The Implications of Food Hardness for Diet in Bats Author(s): L. F. Aguirre, A. Herrel, R. Van Damme and E.

Matthysen Reviewed work(s): Source: Functional Ecology, Vol. 17, No. 2 (Apr., 2003), pp. 201-212 Published by: British Ecological Society Stable URL: http://www.jstor.org/stable/3599176 . Accessed: 10/09/2012 13:27
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Functional

2003 Ecology
17, 201-212

The implications food hardness diet in bats of for

L. F. AGUIRRE*t, A. HERRELI, R. VAN DAMMEJ and E. MATTHYSENJ
*Centro de Biodiversidad Genetica, UniversidadMayor San Simon, PO Box 538, Cochabamba,Bolivia, and y 1, $Universityof Antwerp,DepartmentBiological Sciences, Universiteitsplein B-2610 Belgium

Summary 1. Neotropicalbat communities characterized a broadspeciesdiversity, are which by of can be achievedand maintained resources. only throughpartitioning the available 2. Herepatterns trophicresource withina singleneotropical of utilization savanna bat are investigated.Moreover,the physicalpropertiesof food items (i.e. community its foodhardness withfood size,andwhether differsbetween items hardness), variation are consumedby the batsin this community investigated experimentally. withthe sizeof thefood item,andthat 3. Theresultsshowthatfoodhardness increases distinctdifferences exist in the amountof forceneededto crushdifferentfood items (beetlesvs otherinsectsvs fruits). 4. Using previously dataon biteforcesfromspeciesin the samecommunity published it is exploredwhetherfood hardness mayplaya rolein shapingthe dietsof the batsin The indicatethatfood the community. combineddataon biteforcesandfood hardness in can limitdietarydiversity bats. hardness both directlyand indirectly 5. The results also indicate that dietary specialization may potentiallyresult in a in decrease trophicbreadthfor some speciesthroughits effecton bite performance.
Bite Key-words: force,Chiroptera, feeding Functional Ecology(2003)17, 201-212

Introduction
Over the past few decades, severalstudies have demonstrated the broad diversity of bats in the neotropics, with single sites harbouring up to, or even over, 70 species of bats (Koepcke 1987;Timm et al. 1989;Handley, Wilson & Gardner 1991; Medellin 1993; Patterson, Pacheco & Solari 1996; Simmons & Voss 1998; Lim & Engstrom 2001). Such diversity can probably only be achieved and maintained through an intricatepartitioning of the available resources. Both spatial (e.g. where animals forage)and temporal (when animals forage) patterns of resource utilization are clearly major factors structuring these communities (Heithaus, Fleming & Opler 1975; Bonaccorso 1979; Findley & Black 1983;Fleming 1986).Withinthe three-dimensional habitat matrix, bats can specialize on a certain spatial component thereof, thus reducing or even avoiding competition with the rest of the community (Heithaus et al. 1975; Arlettaz, Perrin & Hausser 1997; Kalko 1997; Bernard, Albernaz & Magnusson 2001; Kalko & Handley 2001). Bats have done this to a large degree and consequently slow fliers, hoverers, gleaners and fast fliers, all with a specialized wing design and flight pattern, are typically part of a bat community (see Norberg 1994). Other spatial aspects such as the use ? 2003British Society Ecological tAuthor to whom correspondenceshould be addressed. E-mail: laguirre@fcyt.umss.edu.bo

and selection of roosts and temporal differences in to foragingpatternshavebeen demonstrated be important elements of resource partitioning in bat communities (Heithauset al. 1975;Fenton 1982;Fenton& Rautenbach 1986). Given the considerable diversity of trophic niches in New World bat communities, partitioning of food resources is probably one of the major elements structuring the community. The explosive, adaptive radiation of certain groups such as phyllostomid bats into previously unexplored trophic niches lies, at least partly, at the basis of the tremendous diversity and complexity of bat communities in the neotropics (Dumont 1999; Freeman 2000). Previous studies have documented how neotropical bats can be grouped into feeding guilds such as insectivores, carnivores, piscivores, sanguinivores, nectarivores and omnivores, which partition the available food resources (see for example Bonaccorso 1979; McKenzie & Rolfe 1986; Neuweiler1989;Fenton 1990;Findley 1993;Kalko 1997). Yet, most feeding guilds are represented by several species of bats, all specializing on similar food items. As within insectivorous, omnivorous and frugivorous guilds many species coexist (e.g. a typical community can have five or more frugivorous bats and might have up to 15 or more insectivores, see Heithaus et al. 1975; Aguirre, Hanagarth & de Urioste 1996;Medellin 1993; Dumont 1999; Simmons & Voss 1998), food resource partitioning at these lower levels must take place too.

201

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L. F Aguirre

et al.

