Water Depth‐Dependent Notonectid Predatory Impacts Across Larval Mosquito Ontogeny

Water depth‐dependent notonectid predatory impacts across larval mosquito ontogeny

Dalal, A., Cuthbert, R. N., Dick, J. T. A., & Gupta, S. (2019). Water depth‐dependent notonectid predatory impacts across larval mosquito ontogeny. Pest Management Science. https://doi.org/10.1002/ps.5368

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Download date:01. Oct. 2021 This articleisprotected bycopyright.Allrightsreserved. to handle variations depth inwater differences inresponses emergent.to were Bothnotonectidswere able differedRESULTS:Consumption rates significantly between the predators, andinterspecific quinquefasciatus predators, We enemies quantify vectorspecies. predatory thus the towards impact oftwo notonectid effects, sizeDifferences influencing inprey drivethecould size-refuge efficacyprey ofnatural through predator water towardsinfluence efficiencies 3D-space the column. preywhichutilise often remain differences unquantified. Inparticular, inwater depth within systems aquatic could canmodulate BACKGROUND: strengthContext-dependencies the ofpredatory interactions and ct Abstra Accepted Article BiologyMedical United Centre, 97Lisburn Road, Belfast BT97BL, Ireland, Kingdom Northern 1 2 *Correspondence to *Correspondence Department of Ecology and Environmental Science, Assam University, Silchar- 788011,Department India ofEcology andEnvironmental AssamUniversity, Science, Silchar- Institute for GlobalFoodSecurity, School ofBiological Sciences,Queen’sUniversity Belfast, Water depth-dependent notonectid predatory impacts acrossmosquito larval ontogeny impacts notonectidpredatory depth-dependent Water

article as doi: 10.1002/ differencesbetween version lead to this may been through the copyediting, typesetting, pagination andproofreading process, which This article hasbeen accepted for publication andundergone full peer review but has not

C. quinquefasciatus Anisops breddini Arpita Dalal*Arpita A short running title: Notonectid running predationA short title: against mosquito larvae prey across prey awater depthgradient, functional responses using (FRs). : University, University, India,EmailSilchar, Arpita Dalal, Department ofEcology andEnvironmental Assam Science, 1 , RossN.Cuthbert and ps.5368 prey across prey allinstaryet stages, ratespredation generally were Anisops sardeus 2 , JaimieDick T.A.

, towards four different larval instars of fourdifferent Culex , towards larval

and the Versionof Record. Please cite this : [email protected] 2 andSusmita Gupta

This articleisprotected bycopyright.Allrightsreserved. down Accepted ecosyaquatic contextand abiotic dependencies can strengths interaction alter trophicgroups in between communities andodonatelarvae ephemeral aquatic ecosystems systems 1 Keywords: medically mosquitoes. important Article volumes,habitats ofvaried promotion their inaquatic systems help could CONCLUSION: dependencies whichmodulate impact.their predators towardsimpacts ofnotonectid feeding rates as reduced increased.further depth Ourresults demonstrate substantial predatory andhandlingBoth capturerates times often were at greater greaterandthusmaximum depths, treatmentemergent specific withpotential in groups, implications for were displayed bynotonectids inthe majority FRswere oftreatments; however, TypeIII differential ratespredation specieswere ofthis shallow in waters.most pronounced FRs Type II higher towards early as opposedlate to instarprey. INTRODUCTION

process Predator Predator communit , 1,

which f unctional response, unctional stems, understandings andpredictive contexts ofhowinfluence such via es

such as such as are predation

trait Given notonectids are Given notonectids ,

in turn larvae are -

and density ,

ies influences

often play acrucial role

especially

Culex quiquefasciatus - mediated withtheirprey interactions

top predators

structuring and functioning l acking

capable dispersal betweenof aerial ephemeral aquatic

mosquito, quantify and biotic and context abiotic ,

predaceous insects suchas

. In particular, abiological within context, control in , 5

and the Anisops sardeusAnisops

regulati have the potential have thepotential to structure , biological control, depth,water prey on

at at the of

prey populations prey in was most voracious, and hemipterans,

prey population stability.

ecosystem . 6,

reduce proliferations of proliferations of reduce 7

However, bothbiotic

level coleopteran coleopteran crucial . natural 3,

top 4 In

size -

This articleisprotected bycopyright.Allrightsreserved. examinations beenand have congruent showntobe with Acceptedefficiency) resource thus should be integ mosquito lifelarval stage. history strategies (e.g. B such as including wetlands, worldwide distribution, Articlelymphatic filariasis filariasis, Culicidae) forpotential encounters with humans rangevast of andartificial aquatic natural systems 700,000 people feveryellow including environmental iological control control iological

Mosquitoes Mosquitoes are Functional aquatic insects,arachnids, crustaceans, fish,andbeen have birds bats identified (e.g. predator

West Nile Sandbisvirus,StLouis virus, encephalitis, Rift Valley fevervirus and prey size, towards

, andmaximumhandling time feeding rate is amember of , . especially 17, 18 17, contexts 19 dying ) ,

medically important and particularly using predators r 2

esponses 2 (see rated considerations into could exert could exert a significantontheefficacy agents,influence and ofbiocontrol rice fields

