Posted on Authorea 22 Jan 2020 | CC BY 4.0 | https://doi.org/10.22541/au.157971041.10418309 | This a preprint and has not been peer reviewed. Data may be preliminary. ot aoaoyifcinmdl r sfli eosrtn h nuneo oneto pntecourse the upon coinfection of influence the the in demonstrating outcomes in associated useful predicting are and understanding models for infection essential Laboratory is to species parasite populations commensalism, host. between approach and interactions Understanding hosts that mortality. coinfected infections or immune in asymptomatic morbidity Th1/2 cause from that pathology, e.g. infections & of severe responses Sher, more levels immune respectively(Jankovic, impact varying of macroparasites and helminth macroparasites) types and 2001), diverse life-cycle helminth parasites Yap, qualitatively intracellular size, causing and to by ways, response microparasites in of distinguished variety polarisation single-celled (here a guilds in macroparasites, hosts parasite poses helminth their viruses Different and as and difficult, ecologists. bacteria niche is disease endoparasites, and species helminth for how multiple and challenge Predicting with protozoan significant ecological 2015). coinfected ectoparasites, a al., an populations by et and Graham, through infection Seabloom L individuals for 2007; communities A Fenton, in accounting 2008; & parasite interact the Pedersen Hudson, will within-host is & 2015; species Fenton, Boag, explore species & parasite (Cattadori, to parasite Roode, phenomena De multiple isolated effort Johnson, as with recent 2008; infections coinfection treating a than and been rather ubiquitous, lens, has is There populations wild norm. in infection Parasitic to study longitudinal for need Introduction A explored. are might species associations highlighted. same these is the how interactions from explain of research associations. causality to inter-species laboratory determine suggested of analogous for prevalence are with account the ecological, to Comparisons to need and due the occur. immunological populations, highlights infected including study naturally mechanisms, This studying with potential 5. when and Various including possible role. associations, central as between-guild species, a multiple infections parasite playing involving many hepaticum associations, as of population was negative Calodium range island species and species recorded broad positive helminth an Every of under-studied a from framework the 4. between data a helminths. within a cross-sectional associations hepatic exist used in identify and to We gastrointestinal guilds to found and across domesticus, 3. ectoparasites, arthropod musculus associations host. microparasites, understanding Mus parasites the better blood-borne mouse, characterise and within within-guild research, house to manifest laboratory-based the and might aimed existing We of associations interactions with observed comparisons parasite groups 2. for any on pairwise parasite allowing how depending species, multiple explored. many of model ways spanning commonly While a various network less of interaction in population been infection. wild an interact has upon how can mediated host understanding which be the characterised, might species, of been parasite have state communities multiple the parasite harbour and involved typically species populations the animal Wild 1. Abstract 2020 5, May 2 1 MacColl Fenn Jonathan negative and positive guilds mixed across mouse of house network associations wild a a in of exist communities population Parasite connected: all It’s UNSCAinWl ateSeils Group Specialist Cattle Wild Asian SSC IUCN Nottingham of University 1 hitpe Taylor Christopher , 1 nrwWolfenden Andrew , 1 n a Bradley Jan and , 1 tatYoung Stuart , 1 1 2 aa Goertz Sarah , 1 n Lowe Ann , 1 Andrew , Posted on Authorea 22 Jan 2020 | CC BY 4.0 | https://doi.org/10.22541/au.157971041.10418309 | This a preprint and has not been peer reviewed. Data may be preliminary. ouaino os iehsbe rsn nteiln o vracnuy n a upeetdb an by supplemented was and century, a over feral for A island bedrock. of the exposed consists frequent on and and present hectares vegetation been 45 ruderal of has around Firth tall mice the of of in Reserve patches house W), Nature with of 2.5667deg National grassland N, population coastal designated (56.1833deg May of a of mosaic is Isle a island the The on undertaken Scotland. was Forth, study this for fieldwork The associations. involving these Methods associations to underlying & negative research mechanisms and Materials positive potential population this of discuss wild network connect and a a establish easily parasite, from infections, observed more of data every range to survey broad a parasitological us from ( infections, cross-sectional allow infections improve mouse use individual will will house we of the possible species Here