Posted on Authorea 16 Aug 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.158679916.68461360/v2 | This a preprint and has not been peer reviewed. Data may be preliminary. rwn ubro tde lorvae htbtcmuiisaefre yseismr closely-related more species by formed are communities bat Rautenbach, that remarkably they revealed & sharing as also (Aldridge and group, studies characteristics guilds, a of echolocation feeding such number and same represent growing morphological the bats as Mancina to Echolocating such when pertaining 1987; place. or traits species species same functional numerous cryptic the similar of morphologically in assemblages and dimensions been coexist form elusive niche have rare, forms can ecological processes with related Studying additional working many closely 2009). when but many (Holt, im- especially 1974), most concept challenging, (Schoener, The niche be use Hutchinsonian 1957). can food this (Hutchinson, –known and niche complete habitat resources ecological co-occurring to available to their in described linked limiting of partitioning are dimensions of Resource several ones use competition. along portant differentiate corol- and achieved limited a a overlap be that As have niche can positing reduces 1960). to species ecology (Hardin, that likely use partitioning– in are similar resource principle sympatry their as on central in relying a living species two is species of lary, law, coexistence Gause’s stable the or prevent resources principle, exclusion competitive The Introduction understand fully to samples extensive of need and third, the grounds the highlight while feeding results preferences, that These habitat species foraging generalist. coexistence. distinct bat phylogenetic more most species each their was the of by exhibited species predicted menu species distantly-related Furthermore, As the bat more moths. habitat-driven. related from abundant primarily most-closely inferred summer was temporarily two during we partitioning the few overlap niche relationships, on species, niche that feed prey higher suggesting to showed of differed behavior they techniques traits hunting spring, hunting their small ecological during switched levels. the a prey species inter-specific of such considering bat and array by at intra- three distinct both Even all a at when on niche fall, metabarcoding. trophic fed and their bats DNA of different genus using variation the the using spatial (April-October) while and of details season Indeed, seasonal species studied unprecedented strong breeding identified we bat with entire we Here, closely-related scale, their accessed geographic phylogenetically unexplored. throughout be vastly and and can remain phenotypically sympatry partitioning niche three of resource of trophic of traits diet their mechanisms functional differential underlying Although similar the the highly prey, sharing guilds. of species feeding identification multiple genetic of identical insectivorous assemblages to instance, local For allow pertaining form to principle. typically order exclusion and in Hemisphere competitive dimensions Northern this ecological the challenge some systems from along biological bats differ many must Yet, species coexistence. co-occurring stable of their niche the that postulates theory Ecological Abstract 2021 16, August 2 1 Andriollo Tommy metabarcoding bat DNA cryptic from of inferred habits species feeding differential everyone: for Food uemdhsor auel el il eGeneve de Ville la Li`egeUniversit´e de de naturelle d’histoire Museum tal. et 1 02 Roswag 2012; , oa Michaux Johan , tal. et 2 n aulRuedi Manuel and , 08;Shize ak,20;Vesterinen 2001; Kalko, & Schnitzler 2018a; , 1 1 Plecotus tal. et 08.A 2018). , na area an in Posted on Authorea 16 Aug 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.158679916.68461360/v2 | This a preprint and has not been peer reviewed. Data may be preliminary. edsge hssuybsdo apigrgm oeigteetr eido ciiyo h asin bats the of the activity of variation of seasonal period and entire spatial seasonally, the the vary document covering to to regime known take was are would sampling goal diversity groups first a and prey Our on abundance particular (April-October). based insect on roosts Since study maternity specialization year. this preferred while the close in designed spring), of availability in when we in periods established wider typically other are be moths during would species bats place (for three of low all niche trophic of is the colonies resources that multiple (Gilli´eron hypothesized grounds where we theory, feeding sympatry foraging overlapping of exploit studied area we potentially bats, unique and long-eared proximity a similar highly in the among diet mediated their is competition interspecific how understand To (Alberdi (Gilli´eron occur not region Geneva does the in macrobullaris or P. 2015), Lemaire, al. & Dinaric et the (Arthur in Prealps areas wide French across the species in both coexistence of (Tvrtkovi´c co-occurrence stable the Alps their by preventing contradicted Rutishauser Ashrafi is niche, observation 2018; 2010; ecological this Aizpurua, However, same (Mattei-Roesli, & the scale occupy (Alberdi they regional sympatry (Ashrafi that niche the suggested trophic authors at same several the distributions for parapatric compete essentially taxa of latter (Alberdi diet the moths the that that tympanate indicated on remains extensively feed (Ashrafi orders to of insect diet known the are Whereas 1997). bats Vaughan, long-eared Indeed, 2001). Ashrafi may they 2018; Dietrich (Andriollo that Ruedi, 2015; al. similar hybridize & (Barataud, so et not (Andriollo characteristics are do field species call three but the echolocation all in building, identical Morphologically, identify trios. same to in the found challenging in rarely be more including much vicinity, are close and 2018) in coexist (genus can species species, species bat sister long-eared closely-related austriacus three are P. involving two system (Arrizabalaga-Escudero Europe: bats biological to Western a prey applied here their study explored case of We a traits in functional species sibling or two al. habitat of et to ecology the consumers disentangling multiple (Razgour help of categories ecological characteristics different relate (Qu´em´er´e to to assigned exist were Chang species (e.g., analyses prey species once prey al. (Spitz predators singular itself et of or prey pairs groups the of prey of preferences ecology major the about from considerations derived al. informal usually on is relied information authors no (e.g., or strategies little feeding and specialist Aldasoro from (e.g., Sato 2015; generalist species between distinguish food interacting to of for allows ecology feeding breadth disentangle niche to Lopes used trophic classically the are niche trophic species: the of measurements main Two unclear. identi- remain taxonomic still in bats resolution in unprecedented coexistence provide which techniques, (Alberdi metabarcoding fications (Salinas-Ramos of analyses advent Pres- diet the 2014; on