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Diet of brown-nosed coatis and crab-eating raccoons

from a mosaic landscape with exotic plantations in
southern Brazil
a b b b
Lucas M. Aguiar , Rodrigo F. Moro-Rios , Thiago Silvestre , José E. Silva-Pereira , Diego
c b c d d
R. Bilski , Fernando C. Passos , Margareth L. Sekiama & Vlamir J. Rocha
a
Universidade Federal da Integração Latino-Americana (UNILA), Foz do Iguaçu, Brazil
b
Programa de Pós-graduação em Zoologia, Departamento de Zoologia, Universidade Federal
do Paraná (UFPR), Curitiba, Brazil
c
Programa de Pós-graduação em Ecologia e Conservação, UFPR, Curitiba, Brazil
d
Departamento de Ciências Exatas, Naturais, Educação e Sociedade (DCNES), Universidade
Federal de São Carlos (UFSCAR), Araras, Brazil

Version of record first published: 13 Dec 2011

To cite this article: Lucas M. Aguiar, Rodrigo F. Moro-Rios, Thiago Silvestre, José E. Silva-Pereira, Diego R. Bilski, Fernando C.
Passos, Margareth L. Sekiama & Vlamir J. Rocha (2011): Diet of brown-nosed coatis and crab-eating raccoons from a mosaic
landscape with exotic plantations in southern Brazil, Studies on Neotropical Fauna and Environment, 46:3, 153-161

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 Studies on Neotropical Fauna and Environment
Vol. 46, No. 3, December 2011, 153–161

ORIGINAL ARTICLE
Diet of brown-nosed coatis and crab-eating raccoons from a mosaic landscape with exotic
plantations in southern Brazil
Lucas M. Aguiara*, Rodrigo F. Moro-Riosb , Thiago Silvestreb , José E. Silva-Pereirab , Diego R. Bilskic ,
Fernando C. Passosb,c , Margareth L. Sekiamad & Vlamir J. Rochad
a
Universidade Federal da Integração Latino-Americana (UNILA), Foz do Iguaçu, Brazil; b Programa de Pós-graduação em
Zoologia, Departamento de Zoologia, Universidade Federal do Paraná (UFPR), Curitiba, Brazil; c Programa de Pós-graduação
em Ecologia e Conservação, UFPR, Curitiba, Brazil; d Departamento de Ciências Exatas, Naturais, Educação e Sociedade
(DCNES), Universidade Federal de São Carlos (UFSCAR), Araras, Brazil
Downloaded by [University of Saskatchewan Library] at 17:13 08 September 2012

(Received 21 July 2010; accepted 10 November 2011)

We described the diets of two procyonids, the brown-nosed coati Nasua nasua and the crab-eating raccoon Procyon
cancrivorus, through analysis of stomach contents of road-killed specimens in southern Brazil. We compared them
with previously published dietary information for another syntopic mesopredator, the crab-eating fox Cerdocyon
thous. The landscape of the study area includes native grasslands, forests, exotic tree plantations, and other crops.
Food items were represented by frequency of occurrence (FO) and relative volume (RV). Stomach contents of
23 coatis were analyzed. Animal and plant items were equally frequent, although the volume of plants was greater.
Exotic plant species were consumed more than native plants. Among prey items, invertebrates were more important
than vertebrates, mainly because of the frequency of coleopterans and annelids and the volume of necrophagous
dipteran larvae. Five specimens of raccoons were analyzed, in which animal items had the highest FO and plant
items had the highest RV. Both invertebrate and vertebrate prey had the same FO, although vertebrates had a
higher RV. Our data suggest that these procyonids are opportunistic hypocarnivores, utilizing anthropic sources,
with diet overlap. The coatis also overlapped their diet substantially with the foxes. The stomach-contents analysis
revealed the importance of fruit pulp biomass, soft-bodied animals such as larvae, and also soil, which might
indicate feeding habits such as scavenging and geophagy.
Keywords: Cerdocyon; hypocarnivores; mesopredators; Nasua; omnivores; Procyon