distribution of as model affecting studies. infections in factors of laboratory assessments the range related these and a broad communities, performing as parasite and analogous for of by account understanding informed which and our models to species multi- holistic related model towards directly settings. of a Working more laboratory context of be controlled a population to in wild associations performed within these a studies infections of co-infection in understanding individual networks 2019). our interact al., considering allow et cross-guild will shifting of (Beechler Characterising assemblages, traits importance functional parasite ecosystems. the specific parasites of with illustrate richness negative parasites examples taxonomic favouring distribution, and increasing These or position hosts, taxonomic methods the overall ( transmission the within buffalo African parasites in wild of overlap in communities or infections tuberculosis host, bovine the New competition. most of localised associations were from state positive guilds, associations while result negative and the that primarily Here, types by indicating associations tissue niche, 2019). governed physical across al., a be associate et shared to (Dallas may which shown positive parasites were between being desert found associations commonly Sonoran of species. the non-model majority in of the populations species with wild rodent in 22 explored recently of more Parasites been have communities parasite Cross-guild sengis, lions, voles, and field cross- rabbits, and wild 2010), negative infecting microparasites and al., popu- microparasites positive between et in wild of Telfer and observed Networks in been 2018), 2010). observed have Jolles, al., interactions pairs & species et Beja-Pereira, species (Taylor Luikart, between 2004; Etienne, microparasites Balcha, interactions (Ezenwa, between & helminths competitive Erko, interactions in- and Legesse, include pairwise Mutualist 2010; specific lations Sheldon, 2002). Ferrari, on & al., 2012; Palinauskas, or Bastos, Knowles, et 2010), & 2009; Nacher al., Chimimba, Hudson, Anguelov, et & (Brettschneider, Telfer Rizzoli, guilds Sandra Cattadori, different 2015; of Bennett, under- species & well between Stevenson, Fagir, increasingly Fenton, teractions Boag, Lutermann, is (Lello, 2004; animals interactions within-guild wild Hudson, on coinfected & within focussed in typically exploited have structure niches studies change community coinfection the parasite also stood, and of may response importance They immune the in 2008). While 2012). changes Graham, to al., L due et (A cycle, (Duncan processes processes life host Read, & ecological these parasite’s the Gras, Lamb, direct a of Stancampiano, of Graham, through combination 2015; stages a L 2005), Bashey, between through 2009; Stevenson, Andrea Brun, or in & 2005; Sch¨otzau, & 2010), changes Stearns, Loredo-Osti, Shi, Poglayen, the through competition(Balmer, Morgan, & influence both niche Walker, Segura, can as mediated McCormick, Su, changes such be Louie, these 2005; can and (Chen, Allen, species ways, state & parasite of immune between variety Interactions host’s a in a infections. host other infected their begun of naturally within 2010). populations course environment in al., wild the et types of Telfer alter studies parasite S Parasites have 2019; multiple Ovaskainen, recently incorporating & only Laine, However, communities (Dallas, immunity. parasite animals host multi-species of explore role to the and infection of lpatlsmyurus Elephantulus rcoau cuniculus Oryctolagus u uclsdomesticus musculus Mus Ltrane l,2015). al., et (Lutermann Lloe l,20) n natrpdetprst omnte netn wild infecting communities ectoparasite arthropod in and 2004), al., et (Lello atealeo Panthera Futi-oe ta. 09,i emnhcmuiiso wild of communities helminth in 2019), al., et (Fountain-Jones oepoeascain mn rvlneaditniyof intensity and prevalence among associations explore to ) 2 ycrscaffer Syncerus eefudt hf cross-guild shift to found were ) irtsagrestis Microtus (Sandra Posted on Authorea 22 Jan 2020 | CC BY 4.0 | https://doi.org/10.22541/au.157971041.10418309 | This a preprint and has not been peer reviewed. Data may be preliminary. i Etat N rp unaBoMsahset,UA,adtersligDAue o C.Temi- The PCR. for used DNA resulting extraction the 2-step and a USA), using Massachussetts, clots Bio blood Quanta frozen prevalence Prep, the the only DNA from from therefore (Extracta extracted and was kit and DNA to dynamic, prevalence, move microparasite highly rapidly from record are To fleas Nottingham, numbers As of death, counted. University after species at recorded. and each created was of capture (key number after key total dissecting including visual a and body, a under Myhill), examined using Laura was identified by mouse each photos parasites of all fur with the discoloration). dissection, microscope