Indeed, comparative FRs canbe integrated

from - prey) prey) vectors of deadlyhuman diseases

could have and However, in tropics, the 10, 11, 12 11, 10, Culex pipiens vector are are

(FRs) interaction strengths and ) known tocolonise . Mosquitoes in the

context has effective been borne diseases artificial during are centralare important . 9

with In particular vector

complex - effort

dependencies,

ecological impacts in

more infected onebillionpeople and than container

among consequences for target

species. to s

the derivationthe of which target mosquitoeswhich target their during aquatic . annually and is acompetent and is , 20,

a broad rangea broad of includ , C. pipiens

21 natural candidate enemies. Culex quinquefasciatus in mosquito p s.

parameters

13,14,15 The

such as such as ing . 8 capture

Mosquitoes Mosquitoes

To combat these in urban areasin urban

complex exhibit a to examine relative efficacies

biological control changes in water depthchanges inwater malaria, dengue, are are density opulationmanagement rate aquatic environmentaquatic central vector ofbrancroftian

prey

(i.e. successfully - dependen population

where there high isa

attack components ofFRs Say, 1823 (Diptera:

vectors

n abundant and

rate agent t ika, and ika,

consumer , colonise a colonise over

regulation, regulation, , search , search predators predators . 16, 17, 18 17, 16, s,

This articleisprotected bycopyright.Allrightsreserved. impact Acceptedemploys comparative naturetransient ofsmall where breed. habitats mosquitoes aquatic interactio mosquito prey, influences the strength of (e.g. Although canvarysubstantiallyaccordingto efficiencies establish both natural and artificial ephemeral aquatic structur the Article consumption wh proportional rates impart topreypopulation stability stage adult that iscapableof disease. vectoring context the mosquitoes,of FRsmay TypeII the recruitment reduce through tothe ofmosquitoes consumption lowdensities at due a predatory context,biocontrol identificationthe exhibit ofagentswhich FRsdesirable Type II is contexts. biologicalcontrolbetween in agents andacrossvector control, multiple environmental to their to their populationpotentially 3 1 , 3

of Notonectid 2 . 17, 1817, , 3 28 ns potentialoften failto depthaddress water predatory efficacy of two widespread and abundant widespreadandabundant two 3 , ing of ), 29

Three Three it isunclearit howvariabilities insearch ar Whilst t

which aquatic aquatic s

(water boatm(water form ofFRs (shapes) types

FR heir diet forage through s

communities offtake rates by offtake rates

(e.g. (e.g. that reduces the

is known to notonectid 17, 18 17, e s n) dest ere preyarescarce

by refugia providing , 2 out are voracious predators voracious are 7 ,

abilising effects

(e.g. (e.g. 3 4

notonectid predators, ) waterthe column

such s to

include all include all size classes ofmosquito larvae towards mosquito been has larvae

2 ecosystems decipher theeffect 6

, 2 potential fo

consumers. isparticularly This important for In contrast 7 the have described: been Typeand I,II III. ).

In the In the tropics, t larval ontogenicstage larval .

20 variation , attributable ea within ecosystems 3Daquatic

to where mosquitoes r prey refuge r preyrefuge

. to FR Type II

low prey densities Indeed, examinations of Indeed,

and Anisops breddiniAnisops s s , despite the, despite dynamic and

of are known to his group dominant isoften Therefore,

to water depthonthe high rates ofpreyhigh rates and s

, Type III FR , TypeIII of mosquitoof prey thus successfully , the presentthe study well

owing to low play akeyroleplay

Kirkaldy, 1901 persistence - described biotic ,

s 20, 2 20, predator

may 3 . , 2 . 3 2 0 5 4

in In In in

This articleisprotected bycopyright.Allrightsreserved. 2.2 traditi classonof size adiet laboratory mixture campus bysampling everychanged 10days to were acclimated these conditions waslaboratory maintained at separately Ecology andEnvironmental Science, Assam SilcharUniversity, hand during 31 ( 2.1 2 Notonectidae) (Heteroptera: Notonectidae)and

Accepted Article7.21 ±0.06mm MATERIALS ANDMETHODS 00 Functional responseFunctional experiment Collection of .

sq. onal non These n the The prey, The prey, The focalpredators, size

to attract oviposition.to attract m,~ 62cm depth

in aquaria aglass 20 as notonectids, the 17 su otonectids otonectids were transported - towards fourtowards centrifugal organic cane lump palm sugar made dateor from tree ) were collected from collected ) were larvae of predators andprey predators mmer egg rafts fromc egg rafts artificial crushed

and - monsoon season ) C. quinquefasciatus fed of Irongmara larval larval holding

30 °C(± 2°C)

where

jaggery The mosquito egg The mosquito egg werethenrafts transferredsame tothe ad - matched Anisops sardeus instar libitum , for atleast

25 Lde upon hatching, mosquitoupon hatching, larvae

(

fishery ponds fishery Arya Farm ProductsArya Farm Pvt. Ltd

stages ,

a in source water in source with by trawlingakick Cachar, Cachar, dult dult

- on a 12:12light:darkphotoperiod chlorinated watertap ontainer ontainer

, of m one A. breddini

were collected from

osquito osquito Herrich Culex quiquefasciatus Assam, India

week ( p that werethat ond 1: and chironomid larvae.