? 2003 British Ecological Society, Functional Ecology, 17, 201-212

Although membersof a guild are, at first glance, utilizingthe same prey (e.g. insects or fruits), food whichmayshaperesource itemsdifferin manyaspects, utilizationpatternsof bats. Not only do food items differin spatial temporal and availability (Kalko,Herre & Handley1996b; Kalko,Handley& Handley1996a; and Julien-Laferriere 1999),but the behavioural physof icalproperties the food, suchas its size,evasiveness and resistanceto mechanicaldeformation,can also for similarprey.Recently, differdrastically seemingly showed how fruit hardness has a Dumont (1999) pronouncedimpact on the feeding behaviour,and might ultimately even determine the food items bats. selectedandutilizedby frugivorous phylostomid Otherauthors(e.g. Freeman1979;Sosa, De Asengao & Soriano 1996) suggestan importantrole for prey size and hardnessin prey selectionin insectivorous bats. Moreover,Goldman & Henson (1977) have to how observed certain speciesof batswereunable eat beetles(i.e.werenot ableto crushthem),although they theseprey. Theseobservations werecapable capturing of indicate that food hardnessmight be importantin at use food determining resource andfoodpartitioning levels within lowerorganizational (i.e. guilds), potentially competitionbetweenmembers resultingin decreased of the sameguild. Theoretical considerations Lucas 1979;Lucas (see of & Luke 1984)suggestthat the physicalproperties between'hard-shelled' food itemsareindeeddifferent prey,such as beetles,and otherinsects(Currey1970; Freeman1979, 1984;Lucas, Corlett & Luke 1985). Similarly,fruits can be groupedinto hard and soft propertiesof the categoriesbased on the structural outer skin (Freeman1988;Dumont 1999).However, whereasmany authors have speculatedabout food hardnessand how it may affectprey use, dental and and cranialmorphology, niche segregation (Freeman 1979, 1984; Sosa et al. 1996), the hardnessof food items has rarelybeen tested (but see Dumont 1999). Moreover,the relationsbetweenfood hardnessand size remain unexploredto date, but may have an impactupon food selection(butsee Herrel important et al. 1999,2001).As previously by suggested Dumont of 1999),the assessment the physical properties (1997, could offer of food items (size, hardness, toughness) in of into tremendous insights therange variation cranial and mandibularmorphology and food utilization If amongfrugivores. fooditemsdo showlarge patterns the thereof in differences physical properties, demands of and on thecranial morphology theperformance the are feedingapparatus probablystrong(e.g. Freeman 1979,1988,1995). In the presentstudywe investigate patternsof food resourcepartitioningin a previouslystudied Neowe Moreover, wantedto test tropicalbat community. whetherfood items consumedby the species,at the differences and community guildlevels,showmarked which suchas sizeandhardness, in physical properties of our to improve understanding resource mighthelp

partitioning.We do so by assessingfood hardness for experimentally a widerangeof fooditemsnaturally eaten by bats in the community. Finally,by combining these data with previously published data on bite forces for bats from the same community,we mayplay a rolein shaping explorehow food hardness the diet of different species, and whether trophic specializationcomes at the expense of a reduced et as previously (Aguirre al. trophicbreadth suggested 2002).

Materialsandmethods
STUDY SITE AND SPECIES

We studiedfeedinghabitsof bats froma community Wildlife to Refuge,a seasonally belonging theEspiritu of flooded savannalocated in the central-north the 'Llanosde Moxos'(Beni, Bolivia).The 23 speciesof bats of which dietary data were collected (Table1) constitutemorethan 60%of the total bat community et of the area(Aguirre al. 1996).All dietarydatawere collected theendof thedryseason(August-October) at in whichprecludes fromassessing us seasonal variability or in the surroundings diet. Batswerecaptured inside, of, forestislandswithinthe savannausing mist nets. After capture,bats were evaluatedfor age, sex and reproductivestatus. Adult bats were distinguished from sub-adultsby the degreeof ossificationin the joint metacarpal-phalangeal of the third fingerand females weredetermined directpalpaby reproductive tion of the abdomen.
STOMACH CONTENT ANALYSES

on up Depending theirabundance, to eightindividuals females) perspecies(onlyadultsandnon-reproductive were capturedand preservedfor stomach content wereremoved and Stomachs analyses(seeAppendix). storedin a 75%aqueousethanolsolutionfor analysis. The contentswas removed, placedin a Petridish and analysedunder a Zeiss binocularmicroscope(Carl ZeissInc., Chester, USA). Insectsin the sampleswere in determinedto the family level using descriptions & Borror, Triplehorn Johnson(1989)and by comparcollected collectionof arthropods isonwitha reference the first author at the sites where the bats were by captured.Where possible, the size (length) of the the insectswas determined comparing remainsin by collection. the stomachwithinsectsfromthe reference All prey were classifiedas belongingto one of five lengthcategories(Table2). The stomachcontentsof frugivorousspecies (containingmainly seeds) was in withseedscontained a refidentified comparison by erencecollectionat the BolivianNationalHerbarium. of bats was As the identification diet in frugivorous of basedpurelyon the presence seedsin the stomach, smaller fruits(seedsof large it mightbe biasedtowards fruitsmaynot be ingested).