Introduction The diets of these species may overlap substantially

Coatis (Nasua) and raccoons (Procyon) are mid-sized
procyonids that are among the most abundant car-
nivores in the Neotropics (Gompper & Decker 1998;
Nowak 1999; Zeveloff 2002; but see Beisiegel &
Mantovani 2006; Arispe et al. 2008). Coatis are pri-
marily diurnal and scansorial forest dwellers, which
form social bands constituted by cooperatively forag-
ing females and juveniles; most adult males are typi-
cally solitary, except during the female estrus (Olifiers
et al. 2009). Raccoons are generally solitary nocturnal
forest dwellers, terrestrial or cursorial, associated with
streams, and skillful in manipulating food (McClearn
1992; Gompper & Decker 1998; Nowak 1999).
Brown-nosed coatis (Nasua nasua) and crab-eating
raccoons (Procyon cancrivorus) have an omnivorous
diet (Nowak 1999), consuming mainly fruits, inver-
tebrates, and small vertebrates (Redford & Stearman
1993; Gompper & Decker 1998; Alves-Costa et al.
2004; Beisiegel & Mantovani 2006; Gatti et al. 2006).
with those of other mesopredators, and therefore some
inter-species spatial or temporal ecological segrega-
tion is likely to occur (Gatti et al. 2006). Coatis and
raccoons have generalist diets, and both seem to tol-
erate human-disturbed environments, using garbage
and exotic species as food resources (Valenzuela
1998; Nowak 1999; Rocha et al. 2005; Beisiegel &
Mantovani 2006; Gatti et al. 2006; Hirsch 2009).
The majority of dietary studies for coati and raccoon
species are based on the analysis of scats. Data from
analysis of scats will seldom detect soft-bodied inver-
tebrates or fruit pulp, which are easily degraded during
digestion, and might be revealed only by the analysis
of stomach contents or by direct observations (Rocha
et al. 2008; Hirsch 2009).
Mid-sized carnivores are among the mammals
that are most frequently killed by collision with
vehicles (Ford & Fahrig 2007). Although wildlife
mortality on roads poses challenges to conservation