from Stevens, relied assemblages & species (Patrick such underdispersion within phylogenetic called mechanism a ley chance, by expected than 09 Salinas-Ramos 2019; , tal. et 02.Ti yohsso opttv xlso sas hlegdb h atta h itiuinof distribution the that fact the by challenged also is exclusion competitive of hypothesis This 2012). , 2018). , 96 ugsigta l he pce hr iia oaigbhvo swl Shize Kalko, & (Schnitzler well as behavior foraging similar share species three all that suggesting 1996) , 01 Roswag 2011; , tal. et 08 Riedinger 2018; , tal. et tal. et 05 Salinas-Ramos 2015; , smr itnl eae (Juste related distantly more is , 03 Spitz 2013; , 08.Hwvr uheooia nie osdrec ryseisa ipersuc type, resource simple a as species prey each consider indices ecological such However, 2018). , tal. et ssilcnndt h ihreeain nteesenpr fisrne where range, its of part eastern the in elevations higher the to confined still is tal. et tal. et tal. et 05,i osc (Courtois Corsica in 2005), , 09 lr,21) ept hs ehia dacs ehnssmdaigspecies mediating mechanisms advances, technical these Despite 2014). Clare, 2019; , tal. et tal. et tal. et 08) e,mr ohsiae prahsrligo utvraestatistical multivariate on relying approaches sophisticated more Yet, 2018b). , 01 ot,21;Razgour 2011; Motte, 2011; , tal. et 2014). , 05 Vesterinen 2015; , 03.Tems omnapoc ocaatrz neseiccompetition interspecific characterize to approach common most The 2013). , 04.Tog rmsn,teruei tl iie n o ntneddnot did instance for and limited still is use their promising, Though 2014). , tal. et .austriacus P. .auritus P. 05,wiedeaynceoelpmaue h ee fcompetition of level the measures overlap niche dietary while 2015), , tal. et 09 Chang 2019; , a emr ies si nldsas aype rmother from prey many also includes it as diverse more be can tal. et tal. et tal. et and tal. et 04 Spitzenberger 2004; , 01 Dietrich 2011; , 08,wieohr sdsml et oass habitat assess to tests simple used others while 2018), , 2 .macrobullaris P. tal. et 01,i h yees(Alberdi Pyrenees the in 2011), , .auritus P. 01,mcohsooia dnicto fprey of identification micro-histological 2011), , tal. et tal. et 09 Kartzinel 2019; , tal. et and tal. et 01.A hs w pce exhibit species two these As 2011). , tal. et antb ieetae,suggesting differentiated, be cannot tal. et tal. et .macrobullaris P. 06 n igsae(Entwistle shape wing and 2006) , 06 Rutishauser 2006; , tal. et 00.Sc tde benefitted studies Such 2020). , 00.Te lohv nearly have also They 2010). , 05.Floigteoptimal the Following 2015). , 06.Piso hs three these of Pairs 2006). , tal. et tal. et Plecotus 05 Rutishauser 2015; , tal. et 05 Leray 2015; , hl h third, the while , tal. et 04.Some 2014). , curn in occurring ) tal. et .austriacus P. tal. et tal. et 04,in 2014), , 2012), , 2012). , 2012; , tal. et tal. et et , , Posted on Authorea 16 Aug 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.158679916.68461360/v2 | This a preprint and has not been peer reviewed. Data may be preliminary. osrcin eunigo C rdcs eutpeigo a aa lseigo eune nMOTUs Andriollo in in detailed sequences as of out Library clustering carried arthropods. data, all of raw were COI identifications of assignation this species-level rely demultiplexing taxonomic lacking, get combinations their products, are to primer and PCR marker databases alternative efficient of sequence most As sequencing reference the construction, extensive species. also which three currently for those is 16S) barcode from (typically diet markers the different on of (Zeale measurement bats (Clarke affect long-eared biases severely amplification by Zeale some in consumed detailed exhibit prey setup may PCR they a bats, bp- and al. insectivorous 157 primers of A ZBJ wells. using diet adjacent amplified the that was among and gene contamination barcode al. as procedures COI et such laboratory the bias followed of during systematical fragment protocol any occurred long prevent extraction cross-contamination to DNA plates major The sequencing (Taberlet no individual. limited that was single ensure tag-jumping 1). a to Figure of in used Deagle production 8 were (wPOO; and fecal occurrence 6 the prey issued 1, of were to (numbered Zeale samples percentage equivalent species individual calculate bat These are to different “individual date. used and a six each were by for extracted samples each pellets) also individual totally (1-3 occupied we These guano colonies samples, This of neighboring community individual. mg three the 8 particular from Besides about a animals. and in the colonies of consisting animals maternity capture preference samples” by disturbing to without hunting consumed need sampling the spectrum the repetitive allowed prey without reflecting Such collection weeks. overall therefore guano than two the of samples during rather design community represent colony colony, unobtrusive Such aggregate to a given collected. an from likely guano a members in more of several from consisting therefore by bulk samples” are produced the “community samples pellets from extracted of community taken we collection pellets locations, random guano a 9 of represent the mg and 60 dates about 11 of the of each For identification (from 1). autumn prey Figure and and hibernacula; reared) to sequencing are disperse Extraction, pups adults when and weaned mid-August, are mid-June, to to juveniles mid-April mid-June when (from (from mid-October, spring covers i.e. summer to timespan bats, This mid-August born), temperate mid-October. of are to typical pups activity mid-April of from before seasons week indicating major other (Ashrafi three every other, territory the sampled each same notably and from the roost km each in 2 under feed within placed potentially found could were Preatoni colonies species species 2015; all three all the that from of bats suggested bats colonies that bats long-eared closest brown the The by of colonies). occupied bats (two identifications long-eared and genetic grey 2018) Initial by Ruedi, km-radius colonies), 1). & 18 (Figure an (Andriollo within region monospecific established Geneva were bats long-eared the of colonies in breeding area nine from samples guano collected We sampling Feces Methods and Material occurring aim partitioning final niche Our trophic diet. species. their of bat in mechanisms elusive of identified the and strategies species sympatric revealing feeding prey three by the of these coexistence the indirectly hundreds among species identify infer of understand to to traits was technique to ecological goal was metabarcoding the second high-resolution using The a predators competition. with these interspecific and of simultaneously periods species crucial three all of diet 06 Elbrecht 2016; , 21) lhuhteepieswr pcfial eindt mlf ag ag fatrpd from arthropods of range large a amplify to designed specifically were primers these Although (2011). tal. et 21)wt oictost mrv il sdsrbdi Andriollo in described as yield improve to modifications with (2011) tal. et tal. et 01.Drn h er21,alfehgaowscletdfo la ae sheet paper clean from collected was guano fresh all 2015, year the During 2011). , 09 Tournayre 2019; , .austriacus P. tal. et 08.Adtoal,alDAetat eernol lcdinto placed randomly were extracts DNA all Additionally, 2018). , tal. et tal. et toclne)o yapn ogerdbats long-eared alpine by or colonies) (two 09.Hwvr ste okpriual elwt the with well particularly work they as However, 2019). , 01,i em nieyta uhptnilba would bias potential such that unlikely seems it 2011), , 3 tal. et tal. et tal. et (2019a). tal. et 03 Gilli´eron 2013; , lctsauritus Plecotus 21a.PRblanks PCR (2019a). 