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Ecomorphology offoodpartitioning in bats

Table1. Summary of in Nomenclature followsAguirre al. (1996). et Table morphometrics thespeciessampled thepresent study. entries means. acronyms thespecies composed thefirstletterof thegenusandthefirstthree are The for are of letters thespecies of name.BiteforcedataweretakenfromAguirre al. (2002).SeeAguirre al. (2002)for samplesizesof biteforcedata et et Bat species
Artibeusjamaicensis Carolliaperspicillata Desmodus rotundus Eptesicusfurinalis Eumopsglaucinus Eumops hansae Glossophagasoricina Micronycterisminuta Molossops temmincki Molossus molossus Molossus rufus Myotis albescens Myotis nigricans

Acronym
AJAM CPER DROT EFUR EGLA EHAN GSOR MMIN MTEM MMOL MRUF MALB MNIG

Mass(g)
46-60 18-10 32-50 8-50 30-10 17-30 20-10 9-20 6-90 21-70 26-60 5-60 4-60

Forearm lenth(mm)
63-10 41-80 58-20 38-60 57-10 39-20 33-70 37-20 31-60 41-60 48-40 33-60 33-10

Skulllength(mm)
3-10 22-80 12-50 15-00 23-70 18-20 20-20 19-20 14.60 18-60 20-50 13-60 13-20

Biteforce(N)
24"96 6-65 8-60 7-30

2-25 2-18 8-34 8-40 2-18 1-27

Myotissimus
Noctilio albiventris Noctilio leporinus Phyllostomus discolor Phyllostomus elongatus Phyllostomus hastatus Platyrrhinuslineatus naso Rhynchonycteris

MSIM
NALB NLEP PDISC PELO PHAS PLIN RNAS

10-00
29-70 76-10 37-70 42-50 88-20 25-10 6-00

38-10
59-70 88-50 63-30 66-10 78-30 46"80 33-80

14-40
21-60 28-90 30-20 29-50 38-40 24-80 12-60

2-88
11-91 19-90 21-61 14-78 68-00 25-40
-

Sturnira lilium
Tonatiasyilvicola

SLIL
TSIL

21-60
26-00

42-10
52-50

22-70
26-00

7-72
21.63

Table2. Range of lengths of the insects found in the and bats stomachsof insectivorous omnivorous at the study site. Insect lengths were estimatedbased on a reference of at entries are collection insectssampled thesamesite.Table of the absolutenumber preyof thatsize categoryfoundfor eachspecies Rangesize(mm) Species
P hastatus P elongatus P discolor T silvicola N albiventris E. furinalis M. simus M. temminckii M. ater M. molossus

N
4 3 3 5 8 5 2 1 5 4

1-5

5-10

10-15

15-20
2 1

20-25
3 1 1 3

14 2 5 2 1

64 8 4 4 1 1

2 1 2 11 1 1 2 1

3 1

2 3

of crushing strength fooditemswasmeasured usingan isometricKistlerforcetransducer (type 9203, Kistler connectedto a Kistler Inc., Winterthor, Switzerland) 5995, Kistler Inc.; see Herrel chargeamplifier(type of A et al. 2001fora description theset-up). longscrew with a flattened end (surfaceareaof 3 mm2) free was mountedon the forcetransducer, pushedonto the and failure itsexternal of fooditemuntilmechanical surface occurred. insectsthe resistance different For of parts of the preywasmeasured, themaximalresistance and encountered recorded(usuallyfor the head and was A thepro-thorax). similar was procedure usedto record the resistance mechanical to failurefor fruits.Fruits weretestedat several positionsalongthe fruitand the the as highestforceswererecorded indicating maximal amountof forcean animalwouldneedto reducethat fruit.For Piperfruits,the amountof forceneededto 'strip'the fruitsfromthe stalk,as well as the amount of forceneededto crusha singlefruitwas recorded. ANALYSES To investigate betweenthe food dimenrelationships sions and hardness,all data were loglo-transformed and hardnesswas regressedagainstmass, length or width of the food item tested. All food items were subsequentlyclassifiedas belonging to one of five hard soft categories: arthropods beetles), arthropods, (i.e. hardfruits(figs)andsoft fruits(Piper).Solanum fruits wereincludedinto the analysesas a separate category as no a priori predictionsregardingtheir hardness could be made.To test whetherthesefunctionalfood

FOOD HARDNESS

To investigatethe force needed to crush food items consumedby the variousbats in the community,a
wide variety of arthropods and fruits were collected at the field site to represent the entire variety of prey availableto the bats (Table 3). None of the fruits tested was selected on the basis of ripeness. Our hardness values are thus a general indication of the hardness of that fruit type at the end of the dry season. The linear dimensions (length and width) and masses of all food items were recorded before crushing. Food hardness or

? 2003British
Ecological Society, Functional Ecology, 17, 201-212

204 L. F Aguirre et al.

Table3. Size and hardnessof potentialfood items collectedat the studysite (Espiritu,Bolivia).Many of the food items 2 are measured eatenby the batsin the community Tables and 3) (see Food items
Arthropods

Mass(g)

Length(mm)

Width(mm)

Max. force(N)

COLEOPTERA Scarabaeidae Melolontinae sp. 1 sp.2 sp. 3 Rutellinae Aphodiinae Hydrophylidae Dytiscidae 1 sp. Elateridae Tenebrionidae Cerambycidae Curculionidae sp. Dytiscidae 2 ODONATA sp. 1 sp. 2
ORTHOPTERA

13 25 12 5 8 2 4 5 1 1 1 1 1 8 1 4 2 8 3 3 11 2 11 6 7 7

2-12 +?0-89 0-24+ 0-21 0-10? 0-05 0-94+ 0-13 0-63? 0-13 0-97? 0-06 0-05? 0-00 0-13+ 0-15 0.00 0.10 0.10 0-30 0-30 0-07? 0-03 0-40 0-73? 0-68 0-10? 0-14 0-28+ 0-47 0-08+ 0-03 3-15? 0-64 1-47? 0-23 2-80? 0-00 8-22? 2-80 ? 15-30 0-00 1-03? 0-30 0-33? 0-11

