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This article was downloaded by: [UNESP] On: 06 March 2014, At: 07:02 Publisher: Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House, 37-41 Mortimer Street, London W1T 3JH, UK Studies on Neotropical Fauna and Environment Publication details, including instructions for authors and subscription information: http://www.tandfonline.com/loi/nnfe20 Food habits of the ocelot, Leopardus pardalis, in two areas in southeast Brazil Rita de Cassia Bianchi a , Sérgio Lucena Mendes a & Paulo De Marco Júnior b a Departamento de Ciências Biológicas , Universidade Federal do Espírito Santo , Vitória, Brazil b Departamento de Biologia Geral , Universidade Federal de Goiás , Goiania, Brazil Published online: 10 Dec 2010. To cite this article: Rita de Cassia Bianchi , Sérgio Lucena Mendes & Paulo De Marco Júnior (2010) Food habits of the ocelot, Leopardus pardalis, in two areas in southeast Brazil, Studies on Neotropical Fauna and Environment, 45:3, 111-119, DOI: 10.1080/01650521.2010.514791 To link to this article: http://dx.doi.org/10.1080/01650521.2010.514791 PLEASE SCROLL DOWN FOR ARTICLE Taylor & Francis makes every effort to ensure the accuracy of all the information (the “Content”) contained in the publications on our platform. However, Taylor & Francis, our agents, and our licensors make no representations or warranties whatsoever as to the accuracy, completeness, or suitability for any purpose of the Content. Any opinions and views expressed in this publication are the opinions and views of the authors, and are not the views of or endorsed by Taylor & Francis. The accuracy of the Content should not be relied upon and should be independently verified with primary sources of information. Taylor and Francis shall not be liable for any losses, actions, claims, proceedings, demands, costs, expenses, damages, and other liabilities whatsoever or howsoever caused arising directly or indirectly in connection with, in relation to or arising out of the use of the Content. This article may be used for research, teaching, and private study purposes. Any substantial or systematic reproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in any form to anyone is expressly forbidden. Terms & Conditions of access and use can be found at http:// www.tandfonline.com/page/terms-and-conditions

Food habits of the ocelot, Leopardus pardalis, in two areas in southeast Brazil

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This article was downloaded by: [UNESP]On: 06 March 2014, At: 07:02Publisher: Taylor & FrancisInforma Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House,37-41 Mortimer Street, London W1T 3JH, UK

Studies on Neotropical Fauna and EnvironmentPublication details, including instructions for authors and subscription information:http://www.tandfonline.com/loi/nnfe20

Food habits of the ocelot, Leopardus pardalis, in twoareas in southeast BrazilRita de Cassia Bianchi a , Sérgio Lucena Mendes a & Paulo De Marco Júnior ba Departamento de Ciências Biológicas , Universidade Federal do Espírito Santo , Vitória,Brazilb Departamento de Biologia Geral , Universidade Federal de Goiás , Goiania, BrazilPublished online: 10 Dec 2010.

To cite this article: Rita de Cassia Bianchi , Sérgio Lucena Mendes & Paulo De Marco Júnior (2010) Food habits of the ocelot,Leopardus pardalis, in two areas in southeast Brazil, Studies on Neotropical Fauna and Environment, 45:3, 111-119, DOI:10.1080/01650521.2010.514791

To link to this article: http://dx.doi.org/10.1080/01650521.2010.514791

PLEASE SCROLL DOWN FOR ARTICLE

Taylor & Francis makes every effort to ensure the accuracy of all the information (the “Content”) containedin the publications on our platform. However, Taylor & Francis, our agents, and our licensors make norepresentations or warranties whatsoever as to the accuracy, completeness, or suitability for any purpose of theContent. Any opinions and views expressed in this publication are the opinions and views of the authors, andare not the views of or endorsed by Taylor & Francis. The accuracy of the Content should not be relied upon andshould be independently verified with primary sources of information. Taylor and Francis shall not be liable forany losses, actions, claims, proceedings, demands, costs, expenses, damages, and other liabilities whatsoeveror howsoever caused arising directly or indirectly in connection with, in relation to or arising out of the use ofthe Content.