- prior to theprior to experiment to a Schaeffer, 1849

( 7.04 ±0.06mm

net through thecolumn water

3800 sq. laboratory laboratory

. Bothspecies ( filled 24°41'15.78"N, 92°45'12.21"E)

with continuousaeration ) the Assam University AssamUniversity the

were ad libitum with m, ~70cm depth

prey in the (Heteroptera: reared to the desired

jagger ) . and , andnotonectids

were

D . .

Jaggery isa Jaggery

epartment of with water A. sardeus y

and kept

; pond 2: ; pond2: water

by . The

This articleisprotected bycopyright.Allrightsreserved. densityprey wasused Acceptedhoc Tukeytest FR derivations. overdisper lineargeneralised model (GLM)assuming a to‘waterrespect depth’, species’ ‘predator instarstage prey experimental treatment group eliminate time confounds removed tointroduced Article25, 50,75,100 mm 0.05 stagesfour different instar of cm chlorinated tap water to conducted in density’,respectively,‘prey onnotonectid experimental todiscern design effectsthe of‘water depth’, ‘predator ‘prey species’, instar’ and experimentation ( D3, I To discern FRTo discern types analyseswereStatistical performedin Rv3.3.2. ndividual

~ 4.45 L) ~ 4.45

and number ; 3 s ion rd glass aq glass

individual individual instar, s . ;

‘Prey density’ was density’ ‘Prey included as an effect, individual importance owingtoits in F were conducted for pairwise comparisons of significant effects n

to standardize hunger and -

s . Bothpredatorspecies( = 3per experimental group tests with Type III sums of squares were tests TypeIIIsumswere with employed squares main toderive of effects.

of predator each

3.99 s prey uari

for of

three different arena a , remaining live remaining



across density density (

batches batches wereoftrials experimental with a

, 0.03 mm; 4

l s C. quinquefasciatus ogistic ofogistic regression oftheproportion killedasafunctionprey of s . Controls predators each consistedofprey without at

and each experimental group standard n were species = 3 per experiment= 3per

levels depths: th

prey instar, 4.9 allowed to - A. breddini

surface of area mosquito interaction strengths. Feedingt and were starved quasi . ) enumerated to numbersquantify We employed factorial‘3 a . Following the additionofprey,notonectids

5 cm (

‘prey instar’, interactionand their

(i.e. 1 (i.e. - Poisson familyPoisson  feed D1,

and 0.05 mm 3 al group).al st 5

in isolation for 24h instar, 1.1 Raw consumption was analysed withRaw wasanalysed consumption ~ .

18 cm 2 A. sardeus 4 1.44 L) 1.44 , 37 35, fully

) under s ) under 2

. Aquaria were filled withde . Aquaria , Here, a Here, a significantly negative , owing to residual to residual , owing

a , 10cm randomised fter 

) were for 24h

0.02mm which ix × ( prey prey

D2, 2 separately prior to prior to

× the predators were predators the killed ; 2 .

3 according to according to 4 ~ 2.95 L) ~ 2.95 densities (5,10, densities 6 s

nd ×

using instar, 2.21 instar, 6’ rials were rials were . To water

provided provided

and 15 depth were Post -  ,

This articleisprotected bycopyright.Allrightsreserved. P sardeus Anisops Acceptedquinque mortality 3 differenceswhere significant produce betterindicate fit) Information andfit models Criterion whichminimised information loss model orIII) orcontinuousType II flexible)forms. (i.e. maximumWe used likel Article FR) orFR), TypeIII 1(a insteadoptimized, ameans providing toexamine fitsofcategorical (i.e. is the initial prey density where experimentalallocated time: FR negativeby asignificantly first

RESULTS < 0.001; Fig.1) < 0.001;

was

order term aTypeII indicates b fitting - Survival of Survival was prey

fasciatus dependent capture ( rate implemented is the attack or is theattack capture rate, which,combinedwith the exponent scaling 95% confidence(CIs) intervals around FRs foreachpredator

was thus .

4 0

Where equivocal, FRtypeswere wecomparedmodels candidate using Ak

consumed moredid than prey significantly density larvae . .

attributed topredation W T here were

e that that

then was

and accounts for second second

non

not 100

37, 38, 39 38, 37, N were then

e significant consumptive between differences prey instar stagesby -

significantly affected parametrically bootstrapped is the number eaten. ofprey % in the% in control

ord FR

), er termer FR aType indicates III

, whilst a significantly asignificantly , whilst positive

h non

deduced onthebasis of CIoverlaps

is thehandling time, by notonectids.

- replacement ofprey

(

groups across

by water depth (

Overall

A. breddini

starting starting T Here,

is the totalexperimental time,is the

) all all (i.e. preydepletion)(i.e. ) , )

consumption of

treatments q .

first FR fits can befixed 0 at TypeII (a

A treatment combination F

(AIC; lower (AIC; lower values

overall ( overall 2, generalised formgeneralised ofthe 403 ihood estimation ihood estimation order termorder followed .

( = 1.391 , n

and experimental

= 2000) to F q , gives the 1, 1, C. C. 403 over the ,

= 16.516,

= 0.25 (1) a ike’s ike’s for

, N 0 ) 0 .