*Corresponding author. Email: lucas.aguiar@unila.edu.br

ISSN 0165-0521 print/ISSN 1744-5140 online

© 2011 Taylor & Francis
http://dx.doi.org/10.1080/01650521.2011.640567
http://www.tandfonline.com
 154 L. M. Aguiar et al.
biology (Forman & Alexander 1998; Coffin 2007),
important biological information can be obtained
through appropriate collection of road-killed fauna.
Here, we describe the diets of road-killed brown-nosed
coatis and crab-eating raccoons in a human-disturbed
area composed of a mosaic of exotic and native veg-
etation in southern Brazil. We also compare the diet
of these procyonids with the diet of another syntopic
mesopredator, the crab-eating fox (Cerdocyon thous).
The samples of all three species were collected in the
same study area and period, but the results for the
foxes are described elsewhere (see details in Rocha
et al. 2008).
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Materials and methods
Study site
The stomachs analyzed were taken from adult speci-
mens killed along a stretch of state highway PR-160
that passes through the Fazenda Monte Alegre. This
farm is located at 24◦ 12 42 S, 50◦ 33 26 W, on the
middle Tibagi River in the municipality of Telêmaco
Borba, state of Paraná, southern Brazil. The area
comprises approximately 127,000 ha, of which 37.2%
is native (mixed ombrophilous forest (Araucaria pine
forest), semi-deciduous forest, and natural grass-
lands), and 53.7% is a mosaic of plantations of the
exotic Pinus spp. and Eucalyptus spp., and the native
Araucaria angustifolia. The developed area of the
farm, where orchards of exotic and native fruits are
located close to human habitations, comprised 9.1%
of the study area. The region has a mean altitude
of 885 m and the climate is mesothermic, humid
subtropical, with annual rainfall between 1300 and
1400 mm (Nakajima et al. 1996). Mazama gouazoubira
is the most frequently road-killed mammal, followed
by N. nasua and C. thous (Zaleski et al. 2009).
Data collection and analysis
All road-killed animals were collected fresh, in daily
collections between 2002 and 2008. The stomachs
were removed and frozen to conserve them. The food
items in the stomachs were analyzed in the laboratory
of the Ecological Park of Klabin S.A. in Telêmaco
Borba municipality. Seven coati stomachs were col-
lected prior to the implementation of a logistic scheme
for the purpose of this study, and therefore the food
items for these stomachs could only be quantified
through frequency of occurrence (FO). For all other
stomachs collected the relative volumes (RV) of their
food items were measured. The total volumes of food
items were measured in graduated test tubes by water
displacement (Rocha et al. 2008). Next, the volume of
each food item was measured separately, and the items
were identified to the lowest possible taxonomic cate-
gory with the aid of a dissecting stereomicroscope. The
plant items were identified by comparison with the ref-
erence collection of the herbarium of the Universidade
Federal do Paraná (UFPR), together with the check-
list of regional flora (Nakajima et al. 1996) and other
botanical references (e.g., Lorenzi 2002). Animal items
were identified using keys (e.g., Buzzi 2002, for inver-
tebrates), comparing to reference collections of the
UFPR Department of Zoology, and consulting a
checklist from a local mammal survey (Reis et al.
2005). Results were expressed as relative volume (RV,
item volume ×100/total volume of all stomachs) and
frequency of occurrence (FO, number of stomachs
that contain an item ×100/total number of stomachs)
of diet items. To compare visually the diets of the
procyonids with the diet of the foxes (30 stomachs ana-
lyzed in Rocha et al. 2008), we plotted the FO data
for these three syntopic species in a Correspondence
Analysis (CA) using the program Statistica (Statsoft).
For this analysis, we grouped the food items at order
level, because a preliminary evaluation indicated that
dietary differences were detectable at higher taxo-
nomic levels. We applied the Levins Index to measure
niche breadth, and the Pianka Index to measure the
dietary overlap of the most commonly sampled species
(coatis and foxes).
All items analyzed were deposited in the
Department of Zoology of UFPR. The remaining
animal tissues were stored frozen in the laboratory of
Klabin S.A. for use in future studies.
Results
Stomach contents of coatis
The stomach contents of 23 coatis were analyzed, and
65 food items were identified (44 animals, 19 plants,
soil, and one non-identified item), distributed in
142 occurrences (102 for animals, 36 for plants, three
for soil, one for the non-identified item) (Table 1). The
FO was calculated for all the stomachs, and the RV
included 16 of them. Both animal and plant items were
frequent (100% and 91.3%, respectively) (Figure 1A).
Plants were most important in terms of relative volume
(72.6%), followed by animals (16.2%) and soil (8.6%)
(Figure 1B). Large amounts of soil occurred in three
stomachs. In two of them, soil co-occurred with earth-
worms and termites, but the other stomach, which
contained the largest amount of soil (45 ml), contained
no prey.
The consumption of exotic plant species was
higher (FO = 69.6%; RV = 64.6%) than that of native
plants (FO = 43.5%; RV = 7.4%). Fruits occurred
in 87% of the stomachs and had a relative volume
 Studies on Neotropical Fauna and Environment 155

Table 1. Frequency of occurrence (FO, in %, n = 23) and relative volume (RV, in %, n = 16) of animal and plant items found in
stomach contents of Nasua nasua (coati) in Paraná, southern Brazil.
FO RV