30-62+ 4-13 ? 18-20 2-08 ? 12-25 1-91 24-60? 0-55 ? 18-50 1-12 40-67+ 1-15 ? 13-75 4-92 17-20+ 6-14 11-00 19-00 16-00 19-00 4-80 ? 14-25 2-31 36-00 53-25? 17-17 22-50+ 6-36 18-75+ 11-96 15-00? 1-00 33-50+ 3-54 14-83 0-88 + 23-00? 0-00 27-17+ 2-64 44-00+ 0-00 ? 10-60 2-20 7-97+ 0-64

? 16-00 2-27 9-84? 1-07 6-42? 0-90 13-60+ 0-89 ? 11-63 1-22 19-00+ 0"00 5-25+ 2-50 5-40+ 2-07 4-00 3-00 5-00 10-00 6-00 5-63? 1-30 18-00 6-25? 1-71 4-50+ 0-71 6-25? 2-82 5-33+ 0-58 14-00+ 0.00 ? 16-27 0-90 28-00? 2-74 26-50+ 0"00 34-00? 0-00 0-44+ 0-05 8-91+ 0-76

34-02+ 11.35 7-86+ 3-51 3-47? 1-35 24-78+ 2-00 14-48+ 2-27 20-83? 6-51 2-25+0-31 7-22+ 5-24 3-00 10-00 20-40 8.90 1-60 2-71? 0-69 3-40 7-45? 3-20 2-65+ 0-49 ? 1-75 1-24 1-67+ 0-38 2-35+ 2-05 4-23? 0-54 2-00? 0-00 5-19? 1-99 16-95+ 1-90 0-83? 0-44 4-70+ 1-31

Blattidae Acrididae Tettigoniidae LEPIDOPTERA Noctuidae HOMOPTERA Cicadidae ARACHNIDAE Theraphosidae

Fruits

Ficusobtusifolia Ficustrigona Ficuseximia Dipterixodorata Piperaduncum Solanum wrightii

categoriesdifferedfrom one another,food hardnesswas regressed against food mass (both loglo-transformed), residuals were calculated and used in an analysis of variance, and subsequent post-hoc test (Duncan test). All analyses were performed in Statistica (Version 5-0, Statsoft Inc.)

Results
DIET The bats used for dietary analysis in the present study representeda subsample (62%)of the total community found in Espiritu(Aguirreet al. 1996).Becauseonly mist netting was used to capture bats, aerial insectivores are probably under-represented in our sample. In total, dietary data were collected for 23 out of the 37 species present in the area (Table 1). Although all trophic guilds were included in our sample, carnivorous bats are under-representedin the analysis. From this group, only Noctilio leporinuswas examined. All three individuals contained only fish in their stomachs. Fish

? 2003British Society, Ecological Functional Ecology, 17, 201-212

were strongly chewed and only small, white fleshy remains and crushed fish scales could be recovered from the stomachs. Consequently no estimate of fish size was possible. A large portion of the insectivorous bat community was analysed for diet (68%) and arthropods belonging to two classes and nine orders were represented (see Appendix). More than half the insects eaten by bats in the community were beetles (Coleoptera). Other important items consumed by insectivorous bats were Diptera (especially individuals of the suborder Nematocera, comprising 10%of all arthropods), ants (Formicidae) and crickets (Gryllidae). Molossid bats (Eumops glaucinus, Molossus rufus and M. molossus) and Noctilio albiventriswere the only species to prey on damselflies(Zygoptera).Within the insectivorousguild, Noctilio albiventriswas the species including the largest arthropod diversity into its diet, with coleopterans constituting more than half of its diet (see Appendix). As mentioned, crickets were consumed frequently by bats, with Tonatia sylvicola showing a distinct preference for them (together with scarabaeid beetles). On

205 Ecomorphology offoodpartitioning in bats

the other hand, vespertilionid molossidbats did and not seemto be veryselective atea wideproportion and of theavailable but in spectrum, typically lowquantities.
Among the omnivorous bats, Phyllostomus hastatus

Table Results theregression 4. of on analyses performed the

food hardness data Slope Intercept
r2

ate a large number of arthropods,predominantly of Regressions fooditemmassvs. its lineardimensions beetles(96%). othertwoomnivores, discolor The P and Piper (N = 7) 0-55 P elongatus, 2-02 bothatea widervariety arthropod 0-56* of Length prey Width 0-21 0-64 0-28 Boththefrugivorous nectarivorous and (seeAppendix). Solanum = 7) (N bats in the samplestudiedhere typicallyhad a low 0-22 1-01 0-87** Length of diversity fooditemsin theirstomach Appendix). (see Width 0-92 0-03 0-95**
lineatusand Artibeus Among the frugivores, Platyrrhinus were eating mostly figs. Sturniralilium and jamaicensis Carollia perspicillata, on the other hand, were found
Figs (N = 30)

to eat only Solanumand Piper fruits, respectively.