This articleisprotected bycopyright.Allrightsreserved. depth towards greatest the at depth (Table 1 towards significantly negative were prey ( breddini to intermediate andhigh depths by compared lowor either depths to high 1 experimental treatments terminteraction depth 2 low andintermediate depthsascompared to the depthgreatest (both were most pronounced variations in depth water ( differentialconsumptionHowever, instar the of across wassignificantly prey stages by altered (all were consumed instars (all earlier than notonectids

AcceptedP Article st nd

instar prey < 0.05 A. breddini – P

< 0.001). turn, In level th Type II

1 instars was, however,significantly greater ; Fig. 1b st supplied

and (all ( F

was

3, 3, 1

was FRs were P prey 3 st 403 further indicated emergent indicated emergent further complexities infeeding patterns rd

instar preyby > 0.05). ).

most depth affectedbywater variations instar = 138.336 significantly overall Furthermore, s , there were no first

for consumption of 1 3 ( rd prey prey F

most prevalent acrossmost prevalent F However,

order terms (Table 1).order terms(Table 6, 6, instar prey wereconsumed than less ( 3, 3, F 403 ; 403 , 5, 5, at at shallowestthe depthby Fig. 2 P 403

= 4.802,

A. sardeus (both P<0.05

= 3.044, great <

= 317.798 ignificantly more prey were moreprey co ignificantly 0.001 overall ). Type III FRs only). TypeIII were evidenced A er

(both (both significant ‘water depth×predatorprey significant ‘water species instar’ P

at ; Fig.1 P

< 0.001), whilst P

the lowestthe depth towards

< 0.001). Here, < 0.001).

significant consumptive differences (Table 1 = 0.006 st , P at

instar prey,withsignificantly more prey ; Fig.1a

< 0.01; the

). < 0.001). < 0.001). experimental experimental

However, Type III FRs However, TypeIII were Significantly fewerSignificantly intermediate oppose as to ; Fig.1 ; Fig. Fig. ). Consumption of 2 A. breddini 2

, and highest ). Here, effects ofwater the depth consumption by ). 1 nd

For ). instar prey were consumedmost treatment

On theotherhand,consumption 1 A. breddini st nsumed as higher densitiesnsumedas higher

instars

, and towards 1 P st 4

th < 0.001).

instar prey at groups

instar mosquito prey at

intermediate depths overall overall nd

A. sardeus the the intermediate , capturewere rates between between

the the instar preyby , as indicated by evidenced across each low water depthlow water Conversely, for Conversely, ( , ascompared 2 P nd

= 0.010).

instar prey

towards towards killed killed

A. water of at as

This articleisprotected bycopyright.Allrightsreserved. be to tended atlow reduced (Tab depths instar tended handling attimes to be lowest intermediate depths water 2) (Table 3 quinquefasciatus be to tended than higher significantly than higher all towards preyinstars other ( atthelowascomparedreduced to and significantly reduced variations in depth water ( However, towards capture intermediate depths water similarrelatively among instarstages 3a (Fig. groups were instar prey more similar ( compareddepths to intermediate groups andhigh ( capture rates with increased depths, greater water significantlywith rates reducedat low water treatment ascompared lowandintermediate groups( to t significantly towards lower

Accepted Article owards rd

instar prey, owingtooverlapping of CIs(Fig. low handling times were

Overall, for Overall, For rates were significantly 2 depths (Table 2; depths nd

A. sardeus instars, instars,

prey 2 A. breddini A. breddini nd

in thedeepest treatment group,yet

, whilst theywere reduced toward 2 , capture rates were, capturerates significantlylower towards instar prey,capturerates were A breddini

Fig. , ascompared lowand to highdepths ( Table 2;Table also also 1 st 3g).

instar prey at depthsinstar preyat low water ( , were similaracrosshandling times water depths for 1

not significantly capture rates were significantly greater

Capture ratestowardsCapture 4 intermediate andhighdepths(Table2; Fig. 3h) Fig.

overall 1 towards Table 2;Table at at highthe compared lowdepth to 3f l e 2). Overall,e 2). 4 ).

Capture ratesof3 Capture – d). Fig. 3d) different accordingtowater depth 4a

Table 2;Table – c). generally lowand . Fig. 3e

th Table Table 2; However, for groups, theseprey Overall, were more

instar handling prey times nd th

F instar groups instar preyweresignificantly ig. 3c rd st –

, 3

Tab1e 2; Tab1e 2; reduced reduced instar prey by h Table 2;Table Fig. 3b). ). A. breddini rd

). Capture

Capture ratesof

and 4 similar between similar between intermediate 1 st relatively unaffectedrelatively by at

Fig 3a). th

instar prey at instar preyat

Fig. 3e). .

(Fig. 3). For 3

here

the high the For instar

capture rates were rates towards 4 A. sardeus

A ( rd Table 2; .