extensive following = infections, 5 ectoparasite to assess To decolouration, no helminth = hepatic 0 the (from colouration by infection of hepaticum level analysis. the further measure in To included not was gastroin- and the tapeworm mice) in whipworm the found recorded the was recorded, commonly recorded in- and were animals: Also worms identified large dissected helminth visually . of the and were species of intestines observed Two tracts parasites small forms. testinal all adult caecum, and and larval stomach, microscope both a the including under be status, dissected could infection were surveys testines helminth ectoparasite gastrointestinal survey until To roll paper lost. Brigada, in (Carreno, being age wrapped from Infection discriminating parasites were at prevent length reliable Bodies to foot most order length, the in proved tail 1990). conducted mass length, Castro-Vazquez, lens throughout body Bradfield, continuously eye & mass, grows (Rowe, mice, were it body Rosi, corn age as lenses to in age estimate compared eye length estimate When to ear the to and 2008). used used Later, commonly (Augusteyn, measurements is lifespan formalin. mass resulting animal’s lens buffered an the Eye 10% and 1985). in Swinney, weighed, & stored and Quy, and dried removed at out, were kept dissected eyes then the diagnostics, dissection, microparasite During for nitrogen liquid ethanol. in 80% in stored 1 for and temperature retained room -80deg were condition at clots animal clot point. for resulting to this proxy allowed 4deg body and at a exsanguination, at relative confirmed as during stored the of also used pipetting then of measure was by be hour, a tip obtained animal to were Index, the the whole, samples of Mass a from Blood sex as Scaled distance The population calculate the the 2009). to length, for Green, used snout-vent curve & then (ii) (i) a (Peig were and against were: values grams, scales These taken which in variables mass anus. mouse CO physiological the the increasing External to of using nose mass guidelines exsanguination. the Office by weight, Home confirmed with body was accordance Death in euthanised concentrations. were mice material, trapping, al, Following bedding et as Taylor hay see with locations provided trapping birdseed, of Dissection of mixed day. limits map with the the a baited throughout within were (for and frequently island Traps sites checked the Ugglan the 2019)). and breeding of across al., of coverage bird areas et the for mixture (Taylor different maximise clearance 2019 in a to allowing set order and using traps in terrain, twenty out selected the of were carried transects locations of Trapping were consisted seasonality island. Materials) effort the Trapping Supplementary to live-traps. (see Due Longworth sessions pressure. migration. predation trapping little no Eight owls very with short-eared 1991; be closed, including Noble, to island, & assumed assumed the Nash, is is King, on there Triggs, population raptors) (Berry, The (primarily 1982 predators in 1991). Eday of island Triggs, Scottish 1885; the from Muir, animals 77 of introduction C pnrtr rmtefil.Tegsritsia rc a eoe n trda omtemperature room at stored and removed was tract gastrointestinal The field. the from return upon ) , h ie a iulyasse uigiiildseto n crdbsdo h ee fde- of level the on based scored and dissection initial during assessed visually was liver the C o nte or h lo a hncnrfgda 350RMfr5minutes, 5 for RPM 13,500 at centrifuged then was blood The hour. another for yeoesssp. Hymenolepsis huhti a aeyrcre 5isacsoe 366 over instances (5 recorded rarely was this though , 3 rcui muris Trichuris aoimhepaticum Calodium n h pinworm the and soflammeus Asio as nw as known (also uigtewinter, the during yhcaobvelata Syphacia Capillaria 2 Posted on Authorea 22 Jan 2020 — CC BY 4.0 — https://doi.org/10.22541/au.157971041.10418309 — This a preprint and has not been peer reviewed. Data may be preliminary. n 1 eeifce with infected were 112 and umr falprst pce eoddi h os ouaini hw nTbe1 u f36recorded 366 of Out 1. Table in shown is population with mouse uninfected the in were recorded 127 species mice, parasite data all of (Sup- ecological summary reported A in are power sizes effects statistical all and but Results corrections testing, multiple Bonferroni for of corrected not use Materials). are plementary the models surrounding the ‘ 2004), issues the (Nakagawa, the using averaged to were Due AIC) lowest The the process. (with averaging ‘ model” package model a “best through the selected Any to and simplified power. subsequently were are statistical ‘ level reported, increase infection henceforth of to are Models that excluded datasets models. species complete exclusionary these with these parasites from with of taken repeated pairs to between were taken relationships were variable significant zero and predicted of microparasites