04 Elbrecht 2014; , .macrobullaris P. tal. et 2018) , tal. et (five et , Posted on Authorea 16 Aug 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.158679916.68461360/v2 | This a preprint and has not been peer reviewed. Data may be preliminary. hm nw ofe ngaobtntlkl ob ryduo ybt eg,mtsaddretdbeetles). dermestid and mites (e.g., bats by upon of preyed (21 be species to likely arthropod of represented not were they reference slugs, but as or existing of guano excluded any species rotifers), on were Two to and feed MOTUs match 46 to bacteria not further known algae, did A them) (fungi, (14%) produced dataset. contaminants 125 the total) environmental MOTUs, from in obvious these sequences removed were distinct Of samples and 1’349 (284 MOTUs. representing sequence 883 samples doi:10/XXX), DNA into Zenodo guano clustered on individual were deposited and that data (raw community reads both Illumina of 5’016’988 library sequenced The prey consumed of dataset Curated Dray 1995; Results Hochberg, p-values were & and models Benjamini permutations, these Braak (FDR; 9’999 ter of method 2008; performing rate outputs Legendre, discovery by and & false assessed out, (Dray the was errors carried by statistics I were adjusted table type were fourth-corner table linking inflated these linking avoid the of to the L, order Significance of and in 4) 6) species, (model significance (model prey the combined columns test to the (Dray to and traits species used of 2) ecological was bat and traits statistics (model the fourth-corner species ecological The bat containing analysis between species. the R, relationships bat RLQ containing each of matrices): an of Q, (or spectrum used prey tables issued, we the three is containing traits, following ecological sample the their category a of and which latter structure prey joint hunted This consumed the when the not. summarizes surfaces species, or solid bat night from the (Dol´edec flightless. gleaned at relate or to be flying diurnal must order was strictly In that either species woodlice are or prey they flies as the related night, syrphid mostly entomological whether by 1) as the (Table included such from traits ecological prey 1) also first binary characterizes we (Appendix 18 but diet, species into their habitat, sorted prey from was bats identified to information long-eared all This of 2). of species (Appendix preferences three literature the ecological of main habitats hunting the the gathered indirectly infer to order samples In prey among similarity the explaining of ones traits the presence Ecological identify the to on based attempt (PCA) overlap). an niche analysis in to component (Dray species linked principal ade4 scaling prey directly package simpler multidimensional (i.e., R all a the through of out in carried projected matrix implemented also were absence function We overlap (PCoA) 2007). niche analysis Dufour, Lawlor coordinates pairwise & by principal of proposed the algorithm matrices using (Gotelli randomization species, (MDS) 3 EcoSimR Stati- bat package RA 1959). three the the Morisita, the overlap using 1966; in niche (Horn, tested implemented whereas index was and 1968), overlap Morisita-Horn’s (1980) (Levins, niche the index using of Levins’ calculated significance the was stical using species measured of was pairs predator between each asymptotic of extremely of breadth with estimator Niche minimum dealing Chao2 when the used Colwell indices we 1987; 2013). other season, (Chao, and than richness species better each species (Andriollo for performs bats evaluated long-eared it Deagle spectrum of (wPOO; prey as that sample as preferred given such sample the diets was extracted diversified each in diet for identified weighted the species dataset, total prey describing the of in number occurrence of the percentage by as treated were species Prey niche trophic of Measurement .auritus P. tal. et eedsadda ela hywr etil eodr ryo h aai ete ae ythe by eaten beetles carabid the of prey secondary certainly were they as well as discarded were , 96 Legendre 1996; , eoea reticulatum Deroceras tal. et 97 sipeetdi h akg d4 hsodnto method ordination This ade4. package R the in implemented as 1997) , tal. et and 02 optdwt h otaeEtmtS910(Colwell, 9.1.0 EstimateS software the with computed 2012) , ro vulgaris Arion 4 tal. et tal. et rsn ntredsic ape rmroosts from samples distinct three in present , tal. et 09) oetmt h opeeeso the of completeness the estimate To 2019a). , 03.T iulz ih vrasamong overlaps niche visualize To 2013). , 04.Bt emtto fetr rows entire of permutation Both 2014). , tal. et 08.Ti ehdfor method This 2018). , tal. et tal. et 2014). , 2012). , Posted on Authorea 16 Aug 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.158679916.68461360/v2 | This a preprint and has not been peer reviewed. Data may be preliminary. ote16idvda ape nlzd(iue2) si xetdfrsc nqa apigeffort. sampling unequal such compared absolute for 3), In expected (Appendix is 3). samples as (Appendix community 2a), in analyzed 96 (Figure retrieved samples the analyzed were community samples in orders the individual identified arthropod 186 in were the recovered in species to proportions species prey similar bat less very species however, three predator and numbers, the one pattern spring, from of or from seasonal diet season ones same the one the The from than considerably varied Lepidoptera another. for dipterans more to 91% consumed exhibited and of invariably 80 proportion autumn (73, the and while species summer bat lepidopterans in long-eared estimates, gathered wPOO three samples to all According in prey species. austriacus preferred bat included P. each most species of the samples prey 54 far of by Only shared 20% represented than These bats. more these in 2a). all detected (Figure were of species detect diet to bat the albipuncta ( necessary three of Mythimna moths been diversity the have common extreme by would very the shared 66) nine stresses were of which MOTUs (instead richness, prey species species (12%) per total samples the individual of 220 95% potential to the 98 of 66% that of to indicated of 52 diet represent diet the only the numbers richness, these colonies, prey that neighboring suggest curves three accumulation species. the from bat extrapolations in long-eared gathered three samples the austriacus individual of P. of the diet samples on the fecal in focusing (Merz, in proportions When local comparable detected in in only represented were recorded were spiders) already orders most were arthropod and samples woodlice fecal scorpionflies, ans, these