Taxon Items found n % %

Animals
Platyhelminthes N.I. Entire individual 1 4.3
Nematoda N.I. Body fragments, entire individuals 5 21.7
Annelida
Clitellata
Hirudinoidea spp. Entire individuals 2 8.7
Oligochaeta spp. Body fragments 10 43.5 1.5
Mollusca
Gastropoda spp. Shell fragments 2 8.7 0.3
Arthropoda spp.1 3.1
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Arachnida
Araneae
Lycosidae
Lycosa sp. Abdomen, legs 1 4.3
Araneae spp. Abdomen, cephalothorax, legs 4 17.4
Opiliones spp. Entire individuals 2 8.7
Myriapoda
Chilopoda spp. Entire individuals, legs, segments 2 8.7
Diplopoda sp. Segment of trunk 1 4.3 0.1
Insecta
Blattodea sp. Wings 1 4.3
Coleoptera
Cerambycidae Abdomen, body fragments, eggs, 5 21.7
legs, wings
Ctenoscelis sp. Abdomen, eggs, wings 1 4.3 0.1
Coleoptera spp. (adult) Abdomen 5 21.7
Coleoptera spp. (larvae) Entire individuals 8 34.8 0.1
Dermaptera spp. 2 8.7
Diptera sp. (adult) Head 2 8.7
Diptera spp. (larvae) Entire individuals 6 26.1 5
Hymenoptera
Formicidae Entire individuals 1 4.3
Camponotus sp.
Formicidae spp. (adult) Entire individuals 2 8.7
Formicidae spp. (pupa) Entire individuals 4 17.4
Hymenoptera spp. Antenna, head, legs, wings 6 26.1
Isoptera
Termitidae sp. Entire individuals 1 4.3
Isoptera sp. Entire individuals, head 1 4.3 0.1
Lepidoptera sp. (adults) Wings 1 4.3
Lepidoptera spp. (larvae) Entire individuals 5 21.7
Odonata
Zygoptera sp. 1 4.3
Odonata sp. (nymph) Abdomen 1 4.3
Orthoptera
Proscopiidae sp. Legs, wings 1 4.3
Tettigoniidae spp. 2 8.7
Orthoptera spp. Legs, wings 4 17.4 0.1
Insecta (adult) spp. Abdomen 2 8.7
Insecta spp. (larvae) Entire individuals 1 4.3
Invertebrate N.I. Entire individuals 1 4.3
Aves spp. Beak, feathers, legs 2 8.7 2.5
Rodentia
Muridae
Mus musculus2 Entire individual 1 4.3 1
Cricetidae spp. Feet, incisor, hair, mandible, molars 2 8.7 1.8
Rodentia sp. Hair, tail 1 4.3 0.5
 156 L. M. Aguiar et al.

Table 1. (continued)
FO RV

Taxon Items found n % %

Plants
Cucurbitaceae
Citrullus sp.2 Seeds 1 4.3
Lauraceae
Persea americana2 Pulp 2 8.7 9.5
Bombacaceae
Chorisia speciosa Seeds 1 4.3
Piperaceae
Piper aduncum Seeds 1 4.3
Piper sp. Pulp, seeds 1 4.3
Moraceae
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Ficus sp. Pulp, seeds 1 4.3

Morus nigra2 Pulp, seeds 2 8.7 9.7
Rhamnaceae
Hovenia dulcis2 Pulp, seeds 1 4.3
Rosaceae
Eriobotrya japonica2 Bark, pulp, seeds 9 39.1 28.9
Solanaceae
Vassobia breviflora2 Pulp, seeds 1 4.3
Magnoliopsida N.I. Leaves 2 8.7
Arecaceae
Syagrus romanzoffiana Pulp, seeds 6 26.1 3.8
Poaceae N.I. Leaves 1 4.3
Musaceae
Musa sp.2 Pulp 1 4.3 3.2
Liliopsida N.I.1 Leaves 1 4.3
Liliopsida N.I.2 Leaves 1 4.3
Magnoliophyta N.I. Seeds 1 4.3 0.1
Pinaceae
Pinus sp.2 Leaves 1 4.3
Metaphyta N.I. Leaves 2 8.7 0.5
Persea americana + Eriobotrya1 13.3
japonica
Piper sp. + Vassobia breviflora1 3.6
Non-identified item 1 4.3 2.6
1 Items which were formerly grouped for RV (because the pulp or the small seeds were mixed and it was not possible to measure them separately
in these samples), and later identified to lowest taxonomic level for FO.
2 Exotic species.