This article may be used for research, teaching, and private study purposes. Any substantial or systematicreproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in anyform to anyone is expressly forbidden. Terms & Conditions of access and use can be found at http://www.tandfonline.com/page/terms-and-conditions

Studies on Neotropical Fauna and EnvironmentVol. 45, No. 3, December 2010, 111–119

ISSN 0165-0521 print/ISSN 1744-5140 online© 2010 Taylor & FrancisDOI: 10.1080/01650521.2010.514791http://www.informaworld.com

NNFEORIGINAL ARTICLE

Food habits of the ocelot, Leopardus pardalis, in two areas in southeast BrazilStudies on Neotropical Fauna and EnvironmentRita de Cassia Bianchia*, Sérgio Lucena Mendesa & Paulo De Marco Júniorb

aDepartamento de Ciências Biológicas, Universidade Federal do Espírito Santo, Vitória, Brazil; bDepartamento de Biologia Geral,Universidade Federal de Goiás, Goiania, Brazil

(Received 18 January 2010; accepted 4 August 2010)

The objective of this study was to compare the diet of the ocelot at two sites in southeastern Brazil: the small(957 ha), isolated Caratinga Biological Station (CBS), Minas Gerais and the large (>44,000 ha) contiguous area,comprised of the Vale do Rio Doce Natural Reserve (VRDNR) and the Sooretama Biological Reserve (SBR). Wecollected 60 scats in CBS from January 1997 to July 2000. Small rodents, small marsupials and primates were themost important items in terms of frequency of occurrence. In terms of biomass consumed, the brown howlermonkey (Alouatta guariba) was the most important item. In the VRDNR/SBR we collected 77 scats from April1995 to September 1996 and from January 1999 to September 2000. The main food items were armadillo(Dasypus sp.), small rodents, teju (Tupinambis merianae), and small marsupials. In VRDNR/SBR the ocelot had amore diverse diet, probably reflecting the diversity of prey species found in this area. The occurrence of ocelots inCBS indicates the adaptive flexibility of this felid to forests fragments, probably facilitated by the high biomass ofpotential prey – in this case, the primate Alouatta guariba.

Keywords: diet; Felidae; Atlantic Forest; Leopardus pardalis; ocelot; Brazil

Introduction

The ocelot, Leopardus pardalis (Linnaeus, 1758), isone of eight species of felids that occur in Brazil. It isfound from extreme southern Texas to southernBrazil and northern Argentina. It is a mid-sizedcarnivore (8–15 kg) with a solitary and territorial life-style (Ludlow & Sunquist 1987). Although legallyprotected, ocelot populations continue to be threat-ened by deforestation, fragmentation, and conse-quent genetic isolation (Oliveira 1994; Oliveira &Bianchi 2008), especially in the Atlantic Forest ofBrazil, where there has been a loss of about 92% ofthe original vegetation (Evolução... 1993). As a car-nivore, the ocelot may be more directly affected byfragmentation and habitat loss, since it putativelyneeds large areas of use. Thus, there are few Brazilianconservation areas which effectively protect popula-tions of this species (Redford & Robinson 1991;Oliveira 1994).

Studies on the diet of ocelots are more commonthan those about other Neotropical felids. However,information about its diet is incomplete for most ofits distribution (Mondolfi 1986; Emmons 1987;Ludlow & Sunquist 1987; Konecny 1989; Sunquistet al. 1989; Crawshaw 1995; Chinchilla 1997; Morenoet al. 2006; Bianchi & Mendes 2007), or is based onnon-standardized data collection (Bisbal 1986; Facure& Giaretta 1996; Ximenez 1982). In the Brazilian

Atlantic Forest, the ocelot was studied in detail onlyin Foz do Iguaçu, Paraná (Crawshaw 1995). Thispaucity of information limits understanding the influ-ence of habitat fragmentation on ocelots. Small mam-mals were the main items consumed in all previouslocations studied, and these were consumed accordingto their local abundance (Bisbal 1986; Emmons 1987;Ludlow & Sunquist 1987). Forest fragmentationcould affect prey abundance. We examined the diet ofocelots in a large intact forest and a small forest frag-ment, so as to identify their primary prey in each areaand compare any differences in their diet.

