollections, itwas
foundthatfemaleC. puella carried a significantly heavieraverage parasite load
thandidmale C.puella (Mann-Whitney testT=2393,n(male)= 54, n(female)
= 58; P<0.()01)(Tab. II, Fig. 2).
X2testsindicatethatthisfemalebias occurs in A. cuspidator (P<0.01;n= 127
mites on 9 C.puella and87miteson S C.puella), A.bicuspidator (P<0.001;n=
75& 10),andA. claviger(P<0.001;n= 55& 1).Therewasaslightly greaterpre-
dominanceofA.maculatoronfemales(101)compared tomales(93)butthiswas
not significant. InclusionofA. cuspidator, A. hicuspidalorand A.maculatordata
fromprevious collections(268)(there were nofurtherindividualsofA.claviger),
bringingthetotalnumberofmitesinthedatasetfrom549to 817, madenoma-
terialdifferenceto thefirst two species. A. cuspidator(P<0.01; n= 186miteson
9 C.puella & 135miteson 6 C.puella) andA. hicuspidalor (P<O.OOI; n=91 &
10)but nowindicatedastrongfemalepreference by A.maculator(P<0.001;n=
230& 109).
SEASONAL DIFFERENCES
Seasonaldifferenceshavealsobeenexaminedby groupingcollectiondatesun-
derfourheadings, endof May, midJune,early Julyandmid July. Although only
limitedinformationisavailable atpresent, itdoesappearas though thenumber
TableII
Location oflarval arrenuridsonmaleand female Coenagrionpuella
CC..ppuuecllllaa TThhoorraaxx TT//AA AAbbddoommeenn TToottaall
11//22 22//33 33 11 22 33 44 55 66 77 88 99
FFeemmaalleess 44 114411 222255 44 88 00 1177 2299 1100 99 11 444488
MMaalleess 99 5577 114444 33 44 00 00 00 00 00 00 221177
TToottaall 1133 119988 336699 77 1122 00 1177 2299 1100 99 11 666655
ArrenuruslarvaeonCoenagrionpuella 197
ofspecies ofArrenurusincreaseas theyearprogresses.InlateMay only A. cus-
pidator and A. maculatorwere found.By mid June fiveof the six species were
present inthecollections, only A. bicuspidator being absent. By early Julyall of
thespecies were recorded.
SIZEDIFFERENCES OFTHEPARASITES
Themedianmitesizes fall intotwo clearly definedgroups(Tab. III).
Thesize dataforthe A. affinis complex andforA. bruzeliiare excluded from
furtheranalysis sincetheformercontainsmorethanone species andthesample
sizeofthelatteris very small.Thesizerange ofeach ofthe remainingfourspe-
ciesis very wideandthereis considerableoverlapbetweenthem(Fig. 3).
The dataare not normally distributedand hence Dunn’s Pairwise Multiple
Comparison Methodwasusedintheanalysis. Itrevealedasignificant difference
in thesize distributions atP<0.05 forall pairwise comparisons except A. cuspi-
datorandA. maculator(Tab. IV).Thesimilaritybetweentheselasttwo species is
clear(Fig. 3)with73%ofA. cuspidatorand85%of A.maculatorbetween200|am
and3()0(.iminlength. Although 61%of A. clavigerarealsowithinthisrange,27%
arewithintherange300-400pm compared withonly 7%and6%ofA.cuspidator
and A. maculatorrespectively. A. bicuspidator standsout with41%inthe range
450-500pm andafurther 15%and 12%inthe400-450pm and 500-550|im ranges
respectively.
Fig. 2.Number ofmites(allspecies)oninfectedCoenagrionpuellafemales andmales
198 R.A.Baker, P.J.Mill &A.Zawal
Table111
Sizes ofthe596 Arrenurus mitesmeasured. Allmeasurementsarein micrometres
AArrrreennuurruuss nn MMeeddiiaann MMeeaann MMiinniimmuumm MMaaxxiimmuumm
mmaaccuullaattoorr 119944 224466 225522..99 118844 550044
ccuussppiiddaattoorr 221144 224499 227744..22 117700 554422
ccllaavviiggeerr 5566 225533 330000..99 222288 553366
AAJfffiinniiss--ccoommpplleexx 3366 441133 441144.,33 222288 664444
bhiiccuussppiiddaattoorr 8855 445566 442255,,33 222266 552288
bbrruuzzeelliiii 1111 551122 448888..55 332244 661144
Interestingly, inclusion of A. bruzelii in the Pairwise Comparison indicates
that itis not significantly differentin size fromA. bicuspidator, but this should
be viewed withcautiongiven the smallsample sizeoftheformer species.
