Predator Size and Prey Size–Gut Capacity Ratios Determine Kill

Home , Surplus killing

Medicine, Ghent Univ., Heidestraat 19, BE 19, Heidestraat Univ., Ghent Medicine, 1 Janssens J. P. Geert Cuype Annelies De mammals carnivorous terrestrial size prey and size Predator c Please [10.1111/ Record. of Version the and version this between differences to lead may which process, proofreading and pagination typesetting, copyediting, the through been not has but review peer full undergone and publication for accepted been has article This date: Decision Email: Belgium. Merelbeke, Heidestraat Univ., Ghent Medicine, ofVeterinary Faculty Ethology, and Genetics author: Corresponding 5 Germany Mainz, Mainz, Univ. 4 3 2

Accepted ArticleFlor Gutenberg Johannes Paläontologie, Analytische und Angewandte für AG fürGeowissenschaften, Inst. London,UK London, Societyof Zoological Zoology, Inst.of Animals Zoo for Clinic Veterinary Facultyof Ethology, and Genetics Nutrition, of Dept AnimalNutrition, Laboratoryof isbad Quaternary Research Dept, National Museum, Bloemfontein, South Africa South Bloemfontein, Museum, National Dept, Research Quaternary isbad oik .0 19 5488 - Jul 1 1 , Marcus Clauss Marcus ,

, Exotic Pets and Wildlife, Univ. of Zurich, of Univ. Wildlife, and Pets Exotic , Annelies De Cuype, Laboratory of Animal Nutrition, Dept of Nutrition, Nutrition, of Dept Nutrition, Animal Laboratoryof Cuype, Annelies De

– [email protected] gut capacity ratios determine kill frequency and carcass production in production carcass and killdetermine ratios frequency capacity gut

2 , Chris Carbone Chris , - 9820 Merelbeke, Belgium Merelbeke, 9820 3 , Daryl Codron Daryl

Zürich, 4,5

, An Cools , Switzerland t ti atce s doi: as article this ite 1

19, BE 19, , Myriam Hesta Myriam ,

- 9820 1

and and - physiology. The model predicts a negative relationship between predator size and kill frequency for largefor kill and size frequency predator between negativerelationship a modelpredicts physiology.The that the dichot indicating spectrum, bodysize prey oflarge and is below, feeders prey small majorityof Although the massprey (iMprey). corrected pack and (C) capacity stomach prey small groups, functional maincarnivore Two gut capacity. carnivore and ofprey consumption partial size, pack mass, developed We groups. functional carnivore to adapted insufficiently models are existing and rare are data frequency fieldkill Current ofecosystems. predator of part fundamental a frequencyis kill Carnivore Abstract Accepted words: predator Key gut capacity. large theirrelatively to due activity hunting in reductions distinct canwhoachieve carnivores, large for so is particularly This hunting. to related activities reduce thus which can carnivores, Articleprey large allow, conditions ecological Where consumption. carcass partial and kleptoparasitism, suchas scavenging, processes ecological other drives or frequencies kill reduces gastriccapacit full to haveeat to necessarily not do carnivores Large clearance. gut and digestion fast requiring day, per repeatedly capacity theirfull to eat may to have carnivores small capacity, estimatedstomach to requirements prey carnivore prey - feeders. However, for small prey small for However, feeders.

- feeders versus large prey large versus feeders – ‘This article is protected by copyright. All rights reserved.’ prey size r preysize atio, gut capacity, kill frequency gut kill atio, capacity, - feeders above a body massofbody10 a above feeders

- feeders, this negative relationship was absent. When comp When was absent. relationship negative this feeders, a kill frequency model accounting for carnivore mass, prey carnivore for model accounting killa frequency y per day, or do not need to eat every day, which inwhich turn everyday, eat to notdo need yor day, per - feeders, were established based on theon relationsh based wereestablished feeders, omy is rather linked to bodylinkedsize to is rather omy – prey interactions, which are important shapers shapers whichimportant are preyinteractions,

– 20 kg, both occur across the whole the across kg, occur both 20 - feeding appears att feedingappears - related ecology than ecology related

ip between between ip ractive for ractive

aring aring - et al. 1999 al. et level aninterspecific at feedingrelationships usedappr commonly issizea prey to size Relatingpredator ecosystems marine and terrestrial studying Pete interactions intraguild and predation both through structure, community top ofthe drivers important are carnivores Terrestrial Introduction Accepted energeticrequirements and size prey carnivore on rangebased carnivore broad a for Apart 2015 do and wolves) killedby prey consider only Others observations. direct thanon (e.g. carnivores frequ killgive publications methods.Some missedfieldby therefore entirelyand mostlyconsumed are preysmall items fact that methodsthe and field intensive labour ofthe because frequen kill offield Reports capacity. stomach maximum and requirements energy of scaling in the discrepancy namelyputative a physiology, carnivore of aspect another to attention draw to frequencies ofkill thefor calculation model usewea rates encounter and detection on and hunting of the energetics on focused have consequences ti active frequently lesskill hunt to and need carnivores mass theirown Articleexceeding or to equal vertebrates largeopt for to tend (>20kg) carnivores largewhereas invertebrates) sm veryon specialize to tend (<20kg) carnivores kg 20 ofabout massthreshold body a at feedingoccurs preylarge d rives the choice for a specific prey size specific a for rives thechoice me with body size in herbivores in sizeme withbody rson 1994, Ritchie and Johnson 2009) Ritchie Johnson and 1994, rson ).

from field kill frequency observations, efforts have been made to estimate carnivore kill frequency kill carnivore made estimate to been have efforts observations, frequency kill field from

(Jeschke 2007, Rizzuto et al. 2018) al. Rizzutoet 2007, (Jeschke

, Tucker et al. 2016, Codron et al. 2018 al. et Codron 2016, al. et , Tucker

not consider all prey species killed by the carnivore (e.g. (e.g. by carnivore killed the species preyall not consider 1980 cy data are scarce and almost exclusively available for large carnivores, carnivores, largefor available almost and scarce exclusively are data cy ; feral cat) but derive the data from the data but derive cat) ; feral ; Siegel200 . Correspondingly, it has been calculated that larger terrestrial that larger calculated been has Correspondingly, it . (Pawar et al. 2012, Carbone et al. 2014) al. Carbone et 2012, al. (Pawaret (Peters 1983, Carbone et al. 1999, 2014) Carbone 1999, al. et 1983, (Peters

. . Predator and have more daily sleep (in contrast to decreasing daily sleep sleep decreasingdaily to more(in contrast havedailysleep and (Cohen et al. 1993, Heithaus 2001, Carbone et al. 2014) al. Carbone et 2001, Heithaus 1993, al. et (Cohen 5 (Rosenzweig 1966, Holling et al. 1976, Paine 1976, Carbone Carbone Paine 1976, 1976, Holling al. et 1966, (Rosenzweig ). The explanations for the prey size dichotomy and its and dichotomy sizethe for prey explanations ). The (Peters 1983, Vézina1985) 1983, (Peters all prey (vertebrates and a considerable amount of considerable a and (vertebrates prey all ) . For example, it is known that carnivore bodysize carnivore that known For itisexample,. - prey relationships are considered fundamental for fundamental considered are preyrelationships - down control of ecosystems and the shaping of shapingof the and ecosystems of control down - specific kill frequencies (e.g. the number of moose of number the (e.g. frequencies kill specific (Carbone et al. 1999, 2007, Rizzuto et al. 2018) al. et Rizzuto 2007, 1999, al. et (Carbone oach to describe community interactions and and communityinteractions describe to oach (Carbone et al. 1999, 2007) 1999, al. et (Carbone caloric requirement estimations rather rather estimations requirement caloric (Terborgh 1992, McLaren1992, and (Terborgh

and can generally be less canbe generally and . . Inthe c present encies for smallfor encies : a switch : a from to small Zimmermann et al. al. et Zimmermann (Peters 1983, Vézina 1983, (Peters

ontribution, ontribution, . Small .

prey can feed selectively, meaning that they can afford to consume highly digestible body parts suc parts body consumehighly digestible to afford meaningcan they that selectively, feed preycan frequency kill infor previous accounted not been have factors Several oftheir top on preysurplus canproduce 'lazy' they be since to afford can Largercarnivores hunting. oftheir day portion significant invest a to thus need and frequencies kill higher require carnivores that smaller models predict These preybody (ii) and size predator with (i) frequency in kill decrease a predict estimates frequency kill Typically,these predator. ofthe size prey byaverage the 1985) Acceptede from gut capacity differentiate to it impossible makes caloric from werederived requirements ca prey large a isconsidered the carnivore gut capacity, the exceeds preysize smalla prey isconsidered the carnivore preysize, exceed to is assumed capacity prey sizes larger at intake thewhich biomass limits capacity, theby gut constrained will be will consume muchcarnivore how a and selectively how questionof werea carnivores switches), whichsize prey withat threshold a vs predator, small (large mass body carnivore on groups functional in carnivore ofbasingdivision the Instead size. hunting pack of an estimate and in prey, requ energy maintenance specific the carnivore species, carnivore per mass prey most common ofmass, the average carnivore on modelbased frequency wekilla so, develop we wante Therefore, 2006) 1.6 kg 0.8 4 of cat a for frequencies kill of estimates unrealistic example,to for may lead, notexceed does or preydoes whose separately predators not considering capacity; intake Articleskin,bones) Anwar 2004, Bothma Coertze and 1996, (Mills 2015) al. et Bosch organs muscles and pacity expands the approaches used in the models ofmodels the in used the approaches expands pacity (Peters 1983) (Peters . In order to do so, simply stated, data on p on simply data so, stated, to do Inorder . .