Jong in (de inventories identified (Blattodea, faunal (Andriollo orders species by national represented detected Most only or other were Hy- species. while 2012) rep- Trichoptera) Hemiptera, of most species, and handful respectively). the 17 Raphidioptera, MOTUs, a to expected, Psocodea, accidentally 193 14 as Orthoptera, or and included Mecoptera, and bats (392 Dermaptera, Neuroptera samples) by Diptera and individual targeted and Coleoptera and Lepidoptera be community menoptera, were to (both orders mm) spectrum insect (3 resented prey enough complete occur, large the do possibly In they is since spiders, it (Vesterinen dataset 3%, 14 as grooming this and kept included during in 7% consumed also Kaˇnuch also retained (2%, (e.g., was but were the level analyses springtail Woodlice MOTUs), to order guano The (668 species. taxonomically bat or insects springtail in identified family a rarely, mostly large were genus, in albeit were one MOTUs the occurred and prey distinct to which woodlice Hunted assigned these of four were of (46%) 1). ones 319 (88%) Appendix remaining retained, 602 respectively; were the of MOTUs while total distinct level, A 687 species individual of the sample. and total guano colonies a nine single for colonies, samples community three the for including prey, samples consumed of dataset curated the In bats long-eared of diet Molecular samples. guano small particular these on these feed in from mostly identified absent would these were pipistrelles species that whereas that prey of bats), 1993), indicated long-.eared most inspection however, of Indeed, visual (typical (Swift samples, flies A dipterans prey. large fecal or exogenous contaminated bats. moth to were potentially long-eared samples minimally five with contributed these occasionally bats in roosted non-targeted have composition may prey species the these that Galan suggesting also (see pipistrellus species prey Pipistrellus actual their not bats, tal. et and and 09) neetnl,anme fdunlo o-yn rhoos(awg,orthopter- (earwigs, arthropods non-flying or diurnal of number a Interestingly, 2019b). , tal. et .macrobullaris P. .macrobullaris P. , 95 n eoie nlrebels(evn&Lbi,20;Rbno Stebbings, & Robinson 2008; Libois, & (Kervyn beetles large on serotines and 1985) , yhmapallens Mythimna and .auritus P. goi exclamationis Agrotis peiu serotinus Eptesicus epciey,floe ydpeas(,1 n 5;Fgr c.Guano 2c). Figure 15%; and 13 (4, dipterans by followed respectively), , nldd30 7 n 5 ryseis epciey(iue2) Chao2 2a). (Figure respectively species, prey 157 and 171 300, included en h otudrsiae Fgr b.Teeetaoain also extrapolations These 2b). (Figure underestimated most the being tal. et , otapronuba Noctua 2013). , tal. et , a eetdi v fte24fclsmlsanalyzed samples fecal 284 the of five in detected was , goi ipsilon Agrotis tal. et tal. et 04,o eelkl eetclnzr o h country the for colonizers recent likely were or 2014), , 5 05 Leelapaibul 2005; , 08.Uepcel,teDAo w a species, bat two of DNA the Unexpectedly, 2018). , , oohl noctuella Nomophila , uorpagamma Autographa tal. et 09 Rydell 2009; , and .auritus P. etac-nigrum Xestia , oldiaambigua Hoplodrina tal. et hl other while , .auritus P. .auritus P. 2016). , that ) , , , Posted on Authorea 16 Aug 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.158679916.68461360/v2 | This a preprint and has not been peer reviewed. Data may be preliminary. natrpd iigi pnhbtt n ndyadmutiosaes(.. ces lps.Finally, habitats slopes). closed screes, of typical (e.g., prey to areas so negatively mountainous and and species, prey dry 1). ubiquitous (Table to in associated and positively only habitats was open in austriacus P. living arthropods on egrw)rte hnfo pno onanu ra sc smaosadlws lps cesand screes (Table prey slopes, of lawns, flightlessness or and diurnality meadows to as associated significantly (such and areas positively mountainous 1). also bat was or one and open from areas) differed from rocky 0.05) than < 2). few rather (p (Appendix for significantly hedgerows) literature or prey another. the family of to in traits or found species ecological order be that the the could indicated to to statistics traits only identified Fourth-corner ecological identified MOTUs specific (88%) MOTUs no 605 either which for were for estimated items species be non-evaluated prey could The arthropods of level. consumed species traits the ecological of preferences the Habitat and species predator between Relationship 0.01). pair < (P The autumn. and and 0.62). seasons: (0.28) over- summer three niche during the across those trophic value (mean than of lower Measures indeed 4b). were ritus (Figure spring season during austriacus and species P. species bat among among unequal lap 4). were (Appendix similarities autumn Dietary in collected samples three guano in the as retrieved by such eaten noctuids summer frequently noctuella whereas were samples, similarities that blanda These spring phenology drina 4). the seasonal (Appendix of beetle marked samples characteristic June a of the were with instance, composition For prey species species. dispersed the insect bat more on by was conducted colony driven the PCA each that were of from the appears diet those in the it made recovered particularly when comparisons 4a), also representation, spring same-species (Figure MDS than in season obvious roosts the given less bat a in however, within different was, similar This the more seasons. were among across bats overlap long-eared niche of of species three measure the to According overlap niche species) trophic per in colony Variation analyzed one in of higher effort also same was a not of (results period with colonies samples sampling (22.7 of (i.e. community the number samples or higher all individual individual the a throughout on for to of restricted due measured focused amplitude only we index a specific not whether Levins’ consumed is Regarding or predators result 3a), This the seasons. (Figure shown). that season fall implies the and of This summer spective the decreas- species. actually during bat 2010). was arthropods three diet, breadth (Altermatt, niche of the consumed summer autumn 3b), of range for in (Figure and increase year level taxonomic peak spring an family the abundance during the to throughout three and at low due diversity identified ing all mostly indeed prey for species was was considering however, changes whose when bat pattern, seasonal moths, Indeed, long-eared This of geometrid each 3a). pattern and of (Figure same noctuid breadth summer the niche during exhibited higher trophic species and The prey of species. terms bat in diversity dietary breadth The niche trophic the of Variation .macrobullaris P. h ags auso rpi ih vra eeosre between observed were overlap niche trophic of values largest The . ± .auritus P. .)or 8.4) n h hepialid the and .macrobullaris P. eems bnati ape olce uigsme.Telt-umrcrambid late-summer The summer. during collected samples in abundant most were hwdn akdpeeec o pcaie ry n orhcre ttsisidctdta it that indicated statistics fourth-corner and prey, specialized