Conversely, C. instar prey, instar prey, breddini

, and were In turn,

( depth A. sardeus

Fig. 4d), were were st , 2

Fig. 3e). Fig. 3e). were nd ,

4 and th th

yet yet

This articleisprotected bycopyright.Allrightsreserved. destabilisingpotentially present study, agentscontrol 4 factorspotentially and abiotic influencingthe impact impact pro 4 4). times of magnitudesresponse were thus prey were reducedgenerally at the depthhighest treatment 2). (Table times by at were significantly shorter low compared intermediate to depths. wereinstar prey, similar handling times towards 1 depths magnitudes significantly than longer

Accepted3 Article

DISCUSSION In thisregard, FRs can act liferation andemergence of handling times were not s For Biological control ofmosquitoBiological

of predator C. quinquefasciatusC. A. sardeus significantly

A. sardeus were significantly were significantly reduced A. sardeus in controlling st

instar prey, as comparedinstar prey,as the to high s

were significantly different not depths across water (Fig.

on prey , handling times different

Type II Type II most

was identifiedas significantly greater thansignificantly greater other preytypes all (Fig. 4e

larval diseaselarval as aframework have been wereprey generally similar between the two predators

other prey groups preygroups other (Table Fig. 2; 4a

medically important across depth treatmentsacross depth significantly FRs between most were prevalent es were significantly shorter at at were significantly shorter the andintermediate low for this preyfor this shown tobe at intermediate andhigh

using natural enemies

vectors a more voracious reduced for4 in deciphering thein deciphering predatory of potential biological

stage mosquito highly

aquatic ecosystems of consumers within ecosystems

depth (Table 2;depth (Table Fig.

( Table 2; Table

(Fig. 2 th

context instar prey(Fig.2e andling times predator than species can be effective can be to mitigate the a depths Fig. 4h) – –

d). For 3 d - the predators the dependent ).

Accordingly, (e.g.

rd (Table Fig. 2; . ,

17, 18 17, instar prey, handling however A. breddini for 4 19, 4 19, 4e). 4e). 4g), however were , , 1

3 – – ). with bothbiotic 2 th Towards 2 , present the h). h). , 34

However, the instar prey instar prey , 43

Handling Handling Functional FR 4 overall th

. During the During the .

4f)

2 instar Whilst 7 , 3 ,

2, 2, nd but (Fig. 3

4 , 4

2 , This articleisprotected bycopyright.Allrightsreserved. low densiti mosquito preyat refugia. FRsType III Accepted traitsselectivity betweenmultiple coexisting mosquitosizeclasses. population reducing manyIndeed, predators unabletohandleresourcesare relatively bigorsmall which present study competition inalleviation, producing larger turn adults and betterthat are vectors. isbeneficial, givenontogeny thatsize effects refuge in large facilitatemosquito preycould of selection control habiological to agents whichare able mosquito larvae a indicating Article notonectids typically instar prey were most consumed overall. with handling timesandthus maximum longer rates feeding lower forthesegroups. Moreover, earlier instar tendedprey impactedto be more latethan greatly mosquitoes, instar earlymosquitomarked instars towards preypredator speciesand sizes, effects found also study emergent evidenceprey for TypeIII FRs with stabilizing congruent population . exhibits

Within settin laboratory 2 Although interactivecomplexities were present depths,in ourresults betweenwater 5 , conform to 4 that 7

displayed Therefore, Therefore, predation by whilst could could identifies

population higher re particularlyre high, androbust to depth variations in water prey and size have - level offtake rates Type II FRsType II

capture rate Type II Type II

notonectids notonectids as stabilis - destabilis es, and Type III gs, p A. breddini ing consumptive

(e.g. s

towards instar prey late ing predatory impact effects redator effective , wherein , wherein consumedpredators indeeper less 3 . 4 2 5 , 3

, 4 3, 3,

- on prey that areon preythat A. sardeus FRs

on hand,the other 4 prey interactions However, further researchisrequired to quantify 3 ). differences between were depths particularly

In thepresent study,although

, mosquito predators, and mosquito predators, particularly

A. sardeus

has the potential forhas the potential impact high s ndle mosquitoes larval throughout their

from speciesof uponlowdensities this ,

associated

mostly showed Type II FRs TypeII mostly showed and so with may allowmosquito better

invertebrate

minimiz In contrast toType FRs, II

with low es , potentially , potentially both

refuge effects consumers - density 4 4 of

waters

Thus, the Second A. sardeus the the on , prey prey

focal larvae . . T .

he 2 5 , 3

This articleisprotected bycopyright.Allrightsreserved. insects aquatic generally inshallowwateroften reduced Acceptedcapture rates were longest profound in in the ofintermediateprey which maximumgreater feeding rates rate Articlefor mosquito larvae bythisspecies atlow densities. prey capture rate present study mosquito prey strategies predator where the alternativeencounters prey and exhibits frequency sizes to persist higher

2 most prey profitable for predators. s nd . to prey whil sizes, generally

However, empirically, Type III FRscould displayed followed by

In many cases in

water depthwater via increased (see the s of these

. were also mostly elevatedwere also indeeper whilst,conversely, waters, l

ow densityincontainer refugia lowest water depth lowest , whichmayminimize effects such 49 , suchas 48 decreased byreductions A. sardeus , 5

). similar consumptive traits

0 water depth Although , 5

with prey size. increasing 1 rd relative relative , 5 s , capturerates of

, instar 2

st destabilizing predatory

odonate

handling timewere

compared to , s

size (i.e.