Glossophaga soricina seemed to be an exclusively

nectarivorous speciesat our field site at the time of study,as no otherfood was foundin its stomach.The last speciesin the community, CommonVampire the
Bat Desmodus rotundus, was mainly feeding on

0-41 Length Width 1-33 Beetles(N = 81) 0-27 Length Width 0-34 Otherarthropods = 29) (N

1.10 -0-45 1-42 1-15

0-94** 0-93** 0-69** 0-74**

Length Width

0-33 0-24

1-52 0-99

0-53** 0-51**

blood, althoughan intactsmall(<2 mm)waterbeetle (Dytiscidae)was foundin the stomachof one of the As individuals examined. the beetlewas not chewed,it mighthavebeeningestedaccidentally.
PREY DIMENSIONS

Although all bats (with the possible exception of for chewtheirfoodthoroughly, arthropods, Desmodus) the sizeof thepreyitemcouldoftenbe estimated using a reference collectionof arthropods capturedat the same site. Whereasvespertilionid bats (Myotis sp.,
Eptesicus sp.) typically ate small prey, Phyllostomus hastatus and P elongatus consumed mostly medium

of vs. Regressions fooddimensions foodhardness Figs -0-79 Length 1.12 1-22 Width -0-92 Mass 0-45 0-46 Beetles 1-91 -1-47 Length Width 1-45 -0-44 Mass 0-67 1-32 Otherarthropods 0-73 -0-57 Length 0-41 Width 0"05 Mass 0-49 0.18

0-54** 0-34* 0-48** 0-73** 0-63** 0-82** 0-46** 0-07 0-15*

at at significant the a = 0-05level;**significant *Regression the a = 0-01level.

to largeprey(Table Noctilioalbiventris 2). apparently of utilizedtheentirespectrum preysizeswitha dominance of smallpreyin its diet. The two molossidspecies for whichpreysize could be estimatedseemedto eat intermediate-sized as well as some verylarge prey Both prey(notethatall largepreyweregrasshoppers).
Tonatiasylvicola and P discolorhad intermediate-sized

Several preyin theirstomachs. capArtibeusjamaicensis, tured in mist nets carriedfigs which were recovered and measured, allowingan estimateof theirsize. On average, Artibeus carried figs of + 33 g and 7.4 mm in diameter(N = 7). Although there 26.5 + 2.9 thus seemsto be a generaltrendfor the largerbats to eat largerprey,some of the smallerspecies(e.g. M. also molossus) atelargepreyandsomebig speciessuch as N albiventris ate verysmallprey(seeTable2). also
FOOD HARDNESS

masswasthebestpredictor 4). (Fig. 1,Table Forbeetles, of hardness(R2= 0-82), and the loglo of hardness increasedstronglypositivelywith the log10of mass 1-32;slope:0-67).For all otherarthropods (intercept: combined, a significant positive relation between hardness mass,and hardness preylengthwas and and Here prey length was the best predictorof present. hardness -0-57, slope:0-73;R2= 0-46).For (intercept: and figs,a similar highlysignificant positive relationship was observed between hardness one handandmass on and fig diameter the otherhand,withthe diameter on of -0-79, beingthe bestpredictor hardness (intercept: slope: 1-12;R2= 0-54).As the sizerangesof the Piper andSolanum collected weresmall, regression no analysis was performed. An analysisof varianceperformed the residual on hardnessdata (hardness regressed againstmass,both indicated foodcategories that differed loglo-transformed)
significantlyfrom each other in hardness(F4,149 = 57-01;

? 2003British Ecological Society, Functional Ecology, 17, 201-212

Food hardnessestimatesindicatedthat within each food group, significantrelationshipsbetween food itemmassandits lineardimensions wereobserved (i.e. longer or wider items being heavier;see Table4). for Moreover, all preygroupsa significant relationship the between sizeof thefooditem(asindicated either by was mass,lengthor width)and its hardness observed

P < 0-001).Post-hoctests indicatedthat beetleswere significantlyharder than all other prey (see also Table3). Solanum fruitsweresofterthan beetlesbut, harderthan all other quiteunexpectedly, significantly food categories. Hardfruits, figs, including weresimilar to the non-coleopteran arthropodsin hardness,but softer than Solanumand beetles.Piper significantly

206 L. F Aguirre

100 (a)

et al.
z

10.

o
"

"o

El
tarantula
0 .1 . . . ....

0.1

1 Foodmass (g)

10

100 (b)

10
C DO

o o 1 1

K>

c:.,.

0 00

Piper 0.1

10

Foodlength(mm)

100

100 (c)

10
Ino

K 0
[]

000
0"1

l 10 Food width (mm)

Fig. 1. Relationship between food hardness and food dimensions. Fooditemswereclassified belonging one of as to five categories based on their hardness: beetles (white diamonds),other arthropods (blackhexagons),figs (black and circles),Piper(whitesquares) solanum(whitetriangles). For all categoriesfor which a size range was tested a betweenfood size (as indicatedby significantrelationship was mass,lengthor width)and hardness observed(see also Table4). Note how tarantulas very soft for their mass. are For Piper, the length of the frutescencewas measured, the to explaining low hardness lengthratio. fruits were significantly softer than all other food items tested (see also Fig. 1).