DISCUSSION
Thepresent papersupplements earlierwork bythe presentauthors(BAKER
etal., 2007) usingadditionalinformationand seeksto analyse aspects ofthebi-
ology ofthemites andtheirrelationship with C. puella, giving particularatten-
tionto theidentificationanddistributionofthespecies ofArrenurusonits host,
choicesbasedonthesex ofthehost andsize differencesoftheseparasitic larval
mites.
Fig. 3. Frequencysizedistribution offourspecies ofmites foundonCoenagrionpuella
ArrenuruslarvaeonCoenagrionpuella 199
Itis clearfromthe present Table IV
study that, in the case of C. Pairwise MultipleComparisons(Dunn’sMethod)betweenthe
puella, themajorityofmites sizes of four species ofarrenurid mites. SignificanceatP=
<0.05. Themultiplecomparisonsonranksdonotinclude an
occurontheposteriorhalfof adjustmentforties
the thorax, betweenthe sec-
ond and third pair and be- AArrrreennuurruuss ccuussppiiddaattoorr bbiiccuussppiiddaattoorr mmaaccuullaattoorr ccllaavviiggeerr
hind the third pair of their
host.They arefoundinsmall- ccuussppiiddaattoorr YYeess NNoo YYeess
bbiiccuussppiiddaattoorr -- YYeess YYeess
er numbersontheabdomen. mmaaccuullaattoorr -- YYeess
Occasionally, butinverylow ccllaavviiggeerr -
numbers, mitesarealsofound
atothersitessuch as theheadandbetweenthefirstand secondpair oflegs. The
meanintensityoftheparasitesininfectedC.puella was8.8, agreeingcloselywith
afigureof8 parasitic larvaeperindividual,in work by GRANT & SAMWAYS
(2007).
The Arrenurus species appear to be generalist parasites, each species being
foundonanumberofgeneraofhostdamselflies(DAVIDS, 1997;BAKERetal.,
2007). DAVIDS(1997), forexample, recordedthe larvaeof A.bruzelii oneight
differentzygopteranhosts,andrecordedninedifferentspecies ofArrenurusfrom
C. puella. However, DAVIDS(1997) reviewrelies, to an extent,on olderrecords
whichinsome cases maynot haveprovided adequate descriptions of the larvae
involved.Zawal’s recent descriptions ofthelarvalspecies give moredetailedac-
counts. Inanearlierpaper,BAKER etal.(2007)recorded fourspecies ofArren-
uruson C.puellabutspeculated that,with furthercollections, morespecies were
likely tobefound.ThepresentstudyaddsA.clavigerandtheA.affinis/neumani/
vietsi-complex to thatlist.Themost commonspecies foundare A. bicuspidator,
A. cuspidator and A. maculator, which together make upover 80%of thetotal
ofidentifiedmites.
Although arrenuridmitesdo not havesingle species hosts, theirchoiceof at-
tachmentsiteis clearly definedin someofthespecies studied. Inthesecases re-
source partitioning may occur. Thus A. claviger is foundmainly on the middle
segmentsofthe abdomen, A. maculatorselectsthe regionbehindthe thirdpair
oflegs andA cuspidator is foundinroughly equal numbersbetweenthesecond
andthirdpair oflegs andbehind thethirdpair. Sitespecificity has beenrecord-
ed forotherspecies of Arrenurus. MITCHELL(1968) described two species of
Arrenurus larvae from North Americaattached to Paracercion hieroglyphicum
(formerly in Cercion), A. mitoensis onthe thoraxand A. agrionicola onthe ab-
domen.In MITCHELL’swork(1968) the incidenceofthethoracic parasitewas
low butthe abdominalparasite was abundant, with98%ofthe hosts infected.