ppointed to one of two functional groups based on the relation of gut capacity to prey size. The prey The to size. ofgut capacity relation theon groupsbased functional oftwoone to ppointed . P redators might not be able to fully consume large prey, due to the limitation of their own oftheir limitation the prey,to due large fully to able might consume be not redators

times per day, rather than the 'multiple times' considered realistic for cats cats for realistic considered times' 'multiple the than times rather day, per , whereas predators that kill comparatively small prey will consume their prey entirely their will consume small prey that kill comparatively whereas , predators (Hornocker 1967, Bowland and Bowland 1991, Stahler et al. 2006, Gidna et al. 2014, Gidna2014, al. et 2006, al. Stahleret 1991, Bowland Bowland and 1967, (Hornocker d to explore the relationship between kill frequency and carnivore body size. In doing Indoing size. body carnivore and frequency kill between therelationship explore to d ‘This article is protected by copyright. All rights reserved.’ energetic maintenance requirements requirements maintenance energetic food food intake data of mainly captive animals animals mainlycaptive ofintake data

et al. 2011) al. et redator daily food intake (or ingestion rate) are divided divided are ingestion rate) (or daily intake food redator size for a range of mammalian and a and mammalian rangeof a for size nergetic requirements. requirements. nergetic Peters (1983) Peters

including less digestible body includingless digestible

(Chakrabarti et al. 2016) al. et (Chakrabarti (Jeschke 2007) (Jeschke

and and irements and metabolisable energy metabolisable and irements modelling Vézina(1985) - feeder. The inclusion ofgut inclusion The feeder. According to the functional to According . -

. Predators taking large taking Predators . feeder. Whenever the Whenever feeder. (Farlow 1976) (Farlow

. Whenever the gut Wheneverthe . (Vézina 1985) (Vézina vian carnivores. carnivores. vian

where

intake capacity intakecapacity parts (i.e. fur, (i.e. parts (Bradshaw (Bradshaw energy , which , has

mass and gut capacity. and mass mass. requi energy higherthan scaled predator thefor individual available mass prey the mass;inif contrast, predator withincreasing occur would in frequency reduction kill no then energy requirements, to, similar or lowerthan, scaled thefor individ available mass prey mean thewas if that hypothesis working Our non and vsusecomplete selective and incomplete madeconcerning are adjustments prey group, energetic Accepted(th kill/x days) (1 frequency (N ofanimals) (number size pack (%)), items ofprey number species/total certain ofa items (= offrequencyoccurrence or relative (%)) other or ofscats number species/total certain a preyof items (=identified ofoccurrence frequency on based most prey frequent sex, samples,carnivore live remains, kill analysis (scats, diet for used methods location, study species, carnivore from publication: each wereextracted following data The (2014) feeding 'sub canbe prey small several marineenvironment prey;the in invertebrate and vertebrate ofterrestrial dispersal in the difference (e.g. feedingtime and search termsof Clearly,thes carnivores. aquatic and carnivores, omnivorous and feeders 2014 wereexc invertebrates strategies foraging (e synonyms possible gu kill frequency, size, pack (preyfactors size, associated habit feedingon second the and plural) and singular name, common (latinname, ofthe Carnivora order the on based terms,one sear literature The carnivores. Articleto 2014 scholar Google Pubmed and ofKnowledge, Web using performed was review Aliterature set( Data methods and Material - selective useofprey. selective ). ). We expected that these results would depend on the difference in the relationship between prey prey between the relationship in theon difference woulddepend results that these expected We ). The latter was done given the difference in foraging strategies between terrestrial vertebrate terrestrial between strategies foraging in wasthe difference given latter done The

whe December Supplementary Supplementary reas this is not possible in terrestrial environments; cf. cf. environments; in terrestrial not possible isthis reas , 2015, 2015,

as well as carnivores carnivores wellas as ‘This article is protected by copyright. All rights reserved.’ .g. nutrition, feeding habit feeding .g.nutrition, luded (based on the quantitative dataset on mammalian diets of mammaliandiets on dataset theon quantitative (based luded

rements, then a reduction in kill frequency would occur with increasing predator predator withincreasing would occur frequency kill in reduction a then rements, to identify potentially eligible studies reporting feeding habits of wild of habits feeding reporting studies toidentify eligible potentially material Appendi

e 'real' e ch was conducted following wasch conducted or actually observed observed actually or

views, gastrointestinal content, regurgitation), number of number regurgitation), content, gastrointestinal views, whose ces

diet consists mainly (≥ 50 %) of vegetation and/or and/or of vegetation %) mainly 50 (≥ consists diet and 2 and

Aquatic carnivores or carnivores that depend on aquatic aquatic on depend that carnivores or carnivores Aquatic )

kill frequency kill Leenaars et al. (2012) al. et Leenaars dued' in the same hunting bout by filter by bout hunting same the dued'in Pawar et al. (2012) and Carbone et al. al. et and Carbone (2012) al. et Pawar ) and maximal gut capacity maximalgut capacity and ) e foraging strategies differ in differ strategies foraging e ual predator at a killa at ualpredator

by using two search twobysearch using t capacity

Wilman et al. al. et Wilman

identified prey identified ,

from January fromJanuary ) pack , including all including , ), kill ), - prey - jubatus ( gastrointestinal in the sustained be that can contents maximal in the role hence decisive playinga is dispersed, food which from reactor' 'batcha as acts stomach the carnivore since reliable considered be can estimates these Nonetheless, gut) reporte total capacity not (and maximalstomach the from stem usedhere estimates tract thegastrointestinal of content total the measuring (C). event) (kg/carnivore/feeding Kill frequency modelling modelling Kill frequency feedinggroup. the concerning number N hence frequency, killvs bodysize study to the relationship in order generalize has to one Additionally, 1989) Gittleman group foraging group, (population theon context depending candiffer required. are generalisations hence and frequency, initial aim the However, bycarnivore. the Zalewski calculated. mass(M prey common by carnivore the hunted preyspecies were not taken period) time same the preyin taken ofother irrespective wolves, killedby moose theof number (e.g., species prey specific a applyto only fromt o Actually thesee acquisition, data informationon dat frequency kill and gut capacity maximal pack size, For doubling ofinformation. avoid fromto the dataset wereexcluded reviews in these Hayw 2005, Kerley and (Hayward

Accepted Articlecrocuta Crocuta pack data obtained in this dataset are of are dataset in this obtained data he publication itself (i.e., dividing the reported frequency with pack size). Kill frequencies that that frequencies Kill withsize). pack frequency dividingreported the (i.e., itself hepublication ) and the African wild dog ( dog wild the and African ) 2005 bserved kill frequ kill bserved We are aware that environmental conditions (such as season or geographical location) (e.g. (e.g. location) geographical or as season (such conditions environmental awarethat are We ) or the sex within dimorphic species (e.g. (e.g. species withinthe sex dimorphic or ) . However, many reports describe pack size as is, without differentiation ofgroups. without differentiation as is, size pack manydescribe However, . reports ), the tiger( ), ‘This article is protected by copyright. All rights reserved.’ prey

), the average N average the ), ency data ency Panthera tigris Panthera

tract tract T ard 2006, Hayward et al. 2006ab al. Hayward et 2006, ard a, additional literature searches were conducted. For more detailed detailed more For wereconducted. searches literature additional a, Lycaon pictus Lycaon ypically, in herbivores, gut capacity estimates are obtained by obtained are estimates gut capacity ypically,herbivores, in (Hume2002) (Kroshko et al. 2016) al. et (Kroshko

were corrected for the pack size of the carnivore species, obtained obtained species, carnivore ofthe sizethe for pack werecorrected

S an averaged kind, with the preference, if reported, of the pack size ofsize the pack withif reported, thekind, preference, anaveraged upplementary material Appendix 2 Appendix material upplementary Pack was to assess the effect of carnivore body size on kill kill on bodysize carnivore ofeffect the assess to was ), the leopard ( the leopard ),

and the average the average and ), we included the reviews of Hayward an the reviewsofHayward we ), included . We are We into account since these estimates did not consider all not consider did estimates these since account into For the lion( For (Parra 1978, Müller et al. 2013) al. et Müller 1978, (Parra Zalewski Zalewski also . Panthera pardus Panthera Per carnivore species, the average of the average species, carnivore Per actually observed kill frequency kill actuallyobserved aware that pack size is a general term and termand general a is size pack awarethat Panthera leo Panthera 2007 c, 2012) c, , feeding group, breeding group, feeding , ) can affect the prey size hunted size canaffect ) theprey . Therefore, publications used publications . Therefore, ), the cheetah), ( .