Discussion
? 2003British Ecological Society, Functional Ecology, 17, 201-212 DIET ANALYSIS The dietary data gathered here indicate that the bat community at Espiritu can be grouped into a number

of distinct guilds as previously described for other Neotropical bat communities (Willig 1986; Medellin 1993; Kalko 1997). Most species seemed to belong to a single dietary guild. However, it should be noted that sample sizes were low for some species and that samples were only taken at a single time period (end of the dry season). Consequently,as it has been documented that the availability of fruit and insect prey changes seasonally (e.g. see Bonaccorso 1979), we probably underestimated the true trophic diversity of the species in the community. When examining published dietary data for the specieswithin the frugivorousguild, it becomes clear that these species in general are not as selective as suggested by the data gathered in the present study. However, despite the much wider variety of food items typically consumed by these bats, the fruitsfound in their stomach in the present study are a predominant part of the diet (see Bonaccorso 1979; Bonaccorso & Humphrey 1984). Insect eaters such as T sylvicola also seem to be more generalized in certain areas compared with our study site (Humphrey et al. 1983; Kalko et al. 1996b). Although the mollosids are also true insect specialists (e.g. Freeman 1981a), they consumed a wider variety of insect prey than typically reported. Whereas M. molossus is sometimes referred to as a durophagous species eating largely Coleopterans (Freeman 1979, 1981a, 1981b), the individuals in this study ate a wide variety of prey, including ants, flies, Zygoptera, Orthoptera, Hemiptera and Homoptera, with Coleoptera comprising only about 30% of the total prey. Unexpectedly, we found only insect prey in the stomachs of those species generally considered omnivores (P hastatus, P discolor and P elongatus). Nevertheless, other studies have demonstrated that in many cases species such as P hastatus predominantly consume fruit, pollen and insects (Humphrey et al. 1983; Kalko et al. 1996b; Kalko & Condon 1998). Noctilio albiventris seems to be a strict insectivore eating a wide variety of insects (see also Kalko et al. 1996b) and N. leporinus was eating exclusively fish at our site. The presence of an insect in the stomach of one of the D. rotundus was rather surprising and may have been incidental (but see Arata, Vaughan & Thomas 1967). The overall pattern that seems to be appearing by comparing our data with previously published accounts on diet in the same species is that at our study site at the end of the dry season trophic niches seem to be more restricted. As food availability may be limited at this time, competition for resources, and thus also the partitioning thereof, can be expected to be strongest. Additionally, at these periods of food limitation, selection on functional properties that may reduce competition are expected to be strongest. Thus any limitations imposed by food hardness on diet should be reflected by an interaction of diet, food hardness, food item size and bite forces for the species in the community.

207
Ecomorphology offoodpartitioning in bats

FOOD HARDNESS

AND SIZE

IMPLICATIONS

OF FOOD HARDNESS

? 2003British Society, Ecological

Functional Ecology,
17, 201-212

in the Interestingly, datagathered the presentstudyas for well as datapreviously gathered fruitsby Dumont lineardimenthat (1999)indicate food itemsof similar in sionsor massmaydifferstrongly theforceneededto reducethem. As has been demonstrated Dumont by moreforce (1999),freshfigsarefairlyhardandrequire Our to crushthanotherfoodssuchas papaya. datafor unexrather food itemsindicatethatSolanum, natural for is evenharder a givenmass.Piper,on the pectedly, a otherhandcan be considered soft fruit,whichdoes a not require lot of forceto be reduced.Interestingly, a thatCarollia, Piperspecialist, Dumont(1999)showed showed no behaviouraladaptationsin its feeding utilizehard that behaviour wouldallowit to efficiently fruitssuch as figs. Fruithardness mightthus play an bat frugivorous guilds. importantrole in structuring Although the degree of ripeness of fruits was not in assessed the present study,thismayhave specifically As effecton fruithardness. batstypically a considerable fruitsto unripeones,it wouldbe mostinterripe prefer with this estingto testwhether is correlated differences in hardness. variationin food However,there is considerable hardness also within arthropods.As suggestedby beetlesare previousauthors(Freeman1979, 1981a,b) for thanotherarthropods a given muchharder indeed hardness increases much size. Additionally,beetle more rapidlywith mass than for any otherfood item tested(seeTable4). Fora smallinsectivorous nonbat, will arthropods be muchmorebeneficial coleopteran work to reducefor a as they requireless mechanical givensize (butsee La Val& La Val 1980).Indeed,our will data indicatethat, whereasnearlyall insectivores include coleopteransinto their diet, only the larger and speciesof bats,suchas the threePhyllostomus N. will prey on large(>15 mm) coleopterans albiventris, from recovered thestomachs Thelargeinsects 4). (Table of T sylvicola, M. rufusand M. molossuswere all the also 2). (Table Clearly within insecnon-coleopterans food hardnessmay play a role in prey tivorousguild in selectionand ultimately trophicnichepartitioning. relationand Forboth arthropods fruits,significant of shipsbetweenthe dimensions the food itemand its hardnesscould be demonstrated. Biggerfood items will thus typicallyrequire highercrushingforces,and fromthe dietin specieswithlow maythusbe excluded bite forces.Moreover,the scaling of food hardness with food dimensionsmay be differentfor different food types (Table Fig. 1). Interestingly, 4, amongfruof a correlation batsizewithfruitsizehasbeen givores, demonstrated (Bonaccorso1979;Kalkoet al. 1996b). in As previouslygathereddata for bite performance bats (Aguirreet al. 2002) indicate that bite forces withbat size,thisimpliesthatherefood hardincrease bat between size this nessmightbe driving relationship and food size. Big fruits mightjust be too hard for smallbatsto chew.