Theseresultsare incontrast to thepresentfindings. BOTMAN etal. (2002) also
foundspecies specific selectionsitesintwo species ofArrenurusonthe damselfly
Ischnuraposita. A. majorwas attached to the thoraxand abdominalsegments
200 R.A.Baker,P.J.Mill& A.Zawal
oneto three,whereasA.americanuswas attachedtoabdominalsegmentsfiveto
eight. ROLFF(2000) working with C.puella andC. hastulatum, andANDRES
& CORDERO (1998) working onCeriagrion tenellumdemonstratedaclumped
distributionofmiteson theirhostsbut notin allindividuals.It seemsclearthat
eachspeciesofmitehasaclearlydefinedattachmentsiteasSMITH(1988)records.
Although initialcontact couldbemadeatanalternativesite, themitestakeupa
characteristicfinallocationontheirhost, depending on thespecies, anddo not
behaveinarandom manner.
The Arrenurusspecies describedhere showedpreferences intheirattachment
sites. Choiceof differentsites mayreduce interspecific competition forfeeding
and growing space. MITCHELL(1968) foundthat the response to miteover-
crowdingwas to moveto new sites. Inthepresent study, A. maculator, although
typically occurring at the posterior end of the thorax, is also foundon the first
abdominalsegment,suggesting thatovercrowding on itsfavouredsite maylead
to occupation ofan adjacent area.A. claviger, althoughinvariably foundonthe
abdomen, is occasionally foundinsmallnumbersfurtherforwardonthethorax.
SMITH(1988) statedthatectoparasitic mitesassociate withhosts already car-
ryingmites. Ithadbeenassumedthatclumpingofthesamespecies atattachment
sitesis theresultofthesynchronization oflarvalactivity andshouldresultinlar-
vaeof similarsize being found.However, in thepresent study, there are consid-
erablesize differencesbetween individualmites ofthesame species, whichcould
reflect theperiod oftimethey havebeenattachedto thehost,ifindeedthey ar-
rive atdifferenttimes.Itisthought thatthelarger specimens werefeedingorhad
fedandwereaboutto detachandthesmallerspecimens, recently arrivedattheir
sites, were producing stylostomes in orderto startfeeding.
ZAWAL,inhismorphological descriptions (2006a,2006b,2006c,2006d,2007),
found unengorged larvae withthe following lengths: A. maculator200-210, A.
cuspidator 199-204,A. bruzelii212-228,A.affiniscomplex 224-244, A.bicuspida-
tor228-252and A.claviger 234-260pm.Thelargest are thereforeA. claviger. A.
hicuspidatorandtheA.affinis-complex. Inthepresentstudy, whichincludedboth
unengorged andengorged larvae andhenceproduced muchwidersizeranges.A.
maculator A.cuspidatorand A. claviger had lowermedianandmeansizes than
,
thoseintheA.affinis-complex, A. bicuspidatorand A. bruzelii.Of note isthatA.
claviger was the largest of the unengorged mitesrecorded by ZAWAL (2006d)
butdid notshow thelargest engorged individualsinthe presentstudy.
Itisknownthat mostlarvalmitesremain ontheirodonatehost forup to 20
days(CORBET, 1999)butsomedonotreachmaturityanddieon thehost,most
likely dueto overcrowding and/or theinability to feed andthis may inpart ex-
plain thedifferenceswefound.
CORBET(1999) inhisreviewstates that,“mitesinfestmaleand femaleOdo-
nata apparently withoutbias”.Thisiscertainly supported byanumberofstud-
ies, forexample, ROLFF(2000)andMcKEE etal.(2003) However,incontrast.
ArrenuruslarvaeonCoenagrionpuella 201
ROBB&FORBES (2006),working withLestesdisjunctus, foundfemalesexbias
inparasitism by larval A.pollictus; LAJEUNESSEetal.(2004) foundsex biases
in parasitism by themiteLimnocharesamericanainfield surveys andFORBES
etal.(2004), intheirfindingsinvolving recapturetechniques, reported asituation
inA.planus inwhichmaturefemalescarriedmoremitesthanmaturemales.Ma-
ture malesalso hadfewermitesthannewly emergedmalesnotrecaptured. Inthe
present studyon C.puella, theauthorsfoundsignificant differencesinnumbers,
withmitesapparently preferingfemales to males.Since thefemaledamselflyre-
turns to water tooviposit, malesflying mainly over vegetationadjacent to water,
it wouldbeof someadvantage forthe mite to showafemale-biasedparasitism
inordermoreeasily to return to water to continuetheaquatic partoftheirlife
cycle.
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