for preference marked no showed and .macrobullaris P. ossetyehbtdabodrtohcncethan niche trophic broader a exhibited consistently .macrobullaris P. .auritus P. xiie poiepeeecsi em ffeighbtt,a tpee preferentially preyed it as habitats, feeding of terms in preferences opposite exhibited roi sylvina Triodia 03) l auso vra eesaitclyhge hnepce ychance by expected than higher statistically were overlap of values All (0.30). eddt osm ryise rmlwadcutrdhbtt (woodlands, habitats cluttered lowland from issued prey consume to tended (22.0 ytmtclyehbtdgetrnceoelpta hyddwith did they than overlap niche greater exhibited systematically ± 9.5). hztou aestivus Rhizotrogus Plasy 04 Robineau 2014; (Poltavsky, .auritus P. .auritus P. 6 xiie uhlwrnceoelpwith overlap niche lower much exhibited .auritus P. hssaoa aito fde iiaiywas similarity diet of variation seasonal This . n h pigmoth spring the and (34.4 .austriacus P. .auritus P. tal. et .austriacus P. ± 07 eeacrigymostly accordingly were 2007) , .)cmae to compared 9.2) omatrapezina Cosmia vlae Fgr ) sthe as 1), (Figure evaluated and oshlelslupulinus Korscheltellus .macrobullaris P. and .macrobullaris P. .austriacus P. .austriacus P. and Nomophila Hoplo- .au- P. irre- Posted on Authorea 16 Aug 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.158679916.68461360/v2 | This a preprint and has not been peer reviewed. Data may be preliminary. fho ohseisapae ntemn faltreln-ae a pce Fgr a u nlddseveral included but 2a) (Figure species bats bat a long-eared these Only three 1). of all (Appendix diet of feces menu the their the in that in identified appeared indicated MOTUs species prey arthropod moth Despite distinct of of resemblance. 680 fifth identification phenotypic over high species-level with corroborates diversified, their moths, much also from was observation on predicted this specialization strategies menu, high hunting specialized ones of their highly similarities this the overall share with high bats (Zeale the long-eared consistent primers all selected were Ashrafi As the assessment. 2002; of than proportions diet biases (Andreas, less recovery These feces that represented 2c). of suggesting groups 1990), analyses (Figure (Alberdi arthropod micro-histological detected studies other and (73- metabarcoding species all 2018) bulk while previous prey the level (4-15%), in represent the ordinal recovered dipterans lepidopterans the of by that to percent followed here identified confirmed diet, four are We their moths. resources of tympanate prey 91%) of when hunters diets specialized similar are insectivorous very of composition exhibit diet species the (Vesterinen studying many for essential since be diet bats, to proven their been have of techniques High-resolution components bats these major of share strategies bats hunting and Sympatric niche use trophic traits. habitat their ecological in partition prey also differences of but Furthermore, analyses diet, year. by menu their experimental the highlighted their of this are of resolution most from periods high share obtained bats critical results with closely-related Experimental at access activity. these to of that and season indicate vicinity, entire design an geographic throughout narrow arthropods Emrich in of living (e.g., bats carried species long-eared been distantly-related cryptic have on species focused sympatric al. either of all diet the but partitioning Vesterinen unravel niche metabarcoding, trophic to of DNA attempts mechanisms using Previous investigate to resolution. models fine-grained ideal with constitute species cryptic Co-occurring Discussion noctuids the including screes, and slopes sunny and or lawns and meadows and dry as such areas nous of diet the Finally, for typical brassicae species Mamestra Moth areas. urban in found of autumnalis diet the in chloropid identified prey ubiquitous The (e.g., species to aprilina Dichonia associated (e.g., species detected also Moth and were bats. tachinids by Forest associated woodlands. also eaten was in diet living Its flies. tipulids albicans syrphid several as and such arthropods arthropods neuropterans diurnal consumed to or the that psocopterans) earwigs, indicated 5) beetles, ground (Appendix analysis with RLQ associated The the most of examination detailed more A species prey Typical 08 Razgour 2018; , rohl domestica Bryophila yhoe obscuraria Nychiodes prptr brumata Operophtera and huaoyanotata Thaumatomyia tal. et ulltn etseisfrctl n oss n h calliphorid the and horses, and cattle for species pest pullulating a , tal. et lcradelecta Eloceria rhp xylosteana Archips 08,o egahclydsato ielmtdsmlns(.. Arrizabalaga-Escudero (e.g., samplings time-limited or geographically-distant on 2018), , , 08.A xetd hswspriual rei h aeo ogerdbt,which bats, long-eared of case the in true particularly was this expected, As 2018). , tal. et .macrobullaris P. iicacitrago Tiliacea htcnb on navreyo pnhabitats. open of variety a in found be can that .auritus P. 01.Tepeetsuyi h rto t idt hrceietede fthree of diet the characterize to kind its of first the is study present The 2011). , n h geometrids the and , . omnyfudi odad rubnaes otis(e.g., noctuids areas, urban or woodlands in found commonly ) ,atog hs aaii iscudb eodr rycnandi ohlarvae moth in contained prey secondary be could flies parasitic these although ), and htlvsi giutrlfils edw n rslns h muscid the grasslands, and meadows fields, agricultural in lives that aclsmlsicue aynnflig(pdr,wolc,cockroaches, woodlice, (spiders, non-flying many included samples faecal and a hrceie ymn ryseistpclyascae omountai- to associated typically species prey many by characterized was yi pomonella Cydia Amphipyra .auritus P. .austriacus P. npo furvata Gnophos .austriacus P. p. yia fdcdoswolns n ettortricid pest and woodlands, deciduous of typical spp.) 7 hiigi rhrso woodlands. or orchards in thriving ) nlddgoerd (e.g., geometrids included tal. et nlddfle rmcliae ad,sc sthe as such lands, cultivated from flies included nlddtenoctuids the included 02 Razgour 2012; , , eitl chrysoprasaria Hemistola tal. et tal. et 01 a iie mato the on impact limited a had 2011) , 01 ot,21;Robinson, 2011; Motte, 2011; , tal. et olnapediculata Pollenia aarn clavipalpis Caradrina apormabilineata Camptogramma 01 Vesterinen 2011; , , oim radicaria Horisme nrhamunda Anorthoa uo aquilina Euxoa tal. et Cyzenis notably 2013; , Musca tal. et and et , , Posted on Authorea 16 Aug 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.158679916.68461360/v2 | This a preprint and has not been peer reviewed. Data may be preliminary. nErp,adwt h togevrnetladtohcseilzto of specialization trophic bat of and long-eared distribution environmental three strong geographic the the wide with in the and observed with Europe, consistent breadths in was niche result trophic in this different while higher the was on which lights species, shed also results These habitats while closed grounds, preferences. to hunting habitat one distinct negative of a exploration species and the insects sister through ubiquitous two to The link positive 1). a (Table of