), the spotted hyena ),the spotted

. The carnivore . Thecarnivore d in literature. in literature. d Acinonyx Acinonyx d collaborators collaborators d

were

group; group; most KF ~ M KF ~ and Q M of theon scalingrelationships Based as Q calculated energy maintenance M on based wasmodeldeveloped (KF) frequency kill Atheoretical Acceptedlupus givenbyvalue average as 70% consumed prey large For skeleton). portionsof skin and inedible to (due edible wereconsid kg 5 of> preywhereas items completely be edible to werekgconsidered 5 < items given by from data weight calculated kJ/kg fresh 5348 at estimated was as M item prey (M ofan individual iM ofC < 1% where those insects), on preying theirgut c relationshipofM The nevertheless. size pack for corrected be that Note M KF ~ therefore and iM (iM predator the individual for available prey of theamount ArticleN mass withpredator scales size t modifications several capacity, gut and feedingselectivity about considerations our and hunting, ofpack occurrence given the However, pack pred prey prey prey

the assumption that the energy content of prey is directly proportional to prey mass, we wouldweexpect prey mass, to proportional directly is prey of content energy the that theassumption

prey

~ M ~ ). Large prey predators were assumed to only consume the equivalent of their gut capacity C per C per gut capacity their of theequivalent only to consume were assumed Large prey). predators ~ M ~ ~ M ~

(or 'large prey (or ~

pred pred M is larger than C, however, C becomes one C becomes C,however, than islarger

pred pred a scaling of pack size with body mass may not be expected, but data for individual species must species individual for but data expected, not be maymasswith body size ofpack scaling a pred apacity C. We divided predators into those whereintothose predators divided C.We apacity pred (q (q - - q n p+n) p) p

- pred n)

(Mills 1990, Stander 1992, Caro 1994) at a 1994) Caro 1992, Stander 1990, (Mills / E - ‘This article is protected by copyright. All rights reserved.’ (Q requirements feeders' who cannot consume their average prey in one meal). preyone in theiraverage consume cannot who feeders' prey prey Bosch et al. al. et Bosch .

~ M ~ pred prey

o this simple concept need to be applied. Under the assumption that pack the assumption Under applied. be to need simplethisconcept o p ) is smaller than C,M than is smaller )

needs to be established for several groups of predators, in relation to into relation ofpredators, groups severalfor established be to needs 2015,

pred

) and metabolisable energy in(E energy prey metabolisable and )

ta iM king into account t kingaccount into prey

< C (or 'small prey C(or < factor constraining KF. The E The KF. constraining factor prey

is the mainof driver the is prey iM E ) scales to ) he selective feeding of wolves wolves feedingof heselective prey prey

< 1% of C (i.e., predators mainly predators ofC (i.e., 1% <

of pred - feeders, prey was considered to be be to was considered prey feeders, - feeders'), and those where C< where those feeders'), and 8048 kJ/kg fresh weight kJ/kg (the fresh 8048 , M Plantinga et al. (2011) al. et Plantinga

prey , carnivore specific specific carnivore , prey KF ). For each species, KF is KF is species, each For ). prey

in our dataset. As soon indataset. our

As long as the mass As as the long for small prey small for

Canis Canis , and prey and , ered 95% 95% ered - feeders feeders

mainly animals captive is latter M Followi frequencies. killmaximum minimumand theoretical outline to moreday), one than kJ/kg, over 5348 at with consumption mode,95% i.e. selective non a prey (in oftheir consumption complete a kJ/kg) or 8048 C of at amount the eating mode,i.e. single a on werebased either KF estimates group, this day. For example for the range of kill frequencies available to large carnivores with the option to switch between switch to between option with the carnivores large to available frequencies kill of thefor range example prey(f ( bear the polar we Because considered applied. be to is extrapolation the which to thespecies include on was based C) (for species using as calculated results regression ( moreappropriate is regression by another, predicted shall be However an average in involved deriving Vézina1985) isimport data caper package using the likelihood, maximum signal phylogenetic with the (PGLS) least squares generalized in phylogenetic analysed phylogen werea to linked data dataset, ofthe structure phylogenetic generalized scalin Evaluationsof FMRofanimals. the mayseasondetermine and habitat diet, suchas taxonomy, factors other mass, body Next mass. to body it to scaling by (FMR) rates metabolic field of assessment by physiologicallyfrom energy and expenditure from not (as intake in Farlowthe dataset used

AcceptedUrsusmaritimus Article pred Peters Peters 0.85 ish) and large prey (seals), we excluded this species from scaling analyses, and used it as anit as used and scalinganalyses, from wespecies this excluded (seals), prey large and ish) ng

advantageous since data are on based exclusivelyfree d , because when scaling allometries are not considered predictive equations where one measure one where predictiveequations not considered are allometries whenscaling because , - 1 Nagy et al. (1999) al. et Nagy . Hence, maintenance energy requirements are based on field metabolic rates (FMR). The The (FMR). rates metabolic fieldon based are energy maintenance requirements Hence, . 1983 least squares ( squares least , and with PGLS results, and is also justified because due to the methodological steps steps methodological the to due because justified is also results, withand and , PGLS ant for a comparison with previous work that is also based on linear regression ( linearregression on based also workis that withprevious comparison a for ant

) as an extreme example of a predator that might switch between comparatively small small comparatively between thatswitch might predator ofa example as an extreme ) and and ‘This article is protected by copyright. All rights reserved.’ g relationships were performed using linear regressions for log for regressions usinglinear wereperformed grelationships Vézina G (Farlow 1976,Peters 1983, Vézina 1985) LS) in R using the package nlme nlme theusing package in RLS) , we parameterize the relationship ofQ relationship the , weparameterize G but shall but shall LS scaling, because PGLS because scaling, LS 1985 Warton et al. 2006 al. et Warton M prey ). , the uncertainty in the the in the uncertainty , the R the be analysed for the best fit to the data, fitthe to thefor best analysed be One must hereby acknowledge that Q mustOneacknowledge hereby

smatr smatr (Orme2010) al. et package package ), we also report standardised major axis (SMA) axis major standardised wereport also

scalings are based on phylogenies that do not thatdo phylogenies on based are scalings (Warton et al. 2012). al. et (Warton M (Pinheiro et al. 2011) al. et (Pinheiro prey - day feeding on their prey (in a sel(in a prey their feedingon day .

data is higher than in the the isin higherthan data Using GLS regression to analyse theanalyse to regression UsingGLS pred etic tree tree etic

ranging animals rather than

= b M b = , deriveand (Fritz et al. 2009) al. et (Fritz pred

pred (standardised) (standardised)

Extrapolation to other other Extrapolationto

q is based on theon isbased

as Q . To account for the accountfor . To - transformed data in data transformed

pred λ

= 791kJ 791kJ = estimated by estimated M major axis major pred Peters 1983; Peters , also and

data. ective ective -

A total of 456 Aof456 total characteristics Carnivore Results 2018) originald The KF. our graphsrepresenting in the wereincluded KF comparison, the For groups. carnivore functional two significantly different from0). significantlydifferent in and GLS t averages, Usingspecies observed Actually M graphlinking the but in not (Fig. 1a), relationship M the graphdisplaying inas an outlier the appeared dog the bush Therefore, member). anby individual consumed be to large too been have would that prey killing was the pack C(i.e. dog bush and ( ( shar weresupposedly iM C than larger had species hunting pack Five day. inanimal one prey whole the not consume [0 to nominally scaled size Pack hunters. weresolitary species 63 and hunters werespecies pack 12 wereavailable, which data pack for species carnivore 75 Of the [ extrapolate function theallometric fo available kg to 387.5 0.1375 from weight ranged Carnivore species. 75 for obtained only could be eight data frequency 1.06;1.34 Acceptedrufus Canis Martes flavigula Article ;1] M ;1]

carnivore species species carnivore .

] . pred

r 9 species ranging ranging species 9 r C for all carnivore species carnivore Call for 4 4.15 [ 4.15 , 2.4 pack members), Ethiopian wolf or simien jackal ( jackal wolfsimien Ethiopian or members), pack 2.4 , 4 [0.4

( ata studies

Speothos venaticus Speothos points and 22 22 and points 35

;0. , 2 pack members), themembers), golden/ pack 2 , compilation is compilation 0.57;30.53] M 0.57;30.53]

kill frequency data were found for 11 species weighing between 11.2 kg and 175.5 kg. kg 175.5 and 11.2 weighingbetween species 11 for were found data frequency kill 54 ] ‘This article is protected by copyright. All rights reserved.’

ing prey that each individual could have eaten moreof eaten have could thatindividual each ingprey SMA. (listed in in (listed in GLS GLS in (listed in Appendix 1) Appendix1) in (listed he

actually observed actually Of the 12 pack hunting species, 7 species had a C < iM C< a had species 7 species, hunting pack the 12 Of stomach

in body mass inbody [with 95% CI] of C = 0.09 [0.06;0.14] M [0.06;0.14] 0.09 ofC = CI] [with95% Appendix1

pred available from the Dryad Digital Repository: doi link (De Cuyper et al. al. et Cuyper (De link doi Repository: Digital Dryad from the available , 11 pack members). Of these, only the bush dog had a higherM a had dog bush only the Ofmembers). pack these, 11 , - 0.81 [ 0.81

; SMA ; SMA capacity data capacity Actuallyobserved - 1.45; ) - 0.46] , 513 prey 513 size , from 0.19 kg to 150.0 kg; these data were used to determine to wereused kg; data these 150.0 kg to from0.19 yielded yielded

were included wereincluded kill frequency kill

in SMA, A

. sian jackal sian Data on the maximalthe on Data

points were incorporated in the database. in the database. incorporated were points pre a similar result of C = 0.09 [ result 0.09 ofC = similar a d

to iM to

and did not show a phylogenetic signal ( signal not show phylogenetic did a and kill frequency data were obtained for 11 species. species. 11 for wereobtained data frequency kill models of models data points data 1 [1;1] M [1;1] 1

scaled scaled in the prey size database. Pack size data data size Pack database. size inthe prey ( prey Canis aureus Canis

(Fig.1b). Can to to Peters (1983) Peters pred , 182 pack size data size pack 182 , 1.11 [0.20;6.18] M [0.20;6.18] 1.11 issimensis pred stomach 0.14 [0.05;0.24 0.14