studies haverepeatedly to foundnotonectids studies preferentially select of mosquito prey. s . . . s Further On the hand, other h This , belosomatid

and for result 5 2 3 A. nd

in water depth in water A. breddini Given both

the and sardeus

(see (see is

largest prey

- generally c 3 consistent work with previous onother notonectid style habitat s This corroborates with result

apture rate capture rate rd impact and speciesofnotonectid other 3

This also also 0 instars ofmosquitoinstars larvae)

although ). surpassed of those

According to the optimalAccording foraging theory, tothe further further andling andling times be intertwined behaviours, withswitching Capture rate suggests

on low canbedeemed densities prey and greater preysize

shorter , s and handling times of indicat s ; yet, s

indicate higherpredationpressureon for by

for specific A. breddini

this was not assessed unimodal ing maximum feeding rates s of small

of A. breddini A. breddini - A. breddini dependent predation

water depthwater - sized response ofresponse

were other predatoryother .

are handling handling times were For

prey s,

considered to beconsidered to .

greater A which display were A. sardeus The higher The higher

were .

, indicating s sardeus

and prey

in the in the capture typically typically

highest highest mostly , mo

st s

This articleisprotected bycopyright.Allrightsreserved. species can remain mosquito control arri ofpresence notonectids in colonise abletoaerially are aquatic particular abdomen enemiesnatural mosquitoes, of or natural usedin thedepth ofwater controlling mosquito larvae inacontainer style habitat 49 that of species 3 were identified size (prey impact of am subdue andconsumeitems prey such the ofthebe dueto increasing ability p attackhigher rate

Accepted Article3, , 5

5 bush, greater search areas maybush, greater enhance 0 val ofthese predators oviposition events after 4

, 5 notone , 57 1 , 5 , Our findings corroborate with the with the Our findingscorroborate ofpreviousresults notonectid workwhere predators 58 2 , ).

, in the urban , in urban the environment

Similar toour finding A. A. artificial allowing and densities Anisops

ctid predators This breddini

as potential biocontrol agents further further

s across across landscapes

and handling time were higher were higher in lower volume

aquatic under water water under them tobelesssusceptible qualitiesto than poor water groupsother

and ) factors. demonstrate

preferentially consumepreferentially

A. sardeus present study aquatic systems bodies s n for

, ot

the rate ofconsumptionthe rate of mosquito larvae by where mo onectids . much systems s 61

where aquatic systemswhere aquatic

were dependent both on physical (water depth) onwere dependent (water both physical and the biocontrolthe efficacy of notonectid

(see (see

Furthermore for larger instar redator to redator to larger prey detect

experiment, longer longer 56

capture efficiencies. can

carry ). squitoes where other predatorswhere other cannot persist.

for G

lead s iven both predators captureiven predators both theirprey by

dur

mosquito larvae overother alternative prey mosquito larvae of via via an ,

Anisopinae ( to oviposition to oviposition avoidance by ations water.

air bubble notonectids may successfully aerial dispersal

stages with multipleprey types 3 than 2 are often are

Previous studiesPrevious

of mosquitoof larvae

(atmospheric oxygen)(atmospheric Notonectinae Thus, the Thus, the predatory overall s

ub under under laboratory colonise. , coupledwithalongertime to -

family ofNotonectidae) could enable efficaciouscould enable highly

be effectively usedbe effectively insmall

predators in predators Unlike many other manyUnlike other

polluted sub have also . . mosquitoes, a 5 C - 6 4

family 0 c , 5 onsidering the the onsidering

conditions ongeneric Although the 5 ,

This might notonectids notonectids on their . shown 59 species biotic

(e.g. .

2 ,

This articleisprotected bycopyright.Allrightsreserved. productivity. Accepted3 Ecol. Evol. 2 ofNew Press University England,Hanover 1 REFERENCES code financial (DST) for providing supportthrough Article facilities.for providing laboratory We are ACKNOWLEDGEMENTS predators research varyinglevelsof ontogeny,waterdepthacross prey and density. breddini canas they neutral depthattain at buoyancy water

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Nam YenNT, VS, DucHM,TuTC,Thang BM, Bolker Rogers D, Real LA, Pritchard DW, RA,Bovy HC Paterson Lenth R Core Team Dick JT,Alexa Cuthbert RN, Dalu

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emmeans: Estimated MarginalMeans, Least aka based controlbased of The kineticsof response. functional

Random and models. search insectpopulation https://CRAN.R

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86 bbmle: Maximum forGeneral Tools LikelihoodEstimation. RPackage ,

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This articleisprotected bycopyright.Allrightsreserved. 51 by predatory aquaticinsects. Accepted50 mosquito predators. 49 with naiads preyalternative available. 48 dynamics. 47 via conspecific prey a heterospecific 46 (1998) scyphomedusa Article45 vectors. and Lounibos LP, 44 comparative functional responses. and nativepredatory freshwateramphipods under change environmental arerevealed by 43 intermediateinvasive predator. Bovy JC, 42

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and BoukalDS, and

and SullivanB, Chrysaora quinquecirrha Ribeiro GSRibeiro

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ill D, ill Dick JTA,Emmerson MC,

Johns and Kishida O, and Kishida

She's a femmelow She'sfatale: a Aquat. Ecol on DM, 38 ,

Maciel

: Who Acomparative whoma pool ? eats ofprey in study selectivity PLoS ONE

297

Switching andsigmoidSwitching by functional response curves damselfly Vulnerability of the copepod Vulnerability ofthecopepod J Anim Ecol .