of 72.1%, resulting mainly from the consumption of
Eriobotrya japonica, Morus nigra, Persea americana
(all exotic), and Syagrus romanzoffiana (native). Exotic
fruits comprised the bulk of the plant items con-
sumed (FO = 56.5%, and RV = 64.6%). Leaves had
a relatively high frequency (FO = 26.1%), but low vol-
ume (RV = 0.5%). Needles of Pinus occurred in one
stomach.
Among animals, invertebrates (FO = 95.7%; RV =
10.3%) occurred more frequently in stomachs than
vertebrates (FO = 21.7%; RV = 5.8%), mainly
due to the frequency of arthropods (FO = 91.3%;
Figure 1), particularly Coleoptera (FO = 60.9%),
Orthoptera (FO = 30.4%), Araneae (FO = 30.4%),
and annelids (FO = 52.2%), and because of the vol-
ume of necrophagous dipteran larvae (RV = 5%).
Vertebrates were represented by rodents (FO = 17.4%;
RV = 3.3%) and birds (FO = 8.7%; RV = 2.5%).
Domestic trash (paper) was found in one stomach, and
nematodes occurred in five of them.
Stomach contents of raccoons
Of the five raccoon stomachs analyzed, 18 items
(14 animals and four plants) distributed in 22 occur-
rences (15 for animals and seven for plants) were
identified. Animals had higher FO (100%) and plants
had higher RV (74.8%), because of the high consump-
tion of S. romanzoffiana fruits (FO = 60%; RV = 72%)
(Table 2). The consumption of exotic fruits (E. japon-
ica, FO = 20%) and leaves (FO = 20% and RV =
0.3%) was low. Both invertebrate and vertebrate prey
had FO = 60%, although vertebrates had a higher RV
(22.9%).
 Studies on Neotropical Fauna and Environment 157
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Figure 1. (A) Frequency of occurrence (FO) for the main diet items of Nasua nasua (coati) in Paraná, southern Brazil. Bars
represent the frequency of higher item categories, and the frequency of their most frequent sub-item. (B) Relative volume (RV)
for the main items of the diet. Bars represent the volume of higher item categories, and the volume of their most representative
sub-item.
 158 L. M. Aguiar et al.

Table 2. Frequency of occurrence (FO, in %) and relative volume (RV, in %) of animal and plant items found in stomach
contents (n = 5) of Procyon cancrivorus (crab-eating raccoon) in Paraná, southern Brazil.
FO RV

Taxon Items found n % %

Animals
Arthropoda spp.1 1.9
Aeglidae
Aegla castro Cephalothorax, cheliped, gill, legs 1 20 0.3
Araneae sp. Abdomen, legs 1 20
Anisoptera (nymph) N.I. Entire individual, head, mouthparts, segments 2 40
Zygoptera (nymph) N.I. Abdomen, gill 1 20
Odonata (nymph) N.I. Abdomen 1 20
Blattodea N.I. Legs, segments, wings 1 20
Coleoptera sp. Elytron, legs, wings 1 20
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Ensifera (nymph) sp. Abdomen, genital system, legs, wings 1 20

Orthoptera N.I. Leg, thorax, wings 1 20
Pterygota N.I. Pair of wings 1 20
Insecta N.I. Larvae 1 20
Vertebrates
Amphibia sp. Bones of legs, foot, muscles, skin, vertebral column 1 20 1.2
Osteichthyes Bones of rib 1 20
Rodentia sp. Entire individual 1 20 21.7
Plants
Rosaceae
Eriobotrya japonica2 Seeds 1 20 3

Arecaceae
Syagrus romanzoffiana Entire fruit, pulp, seeds 3 60 72
Poaceae sp. Leaves 1 20 0.3
Metaphyta N.I. Pulp, stick 2 40 2.5
1 Items which were formerly grouped for RV (because it was not possible to measure them separately), and later identified to lowest taxonomic
level for FO.
2 Exotic species.
3 Data lost.