Materials and methods

Study sites

The Vale do Rio Doce Natural Reserve (VRDNR)includes 21,800 ha and is contiguous to the Sooret-ama Biological Reserve (SBR), of 24,150 ha, which isadministered by the Brazilian Environmental Agency(IBAMA). Both conservation units constitute thelargest remaining forest patch in the state of EspíritoSanto and are considered here as a single study area(19°03′S, 40°05′W). Climate is tropical hot andhumid, with an average annual precipitation of 1.051mm and an average annual temperature of 23°C. Thepredominant vegetation is tropical rain forest of theTertiary tablelands (“Mata de Tabuleiros”), which is

*Corresponding author. Email: [email protected]

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a semi-deciduous, mesophytic forest formation of theAtlantic forest domain. The original vegetation of theVRDNR is typically forest (Vicens et al. 1998). Thevegetation at SBR also consists primarily of forest,though swampy vegetation is found at the edges ofrivers and streams. The mammal fauna of theVRDNR/SBR is nearly intact, including jaguar(Panthera onca), puma (Puma concolor), margay(L. wiedii), oncilla (L. tigrinus), and jaguarundi (Pumayagouaroundi), as well as other carnivores, such asotter (Lontra longicaudis), crab-eating fox (Cerdocyonthous), raccoon (Procyon cancrivorus), coati (Nasuanasua), and kinkajou (Potos flavus) (Chiarello 1999).

Caratinga Biological Station (CBS) (19°50′S,41°50′W, 957 ha), Minas Gerais, Brazil, is a PrivateReserve of Natural Patrimony (Fonseca 1989; Castro2001). Vegetation fits the low-montane Atlanticpluvial forest, with characteristics of semi-deciduousto deciduous forests. Climate, based on Köppen’sclassification, is tropical sub-hot mild humid (AW),with an average temperature of 16.5°C in the coldestmonth (July) and 23.9°C in the hottest month(February). The dry season presents 132.8 mm of pre-cipitation, whereas the wet season presents 839.2 mm(Ferrari 1988). The reserve is surrounded mainly bypasture. There is no information on the population ofocelots or any other carnivorous mammal in theregion, although many species have been seen byresearches in the study area, such as C. thous andL. wiedii; likewise, several scat samples of domesticdogs have been collected.

Data collection and analysis

We determined the diet from fecal samples opportun-istically collected along roads and trails. Field collec-tion occurred monthly from April 1995 to September1996 and from January 1999 to September 2000 atVRDNR/SBR; and in the period from January 1997to September 2000 at CBS.

Because large prey (such as paca or monkey) canbe consumed by an ocelot in several meals and resultin several scats, fecal samples collected in the sameday and containing the same large prey were not con-sidered independent feeding events, thereby ensuringindependence of samples.

Scats were identified as being from ocelots basedon track presence near them, as well as the presenceof ocelot hairs that were likely ingested during self-grooming. The hairs were identified through cuticleand medullar patterns (Quadros 2002). The scatswere oven dried and washed on a sieve. The screenedmaterial (e.g. hair, teeth, and scales) was identified bycomparison with the material deposited at theZoological Collection of Prof. Mello Leitão Biology

Museum (Santa Teresa, Espírito Santo State), throughanalysis of the guard-hair microstructure (Quadros2002), or by consulting a specialist.

We analyzed the importance of each kind of preybased on its frequency (proportion of a prey item inrelation to the total number of items, in percent) andas the frequency of occurrence in the scats (propor-tion of scats that contained a particular prey type, inpercent) (Crawshaw 1995). For estimating the numberof vertebrate taxa used in the diet, we used a Jack-knife procedure (Colwell & Coddington 1994), whichcorrects for sub-sampling bias, allowing estimation ofconfidence intervals, and hypothesis testing. Weevaluated Jackknife using the EstimateS program(Colwell 1997), which also produced a collector curvefrom the output of the Jackknife analysis. We appliedChi-square tests (Zar 1999) to frequencies of occur-rence of taxa in relation to a given site.

To correct for differences in size and weight ofprey in the diet, we estimated biomass consumed foreach taxon. Average weight of each prey was multi-plied by the number of individuals found (evaluatedbiomass = prey’s average weight × number of individ-uals in the scats). We considered the presence of anitem as a single individual except in the case of teeth,where the minimum number of individuals was esti-mated from the number of teeth found in the sample.The relative importance of each prey is expressed as apercentage in relation to the weight of all the items(Crawshaw 1995). We obtained the estimated weightof prey taxa from the literature (Fonseca et al. 1996;Emmons & Feer 1997) or from data on species depos-ited in the collection of the Prof. Mello LeitãoBiology Museum. Biomass for prey larger than 10 kg(Brachyteles hypoxanthus, Mazama sp. and Pecaritajacu) was estimated as 30% of adult mass.