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Biol Stabilization of andnonpermanent chaotic food ‐ 303 De :

predator. : 167

96 J AnimEcol ‐

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and LucyFE, : effect of : andbehavior. preysize – – 7 103 176 :

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This articleisprotected bycopyright.Allrightsreserved. inCacharlocated District, Assam, India. Aand60 Dalal GuptaS, Accepted backswimmerthe (Notonectidae, 59 MatthewsPG animal drinking troughs in NewZealand. 58 242 common 57 Oceanogr. 56 Implicationslarvae: for biological 55 Articleheteropteran bugson 54 NewYork,Press, pp.98 Ecosystems, ed.by from tocommunities, individuals in 53 regulationthe mosquitoes. ofwetland 52 and alter predationthe of

Zuharah Zuharah WF Saha N,AdityaG,BalAand MAScott Saha N,AdityaG,BanerjeeS Saha N,AdityaG,BalAand Saha N,AdityaGandGK Woodward G – 257 native prey.

heteropteran bugsheteropteran ofEastCalcuttaWetlands, Kolkata (2007b) 28

and : 352

and Lester PJ, and Lester Buenoa fuscipennis Buenoa

and . and Seymour RS, Murdoch, WW, Murdoch,

- 366 Hildrew J. Vector Ecol. Warren P, Culex quinquefasciatus

(1983) -

A comparative of study the aquati 117

AG (2007)

. The influence predators ofaquatic onmosquito abundance in

Body predatory and size interactions infreshwaters: scaling Saha GK Saha GK

Raffaelli DG Raffaelli

and SahaGK, Anisops

Selective predation bytheSelective backswimmer, Notonecta. control. , Prey preferences ofaquaticPrey preferences p , insects:

Haemoglobin as a buoyancyHaemoglobin asa regulatoroxygen and in supply

38 (Heteroptera: onmosquitoimmature notonectidae) stages

Body Size: Body Size: The Structure andFunctionof Aquatic Med :

215

Turk J Zool ). J Vector J Vector Ecol , ,

Biol A comparative study of ofthreeA comparative predation aquatic Comparative studies onfunctionalComparative responseof Vet J Exp Biol –

223 larvae.

and Edmonds

Entomol

Control Predation potentialPredation ofodonates onmosquito

(2013) Limnology 40

211

: 35 28 .

392 : : : c insect diversity oftwo ponds

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1 1 - - – Brown - 401 9 8 , India. , India.

8 353 (2014) (2012) - :73 3799

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– (2010) R, Cambridge University R, Cambridge University 80 Int Int Rev Hydrobiol

. .

(2008)

. (2007a) otential otential implications for

Limnol.

92 :

This articleisprotected bycopyright.Allrightsreserved. Accepted Article meta 61 Vonesh JR and Blaustein L, Vonesh JRandBlaustein - analysis and implications for vector contr andimplicationsforvector analysis Predator - induced shif ol. Isr

ts in mosquitots in oviposition site J Ecol EvolJ Ecol , 56 :

263 - 279

(2010). selection: a selection: a

This articleisprotected bycopyright.Allrightsreserved. Acceptedsardeus Anisops breddini Anisops ArticlePredators FRs, forTypeII presented first whilst orderterms andsecond presented are FRs. forTypeIII sardeus Anisops Table 1.

Functional response (FR)typesresulting fromwithregression, first and second terms logistic order of

D2 D3 D2 D1 Depth D1

against four instars (1

0.067 *** - - - - 1 1 0.002 0.026 * 0.012 *** 0.006 * st st

order instar

- - - - ** 0.0003 order 2 0.0005 nd

–

4 th III II II II III types FR ) of

Culex quiquefasciatus

- *** - 0.013 * - *** - order 1 2 0.054 0.016 0.006 0.014 st nd

instar

- - - - - 2 0.00007 nd

order

three different levelofwaterdepth.three order First terms are II II III II II types FR

- *** - *** - - *** - 1 3 0.026 0.024 0.01 0.003 0.108 st rd

order instar

- - - - *** 0.0009 order 2

II II II II III types FR

- *** - *** - *** - *** - order 1 4 Ansiops breddini Ansiops 0.033 0.019 0.018 0.02 0.013 st th

instar

- - - - - 2

order

and

II II II II II types FR

This articleisprotected bycopyright.Allrightsreserved. Accepted Article* =P<0.05,**0.01,***0.001. D3