1.19 [1.07;1.30] 1.19 , 2.5 pack members), red wolf red members), pack 2.5 , -

the yellow throated marten throated yellow the

capacity (C) were (C) capacity 0.06;0.15

, 5.7 pack members), pack 5.7 , and and ]

prey inGLSin 1 butto

, , i.e. the pack could could the pack i.e. , Vézina(1985) pred which was used to usedto was which pred -

M points, 56 kill 56 points, ] M ] - 0.48 [ 0.48 prey Seventy pred prey

- 0.91; λ 1.20 , i.e. i.e. , prey

not -

0.04

than than - ]

con prey large for 0.5 smallfor prey was 0.05 which (intercept), factor the scaling in wasdifference large a there prey predators large and small both of interval confidence in the 95% was included scaling linear size, prey relative on based groupsindividually whencarnivore Incontrast, considering 1). (Table confidenceinterval 95% the in linearity exceeding also 1.0 than larger exponent withmassscaling a predator to scaled mass prey species, carnivore Acrossall Predator Accepteda be and winter, in feeder selective 2013) Rockwell and Gormezano Romberg2007, and f mixed,geesee.g. and = prey summer bound, land winterseal; = prey bound, ice ( bear polar The mass). body (here, mainvariable oneon based predictions of exponent the resulting of0.85 scaling expectation a leading to n0.44, = and 1) (Table 2.34 = p values are corresponding the InSMA, prey. of content energy and vs.incomplete) a by introduced scatter data additional the to due exponent are of exponent resulting the closelyto corresponds which an 1) (Table of use our of the regression to due 0.85 = q wholethedataset, to correspond theoretically should exponents scaling the mass, withbody size of pack evenis that approach ofour consistency ofthe check Astringent days 2). 29 (Fig. killone every and day per waskills two between predators prey large for range frequencies of kill The 2). was shallower which day.prey large Incontrast, per kills 12 mean± of7 a had and Articlemass withbody ofKF scaling not havesignificant a did feeders and PGLS of with an exponent scaling steep verya had completedataset the in KF scaling overall The outcomes model Kill frequency fidence intervals (Table 1). (Table intervals fidence - prey mass scaling mass prey - 1.51 in SMA 1.51 d n = 0.14. Therefore, we would expect KF to scale, in scale, to KF weexpect would n0.14. Therefore, = d Ursusmaritimus

in GLS and steeper in steeper and GLS in

(Table 2). However, when considering predator groups individually, small p small groupsindividually, whenpredator considering However, 2). (Table - in GLS (and of similar magnitude in magnitude SMA) ofsimilar (and GLS in 1.51 in Table 2, underlining the fact that SMA cannot be used for deriving usedfor be cannot that SMA fact underlining 2, the in Table 1.51

) is an example of a species that, depending on its seasonal eco seasonal its on that,depending species ofa is an example ) non - selective feeder with a higher kill frequency in summer (Fig. 2). summer in frequency withhigherkill a feeder selective

SMA

i n GLS and PGLS (but not in SMA; Table 1). Table SMA; in not (but PGLS n and GLS -

if complete consumption of prey was assumed (Table (Table was assumed prey consumptionof complete if

2.34 + 0.44= + 2.34 , might have a very low kill frequency / be a verya / be frequency kill might , low have very a - 0.53 in Table 2. 2. in Table 0.53

- djustment factors for carcass use (complete use (complete carcass for factors djustment Nagy et al. (1999) al. et Nagy feeders had a significant negative scaling, scaling, negative significant a had feeders in GLS, PGLS or SMA SMA or PGLS GLS, in - though 1.05, which deviates substantially from which substantially 1.05, deviates G , with no overlap in the 95% 95% in the with overlap no , LS, to 0.85 LS,0.85 to

Deviations from the expected expected Deviationsfrom the t here was , and in and , - ish; ish;

1.56 + 0.14 = 0.14 + 1.56 no significant scaling scaling significant no

Russell 1975, Dyck 1975, Russell KF ~ M KF ~ ( G Table 2, 2, Table LS p = 1.56 1.56 LS= p - feeders and and feeders - 0.66 However, pred type (i.e., - (q 0.57, 0.57,

Fig. 2), Fig.2), in - p+n) rey . In . 0

- Although the general output for the whole body mass spectrum is a negative scaling of scaling negative a spectrummass wholeis thebody for output general the Although Discussion full their to theoreticallyeat could predators large upwards, kg 30 mass ofapproximately frombody a basis; daily a on items ofprey number than day moreper prey ingest incan, theory, upwards kg from 4 approximately predators large whereas day), a ofless than clearance havestomach a must also against M wellestimated as the meetdemandsas energy to preyrequired of amount the Whenplotting Acceptedin the diffe use,evident they sizethe prey categories. predator impact between ecological in main but itsrelevance consider superior, modelbe to our not wedo approaches, ofofthese all broadness the Given frequencies. kill carnivore ofestimates previous of fromthose ways relevant in differ yieldsthat model our results 2, Fig. from As evident con must be that group latter this is It prey instantaneously. their consume completely cannot they because kleptoparasites) or scavengers of widercommunity the or themselves, preysurplus a (for of create themselves, for consume will necessarily whatthey hunt only thosethat into classified be is that can they predators impact of thefor ecological implications major 10 above were 'smallprey predators well the was there However, strategy. apparent also predators somesmaller because size, ofbody function a not necessarily is gut capacity instantaneous own smaller their than or focus larger prey on predators instantaneous than their smaller is preyonthat size of a focussed inpredators not evident withis size body relationship findingsof general the to modelsimilar output, a is bodysize Articleprey (large gut capacity instantaneous their exceeds prey that subdue that,average, on ofpredators group thefor While behaviours. hunting and dailyactivitybudgets these pr predators, mammalian terrestrial by selected sizethe on prey depending that demonstrate results our bodymass, carnivore sidered facilitatorsof sidered - pred 20 kg, predators mainly go for comparatively larger prey larger mainly comparatively for go kg, predators 20

(Fig. 3), it appears that small predators have to eat moretheirthan eat have to thatpredators small it appears 3), (Fig.

gut capacity (small prey (small gut capacity ‘This article is protected by copyright. All rights reserved.’ additional - feeders' that hunt for prey well below their instantaneous gut capacity, whereas gut capacity, instantaneous welltheir below prey for hunt that feeders' edators may vary distinctively in their kill frequencies, and hence their hencetheir and frequencies, kill in their may vary edators distinctively gut

trophic trophic -

- described general pattern that below 10 that below general pattern described every once only size) ofsufficient preyhunt (or capacity feeders). Notably, our dataset showed that whether or not or whether that showed dataset Notably, our feeders). rent 'preferred' prey species for a given predator givena predator for species 'preferred'prey rent interactions In particular, given the flexibility of predator species inspecies predator of flexibility given the Inparticular,

required, and thus might not need to hunt the same same hunt to the need thus not might and required, .

and those that potentially (but not necessarily) necessarily) not (but thosethat potentially and emphasizing - feeders) a dec a feeders) Peters (1983) Peters (Carbone et al. 1999) al. et (Carbone ly pursue such a large prey large sucha ly pursue

a dichotomy in potential potential in dichotomy a

rease in kill frequency with in frequency kill rease and and - 20 kg of body mass, more mass, kg ofbody 20 gut Vézina(1985)

capacity per day (i.e., day (i.e., per capacity

kill frequency kill . . Oneofthe species species gut ,such a -

feeding other day. other

capacity capacity

claim

to to

0.74; meal 1.45; ( exponent its scaling of interval theconfidence large feeders, observed actually theexact magnitude an exact lawsof biological are study present studies observational original between Acceptedlarge of strategy pursuea carnivores larger these usedif canonl potential this Evidently, events. eating) even (and hunting between intervals larger allows quality lower againstbuffer a not does requirements and gut capacity between discrepancy moreofthem disproportionately ingesting just potentially allo thus requirements, higherthan and one, the on gutof capacity scaling in the model,discrepancy ofa incarnivore our situation assumed The given. marten Japanese mustelids(e.g. in whole Athours. published 24 is < retentiontime) or digesta passage smalla for as Correspondingly, needs. theirgut capacity w eat, to have would carnivores smaller meansthat also this carnivores, M with larger asymptotically intake increases food that observation accounts thefor experimental approach that requirement energy and C) gut capacity Vézina1985) 1983, (Peters Articlemodels previous in, the stated explicitly but not was to, that implicit factor most important the Possibly 1981 Sunquist prey large towards observations inbehavioural bias unintended to (due studies preyobservational in ofthe populat characteristics specific either prey - -

0.37] 0.46] requirements on the other hand, resembles a concept in herbivores where gut capacity also scales wherealso gut capacity herbivores in concept a hand,resembles other the on requirements and and (Wachter et al. 2012, Chakrabarti et al. 2016) al. et Chakrabarti 2012, al. et (Wachter

, respectively , )

included the scaling for large prey large for theincluded scaling scaling exponent depending on the original studies considered. studies original the on depending scalingexponent M - bodied large prey large bodied pred ). - 0.22 [ 0.22