The fact that largerbatsdo seemto selectlargerprey, bats andthatonlythelarger willeat largecoleopterans suggeststhat food hardnesshas majorconsequences before food processing for diet selection.As extensive is feedingbehaviour swallowing a partof chiropteran in reduced the stomachsof the (all preywerestrongly in animalsexamined the present study;see also Kallen De & & & Gans 1972;Czarnecki Kallen1980; Gueldre De Vree 1984, 1988;Dumont 1999),it is likelythat food hardnesswill place constraintson the diet of more harderpreywill require the animals.Moreover, & foodprocessing Gueldre De Vree1984, extensive (De moredemanding. 1988)and will thus be energetically moretime will probably Moreover, require largerprey to chewwhichcannotbe spenthuntingfor moreprey, or for otheractivities.s That food hardnessis ecologicallyrelevant,and may in fact be limitingthe dietaryscope, in bats is in illustrated Fig. 2. Here,the bite forcesof the bats are (see Aguirreet al. 2002)in the community superthe Additionally, graphs. imposedon thepreyhardness preyfoundin the stomachis indicatedon the largest at among figure.Whereas firstsight, little correlation the maximalfood item size, its hardnessand the bite forces of the frugivoresin the community seems bats in and present,for the insectivorous omnivorous the communityclear and stronginteractions among food sizeandbiteforcecanbe maximal food hardness, omnivores for observed. (P Especially the two smaller and P discolor),the maximalsize of prey elongatus well foundin the stomachcorresponds to the maximal eat size of beetlethat they could potentially basedon their bite forces.Also for some of the insectivorous bats (e.g. Tonatia,Myotis) the largest size of prey foundin the stomachseemsto be determined food by bats hardness bite force.Otherinsectivorous such and seemto as the two Molossus speciesand N. albiventris be eatingbiggerpreythanpredicted theirmaximal by the bite capacity. However, largestpreyclassesfound in the stomachs were non-coleopteranarthropods softerfor their size (see black whichare significantly hexagonson Fig. 2). Only for the largestspecies(P not does food hardness seemto be imposing hastatus) limitsto diet selection. werefoundamongsize,hardWhyno correlations cannotbe answered ness and bite forcefor frugivores is here.Eitherfood hardness not by the datagathered preysize at all in these animalsor other determining factorssuchas the energyor time spentchewingprey interactwith preyhardness makingit less favourable to eat the largestfruits that could still be physically Oneotherpotentially reduced chewed). important (i.e. fruit size might be the energetic factor determining such of requirements flight.Frugivores asA.jamaicensis themoverconsidoften takefruitsand will transport et beforeeatingthem(L. F.Aguirre al. erable distances of As observation). the transport heavyfood personal

208
L. F Aguirre et al.
25 20

(a)
60 A. jamaicensis + P. lineatus 50

(b)
0 o ?
Z 40

15 .

a ( 10 S U. bilobatum Ssoricina20 _2

30 o Tonatia

S. C. perspicillata 5.60 li,,u'rn+

G. soricina 0 2 4 6 8 10 12 14 16 18 oMyotis 0

0 o

10

20

30

40

50

60

70

80

Fruit (g) mass

60-

Insect length (mm)

40
a '2 30 20
0 20P. >

P. hastatus

discolor
P. elongatus

20

40

60

80

100

120

140

Foodlength(mm)

of food dimensions bat bite forces(see Table1). (a) The hardness the fruits and betweenfood hardness, Fig. 2. Relationships in measured relative theirmass.Superimposed to (solid upon this graphare the bite forcesof the frugivores the community and the eatencouldbe determined is represented horizontal by lines).Forone species(A.jamaicensis) sizeof thefruitsactually it food hardness does not seemto be limitingdiet in some of the largerfrugivores, clearly the dashedverticalline.Whereas to and seemsto be restricted the smallest least The restricts in someof the smaller diet speciesG soricina species. nectar-eating are bats in the hardfruits.(b) Insecthardnessvs insectlength. Superimposed bite forces(solid lines) for the insectivorous fromthe stomachs vertical maximal and lines).In contrastto (dashed lengthsof the preyrecovered community the estimated determined the hardness for the whatwas observed frugivores, maximal by preysizeconsumed theseanimalsseemslargely by are and Whereas seemsthatboth Noctilioalbiventris bothspeciesof Molossus eatinglarger it of beetles. preythanpredicted by to of and linesaresituated therightof theintersection beetlehardness thebatbiteforcecurves), vertical theirbiteforces(dashed between 2 to above20 mm.(c)Therelationships thedietdata(Tables and4) showhowthesespeciesswitch softerpreycategories bite vs food hardness food lengthfor all preytested.As in the othertwo graphs forces(solidline)andthemaximalsize(dotted bats Whereas largest the for are (P species hastatus) line)of preyin thestomach superimposed theomnivorous in thecommunity. fromthe stomach(whichwas a beetle)is clearly for can eat all preyavailable, the othertwo species,the largest preyrecovered Whitediamonds, blackhexagons, otherarthropods; black of and determined theinteraction preyhardness biteforce. beetles; by Solanum. tested.As in theothertwographs forces(solidline)andthe bite whitetriangles, circles, whitesquares, prey Piper; figs; bats the for Whereas maximalsize (dottedline) of preyin the stomachare superimposed the omnivorous in the community. for the fromthestomach can (which (P hastatus) eatallpreyavailable, theothertwospecies, largest preyrecovered largest species blackhexagons, of and determined the interaction preyhardness biteforce.Whitediamonds, wasa beetle)is clearly beetles; by whitetriangles, Solanum. blackcircles, whitesquares, otherarthropods; Piper; figs;