- 0.026 ***

- ***

*** - *** 0.046

- *** 0.002

*** - *** 0.018

This articleisprotected bycopyright.Allrightsreserved. Accepted Article depth levels, alongsidetheir significance. sardeus Table 2. * =P<0.05,**0.01,***0.001. breddini Anisops Predators sardeus Anisops

towards fourinstars (1

Capture rate( Capture

D3 D2 D3 D2 D1 size Depth D1

) and handling) and time ( *** 2.37 *** 0.082 *** 1.449 *** 0.838 *** 0.706 *** 0.034 b 1 st

st –

instar

4 th *** 0.025 *** 0.01 *** 0.009 * 0.007 0.002 *** 0.011 ) of

Culex quiquefasciatus

*** 1.88 *** 0.258 *** 0.275 *** 0.155 *** 0.043 *** 1.65 h b ) parameters of 2

instar

*** 0.017 *** 0.016 *** 0.018 *** 0.014 *** 0.011 ** 0.005 h

* 0.036 *** 0.881 *** 2.41 *** 2.694 *** 1.365 *** 0.84

Anisops b 3

larvae larvae

instar *** 0.025 0.004 *** 0.021 *** 0.024 *** 0.015 0.007

h breddini at

three different water

*** 0.609 * 5.02 *** 4.726 ** 3.776 *** 0.967 *** 1.08

b 4

and

instar *** 0.045 *** 0.082 *** 0.066 *** 0.057 *** 0.062 *** 0.07 Anisops h

This articleisprotected bycopyright.Allrightsreserved. Accepted ( 95% CIs bootstrapped represent bars and estimates time handling are Points D3). D2, (D1, depths water e shapes, 4. Figure bootstrapped 95%CIs( at e dots, shaped (square 3. Figure ArticleD2 (roun points, (squared D1 residuals: raw are Points areas. shaded the are intervals confidence 95% ( Bootstrapped depths. water lines) dotted (D3, high and lines) dashed (D2, medium (1 stages instar prey four 2 Figure (1 stages sardeus Anisops 1 Figure Figure legends

he wtr ets D, 2 D) Pit ae tak ae siae ad as represent bars and estimates rate attack are Points D3). D2, (D1, depths water three st . Functional responses of responses Functional . d shaped points, andD3(triangle shapedpoints, ◌) . Mean .

n Handling times of times Handling – – atr rts of rates Capture

= 2000). 4 ) gis fu dif four against h) th ) of

raw ge clue br e bar, coloured (grey Culex quinquefasciatus

consumption (± SE) of SE) (± consumption

– n

h) against four different instar stages (1 stages instar different four against h)

= 2000). st – nsp breddini Anisops Anisops breddini Anisops

4 th eet ntr tgs (1 stages instar ferent of )

Anisops breddini Anisops ue quiquefasciatus Culex –

at h)

three depths increasing water (D1

cos l fu densities four all across Anisops breddini Anisops (round shapes, a shapes, (round rud hpd os a dots, shaped (round

(a (a st

– –

d) and d) 4 th

△ larvae across low (D1, solid lines), solid (D1, low across larvae of ) )

– .

st Anisops sardeus Anisops (black coloured bar, a bar, coloured (black

d) and d) ue quiquefasciatus Culex –

th – ) of )

oad fu pe instar prey four towards ) and d) Anisops sardeus Anisops Culex quiquefasciatus Culex – nsp sardeus Anisops

D3). (e –

h) towards towards h) n

–

= 2000) =

(square d) and and d)

three three • ),

This articleisprotected bycopyright.Allrightsreserved. Accepted (1 stages sardeus Anisops 1 Figure Article st .

– Mean Mean

4 th ) of

raw raw ge clue br e bar, coloured (grey Culex qui consumption (± SE) of SE) (± consumption n quefasciatus

–

at h )

three across all prey densities densities prey all across Anisops breddini Anisops increasing increasing water depthwater

(black coloured bar, a bar, coloured (black towards s

(D1

–

or ry instar prey four D3) .

–

d ) and and )

This articleisprotected bycopyright.Allrightsreserved. D2 ( Acceptedintervals confidence 95% (D2 medium (1 stages instar prey four Figure Article round shapedpoints,

2 .

Functional responses of responses Functional , da , shed lines) and high (D3 high and lines) ◌ st are the shaded areas shaded the are ) and D ) and –

4 th ) of 3 Anisops breddini Anisops

(triangle shaped points, (triangle shapedpoints, ue quiquefasciatus Culex , dotted lines) water depth water lines) dotted , . Points are raw residuals raw are Points .

(a (a –

and △ larvae ) .

Anisops sardeus Anisops

across s . : D1 (squared points, (squared D1 : B ootstrapp

low (D1, solid lines), solid (D1, low

(e ed –

( h) n

= 2000) = towards • ),

This articleisprotected bycopyright.Allrightsreserved. Acceptedbootstrapped at e dots, shaped (square Figure Article

he wtr depth water three

3 .

atr rate Capture

95% C I s

s – (

n s

h) against four different instar stages (1 stages instar different four against h) D, 2 D) Pit ae tak ae siae ad bars and estimates rate attack are Points D3). D2, (D1,

= 2000) of nsp breddini Anisops .

rud hpd os a dots, shaped (round st

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th – ) of )

) and d) Culex quiquefasciatus Culex nsp sardeus Anisops represent

This articleisprotected bycopyright.Allrightsreserved. Accepted 95% C depth water shape Figure Article s e , I

s ( 4 .

n Handling time Handling –

s = 2000). ) gis fu dfeet ntr tgs (1 stages instar different four against h) (D1, D2, D3). D2, (D1,

of Points are handling time estimates and bars represent bootstrapped represent bars and estimates time handling are Points Anisops breddini Anisops

(round shape (round st

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4 th s , a , of )

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at (square

three three