was relationship diet from significant prey the only by that the consumed in found prey overrepresented we the were of particular, marked hedgerows preferences In of and species. existence bat the woodlands of three suggested that as the diet such of the habitats grounds of hunting analyses closed preferred RLQ the through in prey of differences habitats traits foraging ecological the differential Integrating studies be to comparative could related many it is in whereas bats, partitioning done these Niche been of has cycle as life scales, the time of narrower most partitioning covers on niche Arrizabalaga-Escudero regime (e.g., seasonal focusing sampling Such designs the phenologies. with if insect overlooked detected bats seasonal insectivorous year be marked all the only to strongly of generally can times with more other regions applies on at probably feed temperate resources constraint insect to partitioned from seasonal behavior of on this availability opportunistic suggest relied same when We they the 1996). i.e. whereas showed (Arlettaz, prey, exploitation, species abundant food sibling spring temporally over similar, of and in highly pressure observed locally range Other selective overlap limiting. diverse stronger niche more lowest more of is the result prey a hypothesis, a on partitioning as niche feed evolved autumn of likely to their predictions diversity appear from the absent of with species virtually Consistent terms were bat which in three beetles all both more including year, lower, 3), niche the diet. was Figure trophic of and availability during the 2 overlap. time resources prey Conversely, (Figure niche this invertebrates partitioning. food when in niche At unlimited spring, increase for abundance. nearly temporary during need and provide a the occurred may alleviating in mostly more insects temporarily resulting giving partitioning common time, species 2010) very of three (Altermatt, few, periods all seasons short that of latter abundant suggests menu few the months, pattern the during summer these This in highest during its appeared Instead, among diversity resources. at species different moth overlap also exploit moth the to is niche as bats in counterintuitive bats) these trophic notably somewhat to these increased of opportunities is of then relationship levels prey niche and This spring main understand bats, 4b). during to (the (Figure of lower autumn information was early species crucial overlap and provided niche three Trophic summer and bats. these seasons long-eared across of in variations partitioning menu strong exhibited the species in bat shifts parallel Despite seasonal season highly 4a). given is any (Figure within species partitioning similar bat Niche more within was than year bats, the rather long-eared of predator, of times the species various of three at identity all temporally sampled the of guano this behavior regardless the exploit Clare hunting of to 1996; opportunistic content behavior and (Arlettaz, prey plastic the feeding demands diet a energetic opportunistic their with their an Consistently of meet had 2020). to component species resource bat important trophic three an other abundant the by represent that hunted could indicating opportunistically and diet, be guild to feeding seem taxa different (Arrizabalaga-Escudero widespread from and bats abundant insectivorous These species. bat long-eared ( species noctuid common .macrobullaris P. tal. et nlddmr ryseislvn noe n ok rmutiosaes Habitat areas. mountainous or rocky and open in living species prey more included Agrotis tal. et .auritus P. 08.Teesae ryseiswr atclrycmo ntesummer the in common particularly were species prey shared These 2018). , 08 Roswag 2018; , .auritus P. spp., .austriacus P. Mythimna hncmae oteohrtobtseis(iue3) Indeed, 3a). (Figure species bat two other the to compared when and .macrobullaris P. tal. et spp., eeligbtentoetoeteepten,frwhich for patterns, extreme two those between lying were 8 2018b). , Noctua .austriacus P. p. htwr rqetyetnb l three all by eaten frequently were that spp.) perdt atto hi rpi niche trophic their partition to appeared a oegnrls ihn marked no with generalist more was .auritus P. .macrobullaris P. tal. et cosdvrehabitats diverse across .auritus P. 01 Cohen 2011; , (Alberdi whereas , tal. et tal. et , , Posted on Authorea 16 Aug 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.158679916.68461360/v2 | This a preprint and has not been peer reviewed. Data may be preliminary. iedffrn ogerdbtrot hogotteya,w nyrcre eae ig fbutterflies of of wings visits decayed eleven recorded the lepidopteran only during diurnal Yet, we of 2011). year, presence Motte, the 1991; the throughout Meineke, for roosts 1990; account bat ( Barataud, to long-eared (e.g., invoked different roosts classically nine bat been in has wings of behavior diet the gleaning in is The reported technique hunting rarely This was the 1991). prey Racey, demonstrated non-flying & experiments Captivity whereas (Anderson in flight. bats in common long-eared not more in (e.g., surfaces, of supposedly behavior solid taxa diet gleaning from diurnal all the the bats of in typically of by diet prominent importance gleaned of the more be in number significantly recovered to were A were expected they woodlice) 1). locomotion are but and (Table different prey bats, spiders bats of the beetles, the carabid species on of of (e.g., three also importance composition groups but non-flying the diet or investigated habitats, in flies) we hunting syrphid differences particular, on explain in information to Here provide bats. only type by not used does strategies techniques traits hunting feeding prey differential of analysis through The mediated are be bats could those partitioning Niche when even Arrizabalaga-Escudero bats, be 1999; insectivorous could (Arlettaz, niche European 2006). selection sympatric habitat-driven Racey, in habitat broadly & inferred that partitioning Nicholls and the the demonstrating similar Finally, without resource studies highly habitat. species for of elusive morphologically natural mechanism number of their major ecology growing in foraging a a them the corroborates track reconstructing also actually for or partitioning applied capture be to to need potential strong a thus 02.Adtoal,sxnuotrnseisfudi h un fln-ae aswr o itdamong listed not were bats (Andriollo long-eared province Geneva of the guano area in (Andriollo the the occur from region in nocturnal to known well-studied found known species local this moth 26 species in for the all the neuropteran bats occurrences of of long-eared six fifth new of Additionally, samplers a were diversity than 2012). that good dietary more 17 total retrieved prodigiously the including we be of 2b), sampled, (Figure portion to design small experimental appeared a our uncovered bats only long-eared we Although study, biodiversity. current monitoring the ecosystem In and assessment diversity (Olofsson for rodents Bats or insects spiders reflect as not such therefore animals might other by attics eaten in been retrieved