, either from the same large carcass or several small prey items, to meet their energy meet their to small items, prey several or carcass large the same from either ,

kill frequencies, which were available from the literature exclusively for large for exclusively fromthe literature wereavailable which frequencies, kill - 0.40; ). Differences in the real scaling exponent to model estimates are most likely due to mostto likely due are modelestimates to exponent scaling the real in Differences ). ‘This article is protected by copyright. All rights reserved.’ - 0.04]

suming intake at C, the model yields two killsyieldstwo model C,the at intake suming

for the complete consumption strategy consumption the complete for is gut capacity C. Our model used different estimates for intake capacity (as intake for different estimates modelused is C.Our gut capacity - feeder (a mustelid). This can only be achieved if stomach clearance (or (or clearance stomach if achieved canmustelid).only This (a be feeder Mustela melampus Mustela , we do not suggest that the scaling exponents de exponents the scaling that not suggest wedo , - ions under observation, or an under or observation, under ions feeders in our model ( model our in feeders had different scaling exponents (Fig. 3). Conceptually, this this Conceptually, 3). (Fig. exponents scaling different had wing larger herbivores to subsi to herbivores winglarger (Cla . If we assume this difference in scal in Ifdifference this we. assume uss et al. 2013, Müller et al. 2013) al. et Müller 2013, uss al. et ; Tsuji et al. 2015 al. et Tsuji . Rather, they describe a pattern that mayin differ that pattern a they describe Rather, . prey feeding prey GLS: GLS: ; SMA M - gu . Although larger carnivores could in could carnivores Althoughlarger . pred ), this condition is apparently this is), apparently condition t retention times of 1.96 to 11.75 h to 11.75 1.96 of times t retention ithin the same day, repeatedly to to repeatedly day, same the ithin M

: - When compared to the scalingofto Whencompared 0.48 [ 0.48

- pred M

or, in this case, eating events eating events in case, this or, pred - s - 0.33 [ 0.33 0.91; t

- - on lower quality lower on representation of smaller ofsmaller representation 0. 33 - - 0.04] 0.34;

[ - 0 . ; SMA: - 3

0.31] . Incarnivores, . the 5; rived in the rived - ing to be truefor be to ing 0.

31 for the single for - ]

diet, but diet, and and M pred -

M diets by diets prey y be - pred 0.81 [ 0.81 ; - - 0.51 [ 0.51 - -

- (competing p (competing ofth defence pack and hoarding,caching food other drive could surplus offood the production Alternatively, 2007) (Jeschke 2018) al. Rizzutoet 2007, efforts hunting reduce to them allows prey large of availability the and ecological capacity simensis Canis wolf Ethiopian as the (such items manyprey small demandsingesting by energy meet their theoryalso Acceptedsmall comparatively ofthese number sufficient a obtaining of challenge the to due species prey prey smaller to on theirability in maylimited be predators larger comparison, the in lackofsociality wolf)a or Ethiopian (cf. habitat in prey their large of to due the lowbe availability mayalso prey.This mid Most Mid p digestible tha newprey hunt rather and parts, mostnutritious its only but carcass, consumea not eve to maychoose they allow, conditions and resources Alternatively,if carcass. same the dayson 1990) al. et Phillips 1990, Vander Wall Sunquist 1981, 1981, (eg. bears wolves, not present are caves,la GormanMills and 1997) 1990, (Stander minimized predatorsis withsuperior contact that so hunts oftheir location geographic the time or ofday lo cheetahsuch as predators Subordinate in Articlewhenlynx decreases typically wolverine frequency kill The kills. predator) lynx (obligate reindeer lynx the Eurasian instance, For days. subsequent prey largefor frequencies creases from 1 to 2, i.e. the lynx provides more scavenging opportunities opportunities morescavenging the lynxi.e. provides 2, to from 1 creases sing their prey to lions and hy and preylions to sing their - to - small rge burrows or dense vegetation to safeguard its prey surplus, but less so when kleptoparasites whenkleptoparasites so but less surplus, its prey safeguard to vegetation dense or burrows rge (Rangifer tarandus) (Rangifer - sized carnivores are not limited by their maximal gastric capacity, but by their choice of small choice their by but gastriccapacity, maximal by not limited their are carnivores sized o r - redators that try to steal a carcass from other predators) and scavenging, and could lower kill lowerkill could scavenging, and predators) from carcass other a steal that tryto redators tions sized carnivores show a large variety in kill frequency with our modela with our frequency varietykill in large showa carnivores sized

. (Sunquist and Sunquist 2002, Balme et al. 2017) Balme al. et 2002, Sunquist and (Sunquist in our dataset that feeds that dataset inour

Ursus maritimus Ursus (Stirling and McEwan 1975, Gende et al. 2001) al. et Gende McEwan 1975, and (Stirling ‘This article is protected by copyright. All rights reserved.’ se species that otherwise would help to overcome large prey overcome prey large to help would thatotherwise species se - , feed selectively on the large prey they acquire, and b and theyacquire, prey large the selectivelyon feed , feeders even more since the carnivore could eat from the prey for several for prey fromeat the could carnivore morethe evensince feeders , with the wolverine (facultative scavenger) being more prone to scavenging to more prone being scavenger) wolverine(facultative with , the enas enas ), hyenas ), (Kruuk 1972, Schaller 1972, (Kruuk (Acinonyx jubatus) (Acinonyx on small rodents), the combination of their relatively large gut large relatively their combinationof the rodents), small on . The leopard leopard . The , felids(eg , e carcass to safeg to carcass e (Lynx lynx (Lynx (Panthera pardus) (Panthera . bobcats and tigers), and mustelids and tigers), and bobcats .

or African wild dogs or African ) 1972, Gorman 1998) al. et 1972,

and wolverine and . Thus, some carnivores can feed for severalfor feed can carnivores . some Thus, behavioural uard e uard . Food caching Food .

xcess prey from kleptoparasites from prey xcess

packages. Because small prey Because packages.

often moves prey into trees, intotrees, movesoften prey /ecological processes suchas processes /ecological (Andrén et al. 2011) al. et (Andrén (Gulo gulo) (Gulo

has been described in described been has (Lycaon pictus), (Lycaon ecome 'full and lazy’ and 'full ecome n consume the less theless nconsume (Lamprecht 1978) (Lamprecht

pproach (Fig. 2). (Fig. 2). pproach , may the, choose

both prey upon prey both - wolverine ratio wolverineratio (Harrington (Harrington (Carbone et al. al. et (Carbone .

both both .

n In offered single meals per day, which maycontribu which day, per meals single offered Bradshaw 2006) 1984, al. et (MacDonald birds) rodents, (e.g. mass body withlower preya carnivores. domestic) non and (domestic the feedingofcaptive for implications mayseveral have frequencies kill on Findings preyout items small filter that can marine predators to contrast in canpursue, they size prey in the limited animals are individually, larger comprehended must be but environment terrestrial a out of filtered be cannot items Accepted rodents small than rather rabbits on living cats 2017) al. et (Lumetsberger 2016) different prey on specialized are wheredifferent individuals withinspecies well occur may prey 'small is a carnivore a whether determine than rather but ecologic existcarnivores, in to seems dichotomy functional a Thus, frequencyreduction. killa not result in did and gutby capacity, mostlynot limited variable, the preda exceeds individual prey the typical if frequency kill in reduction a to leading requirements, energetic the exceeds capacity the intakesize, predator thatlarger predictions at corroborates modeloutcomes our Inconclusion, management o inthe part keya therefore is frequency kill prey size, and size predator 2007) Mason and (Clubb th why the reason be could which signal, satiety animportant not receive metdo they but are requirements energetic where their condition satiety to linked putatively that and is gor a after that occurs theirstomach of distension extreme the fromexperiencing ever daily tigers, or suchas lions feeders, 2005) al. et (Altman Article improved weight body digestibility food and that observed was program, it feed/fastingday gorge random a to program feeding from conventional a adapted weregradually lions where study Ina stereotypy). and dietary over to related fromsufferproblems often lion suchas the captivity cats domestic , within ,

individuals over ontogeny over individuals (Laflamme 2006, Bissot et al. 2010, Deng et al. 2013) Deng al. et 2010, al. et Bissot 2006, (Laflamme

of their environment environment oftheir .Inparticular,recen ‘This article is protected by copyright. All rights reserved.’ The wildcat The . Elucidating feeding strategies in the wild such as the relationships between between the relationships wild suchas in the feedingstrategies Elucidating . . Observations deviating from the general pattern, such as a population of wild of population as a such general pattern, from the Observationsdeviating . tor's gut ese carnivores are particularly susceptible to showing stereotypies stereotypies showing to susceptible particularly are carnivores ese (Felissilvestris) (Carbone et al. 2014) al. et (Carbone

- capacity. Kill frequency outcomes for small prey small for outcomes frequency Kill capacity.

(Veasey 2017) (Veasey and hence comparatively small hencecomparatively and t speculations suggest that a practice of feedinglarge of practice thata suggest t speculations (Elbroch et al. 2017) al. et (Elbroch . Nowadays, domestic cats or wild cats kept in zoos are often are zoos wild in kept cats or cats Nowadays, domestic . - feeder' or a 'large prey 'large a feeder'or (Malo et (Malo , te to the increasing problem of obesity in at least inat in ofobesity problem the increasing to te

which makes it necessary to to necessary makes it which

for example is a solitary hunter that mainly focuses o focuses mainlythat hunter solitary isa example for . Hence, these animals might be in a constant constant a in might be these Hence, animals . al. 2004) al. .

, or in individuals between hunting events huntingevents in individualsbetween or , , or a population of wild living dogs ofa ,population or . Non . -

feeder'.T -

ration - domestic carnivores in carnivores domestic

- physiologic factors seem to factors physiologic supply (i.e. obesity, inact (i.e. supply s , may prevent these animals animals these mayprevent , kill his functional dichotomy functional his

several several

ge f carnivores - feeders weremore feeders - feeding event, event, feeding times (Codron et al. al. et (Codron - prey prey

per day per ex situ ex ivity ivity - n .