The nomenclature of the mammals followedWilson & Reeder (2005) and Emmons et al. (2002).

Results

Diet at Vale do Rio Doce Natural Reserve/Sooretama Biological Reserve

Analyses of 77 ocelot scats collected in VRDNR/SBRindicated 37 taxa and 148 vertebrate prey items.Mammals were the most frequent items, occurring in96% of the scats, followed by the Squamata (21%)and birds (7%) (Table 1). Among the mammals,rodents were most frequently consumed order with59.5% of occurrence in the scats and representing31.1% of the total of the items, followed by Edentataand Didelphimorphia with 29.9 and 15.6% of occur-rence in the scats; and 15.6 and 8.12% of the itemsconsumed, respectively. The most important prey

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Table 1. Spectrum and frequency of food items found in fecal samples of ocelots at Vale do Rio Doce Natural Reserve/Sooretama Biological Reserve, ES (VRDNR/SBR) and at Caratinga Biological Station, MG (CBS).

ItemMass

(g)

VRDNR/SBR (n = 77) CBS (n = 60)

NFrequency of

occurrence (%)Frequency

(%) NFrequency of

occurrence (%)Frequency

(%)

Rodentia 46 59.46 31.10 52 86.68 41.94Cuniculus paca 8200 11 14.29 7.43 3 5.00 2.42Dasyprocta leporina 2100 8 10.39 5.41 – – –Rodentia n.i. 50 8 10.39 5.41 6 10.00 4.84Calomys sp. 25 4 5.19 2.70 16 26.67 12.90Phyllomys sp. 260 3 3.90 2.03 – – –Cavia sp. 400 2 2.60 1.35 1 1.67 0.81Mus musculus 15 2 2.60 1.35 – – –Trinomys sp. 180 2 2.60 1.35 1 1.67 0.81Sigmodontinae n.i. 5 2 2.60 1.35 1 1.67 0.81Oryzomyinae n.i. 80 1 1.30 0.68 – – –Guerlinguetus ingrami 200 1 1.30 0.68 2 3.33 1.61Sphiggurus sp. 1100 1 1.30 0.68 10 16.67 8.06Rattus rattus 200 1 1.30 0.68 – – –Necromys sp. 35 – – – 5 8.33 4.03Oligoryzomys sp. 50 – – – 4 6.67 3.23Akodon sp. 53 – – – 3 5.00 2.42Didelphimorphia 12 15.58 8.12 20 33.34 16.14Didelphidae n.i. 100 7 9.09 4.73 6 10.00 4.84Marmosa murina 52 2 2.60 1.35 – – –Didelphis aurita 1000 1 1.30 0.68 4 6.67 3.23Metachirus nudicaudatus 280 1 1.30 0.68 – – –Monodelphis domestica 67 1 1.30 0.68 – – –Monodelphis americana 30 – – – 6 10.00 4.84Gracilinanus sp. 30 – – – 4 6.67 3.23Primates 4 5.19 2.70 16 26.67 12.91Callicebus personatus 1350 2 2.60 1.35 – – –Cebus apella 2500 2 2.60 1.35 1 1.67 0.81Alouatta guariba 5700 – – – 12 20.00 9.68Brachyteles hypoxanthus 4000a – – – 3 5.00 2.42Edentata 23 29.87 15.54 7 11.67 5.65Dasypus sp. 2200 23 29.87 15.54 7 11.67 5.65Artiodactyla 7 9.09 4.73 – – –Pecari tajacu 5700a 4 5.19 2.70 – – –Mazama sp. 5000a 3 3.90 2.03 – – –Carnivora 5 6.49 3.39 – – –Procyon cancrivorus 5400 3 3.90 2.03 – – –Eira barbara 4850 1 1.30 0.68 – – –Potos flavus 2600 1 1.30 0.68 – – –Lagomorpha 4 5.19 2.70 3 5.00 2.42Sylvilagus brasiliensis 1000 4 5.19 2.70 3 5.00 2.42Mammalia n.i. 50 6 7.79 4.05 2 3.33 1.61Aves 10 12.99 6.76 15 25.00 12.10Aves n.i. – 10 12.99 6.76 15 25.00 12.10Squamata 31 40.26 20.94 9 15.01 7.27Tupinambis merianae 1500 13 16.88 8.78 1 1.67 0.81Ophidia n.i. – 6 7.79 4.05 4 6.67 3.23Tropidurus gr. torquatus 89 4 5.19 2.70 – – –Polychrus marmoratus 94 2 2.60 1.35 – – –Lacertilia n.i. – 2 2.60 1.35 2 3.33 1.61Colubridae n.i. – 2 2.60 1.35 – – –Ameiva ameiva 97 1 1.30 0.68 1 1.67 0.81Mabuya cf. agilis 9 1 1.30 0.68 – – –Teiidae n.i. – – – – 1 1.67 0.81Invertebrates – 12 15.58 – 11 18.33 –Plants – 13 16.88 – 20 33.33 –Total number of itemsb 148 100 124 100Total number of taxab 37 27