the for scores data not infection missing with is individuals there model, As excluded. this were For thus of and As factor. uninfected, ordinal series be an individuals. a as uninfected mites), score ‘ excluding infection fur the data, and of abundance helminths function using (gastrointestinal of effect. performed abundance levels fixed were a infection infection of as GLMMs parasite measures included parasite binomial on not of with negative was bearing patterns species it inter-annual significant or so parasite a seasonal and For have any evident, that would become means to session session unlikely trapping each are of that distribution spacing likely irregular is the density), dynamics, it population linear for Although hepaticum proxy generalised a C. binomial as effects. obvelata, Index here of Mass S. (used series Scaled day muris, per sex, a T. transect age, in per were microparasites, variable captures model of condition, response each prevalence of in measure the a incorporated as as variables used (SMI) Ecological was (GLMMs). parasite models each mixed 3.5.3. of version presence R The in performed were analyses statistical All Analysis Statistical no so lost, was mice by minutes Infection 30 with gene for Information). additional an Biotechnology gel of for agarose PCR UK) Center 2% Nottingham, (National amplified parasite, a Bioscience, BLAST successfully apicomplexan (Source from on of on company samples Identity run sequences amplified external reference was Successfully an amplification. against DNA at confirm data comparison sequencing to Amplified presence/absence and light through only ultraviolet confirmed so analyses. was under and in samples visualised recorded, and used reliably 100V, are be at not species could populations these rodent parasites 2018). wild from al., these across et of Taylor detected measures 1981; commonly Abundance Healing, are which 2003; Avsic-Zupanc, genera & and protozoan Trilar, species Petrovec, flagellate on (Duh, the based below), chosen were see sites – instead bacteria detected the and was apicomplexan protozoan related apicomplexan the closely were detection for selected croparasites dredge ucinwsue ocmaealpsil u-oes n hs with those and sub-models, possible all compare to used was function ’ MuMIn C.hepaticum ordinal npam phagocytilum Anaplasma .hepaticum C. )t n pia oesfrec aaiespecies. parasite each for models optimal find to ’) .hepaticum C. accsi dispersa Sarcocystis a o eoddfrtefis 5 iecuh,adbodco aeilfrthese for material clot blood and caught, mice 150 first the for recorded not was akg nRwsue ocet uuaieln ie oe,tkn the taking model, mixed link cumulative a create to used was R in package ’ .muris T. .obvelata S. .muris T. Sarocycstis eemaue na rirr - cl fifcinsoe h ‘ the score, infection of scale 1-5 arbitrary an on measured were rmcoaaiedt a vial o 5 f36mice. 366 of 150 for available was data microparasite or B.microti n u ie.Lcto n rpigssinwr nldda random as included were session trapping and Location mites. fur and , 3.0) fteemc,4 1.8)wr onetdwt both with coinfected were (11.48%) 42 mice, these Of (30.60%). .obvelata S. or and seicpies eunigo C rdc sdsrbdabove. described as product PCR of sequencing primers, -specific .obvelata S. h dniyo hsprst a ute ofimdthrough confirmed further was parasite this of identity The . rmr eefudt culyb rmifcin yanother by infections from be actually to found were primers atnlaspp. Bartonella .hepaticum C. , 4 .hepaticum C. 3.0) 6 eeifce with infected were 169 (34.70%), neto,mdl nwihteeprstswere parasites these which in models infection, seSplmnayMtras.Teepara- These Materials). Supplementary (see aei microti Babesia n es n h neto nest of intensity infection the and fleas, and model.avg Δ AIC atog nfc another fact in (although ucin(ohfo R from (both function ’ < .muris T. hncompared when 2 rpnsm sp. Trypanosoma (46.17%) clmm ’ Posted on Authorea 22 Jan 2020 — CC BY 4.0 — https://doi.org/10.22541/au.157971041.10418309 — This a preprint and has not been peer reviewed. Data may be preliminary. mueatgns a xli eaieascain ewe aaie ucpil osmlrimmune similar to susceptible parasites between associations environments. negative or explain responses which may upon antagonism framework Immune a providing and explored. observed, parasite further structure, were Immunity multiple be community associations might across parasite positive interactions coinfection and parasite of negative many of for picture mechanisms Both with accounting complicated species, in populations. other of a distributed wild one forming importance are coinfected least studying the at species with when confirming parasite associated groups a individual guilds, was highlighting measured determining across coinfection, species in observed was parasite structure Every intensity community and population. parasite prevalence this of infection role determining variable