Acknowledgements interest. of conflict no that have they declare authors The ofinterest: Conflicts 131448). (grant number (IWT) Flanders in Technology and Science through ofInnovation Promotion for Institute by funded was the study This Declarations adaptatio behavioural and fixedthan physiological rather circumstances ecological in sought must be dichotomy ungulates small very mainly on Accepted carnivores. ofterrestrial diet the on Energeticconstraints 1999. al. C.et Carbone, ( dogs ofdomestic behavior feeding thefor basis evolutionary The 2006. S. W. J. Bradshaw, throug components prey of rates Differential passage A.1991. Bowland, and E. M. J. Bowland, Kalahari southern in success increases hunting Moterhood 2004. R. J. Coertze, and P. J. Bothma, nutritio dog optimal for wolves: insights wild of nutrientDietary profiles 2015. al. et G. Bosch, Bisso felid. large in solitary, a kleptoparasitism Caching reduces 2017. A. al. G. et Balme, 2011. al. et B. M. Anwar, An Article lions. captive feedingin fast ofgorgeand effects Nutritionalbehavioral and 2005. al. et D. J. Altman, References manuscript. ofthis versions earlier on comments constructive for reviewers thank two anonymous We drén, H. et al. 2011. Modelling the combined effect of an obligate predator and a facultative predator facultativea and predator predator obligate an effect of Modelling the combined 2011. al. et H. drén, t, T. et al. 2010. Novel dietary strategies can improve the outcome of weight loss programmes in programmes weight loss ofoutcome the improve can dietaryNovel strategies al.et 2010. T. t, ofserval leopards. S40 113: Nutr. client obese 634 Pakistan. Northern tarandus. lynx prey: common a on 47 8: Anim.Sci. Welf. Appl. familiaris ns. – 644.

Felis serval Felis

) and cats ( cats and ) - -

– S54. -

Food habits of the snow leopard Foodsnowleopard habits ofthe

Felis catus Felis Eur. J. Wildl. Res. 57: 1077 Res.57: Wildl. J. Eur.

and black and Lynx lynx Lynx -

(Woodroffe et al. 2007) al. et (Woodroffe J. Feline Med. Surg. 12: 104 12: Surg. Med. Feline J. – 57. – - ). ). – backed jackal jackal backed 760. 43.

J. Nutr. 136: S1927 136: Nutr. J. and wolverine and

Canis mesomelas Canis – , indicate that the underlying cause for thefor cause ,underlying that indicate the 1083. Gulo gulo Gulo Panthera uncia Panthera – –

S1931. 112.

predation on reindeer Rangifer Rangifer reindeer on predation

. -

Koedoe 34: 37 34: Koedoe

(Schreber, in 1775) Baltistan, (Schreber,

Lett. to Nat. 402: 286 402: Nat. Lett. to -

J. Anim. Ecol. 86: 86: Anim. Ecol. J. – 39.

Canis Canis n? n? h the gut h the -

Br. J. J. Br. -

J. J. – Deng, P. et al. 2013. Effects of dietary macronutrient composition and feeding frequency on fasting and and fasting on frequency feeding and macronutrient composition Effects2013. dietary al. et of P. Deng, 2018. A. al. et Cuyper, De Chakrabarti, S. et al. 2016. Adding constraints to Adding constraints 2016. al. et S. Chakrabarti, species. an asocial in living Group plains: the Serengeti Cheetahs1994. of M. T. Caro, ofver diet the influencing factors Geometric 2014. al. C.et Carbone, ofcarnivory. costs The 2007. al. C.et Carbone, Accepted 2009. A. S. al. et Fritz, large cretaceous late ofa dynamics ofthe trophic A consideration 1976. O. Farlow,J. 201 al. et L. M. Elbroch, ( bear wildpolar Observationsofa 2007. S. Romberg, and M. Dyck, al. et 1993. E. J. Cohen, niches predator reflect isotope carbon stable Carnivore al. et 2018. J. Codron, et D. Codron, Article How welfare: in factor carnivore risk as a biology behavioural Natural 2007. J. G. Mason, R.and Clubb, gastrointestinal and quality ofdiet Allometries Herbivorybody size: and 2013. al. et Clauss,M. postprandial hormone response in domestic cats. incats. domestic hormone response postprandial mammals carnivorous in terrestrial production carcass and frequency Polar Biol. 12: 1625 Biol. Polar ( Arctic charr fishing 67 62: 303 enclosures. improve zoos help could differences species analysing dinosaur ecology herbivore and for implications physiology,and consumption. Press. Chicago environments. terrestrial 288. community (Oldman Formation). Formation). community(Oldman African savannas. savannas. African herbivores. and carnivores between –

328. – 78.

al. 2016. Within 2016. al.

J. Anim. Ecol. 85: 660 85: Ecol. Anim. J. Geo Body sizes of animal predators and animal prey in food webs. food in prey animal and predators sizesBodyanimal of - 7. 7. Datafrom:

Integr. Zool. 13: 13: 166 Zool. Integr. – Salvelinus alpinus Salvelinus Stage 1628. graphical variation in predictors of mammalian extinction risk: big is bad, but bigis bad, risk: mammalianextinction of ingraphical predictors variation - - population isotopic niche variability in Savanna mammals: disparity disparity mammals: Savanna in variability niche isotopic population

Ecol. Lett. 17: 1553 Lett. 17: Ecol. -

dependent puma predation on dangerous prey. prey. dangerous on predation puma dependent Predator size and prey size prey and size Predator -

Ecology 57: 841 57: Ecology -

Front. Ecol. Evol. 4: 15. 4: Evol. Ecol. Front. – ) and Fourhorn sculpin ( Fourhorn sculpin and ) 670. – - 179.

PLoS Biol. 5: e22. 5: Biol. PLoS predation through allometric relation of scats to to scats of relation through allometric predation

– 1559. -

J. Nutr. J. – 857.

Sci. 2: e36. 2: Sci. gut capacity ratios determin ratios gut capacity

Ursusmaritimus tebrate predators in marine and in predators tebrate Myoxocephalus quadricornis Myoxocephalus gigantism. gigantism. -

Appl. Anim. Behav. Sci. 102: 102: Sci. Behav. Anim. Appl. . -

Oikos doi link doi Oikos – prey size relationships in in preyrelationships size -

J. Zool. 302: 164 302: Zool. J. -

PLoS One 8: e68714. 8: One PLoS ) successfully ) -

dinosaur dinosaur - -

J. Anim. Ecol. Ecol. Anim. J. University of of University

e kill kill e ). ). – 170. -

Gorman, M. L. et al. 1998. High hunting costs make African wild dogs vulnerable to kleptoparasitism by kleptoparasitism to wildvulnerable African dogs make costs hunting High 1998. L. al. et M. Gorman, trends. living: Carnivoregroup comparative L. 1989. J. Gittleman, 2014. al. A.et O. Gidna, 2001. al. et M. S. Gende, Accepted mammals. of strategies Digestive 2002. D. I. Hume, elk and deer mule upon predation mountainlion of An 1967. analysis G. M. Hornocker, 1976. al. et C.S. Holling, Predator R.2001. M. Heithaus, 2012. al. et W. M. Hayward, ( ofthe leopard 2006c. al. Preyet preferences W. M. Hayward, et W. M. Hayward, 2006a. al. et W. M. Hayward, ( ofthe lion Prey2005. preferences H. I. G. Kerley, and W. M. Hayward, M. Hayward, ArticleUrine 1981. H. F. Harrington, L. J. Gormezano, coarctata Saussure. coarctata review. a Odontoceti): dolphins(suborder conservation. for requirements Ecological Carnivora): arrive? kleptoparasites before prey rapidlyconsumable capture to need the or limitations morphological 3 withlion( the overlap Bay. westernHudson seasonin hyaenas. pp. Hall, Chapmanand evolution. and ecology, behavior, 322 biology. evolutionary human for its and relevance (Tanzania) Park National Tarangire onlythe tropics. in primitive area. primitive 09 – – 322. 167 323:

- W. 2006. Prey preferences of the spotted hyaena ( hyaena the spotted of Prey2006. preferences W.

J. Zool. 270: 615 270: Zool. J. -

Lett. to Nat. 391: 479 391: Nat. Lett. to

and Rockwell, R. F. 2013. What to eat now? Shifts in polar bear diet during diet ice bear the in polar now?eat Shifts to What 2013. R.F. Rockwell, and

– al. 2006b. 2006b. al.

Consumption choice by bears feeding on salmon. salmon. on feeding bears by Consumptionchoice -

Predator size and prey size: presumed relationship in the mantid Hierodula Hierodula mantid in the relationship prey and sizepresumed size: Predator

Ecol. Lett. 12: 538 12: Lett. Ecol. Can. J. Zool. 54:1760 Zool. Can.J. Panthera leo Panthera ecological neo ecological Prey preferences of the tiger tiger ofthe Prey preferences - Prey preferences of the cheetah ( ofthe cheetah Preypreferences Prey preferences of the African wild dog African ofthe Preypreferences marking and caching behavior in the wolf. in the behavior caching and marking - prey and competitive interactions between sharks (order Selachii) and and Selachii) sharks(order between interactions competitive and prey – 627.

- –

Ecol. Evol. 3: 3509 3: Evol. Ecol. 481. ). ). - -

taphonomic study of carcass consumption by lions in by consumption carcass of study taphonomic J. Zool. 270: 606 270: Zool. J. – 549. -

– J. J. 1764.