have Wiklund butterfly could Bohmann but of (e.g., bats, remains pair by wing preyed primer that same suggest the sample we using Conversely, guano bats insectivorous each other in of recovered (Vesterinen lepidopterans species lepidopteran other al. 160 of of of analyses number molecular out high Previous the 1). despite (Appendix identified of was diet sequence the in remains butterfly corroborate also menu their observation, agricultural radiotracking from (typically, this time-consuming inferences and habitats explain indirect extensive species-poor on more to with based (Alberdi comparatively made conducted results data of habitats be These foraging exploitation can species. of observations to bat conclusion previous last due firm this be by no could lands) While it niche. suggest trophic we narrow surprisingly a aaae xs.I loilsrtsta ascnb sdt oio h rsneo economically-relevant of presence the monitor to Mazzi used 2012; be can suzukii reference bats Drosophila good that that as provided assessment, illustrates such biodiversity also species for tool It efficient exist. highly a databases is guano bat of metabarcoding Alberdi 2012; gasurticae Aglais 01.Ptnilapicto iscnb xldda ira eiotrn eercre ntediet the in recorded were lepidopterans diurnal as excluded be can bias amplification Potential 2011). , tal. et tal. et tal. et 08 iigi h aeattics. same the in living 2008) , tal. et Calliphora and 2017). , ao etada cnmccnenfrfutcosi aycutis(Calabria countries many in crops fruit for concern economic an and pest major a , 04 Ashrafi 2014; , 02 Preatoni 2012; , nci io Inachis and .auritus P. ntoocsosudrclne cuidby occupied colonies under occasions two in ) Lucilia Plecotus tal. et tal. et is etr fdsae o ua n ate n h le ri fly fruit alien the and cattle, and human for diseases of vectors flies, 01 Benda 2011; , hnin than a ofimdb h eaacdn prahsnen butterfly no since approach metabarcoding the by confirmed was 01 Razgour 2011; , tal. et Plecotus .austriacus P. 08,o oea l (Alberdi all at none or 2018), , 9 tal. et ee eoddasnl utry( butterfly single a recorded feces tal. et .macrobullaris P. 06,tewidespread the 2006), , tal. et Ades 02 aeo´,18;Bc,1995), Beck, Bauerov´a, 1982; 2002; (Andreas, 06 ole,21) ugsigaanthat again suggesting 2012), Hollier, 2016; , 01.Oridrc,RQapoc has approach RLQ indirect, Our 2011). , tal. et (Ashrafi 01 Vesterinen 2011; , .auritus P. .auritus P. tal. et .austriacus P. tal. et hs ryspecies prey These . 02 Razgour 2012; , tal. et 2011). , hspuiyof paucity This . rynspaphia Argynnis 09 Merz, 2019; , tal. et tal. et tal. et exhibited 2018). , 2018; , 2011; , tal. et et ) , Posted on Authorea 16 Aug 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.158679916.68461360/v2 | This a preprint and has not been peer reviewed. Data may be preliminary. lrde .&Ruebc,I 18) opooy cooainadrsuc attoigi insectivorous in partitioning resource and echolocation Morphology, (1987). I. bats. Rautenbach, & (2019). H. J. Aldridge, Aihartza, combination. primer & molecular U. through Goiti, bats horseshoe A., ONE among Arrizabalaga-Escudero, PLoS allocation U., dietary on Baroja, insight analysis. N., ecological Vallejo, Gaining diet T. I., for M. Garin, M., Gilbert, sequencing Aldasoro, & high-throughput M. using Nielsen, of C., pitfalls Lynggaard, bat S., Resources and Gopalakrishnan, the Promises distribution K., of (2019). the Bohmann, P. modelling O., shaping Aizpurua, niche factors A., the ecological Alberdi, Unveiling multi-scale (2014). using I. vertebrates, Garin, alpine & macrobullaris distributed J. widely Aihartza, O., of long-eared Aizpurua, mountain A., the Alberdi, of ecology foraging The (2012). J. nal.pone.0035692 Aihartza, & O. Aizpurua, bat I., Garin, A., Alberdi, cies led,A ipra .(2018). O. Aizpurua, & A. Alberdi, References also l’ TA financial de the Schmidheiny. manuscript. from Nature Lucie benefitted la the & study discussions de of G´en´erale This Ernst et fruitful Guiana. Direction version French l’Agriculture for the in de earlier Zinger from held school support Lucie an metabarcoding and DNA on Taberlet eighth comments the Pierre during Mariaux useful Marcon, Jean Eric provided Geneva), Fr´ed´eric Coissac, of (MHNG) thanks (University Eric Pawlowski Monod Boyer, Jan entomological Lionel Copenhagen), valuable and of provided (University statistical (MHNG) Alberdi on Antton Hollier expertise analyses. useful (Universit´e John provided de Pigneur Lyon) and St´ephane (CNRS, Lise-Marie and Dray Rapha¨el Lienhard (MHNG) possible. expertise. Covain Charles study Landry, data this raw Bernard Verelst. rendered Illumina Andr´eextraction. with Adrien (Universit´e Li`ege) who Eric Cauwet, helped and de buildings Lucie and work, of laboratory (alphabetically): Vallotton owners during Li`ege) sampling Laurent Sch¨onb¨achler, helped the Tardy, guano thank Cyril with Emmanuel also Rouco, us We Carlos helped who Pralong, people Janik the acknowledge gratefully We Acknowledgments with human of coexistence peaceful and very efforts conservation the promote areas. suppression At to to rural pest argument barns) to pests. in effective contribution or bats these an their attics that as of (typically hope used we buildings consumption be landscapes, man-made can agricultural common use in very intensively roosts maternity bats a their long-eared establish suggesting that respectively, considering samples, and considered diet noctuids of least, species their the 22% 57 of particular, of and fifth In total a 43, feces. a that their that noteworthy in and is identified landscapes, ipsilon it were agricultural data, pests in our found agricultural from typically as quantified insects be of cannot composed suppression was pest (Baroja to species bats pest eared services. spawning on ecosystem (Boyles feed control important to insect provide diet to their shift known Cohen selectively 2019; are particular bats in biomonitoring, species Insectivorous and assessment diversity Besides , lctsmacrobullaris Plecotus 50 ora fAia Ecology Animal of Journal and , 98,2–3 o:10.1093/mspecies/sey003 doi: 26–33. (958), , 19 , 14 eioep armigera Helicoverpa tal. et . 2,3738 o:10.1111/1755-0998.12960 doi: 327–348. (2), rnir nZoology in Frontiers 7,e208.di 10.1371/journal.pone.0220081 doi: e0220081. (7), 00 Kolkert 2020; , tal. et 01 Brown 2011; , eeldwt N mini-barcodes. DNA with revealed , tal. et 56 , htaekont as ao aae ocoswr eetdi 61, in detected were crops to damages major cause to known are that lctsmacrobullaris Plecotus 3,763–778. (3), 11 00 n r upsdt rvn ao giutrllse through losses agricultural major prevent to supposed are and 2020) , 1,7.di 10.1186/s12983-014-0077-6 doi: 77. (1), tal. et 05 Kunz 2015; , 10 Ciotr:Vespertilionidae). (Chiroptera: tal. et LSONE PLoS 01.Atog h otiuino long- of contribution the Although 2011). , ttd e`v n h Fondation the Gen`eve and de Etat ´ , 7 4,e59.di 10.1371/jour- doi: e35692. (4), goi exclamationis Agrotis oeua Ecology Molecular amla Spe- Mammalian , Plecotus Agrotis tal. et , Posted on Authorea 16 Aug 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.158679916.68461360/v2 | This a preprint and has not been peer reviewed. Data may be preliminary. aaad .