Mass (g) = weight of prey; N = number of items; frequency of occurrence = proportion of scats in which the respective item occurred; frequency = proportion ofthe total number of items; n.i., not identified.aThe weight was considered as 30% of the weight of an adult individual.bWithout invertebrates and plant items.

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types in terms of occurrence frequency were Dasypussp. (armadillo) which occurred in 30% of the scats,followed by Tupinambis merianae (teju), whichoccurred in 17% of scats. Large mammals also occurredin the diet, including deer (Mazama sp.), peccary(Pecari tajacu) and other carnivore species (Eirabarbara, Potus flavus, and Procyon cancrivorus).

In terms of biomass consumed, the most import-ant items in the ocelot’s diet at VRDNR/SBRwere Cuniculus paca, Dasypus sp. and Artiodactyla(Figure 1). Although the frequency of occurrence ofthese in the scats was only 9%, together these taxaaccounted for approximately 70% of biomass con-sumed. Small rodents and squamates contributed rel-atively little to the biomass consumed by ocelots inVRDNR/SBR.

Diet at Caratinga Biological Station

Analyses of 60 ocelot scats collected from CBS indi-cated 27 taxa and 124 vertebrate prey items. Mam-mals were the most frequent items, occurring in 100%of the scats, followed by birds (25%) and Squamata(15%) (Table 1). Among the mammals, rodents werethe most frequently found, with 86% of the occur-rence in the scats and 42% of all items, followed byDidelphimorphia and Primates with 33 and 27% ofoccurrence in the scats, and 16 and 13% of the total ofthe consumed items, respectively. Among the rodents,

the most important taxon was Calomys sp., occurringin 26.7% of the scats and in 13% of the total of theitems. Among the primates, Alouatta guariba (brownhowler monkey) occurred in 20% of the scats and 9%of the total items consumed (Table 1).

In terms of biomass consumed, primates were themost important items, especially A. guariba, followedby C. paca and by small rodents (Figure 2).

Cross-site comparisons

The number of the taxa occurring in ocelot scats atVRDNR/SBR as assessed by the Jackknife procedure(51.8; CI = 52.6–51.1) was about 50% larger than atCBS (34.9; CI = 35.5–34.2). These results differed sta-tistically (P < 0.05) (Figure 3). At both sites ocelotsfed predominately on a few prey types only. Smallrodents, small marsupials and A. guariba were con-sumed significantly more frequently at CBS, whileDasypus sp. and T. merianae were consumed signifi-cantly more frequently at VRDNR/SBR (Table 2).

In terms of biomass, rodents contributed most tothe diet of ocelots at VRDNR/SBR and Primates atCBS (Figure 4). In terms of the frequency of occur-rence in scats, Rodentia, Didelphimorphia and Pri-mates were most consumed at CBS, while Edentata,Carnivora and Artiodactyla were most consumed atVRDNR/SBR, although in both areas, Rodentia wasthe order occurring in most scats (Table 2).

Figure 1. Comparison of the proportions (%) of biomass and frequency of prey items found in ocelot scats (n = 77) at Vale doRio Doce Natural Reserve and Sooretama Biological Reserve, ES.

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Figure 2. Comparison of the proportions (%) of biomass and frequency of prey items found in ocelot scats (n = 60) at CaratingaBiological Station, MG.

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Figure 3. Cumulative number of food items (± SD) detected in fecal samples of ocelot at Vale do Rio Doce Natural Reserveand Sooretama Biological Reserve, ES and Caratinga Biological Station, MG, evaluated by Jackknife procedure.