Zool. 253: 53 253: Zool. -

Acta Zool. Sin. 48: 1 Sin. 48: Zool. Acta

Panthera tigris Panthera – 3523. -

– J. Mammal. 87: 1122 87: Mammal. J. 614. Panthera pardus Panthera Acinonyx jubatus Acinonyx 183 Crocuta crocuta Crocuta

– 68. -

–

In: In: 207

Gittleman, J. L. (ed), CarnivoreL. (ed), J. Gittleman, Lycaon pictus Lycaon .

Panthera leo Panthera . -

- Behaviour 76: 280 76: Behaviour –

- J. Zool. 286: 221 286: Zool. J. 19.

Oecologia 127: 372 127: Oecologia

). ). ) and degree ofdietary degree and ) ) (Felidae: Carnivora): (Felidae: Carnivora): ) -

J. Zool. 270: 2 Zool. 270: J. – 1131.

). ). (Canidae: (Canidae:

- in the Idaho the Idaho in

J. Zool. 267: 267: Zool. J.

- –

– Quat. Int. Quat. 231. 288. – 98 382.

– - free 313.

Lamprecht, J. 1978. The relationship between food competition and foraging group size in some larger somelarger in size group foraging and competition food between relationship The 1978. J. Lamprecht, cats. inand dogs obesity managing and Understanding 2006. P. Laflamme,D. Surplus carnivores. by killing 1972. H. Kruuk, C captive in tracing Stereotypic route 2016. al. et J. Kroshko, lazy’. ‘full and When are carnivores 2007. M. J. Jeschke, Acceptedfree Energetics of 1999. A. al. et Nagy, K. Jarman Assessingthe 2013. al. et H. W. D. Müller, wilddog of distribution affecting and density Factors the L. 1997. M. Gorman, L. G. and Mills,M. habi food census and in capture, advances Methodological L. G. 1996. Mills,M. species. two of ecology behavioural comparative hyaenas:Kalahari The L. G. 1990. Mills,M. Wo 1994. R.O. Peterson, and E. McLaren,B. ( rabbits to Afrom rodents diet change of 2004. A.al. et F. Malo, 1984. L. al. et M. MacDonald, Article2017. al. et Re T. Lumetsberger, A step 2012. al. et M. Leenaars, Part A Mol. Integr. Physiol. 164: 129 164: Integr.Physiol. AMol. Part mammalianherbivores. mass in body with gut fill and retentiontime Park. KrugerNational 223 pp. Press, CornellUniversity Africancarnivores. Hyman. 1555 ( 521 4: 201 302: Zool. 24 46: Anim. carnivores. 36: 1283 Pract. styles. hunting and sizes home range 247 Felis silvestris Felis – 277. – 1558. – 562.

In: Gittleman, J. L. (ed), Carnivore behavior, ecology and evolution, Vol 2. 2. Vol evolution, and ecology Carnivorebehavior, L. (ed), J. In: Gittleman,

Conserv. Biol. 11: 11: 1397 Biol. Conserv. Nutrition of the domestic cat, a mammalianca a cat, the domestic Nutrition of - - by evaluating models for estimating prey consumption by leopards. leopards. by consumption prey estimating for models evaluating - step guide to systematically id systematically to guide step – 242. - – ranging mammals, reptiles, and birds. birds. and reptiles, mammals, ranging - – 343.

- 140. Anim. Behav. 117: 197 117: Behav. Anim.

J. Zool. 263: Zool. 401 263: J. lves, moose, and tree rings on Isle Royale. Royale. Isle ringson moose,tree and lves, -

J. Zool. 166: 233 166: Zool. J.

- Bell Principle: Scaling of intake, digestibility, digestibility, ofintake, Scaling Principle: Bell – 1406. -

Oecologia 152: 357 152: Oecologia

arnivora is predicted by species ispredicted arnivora – 407. Oryctolagus cuniculus Oryctolagus – entify all relevant animal studies. animal relevant all entify 244. –

209.

Comp. Biochem. Physiol Biochem. Comp. rnivore. rnivore. - –

Vet. Clin. Small Anim. Small Clin. Vet. 364. ts studies oflarge tsstudies -

Annu. Rev. Nutr. 19: 19: Nutr. Annu.Rev.

Annu. Rev. Nutr. Rev. Annu. ). ). Iswildcat the –

Science - typical typical -

Unwin Unwin - . s in the the sin

J. J. 266: 266: -

Lab. . Pawar, S. et al. 2012. Dimensionality of consumer search space drives trophic interaction strengths. strengths. interaction trophic drives space search consumer Dimensionalityof 2012. al. et S. Pawar, herbivores. in fermentation hindgut foregut of and Comparison R. 1978. Parra, Size 1976. R.T. Paine, D Orme, Accepted( oftigers organization social The 1981. E. M. Sunquist, and Stirling,I. insemi a lions dynamics of Foraging1992. E. P. Stander, cheetah. habitsofNotesthe foraging on 1990. P. Stander, 2006. al. R.et D. Stahler, sleep. mammalian offunctions Clues the to 2005. M. Siegel,J. lion. Serengeti The 1972. G. Schaller, summer in Bay Hudson southwest Bayand of James bears food habits ofpolar The 1975. R.H. Russell, M. Rosenzweig, timecarnivores. in activity scalingof the Foraging reverse 2018. al. et constraints M. Rizzuto, C Johnson, and G. E. Ritchie, 2011. A. al. E. et Plantinga, Articlemixede nonlinear nlme: and 2011. al. et linear J. Pinheiro, Food1990. al. et P. D. Phillips, ecolo The 1983. R.H. Peters, Yellowstone National Park, Wyoming, Wyoming, Park, National Yellowstone autumn. and 247 2: Evol. conservation. cats. nutritionoffor domestic implications 1 38: Res. Brain Behav. 485 486: Nature 205 pp. Press, Institution Smithsonian folivores. ecology ofarboreal The (ed), Interaction. Pisaster 1 to polar bear ( bear polar to – 36. . et al. 2010. Caper: comparative analyses of phylogenetics and evolution. evolution. and ofphylogenetics comparativeanalyses Caper: 2010. al. et .

McEwan, E. H. 1975. The caloric value of whole ringedseals ( whole value of caloric The 1975. H. E. McEwan,

L. 1966. Community structure in sympatric carnivora. carnivora. in sympatric Community structure L.1966. – - 253.

Ecol. Lett. 12: 982 12: Lett. Ecol. – - 489. limited predation: An observational and experimental approach withthe Mytilus approach experimental and Anobservational predation: limited

Foraging and feeding ecology of the gray wolf( gray ofthe feedingecology Foragingand -

Ecology 57: 858 57: Ecology Estimation of the dietary nutrient profile of free of profile nutrient Estimationdietary ofthe gical implications of body size. size. ofbody gicalimplications . N. 2009. Predator interactions, mesopredator release and biodiversity and release mesopredator interactions, Predator 2009. N. . – 6. - caching in timber wolves, and the question of rules of action syntax. syntax. action of ofrules the wolves,and question in timber caching

– 129. ) ecology and hunting behavior. huntingbehavior. and ecology ) -

U –

998. niversity of Chicago Press. Chicago of niversity – USA. USA. 873.

Br. J. Nutr. 106:Nutr. S35 J. Br. -

J. Nutr. 136: S1923 136: Nutr. J. Panthera tigris Panthera - - arid environment. environment. arid ffects models. ffects

South African J. Wildl. Res. 20: 130 Res.20: Wildl. AfricanJ. South -

Cambridge University Press. University Cambridge -

Nature 437: 1264 437: Nature

Can. J. Zool. 53: 1021 53: Zool. Can.J. – ) i ) -

S48. – J. Mammal. 47: 602 47: Mammal. J. R package version Rpackage n Royal Chitawan National Chitawan nRoyal S1926. lupus Canis - roaming feral cats: possible possible cats: feral roaming -

Phoca hispida Phoca Can. J. Zool. 70: 8 70: Zool. Can.J. -

In: In:

– R package Rpackage 1271. – Montgomery, G. G. G. G. Montgomery, 229.

): Lessons from Lessons):

3 ) in ) relation – – : 1 1027. version 612. -

Nat. Ecol. Ecol. Nat. – – – 137. 132. 21.

5 .2: .2: -

van Aarde, R. J. 1980. The diet and feeding behaviour of feral cats, Felis catus at Marion Island. Island. Marion at Felis cats, catus feral of dietfeedingbehaviour and The 1980. R.J. Aarde, van of cost Revisiting the 2016. A. al. M. et Tucker, martens( Japanese in captive time passage Effectsdigesta on intake al. et 2015. offood Y. Tsuji, forests. in ofdiversity tropical Maintenance 1992. J. Terborgh, Cats ofthe World. Wild 2002. F. Sunquist, and E. M. Sunquist, AcceptedDryad. at spreadsheets (Appen material Supplementary material Supplementary Predator 2015. al. et B. Zimmermann, ofmale and diet The sexes? between competition reduce dimorphism size Does A.2007. Zalewski, preyhabits and food in variation seasonal and Geographical A.2005. Zalewski, African wild( dogs 2007. al. R.et Woodroffe, Species 1.0: EltonTraits 2014. al. et Wilman, H. smatr 3 al. et 2012. I. D. Warton, Bivariate line al. et 2006. I. D. Warton, offree diet the assess methodto An2012. al. et advanced B. Wachter, Articlevert terrestrial prey and among size predator between Empirical relationships A.1985. F. Vézina, the over meaningful the prioritize to need the welfare; animal ofpeak Inpursuit 2017. S. Veasey,J. hoarding inFood animals. 1990. S. Wall, Vander surplus killing. killing. surplus widergeog and local at martens pine female martens. conservation. mammals. 257 3: Evol. Ecol. Methods scats. predators. measurable. 123 Res.10: Wildl. AfricanJ. melampus Nepal. Park, -

PLoS One 7: e38066. One7: PLoS -

) and implications for endozoochorous seed dispersal. dispersal. seed endozoochorous for implications and ) - In: Martens and fishers (Martes) in human in (Martes) fishers and Martens In: -

Oecologia 67: 555 67: Oecologia Ecology 95: 2027. 95: Ecology - -

J. Mammal. 88: 181 88: Mammal. J. -

J. Anim. Ecol. 84: 102 84: Anim. Ecol. J.

dix1

- – 259.

an R package for estimation and inference about allometric lines. allometric about inference and anestimation Rfor package – –

- 128. 565. – 2 -

dependent functional response in wolves: from food limitation to to limitation food from wolves: in response functional dependent - 425.