crucial frequent most interactions The parasite of network The of prevalence Discussion the both. of with of population and prevalence this OR=1.811), the in p=0.0264, predicting seen (z=2.22, OR=2.384), fur p=0.0255, the (z=2.421, in fleas with associations ( species both C.hepaticum for models the hepaticum in observed is OR=4.272) p=0.00037, that (z=3.558, latter hepaticum the of C. that both predicting by between significantly observed with former evidenced and association the as positive of prevalence strong association, the a negative with was a There have models. helminths OR=0.456) gastrointestinal of SMI species the with common two relationship The positive a showed also Interactions p=3.2x10 mites Parasite (z=4.659, of age and Abundance OR=1.060) p=0.0113, OR=1.007). (z=2.533, p=0.0107, (z=2.552, age ( fasciatus mite impact of significantly genera to Four found were parameters ecological No OR=1.0167). with of infected Prevalence be to found with were infected mice be to 123 detected. were microparasites p=8.82x10 No associated (z=3.921, positively prevalence was parasite The the of uniform. age. OR=1.011) predominantly p=0.00156, with is (z=3.163, intensity by individuals infection infection p=0.005, and infected of OR=1.020) level (z=2.780, in some score day showed 261) of infection per of prevalence of and out transect density (152 The individuals per population captured host captures stable. Most between fairly of association number negative remains a host the implying the with OR=0.613), established, in associated has established infection negatively once parasites significantly that of suggesting OR=1.001), number p=0.4941, the (z=0.684, abundance parasite p and dramatically age (z=6.038, having days individuals in rare estimate with burdens, The infection distribution, low binomial negative towards a approximate loads. shifted to higher found greatly were infections species both with for distributions Burden parasites. npam sp. Anaplasma .obvelata S. .muris T. .hepaticum C. eefudi h u ftemc.Peaec ffla a on ob oiieyascae with associated positively be to found was fleas of Prevalence mice. the of fur the in found were ) oe =.7,p008,OR=2.648). p=0.0488, z=1.971, model rvlne(=.0,p001,O=185.Teaudneo ie ntefrsoe positive showed fur the in mites of abundance The OR=-1.815). p=0.0211, (z=2.306, prevalence ioilmxdmdlfudtepeaec fteprst nraigsgicnl ihteage the with significantly increasing parasite the of prevalence the found model mixed binomial rvlneas hwdapstv soito with association positive a showed also prevalence accsi sp. Sarcocystis rvlne(=.6,p003,O=.0)and OR=0.207) p=0.0136, (z=2.468, prevalence accsi dispersa Sarcocystis or Myobia neto cr rdcigteaudneof abundance the predicting score infection rpnsm sp. Trypanosoma .musculus M. C.hepaticum < , a togypstvl soitdwt nraigae(=.8,p=0.0129, (z=2.487, age increasing with associated positively strongly was 2x10 Radfordia -16 R106.Teewsn soito bevd oee,between however, observed, association no was There OR=1.026). , ssoni i 1. Fig in shown is fteemc,7wr onetdwt ohparasites. both with coinfected were 7 mice, these Of . hw eaieascain with associations negative shows , eedtce yPRi n ftemc.I 2 ie no mice, 127 In mice. the of any in PCR by detected were Myocoptes atnlasp. Bartonella accsi sp. Sarcocystis 5 & Trombicula -6 OR=1.006). , oe =.2,p000,OR=3.067. p=0.0203, z=2.322, model .muris T. .muris T. atnlasp. Bartonella rvlnewsngtvl rdce by predicted negatively was prevalence ,adoeseiso e ( flea of species one and ), atnlasp. Bartonella atnlasp. Bartonella z251 =.17 OR=1.189). p=0.0117, (z=2.521, bnac z215 p=0.0336, (z=2.125, abundance S.dispersa .hepaticum C. .hepaticum C. rvlne relationship a prevalence, .obvelata S. hc a edue be may which , n 8wr found were 18 and , prevalence h distribution The . .obvelata S. and .hepaticum C. prevalence. Nosopsyllus .muris T. . was C. -5 , , Posted on Authorea 22 Jan 2020 — CC BY 4.0 — https://doi.org/10.22541/au.157971041.10418309 — This a preprint and has not been peer reviewed. Data may be preliminary. hsclsaewihteprstsocp a otiuet h eaieascainosre ewe these between observed association negative the to contribute may occupy parasites the which space physical impact Both may helminth one by Wakelin, caused Effects Roach, environment Ecological 2013; immune the al., species. to other et modulations Nieuwenhuizen of so survival 1991; not and the and Bianco, mice 1988), and against & Bundy, both protection in infection & Cooper, or observed Else, been to than Didier, has susceptibility Bundy, other species to (Lillywhite, nematode individuals factors different humans predispose of by antigens genetics, between as Cross-reactivity determined such infection. study, host, this the in for within accounted environments immune Pre-existing 2007). important response al., Th1-type of et the site from (Kim away a burden. polarise as may