Dietary comparison soft-tissue items (Alves-Costa et al. 2004; Alves-Costa

The CA showed a different graphical distribution
of major food items for each of the three species
(Figure 2). Considering only the two species with
the largest sample sizes (coatis and foxes), the
measurements of niche breadth by the Levins index
were 0.6 for coatis and 0.7 for foxes. These species
had a Pianka index of 0.86, indicating that their diets
overlapped considerably.
Discussion
Methodological approaches
This study reinforced that both procyonid species
are omnivores, consuming plant and animal food
items with similar frequencies. Fruits comprised the
largest volume in the stomachs, emphasizing their
high dietary importance. These findings agree with
those from other studies of stomach contents (Redford
& Stearman 1993), but differ from those based on
the analysis of scats, which usually do not quantify
& Eterovick 2007; but see Santos & Beisiegel 2006).
Such differences could be caused by ecological circum-
stances or by methodological artifacts.
Analysis of stomach contents is certainly a
straightforward approach for diet studies because it
allows a detailed characterization of the food, and
a realistic evaluation of the quantitative importance
of dietary items. However, invasive methods such as
killing of specimens may not appear justified for mam-
mals and birds, and may conflict with wildlife regula-
tion protection. Moreover, it may have limitations for
other kinds of analyses, for example, analyses of sea-
sonal or sex-age classes (Bisbal & Ojasti 1980). The
analysis of scats produces results with a lower res-
olution of identification and quantification of food
items, and determining the number of items or their
frequency of occurrence can produce less precise esti-
mates of how much the items contribute to the diet
in terms of biomass ingested (Bueno & Motta-Junior
2009). However, scats may be easier to collect and do
not harm the animal population studied. Therefore,
 Studies on Neotropical Fauna and Environment 159
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Figure 2. Correspondence analysis of the higher item categories in the diet of Nasua nasua (coati) and Procyon cancrivorus
(crab-eating raccoon) (in this study), and Cerdocyon thous (crab-eating fox) (in Rocha et al. 2008; see Methods). Geo: geophagy,
Myr: Myriapoda, Cli: Clitellata (Annelida), Che: Chelicerata (Arachnida), Cru-Fis: crustaceans and fish, Nfr: native fruits, Efr:
exotic fruits, Ver: vertebrates, Gas: Gastropoda, Hex: Hexapoda.