(OIK fitting methods for allometry.for methods fitting –

193.

- – 05488) 05488) Lycaon pictus Lycaon 112.

- carnivory in mammals. mammals. in carnivory level foraging attributes of the world’s birds and birds world’s the of foraging attributes level - raphical scales. scales. raphical

Universit

as well as the original data as a file of several Excel ofseveral file as a welldata the as as original

- altered environments. Springer, pp. 77 pp. Springer, environments. altered ) can subsist on small prey: implications for implications prey: small can subsist on ) y of Chicago Press. yofChicago - -

Biotropica 24: 2 24: Biotropica

University of Chicago Press. Chicago of University -

Acta Theriol. (Warsz). 52: 237 52: (Warsz). ActaTheriol. - - -

ranging large carnivores based on based carnivores large ranging

Biol Mammal Study 40: 13 40: Study Mammal -

J. Evol. Biol. 29: 29: 2181 Biol. Evol. J. . Rev. 81: 259 Rev. . 81:

size of European pine European of size 83 – 292.

- 291.

–

18. – - Martes Martes

ebrate ebrate 2190.

– South South –

98. 250. -

(a) 1. Table see statistics, ( the bushdog size, pack large preylarge and ( bear polar ofthe ecotypes the indicate two Predator members(iM theofpack number 1 Fig. legends Figure Accepted Article

Relationship between predator mass and (a) average prey mass or (b) average prey mass divided by divided mass preyaverage (b) or mass preyaveragemass (a) and predator between Relationship s are grouped sare

accordingiM their to prey

The dotted line represents y=x (predator mass = prey mass). mass prey = y=x(predator line represents dotted The prey Ursusmaritimus Speothos venaticus Speothos

relative to their stomach capacity C. The linked diamonds diamonds linked C.The stomach capacity their to relative (b)

, see text). Note thatto due Note text). see , ) is an outlier in (a) but not in (b). For For in (b). but not (a) isin an outlier )

its comparatively comparatively its

lines. The bright dotted line indicates the (non line indicates dotted bright lines.The the respective indicating lines withdotted dark kleptoparasites), or scavengers without th over (i.e., completely consumed canbe prey the that assuming frequency kill the and capacity), stomach by points, constrained from(upper the prey meal single a assuming frequency ( bear polar ofthe the two ecotypes indicate diamonds linked C.The capacity theirstomach to relative thefor demands 2 Fig. dark grey line the model by Vézina (1985). For statistics, For (1985). Vézina by model thegrey dark line Repository Digital observed Actually

Accepted ArticleUrsusmaritimus

Relationship between predator mass and the kill frequency (kills per day) per frequency(kills killmass the and predator between Relationship species , see text) see ,

) are indicated as crosses. The light grey line represents the model by Peters (1983), the (1983), modelPeters by the lightrepresents grey line The as crosses. indicated are ) kill frequencies reported in the literature (for sources, see Appendix 1 Appendix see sources, in (for the literature reported frequencies kill ‘This article is protected by copyright. All rights reserved.’ - specific . For large prey predators, the two linked data points indicate the kill kill the indicate points linked the data two preylarge predators, For . average prey items. Predators are grou are Predators items. prey average - significant) significant) see Table 2. Table see GLS GLS regression line for small prey predators. predators. prey small for line regression

ped

according to theiriM according to e course of several days,several of course e

necessary to meet energy to necessary

GLS GLS

and the Dryad the and Dryad regression regression prey

their stomach capacity per day, and above 30 kg, predators can afford to only hunt/eat every second day. second onlyevery to hunt/eat canafford kg, predators 30 above and day, per capacity theirstomach than o passage stomach haveshorter a mass. predator 3 Fig. Accepted Article

Comparison of the scaling of scalingof Comparisonthe of

Theoretically ‘This article is protected by copyright. All rights reserved.’ , below 4 kg, predators have to ingest more than their stomach capacity (i.e., (i.e., capacity stomach moretheir than ingest have to kg, 4 predators below , stomach ne day), whereas above 4 kg, predators can afford to eat lessthan eat canto afford kg, 4 predators above whereas neday),

capacity C and the daily prey mass requirement with requirement mass the dailyprey Cand capacity

* iM M variable Dependent (PGLS) squares least generalized phylogenetic iM C and capacity stomach between theon relationship (depending datasets 1 Table legends Table

λ prey prey

significantly different from 1; **from 1; different significantly

Scaling relationships of prey mass (M mass ofprey relationships Scaling

iM iM < 1%C whole iM iM < 1%C whole Dataset

prey prey

> C > C

prey prey

< C < C

Accepted Article λ

significantly different from 0 and 1 and from0 different significantly 27 40 74 27 40 74 n

prey

) or prey mass available for the individual predator (iM predator thefor individual massprey available or ) PGLS GLS SMA PGLS GLS SMA PGLS GLS SMA PGLS GLS SMA PGLS GLS SMA PGLS GLS SMA Stati stic

0* (0) - 0* (0) - 0.321** (0) - 0.189* (0) - 0* (0) - 0.220* (0) - λ

prey - (0.26;0.78) 0.45 (0.20;0.64) 0.36 - (0.03;0.11) 0.06 (0.01;0.04) 0.02 (0.00;0.09) 0.02 (0.01;0.06) 0.03 (0.00;0.02) 0.01 (0.24;1.17) 0.53 (0.27;0.98) 0.52 (0.20;0.78) 0.39 - (0.02;0.10) 0.05 (0.01;0.04) 0.02 (0.00;0.08) 0.02 (0.01;0.07) 0.03 (0.00;0.02) 0.01 a ) using )

(95%CI)

standardized major axis regression (SMA), axis major regression standardized

- (0.90;1.27) 1.08 (1.00;1.39) 1.17 - (0.24;1.04) 0.64 (1.06;1.88) 1.41 (1.18;2.02) 1.60 (1.04;1.80) 1.42 (1.83;2.60) 2.18 (0.99;1.47) 1.23 (0.98;1.41) 1.19 (1.12;1.55) 1.31 - (0.40;1.22) 0.81 1.52 (1.30;2.18) 1.74 (1.17;1.96) 1.56 (1.97;2.79) 2.34 b

(95%CI) prey (1.15;2.00) ) with predator mass with(M ) predator

- <0.001 <0.001 - <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 - <0.001 <0.001 <0.001 <0.001 <0.001 p

0.003

generalized generalized pred ) according to to according ) least squares ( least squares a

M pred b G

in different indifferent LS) or LS)or * (completeconsumption) iM meal) (single iM iM < 1%C whole Dataset indicated. are ofscavenging/kleptoparasitism) absence an assuming days, several (PGLS) least squares the individua to massavailable prey C and 2 Table

λ prey prey

significantly different from 1; **from 1; different significantly

> C > C

Scaling relationships of kill frequency with predator mass(M withpredator frequency ofkill relationships Scaling

prey

< C

; for large prey predators, the kill frequency assuming a single meal per prey (i.e., constrained by C) or a or C) by constrained prey (i.e., meal per single a frequencyassuming the kill prey predators, large ; for

27 27 40 74 n

Accepted Article

significantly different from 0 and 1 and from0 different significantly PGLS GLS SMA PGLS GLS SMA PGLS GLS SMA PGLS GLS SMA Statistic l predator iM l predator

)

using standardized major axis regression (SMA), generalized least squares (GLS) or phylogenetic generalized generalized phylogenetic or (GLS) least squares generalized (SMA), regression axis major standardized using

pred a - (0.19;0.58) 0.33 (0.36;1.24) 0.66 - (0.96;1.06) 1.01 (0.97;1.07) 1.02 - (1.30;5.36) 2.64 (0.23;1.09) 0.50 (2.71;30.70) 9.12 6.95 (18.24;92.03) 41.19

(95%CI) ) according to to according )

(3.31;14.56)

pred b

in different datasets (depending on the relationship between stomach capacity capacity stomach between the relationship on (depending datasets indifferent b ------( 0.20 (0.93;1.77) 1.29 - - - 0.22 0.22 ( 0.51 ( 0.33 ( 0.33 ( 0.66 ( 0.53 ( 1.51

(95%CI) ( - 0.20;0.61) ------0.40; 0.74; 0.34; 0.35; 1.01; 0.86; 1.87;

------0.04) 0.37) 0.31) 0.31) 0.30) 0.20) 1.21)

complete consumption of the prey (i.e. over over (i.e. the prey of completeconsumption - 0.024 0.024 - <0.001 <0.001 - 0.332 0.332 0.001 0.002 0.002 p