mite which recognised infection 2001), early ectoparasite been to Alarcon-Chaidez, in response & has observed in Wikel tissue detected 2010; been dermal al., have arthropods et the responses parasitic (Burgess Th2-type and characterised against 1990). well protection 2016), (Kupper, to less Watson, activity contribute and immune & may population, Pate, skin this the Baxter, in in (Moats, defined Inflammation an well models. by less laboratory are mediated from ectoparasites be with alternatively observed therefore, Associations may, of part population the this on in effect observed immunosuppressive interaction The proliferation. nematode hepatic similar during environment muris Th1 IFN a and to matured, polarisation worms Systemic the with as infected increasing artificially between(Th2) rats association In the responses. immune in role a playing nematode 2013). be Harrus, could & and immunomodulation Morrick, Karem, of Baneth, Akanbi, 2007; impacts (Kamani, Peripheral al., individual ( coinfected et one rat Kabeya characterised in brown 2001; observed typically between the pathology Mielke, association are in positive & which A reported Hahn, microparasites, 1999). was Regnery, to Regnath, & immunity Ignatius, McGill, protective (Arvand, Dubois, reduce between responses could Th1-type association 2007) by positive al., the et to (Kim contribute may sp. suppression Bartonella or 2010). Chauvin, facilitation & Immune (Moreau Induction abundance norm. its the impacting be could negatively reinfection , by possible 1992). and responses al., Didier, clearance Th2 et Thompson, parasite of Cooper, Needham that (Bundy, indicating 2002; level age, low al., with steady et association a Faulkner to 1987; decrease then Simmons, H life, and & Else, early 1999), during 1999; Gilbert, spike & Grencis, burdens Zain, which & Lewis, to to Finkelman, (Behnke, polarisation contrast age Else, with in and Potten, increase is responses (Artis, This Th2 models 1992). protective infection Grencis, of in environment immunosuppression Th1 by of indicators a characterised be may are population which this in infections, infection low-level of prevalence high pinworm and the between with association infected artificially negative reduced mice the in observed explain are also associations may antagonism Immune early during impacting exhibited responses Th1-type to T.muris T.muris one l(07 bevdapoetv muersos oifcinagainst infection to response immune protective a observed (2017) al et Moon . .muris T. eimsmie polygyrus Heligmosomoides accsi sp. Sarcocystis ako ta 20)fudpstv soitosbetween associations positive found (2009) al et Jackson . oaiaint h niomn bevddrn oesae fthe of stages some during observed environment Th2 a to Polarisation and C.hepaticum udn Keig 91.Tesed nraein increase steady The 1961). (Keeling, burdens S.obvelata .obvelata S. lnrhssinensis Clonorchis infection. r yial on nteceu rpoia oo,s optto o resources for competition so colon, proximal or caecum the in found typically are neto,adcnrbt oapstv soito between association positive a to contribute and infection, ol irp h muoupesv niomn salse by established environment immunosuppressive the disrupt could atsnorvegicus Rattus .muris T. nwl odmice wood wild in .muris T. nlbrtr as hrceie yesnpii n lymphocyte and eosinophilia by characterised rats, laboratory in C.hepaticum netososre nohrwl oet,i hc burdens which in rodents, wild other in observed infections γ favouring , .hepaticum C. C.hepaticum T1 ekn uigegdpsto Kme l,2007). al., et (Kim deposition egg during peaking (Th1) ,tog hswsbsduo nuuulices in increase unusual an upon based was this though ), 6 pdmssylvaticus, Apodemus neto Km o,&Cug 07 negatively 2007) Chung, & Joo, (Kim, infection .hepaticum C. Trichuris .obvelata S. h/ oaiaini bevd ihIL-5 with observed, is polarisation Th1/2 , g eoiincudipoesria of survival improve could deposition egg .hepaticum C. .muris T. siuui teraptera Aspiculuris itiuin bevdi uas in humans, in observed distributions T.muris .hepaticum C. uvvl Kme l,2007). al., et (Kim survival. rvlnesosn significant no shows prevalence rvlneascae ihage with associated prevalence soitdwt eue innate reduced with associated .muris T. and and C.hepaticum atnlaelizabethae Bartonella S.obvelata .hepaticum C. salsigchronic establishing n h intestinal the and C.hepaticum C.hepaticum hc suffered which , C.hepaticum Similar . lnr & ultner, ¨ .muris T. iecycle life n a and and and T. Posted on Authorea 22 Jan 2020 — CC BY 4.0 — https://doi.org/10.22541/au.157971041.10418309 — This a preprint and has not been peer reviewed. Data may be preliminary. neto oeswihmyb vdn ee hl oeta ahgn uha iue r o nlddi this in included not are viruses as such pathogens potential While here. exist evident associations be may exhibiting least which of pairs at models many species infection with explain the association niches. may of most which an