discussions and comparisons of the diet and the for-
aging strategies of these carnivores may be inaccurate
if they are based on only one kind of source (scats
or stomachs) or one kind of analysis. A coherent
approach would involve determining the calorie con-
tent of each food item and calculating its contribution
to the calorie requirements of the animal. So, each
method used to study the diet of an animal has its
advantages, disadvantages and inaccuracies, and con-
stitute a compromise between feasibility, effort and the
quality of the results.
Keeping this in mind, our opportunistic analysis
of the volume of stomach contents from road-killed
animals showed that fruits were more important than
animal matter for these carnivores, classifying them as
hypocarnivores (sensu Van Valkenburgh 2007).
Opportunistic coatis
Coatis proved to be opportunists, as suggested by their
use of exotic fruits and domestic garbage. Indeed,
in this region coatis are frequently seen close to
human settlements, in orchards of exotic and native
trees, in sanitary landfills, and rummaging among
trash (Rocha VJ personal observation). The consump-
tion of the exotic rodent Mus musculus is another
indication of the proximity of coatis to human habi-
tations (Nowak 1999). Although foraging in human-
modified habitats and in garbage may be advantageous
(Fedriani et al. 2001; Hirsch 2009), it also bears health
risks such as the transmission of diseases, obesity
from eating high-calorie food, conflicts with humans,
injuries and road kills (Rocha et al. 2005; Bueno &
Motta-Junior 2009).
 160 L. M. Aguiar et al.
Coatis and raccoons as seed dispersers?
The consumption of large amounts of whole fruits of
S. romanzoffiana was also observed at other, nearby
sites (Hirsch 2009, da Silva et al. 2011). This suggests
that both species have the potential to disperse the
seeds of this important palm, as well as seeds of
other native species in the genera Piper and Ficus.
Efficient seed dispersal by coatis, including seeds of S.
romanzoffiana, has been reported (Peracchi et al. 2002;
Alves-Costa & Eterovick 2007). However, because the
consumption of exotic fruits was also high, coatis
might disperse exotic species with small seeds as well
(e.g., genera Morus, Hovenia, Eriobotrya, Vassobia and
Citrullus), as might the syntopic C. thous (Rocha et al.
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2008).
Uncommon stomach contents
The consumption of most leaves could be acciden-
tal when the animals feed on the ground, although in
the case of Poaceae the leaves could serve to increase
digestion efficiency or provide some nutritional value
(Dietz 1984; Bueno & Motta-Junior 2009). The occur-
rence of exotic Pinus needles associated with the prey
is an indirect indication that these animals forage in
Pinus plantations (confirmed by direct observation,
Rocha VJ), as does the syntopic C. thous (Rocha et al.
2008).
The consumption of soil by coatis is uncommon.
Although the present records may indicate forag-
ing for earthworms and termites, or else accidental
ingestion, the record of one stomach with a large
amount of soil and no associated prey may indicate
the occurrence of geophagy (Krishnamani & Mahaney
2000). Geophagy is common among omnivores, and
is a multifunctional behavior that is presumed to
have a medicinal benefit. In disturbed environments
such as this, the ingestion of soil may help pro-
vide carnivores with minerals (e.g., iron), or help
against parasites (e.g., nematodes), gastroenterolog-
ical illness, or food intoxication (human garbage,
and toxic prey such as myriapods) (Krishnamani &
Mahaney 2000).
The coatis consumed a higher proportion of inver-
tebrates than vertebrates, which is in accordance with
existing information about their natural diet (Clauss
et al. 2006). However, in contrast to other stud-
ies (Redford & Stearman 1993; Valenzuela 1998),
here soft-bodied invertebrates comprised an impor-
tant portion of the coatis’ diet, in accordance with
Hirsch’s (2009) direct observations. Necrophagous
larvae of dipterans had a relatively high RV, but were
never associated with items that indicate scavenging
(dead vertebrates). Scavenging is thought to be rare
among coatis (Hirsch 2009), and although this finding
suggests the possibility of necrophagy, the coatis might
have foraged only on the larvae present in carcasses,
not on the carcasses themselves.
Dietary comparisons
Both coatis and foxes exhibited omnivorous diets that
resulted in broad niche indices. The Pianka index of
food niche overlap between these species indicated that
they shared a considerable portion of their food items.
However, the CA indicated a different distribution of
food items in the diets of the three species. The posi-
tion of coatis was influenced by a high frequency of
earthworms and myriapods whereas the position of
crab-eating foxes was mostly influenced by the con-
sumption of vertebrates, and the crab-eating raccoon
was the only species that consumed fish and crus-
taceans. However, this is only a graphical exploration
of the results, and we cannot determine whether it
might be biased or not.
In sum, the analysis of stomach contents rein-
forced the importance of soft-tissue items such as
fruit pulp and insect larvae in the diets of these
mesopredators. Also, our data reflect the ecological
plasticity (consumption of exotic species and use of
human habitats and garbage) of coatis and raccoons,
which allows them to occupy disturbed and human-
influenced areas. However, whether these habits are
contributing to the high mortality of these meso-
predators on the roads is an issue that needs further
examination. The role of exotic foods in maintaining
these mesopredator populations, and the role of these
animals in the dispersal of these plants are impor-
tant questions to be addressed. As a final recommen-
dation, we suggest that wildlife researchers consider
road-killed animals as a valuable source of biologi-
cal information, especially for those animals whose
diets are difficult to study by direct observation or by
analysis of scats.
Acknowledgments
We thank UFPR, UFSCAR, PUC/RS, and Klabin
S.A. for logistical support. We also thank CAPES for
scholarships to L.M.A., J.E.S.P, and D.R.B.; CNPq
for scholarships to R.F.M.R., T.S., and F.C.P. (F.C.P.,
process 300466/2009-9); and the “Laboratório de
Biodiversidade e Biogeografia de Dipteras” of the
UFPR for larvae identification. We also thank the
SNFE Editors and the reviewers for their help in
improving our manuscript.
 Studies on Neotropical Fauna and Environment 161

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