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Discussion

Diet at Vale do Rio Doce Natural Reserve/Sooretama Biological Reserve and CBS

Despite including a great variety of items in its diet,which reflects the richness of species in the area,

ocelots at VRDNR/SBR concentrated on just a fewtaxa, such as armadillos, rodents and lizards. In thecontext of Roper’s definition (1994), the ocelot is nota specialized carnivore since it feeds on a greatnumber of species, preying by chance on the mostabundant and vulnerable, such as Dasypus sp. andT. merianae. This feeding behavior was also observed inPeru (Emmons 1987), Venezuela (Ludlow & Sunquist1987), Costa Rica (Chinchilla 1997), and Foz doIguaçu (Crawshaw 1995). In that study, Crawshaw(1995) detected also a great importance of Dasypus sp.,T. merianae, and small rodents in the diet. TheNational Park of Foz do Iguaçu (175,000 ha) com-prises a mammalian fauna as diverse as that found atVRDNR/SBR, where rodents and Dasypus sp. werethe most frequent prey in the diet of the ocelot. In thePeruvian Amazon and in Venezuela, rodents werealso the most important item (Bisbal 1986; Emmons1987; Ludlow & Sunquist 1987). In all the locations,small mammals (<1 kg) were the main items con-sumed according to their abundance in the area.

Although Ludlow & Sunquist (1987) consideredocelots as a carnivore specialized on small prey, wefound a different result, since the most importantprey in terms of consumed biomass were armadillos,

Table 2. Differences in the frequency of occurrence of themain items and taxa found in ocelot scats at Vale do RioDoce Natural Reserve/Sooretama Biological Reserve, ES(VRDNR/SBR) and at Caratinga Biological Station, MG(CBS).

Items/taxaVRDNR/SBR (%)

CBS (%) c2 P

Small rodents 30.26 61.67 13.412 <0.00a

Rodentia 59.46 86.68 4.853 0.02a

Small marsupials 15.58 33.34 4.802 0.028a

Alouatta guariba 0.00 20.00 16.670 <0.001a

Primates 5.19 26.67 12.246 <0.00a

Dasypus sp. 29.87 11.67 6.744 0.009a

Tupinambis merianae 16.88 1.67 8.654 <0.00a

Lagomorpha 5.19 5.00 0.005 0.945Carnivora 6.49 0.00 4.098 0.042a

Artiodactyla 9.00 0.00 5.826 0.015a

aSignificant to 5%.

Figure 4. Comparison of the proportional biomass (%) of taxa detected in scats of ocelots at Vale do Rio Doce NaturalReserve and Sooretama Biological Reserve (VRDNR/SBR), ES and Caratinga Biological Station (CBS), MG.

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pacas, agoutis and adult tejus at VRDNR/SBR andprimates at CBS. In Foz do Iguaçu, the mostimportant prey in terms of consumed biomass werenine-banded armadillos (Crawshaw 1995). Crawshawargued that large prey (Mazama sp. and other largespecies) were consumed as carrion, since they aresometimes killed by cars in the park and thus mightbecome an important alternative food source for oce-lots. Notwithstanding, predation on large prey hasbeen attributed to ocelot, primarily large males(Moreno et al. 2006). Estimates of consumed biomassprovide a better criterion to indicate the most import-ant prey than frequency, but the difficulty to deter-mine the prey individual’s age and sex in order tocorrect the assumed body mass can generate large dif-ferences in terms of weight.

Ocelots prey on other carnivores, as has been doc-umented in other sites, such as Foz do Iguaçu, wherethey fed on P. cancrivorus, C. thous, and Galictis cuja;Peru, where they fed on Bassaricyon alleni; and Pan-ama, where they preyed on Procyon sp. and Nasuanarica. However, such consumption is considered cir-cumstantial (Emmons 1987; Crawshaw 1995; Morenoet al. 2006).

The importance of A. guariba in the diet of ocelotsat CBS (see Bianchi & Mendes 2007) is evidenced byboth the high frequency of consumption and the bio-mass of this primate. In this site, the total primatebiomass consumed by ocelots was equivalent to mostof the biomass of all other prey items together. Wehave no knowledge of any other study in whichprimates were consumed by ocelots with such a highfrequency. A similar situation was observed in thePanama Canal zone, where a surprisingly highnumber of sloths and other arboreal prey was foundin the diet of ocelots, given that they have been con-sidered predominantly terrestrial hunters (Morenoet al. 2006).