different model as Interactions in within immune-mediated, observed. in implicated potentially interactions particularly was are cross-guild population reported, associations multiple these host commonly with wild less parasite, this been other in however, one surveyed has, parasite guilds network multiple Every comprehensive Incorporating species. a well-recorded. are in animals parasites coinfected of the naturally in which parasites from among framework Interactions robust a tested. provide links and and hypothesised causal independent further season, Conclusion therefore direct or be are can age infer associations interactions sex, to parasite reported as of such The possible relevantmechanisms factors of models. not number confounding all a was possible including in by of it parameters possibility this ecological data mediated mitigate be and to cull potentially tried physiological may have sectional We relationships variable. observed unaccounted-for cross any some and by from species, 2007; parasite derived Morand, between associations were & explaining Blumstein, data (Bordes, by the animals hosts As between infect directly 2016). which higher passed 2013, experience fleas, typically Dearing, may more & are socialising time Dizney to mites more themselves as spend which burden, expose hosts observed mite while may environment, associations animals the positive in exploratory vibrations underpin detecting more could e.g. that behaviour factor host a here observed 2007), macroparasites as (Flynn, helminth method, three between orally All transmission transmitted guilds. in notable. within typically dissimilarities observed is or are associations condition of similarities from aggregations host from expected different to arise be that linked might also associated fitness, strongly could reproductive parasites be be between account the to to Interactions on into seem found coinfection, take not measure associated not do and only does hosts SMI infection, this the within of between While was significance parasites observed host. the abundance association question the mite into of positive Fur bringing fitness study, the mice. this by study in (Perkins, supported our SMI condition with host is in in This abundance change mite 2017). a than and Gillingham, rather & mediated immune Pascoe, be ( White, may voles interactions bank these that wild suggesting of mites While deer spp. 2013). Wild Antonovics, unclear. & is (Pedersen coinfection glareolus antihelminthic helminth an from with result treatment could this with whether ( coinfected though mice Alterations rodents 2013), grooming. in Chieffi, of recorded levels & been reduced Lescano, have or quality behaviour fur grooming reduced in causing infection endoparasitic with condition, and mites between with associations niche Positive spatial a by for competition caused in liver be the with upon infected impacts (Sk´arkov´a, experimentally Any parenchyma mice 1986). while 1996), man, muriviperae Sarcocystis microparasite the another. of to Prevalence beneficial be & may Lee competition parasite niche D. one antagonistic by (T. While for grounds embedded of the unclear. stage while and is infective unlikely, lining perhaps species gut is these niche the physical between whereas of a for 1980), space competition Croll, intraepithelial that & the Panesar from 1978; nutrients Wright, obtains it 1980), While species. isce rmtegsritsia rc,cifce niiul a ihrbd asi ohinstances, both in mass body higher had individuals coinfected tract, gastrointestinal the from dissected ) eoycsmaniculatus Peromyscus T.muris hwdapstv soito ihbt esi h u and fur the in fleas both with association positive a showed ) .muris T. and .hepaticum C. 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T. ui matwl-hrceie hne oimn tt in state immune to changes well-characterised impart muris sngtvl soitdwt neto with infection with associated negatively is .obvelata S. n mature and 7 S.dispersa ol niaearl o coaaie sidctr of indicators as ectoparasites for role a indicate could r yial on rei h ue,meaning lumen, the in free found typically are .hepaticum C. hwhsoahlgclcagsi h liver the in changes histopathological show aoimspp. Calodium .dispersa S. oolsasp. Toxoplasma xs nsprt ras the organs, separate in exist eni ol potentially could it mean rfre oas to (referred Qerz il Papa, Viel, (Queiroz, .hepaticum C. Capillaria Myodes . Posted on Authorea 22 Jan 2020 — CC BY 4.0 — https://doi.org/10.22541/au.157971041.10418309 — This a preprint and has not been peer reviewed. Data may be preliminary. eclr .R,Besa .S,Bs,P . on .A . osc,E . ercs .S,...Jolles, . . . 