(05.Aosi clg fErpa as pce,ietfiain td fterhbtt and habitats their of study Paris identification, 8, Biodiversit´e, naturelle, vol. species, & d’histoire bats: Inventaires European Collection of behaviour. ecology foraging Acoustic (2015). M. Barataud, et 1758) (Linnaeus, (1990). M. selection Barataud, Habitat (2013). F. Bontadina, Conservation & and R. Biodiversity Arlettaz, K., cryptic M. Obrist, three of K., of Ecker, partitioning M., niche Rutishauser, Trophic S., Ashrafi, (2011). conservation. F. for Bontadina, implications & Switzerland: R. in Research bats Arlettaz, long-eared M., of Rutishauser, species Alps. cryptic A., Swiss Beck, the S., around characters Ashrafi, species morphological bat Multiple long-eared (2010). cryptic R. of Arlettaz, 281 identification & ed.). Pavlini´c, field I. 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(3), (Chiroptera: , 54 Plecotus (1–2), Posted on Authorea 16 Aug 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.158679916.68461360/v2 | This a preprint and has not been peer reviewed. Data may be preliminary. iln,C,Vli,A,Fieg .&Jkbsn .(08.Rdn rdto nhbraigpaokand peacock Dietary hibernating on please: predation five, Rodent for (2008). Table S. Jakobsson, (2018). butterflies. & M. tortoiseshell M. T. Friberg, small Lilley, A., & Vallin, S. C., Wiklund, A. Blomberg, bats. E., boreal I. in A. DNA partitioning Puisto, fecal J., of sequencing E. ( Vesterinen, generation Bat Next Daubenton’s predator (2013). insectivorous N. Finland. widespread Wahlberg, Southwestern the & in of N. diversity V. dietary Laine, the reveals T., Lilley, J., E. Vesterinen, dutdpvle r ie sflos /+ P<00) / P<00) .. o significant. not n.s.: 0.05); < (P -/+ 0.01); < (P /+++ follows: as given are p-values adjusted 1. Table the performed JRM manuscript. TA final the Tables to sampling. contributed the JRM MR conducted TA and data. experiments the analyzed the TA designed work. and laboratory conceived TA MR Contributions Author Accessibility Data DNA for PCR Taxon-specific (2011). G. Jones, & C. D. faeces. Lees, bat A., 0998.2010.02920.x in L. G. prey Barker, arthropod K., R. barcoding Butlin, K., R. M. Zeale, 007-0465-4 • • arxo clgcltat o l eaacddpe pce spoie nteSpotn Information Supporting the in provided is species TBD references prey Zenodo bibliographic metabarcoded assignations: with all taxonomic file for and traits information ecological sampling of complete Matrix with prey of sequences DNA orhcre etbtenbtseisadeooia riso osmdpe.Sgicneof Significance prey. consumed of traits ecological and species bat between test Fourth-corner an ..()+++ +++ (-) +++ n.s. + (-) n.s. + (-) (-) n.s. (-) n.s. n.s. - (-) n.s. (-) n.s. (+) - (-) n.s. n.s. - (+) (-) n.s. n.s. + (-) (-) n.s. (+) n.s. (+) n.s. n.s. (-) n.s. (-) (-) (+) n.s. n.s. n.s. (-) (-) n.s. n.s. (-) (-) n.s. n.s. areas Mountainous (-) (-) - n.s. n.s. (-) Lawns n.s. (+) n.s. - Meadows (+) n.s. (+) Screes - n.s. + areas Dry areas Mesophilous (-)  n.s. Ubiquitous lands Cultivated +++ Lowlands +++ - habitats Semi-open areas Urban Shrublands +++ Hedgerows flightless or diurnal Exclusively forests Woodlands, habitats Closed clgcltat fprey of traits Ecological LSONE PLoS clg n Evolution and Ecology eairlEooyadSociobiology and Ecology Behavioral , 8 1) 818 o:10.1371/journal.pone.0082168 doi: e82168. (11), oeua clg Resources Ecology Molecular .artsP uticsP macrobullaris P. austriacus P. auritus P. rdtrPeao Predator Predator Predator , 8 17 2) 01–03.di 10.1002/ece3.4559 doi: 10914–10937. (22), , 62 , 11 3,3939 o:10.1007/s00265- doi: 379–389. (3), 2,2624 o:10.1111/j.1755- doi: 236–244. (2), ytsdaubentonii Myotis ) Posted on Authorea 16 Aug 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.158679916.68461360/v2 | This a preprint and has not been peer reviewed. Data may be preliminary. esrdtruhu he eid fteya mn h he ogerdbtseis o ahpairwise each For species. bat spring. during long-eared observed three are the overlap appea- among niche of species year values each prey the lowest and of of the color phenology comparison, periods different observed a three the by throughout represent of represented barplots measured diet is inset the species Three in bat symbol. ring Each different seasons. a across by and season colonies different in lected among measure) (a) (Morisita-Horn overlap seasons. niche three Trophic the 4. Figure for (wPOO). measured data index) occurrence (Levins’ weighted level). as variation considered breadth and season. (a) niche species. and seasons. trophic bat different species the long-eared bat Seasonal in each bats 3. for of Figure diet species the species. prey in bat detected (a) found three species. of orders the indivi- bat for number arthropod three in detected the the richness detected indicating of species (wPOO) each prey curves occurrence colony the of one Switzerland. of (top in frequency southwestern diagram gathered inset in Euler their guano Geneva and The Lake of prey tubes). samples the of dual plastic near number area (small Total study 2. samples” the Figure “individual of view or general tubes) a plastic provides large left) by (symbolized samples” of long- colonies of ones colonies of orange nine Sappey; the of (5) of colonies and represent location region. circles Geneva geographic Blue the and (high). in panel) studied (upper bats dates eared Sampling 1. Figure captions Figure .macrobullaris P. ut-ietoa cln MS ftohcnceoelpcluae mn ea ape col- samples fecal among calculated overlap niche trophic of (MDS) scaling Multi-dimentional lps-ns - +++ +++ (-) n.s. (+) n.s. -  habitats Open Slopes Plecotus 8 atgyad()Lez h apigrgm ie ogte ihr“community either gather to aimed regime sampling L´eaz. The (9) and Cartigny (8) : asadcaatrsi fseicsmln periods. sampling specific of characteristic and bats opeedtst ihalpe tm etadietfidt h pce level species the to identified and kept items prey all with dataset, Complete .austriacus P. .auritus P. lvto ntempi hddfo ae(o)t akgrey dark to (low) pale from shaded is map the on Elevation rdtrPeao Predator Predator Predator 1 ain,()pn ui,()Colx 4 Presinge (4) Choulex, (3) Butin, pont (2) Satigny, (1) : 18 6 olxad()Hrac;prl nscolonies ones purple Hermance; (7) and Collex (6) : (b) ryietfidt h aiylvlol (Family only level family the to identified Prey Plecotus (b) xrpltdaccumulation Extrapolated (b) pce,clne and colonies species, rpi ih overlap niche Trophic Area-proportional (c) rprin of Proportions Posted on Authorea 16 Aug 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.158679916.68461360/v2 | This a preprint and has not been peer reviewed. Data may be preliminary. 19 Posted on Authorea 16 Aug 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.158679916.68461360/v2 | This a preprint and has not been peer reviewed. Data may be preliminary. 20 Posted on Authorea 16 Aug 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.158679916.68461360/v2 | This a preprint and has not been peer reviewed. Data may be preliminary. 21