? 2003British Ecological Society,

Functional Ecology,

17, 201-212

items is energetically demanding, this may be limiting the size of prey actually consumed by these animals. That food hardness is an ecologically relevant variable becomes obvious when considering bite forces for some of the smaller species. For these animals, a large part of the available dietary spectrum is not available as they are not able to crush these prey (e.g. Myotis species). These data also illustrate that trophic specializations leading to a reduced bite performance (as in the case of G soricina or D. rotundus;see Aguirre et al. 2002) will have strong ecological consequences. As illustrated on Fig. 2, G soricina can thus only exploit the smallest and/or softest prey available(i.e. Piper and

small non-coleopterans), suggesting that trophic specialization may lead to an obligatory reduction in niche breadth. In conclusion, food hardness in itself may clearly be limiting dietary scope in bats, excluding large portions of the available dietary spectrum from small animals (e.g. Vespertilionids, nectar-eating bats). Food items differ in many aspects, which may all play an important role in determining the selection of food by predators. Based on the data in the present study we would like to suggest that experimental measures of food hardness, although often ignored in bat community analysis (but see Dumont 1999), might further our understanding

209 of thesecomplexcommunities. demonstrated As here, the combinationof measurements food hardness of Ecomorphology offoodpartitioning and in vivobite forcesfor batsfromthe samecommuin bats nity mightproveespecially insightful. Acknowledgements
The authorswould like to thank N. Burgosfor field supportandP.andB. Bauerforallowingus to workin Espiritu.The authorswould also like to thank three for anonymousreferees helpfulcommentson an earlier versionof this manuscript. L.EA. was fundedby the BelgianCooperation Program (VLIR-KULeuvenUMSS). Part of this researchwas made possible through a travel grant from the fund for scientific research,Flanders (FWO-Vl)to A.H.; R.V.D. and A.H. arepostdoctoral fellowsof the fundforscientific Flanders(FWO-Vl). research,

Fenton, M.B. (1982) Echolocationcalls and patternsof from huntingand habitatuse of bats (Microchiroptera)
Chillagoe, north Queensland. AustralianJournalof Zoology

30, 417-425. and ecologyof Fenton,M.B. (1990)The foragingbehavior

animal-eating bats. CanadianJournalof Zoology 68, 411-

422. of Fenton,M.B.&Rautenbach, (1986)A comparison the I.L. behavior threespeciesof African of roostingandforaging
insectivorous bats (Rhinolophus hildebrandti Rhinolophidae, Scotophilus borbonicus - Vespertilionidae, and Tadarida midas - Molossidae). Canadian Journal of Zoology 64,

2860-2867.
Findley, J. (1993) Bats: A CommunityPerspective.Cambridge

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Oo

?. 0 "

i. ~3 O

~ 'o~

Appendix
Prey items consumed by bats in Espiritu Species NLEP DROT GSOR PLIN AJAM SLIL CPER RNAS NALB MMIN TSIL PDIS PELO PHAS MALB MNIG MSIM EFUR (7) (4) (1) (3) (3) (5) (2) (5) (2) (8) (1) (6) (4) (7) (4) (3) (8) (3) ARTHROPODA EPHEMEROPTERA ODONATA Zygoptera ORTHOPTERA Blattidae Gryllidae Indet HEMIPTERA Corixidae Belostomidae Reduviidae Indet HOMOPTERA Cercopidae Cicadelidae Delphacidae Fulgoridae Indet COLEOPTERA Carabidae Dytiscidae Hydrophylidae Staphylinidae Scarabaeidae Elateridae Cucujidae Chrysomelidae Curculionidae Indet (Continuedoverleaf) 5 2

2 1 1 2

8 1 1 17

1 2 1

10 1 5 8 33 11 20 5 7 3 1 2 6 1 2 2 3 44 2

1 1 4 2 6 1 1

6 3 2 3 1

1 3 4 3

6 48

2 3

1 1

30

1 1

7 4

1 1 16

t S

t. z 2e

Appendix Continued Species

NLEP DROT GSOR PLIN AJAM SLIL CPER RNAS NALB MMIN TSIL PDIS PELO PHAS MALB MNIG MSIM EFUR M (2) (7) (4) (1) (3) (3) (5) ( (2) (5) (8) (6) (1) (7) (4) (3) (4) (8) (3) LEPIDOPTERA DIPTERA Nematocera Brachycera HYMENOPTERA Formicidae Indet ARACHNIDA Fruits Ficus eximia Ficus sp. Solanum wrightii Piper aduncum Piper sp. Indet (Cecropia?) Fish Blood Nectar Unidentified 3 8 3 6 1 1 2 4 1 2 3 1 1 3 4 1 1 1 18 2 40 3 1 2 1 2 1 1 1 5 3 1 1 5 5 15 1

Arthropod orders are indicated in capitals, families in lower case. Sample sizes are indicated in parentheses under the species abbreviation. Table entries are absolute nu stomachs. For fruits, fish, blood and nectar table entries are the number of stomachs containing those food items. See Table 1 for a list of species abbreviations.