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(8.e2395), , 73 International 6,3531– (6), rnsin Trends Ecology Posted on Authorea 22 Jan 2020 — CC BY 4.0 — https://doi.org/10.22541/au.157971041.10418309 — This a preprint and has not been peer reviewed. Data may be preliminary. iue1. Figure FIGURES 4017(01)00556-8 fec ro niae ed rmtepeitrvral oteotu ftemdl hcns fteline the of Thickness 1. model. Table the association. of the output ***p[?]0.001 of the **p[?]0.01, direction value to *p[?]0.05, The ratio variable odds predictor association. the the red positive to from Blunt, indicate proportional leads intensity. arrows is indicates or pointed values) arrow black, each log and (representing of association abundance negative parasite indicate appropriate, arrows where and prevalence, of 1. Fig ectopar- of characterization molecular toward Progress (2001). immunity. J. host F. of Alarcon-Chaidez, modulation & asite Laboratory K., Albino the S. in Wikel, muris Trichuris Scotland. of May, Stages Larval of Early Isle the the of Mouse. 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(8), ewr fpstv n eaieascain ewe aaieseis ae ngnrllna models linear general on based species, parasite between associations negative and positive of Network ora fHelminthology of Journal , 225 3,4948 o:10.1111/j.1469-7998.1991.tb03828.x doi: 449–468. (3), , 43 eeiayParasitology Veterinary 34,4746 o:10.1017/S0022149X00004995 doi: 427/436. (3/4), 13 , 101 Parasitology 34,2527 o:10.1016/S0304- doi: 275–287. (3–4), , 135 7,7771 doi: 767–781. (7), ucinlEcology Functional Science , 330 Posted on Authorea 22 Jan 2020 — CC BY 4.0 — https://doi.org/10.22541/au.157971041.10418309 — This a preprint and has not been peer reviewed. Data may be preliminary. aaieNceCaePrevalence Clade Niche muris Trichuris Parasite Helminth Gastrointestinal Helminth Gastrointestinal eaoa3.3 iet–eggs – Direct 30.33% Nematoda embryonate Eggs 46.17% Nematoda 14 2007) (Flynn, faeces. host with eggs release and colon and caecum the of lumen the in reside adults and host, the in hatch Eggs water. or food contaminated from or perineum, the from host another by ingested are and host, of region perianal on embryonate 1969) Wakelin, 1980; Croll, & Panesar 2007; (Flynn, faeces. host the with eggs release and epithelium in embedded remain worms Adult maturity. reaching until moult and caecum, and colon of mucosa in establish larvae stage First ingestion. upon hatch and environment in Summary Cycle Life Posted on Authorea 22 Jan 2020 — CC BY 4.0 — https://doi.org/10.22541/au.157971041.10418309 — This a preprint and has not been peer reviewed. Data may be preliminary. atnlasp. Bartonella Prevalence Clade Niche Parasite Microparasite Bloodborne embryonate Eggs 58.23% Nematoda Helminth Hepatic atra4.2 Transmission 47.12% Bacteria 15 2012) Dehio, & (Harms ectoparasites. arthropod the by bloodfeeding later during occurs transmission Further intracellularly. reproduce and erythrocytes invade they where bloodstream, the enter bacteria The faeces. vector’s the from potentially ticks), and fleas (typically vector arthropod an by host the on feeding during occurs 2009) Lee, W. C. 2007; (Flynn, body. host the of decomposition or necrophagy via death host’s the upon released are Eggs liver. the in females gravid leaving reproduce, and adults into develop Larvae system. portal hepatic the via liver the enter and mucosa the penetrate larvae Infective ingestion. following intestine the in hatch and environment in Summary Cycle Life Posted on Authorea 22 Jan 2020 — CC BY 4.0 — https://doi.org/10.22541/au.157971041.10418309 — This a preprint and has not been peer reviewed. Data may be preliminary. rmiuasp. Trombicula sp., Myobia sp., Radfordia fasciatus Nosopsyllus Prevalence Clade Niche dispersa Sarcocystis Parasite ectoparasite Arthropod ectoparasite Arthropod Microparasite Bloodborne ie Aai 44%Prst gshatch eggs Parasite 84.42% (Acari) Mites (Siphonaptera) Fleas 6.90% Ampicomplexa 16 41%Eg r adon laid are Eggs 34.15% esrt,1977) Weisbroth, & Friedman 2007; (Flynn, hosts. between contact direct through occurs transmission and mice, of the fur the in occurs Reproduction adults. become to stages nymph and larval through develop they where host, the on 2007) (Flynn, occurs. reproduction where detected, is movement when host the of fur the onto jump and pupate, and moult Larvae hatch. they where environment the into fall and host, the of fur the Sk´arkov´a, 1986) Kol´aˇrov´a, 2007; 1986; (Flynn, owls. by host the of predation following cycle life their complete and tissue, skeletal in develop cysts Bradyzoite blood. the through disperse which sporozoites, ingesting upon infected becomes and host, intermediate M.musculus Summary Cycle Life sthe is Posted on Authorea 22 Jan 2020 — CC BY 4.0 — https://doi.org/10.22541/au.157971041.10418309 — This a preprint and has not been peer reviewed. Data may be preliminary. cosaltapn esosadasmayo h aaieslf cycle. life parasite’s the of summary a and sessions trapping all across 1. Table al falprstsdtce in detected parasites all of Table M.m.domesticus 17 nteIl fMy nldn eoddprevalence recorded including May, of Isle the on