The high consumption of the brown howler mon-key in CBS may be related to the high densities of thisprey species and its relative vulnerability. Populationdensity estimates of brown howler monkey in CBSare 0.9–1.4 individuals/ha (Hirsch 1995). Brownhowler monkeys in CBS are found in groups of aboutsix individuals (typically one male, two or threefemales, and cubs) and they have young all yearround (Strier et al. 2001). Their main defense strategyis to take refuge in secluded places in the foliage andto perform agonistic vocalization (Mendes 1989).While resting, they stay at a height of 15–20 m, butthey use all strata (0–20 m) for feeding purposes, andfrequently come to the ground for drinking (personalobservations). Nevertheless, we cannot discard thepossibility that the predation on brown howler mon-keys was an isolated event, performed by a single

individual, despite the fact that its consumption wascontinuous, for almost 3 years, and relatively evenlydistributed within the years.

Isbell (1990) observed a sudden short-term increasein mortality of vervet monkeys (Cercopithecus aethiops)due to leopard predation in Amboseli National Parkin Kenya. Isbell (1990) argued that this may havebeen due to an increase in the presence of the preda-tor and an individual preference for particular preyspecies, perhaps as a result of earlier experience withthat prey (Caro 1980).

Cross-site comparisons

We observed a significant difference in the use of preytypes in the two areas, probably reflecting differencesin resource availability. The high consumption ofsmall rodents at CBS can be related to the great abun-dance of this group in the area. Fonseca & Robinson(1990) found a greater abundance of rodents in CBSthan in two other areas, one of 70 ha, and another of36,000 ha. Forty-six percent of the rodents observedin the scats at CBS belonged to only three species(Calomys, Necromys, and Akodon) adapted to openand disturbed areas. This may be a consequence offorest fragmentation. Fragment edge is proportion-ally greater as the fragment size decreases, enablingthe dominance of species of open or disturbed areasin a larger proportion of area in small fragments, thanin the large ones (Pardini 2001). Additionally, preda-tors such as ocelots will spend proportionately greatertime in edges as fragment size decreases, resulting inthe taking of more edge-dominant prey. The lowerconsumption of small rodents and small marsupialsat VRDNR/SBR can also be associated to the abun-dance of energetically more rewarding resources, suchas paca, agouti, armadillo, and teju.

These results suggest that ocelots always maintaina set of predominant items, which represent the great-est part of its diet, but opportunistically increase thenumber of rare items that are used where the fauna isricher, such as at VRDNR/SBR. Indeed, the diversityof prey types at VRDNR/SBR was such that wedetected species which had not been previouslyrecorded from the site with the use of traps (Palma1996) or through visual censuses (Chiarello 1999).Thus, the analysis of felid scats and perhaps those ofother carnivores can be an important complementarytool in the survey of the fauna, particularly of specieswhich occur at low densities (Camardella et al. 2000).

Home ranges of ocelots vary from 1.2 to 31.2 km2

(Navarro 1985; Tewes 1986; Ludlow & Sunquist1987; Emmons 1988; Konecny 1989; Crawshaw &Quigley 1989; Sunquist et al. 1989; Laack 1991;Bianchi 2009). As such, the total area of the CBS

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(9.6 km2) is seemingly too small for the maintenanceof a viable population of ocelots, which probably usethe landscape as a whole, moving between forest.However, the brown howler monkey presence at CBSin high densities may facilitate the persistence of oce-lots at the site. Thus, some fragments – althoughapparently small for the maintenance of viable felidpopulations – could be important refuges and localstocks of prey (Chiarello 2000). On the other hand,the pressure by human hunting on the populations ofspecies that are important items in the diet of ocelots,such as armadillos and pacas, can indirectly threatenocelot populations, even in protected and relativelylarge areas as the VRDNR/SBR.

Acknowledgements

We would like to thank the “O Boticário” Foundationof Nature Protection and the FACITEC for financialsupport. We are grateful to the Prof. Mello Leitão Biol-ogy Museum for technical support, Dr. Albert DavidDitchfield, Natalie Olifiers and Dr. Matthew Gompperfor valuable comments and kind help with the final Eng-lish version, and Juliana Quadros for help with the tech-nique of hair slide preparation. We are especiallygrateful to Patricia Guedes and Alexandre Christoff forthe identification of small mammals.

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