Studies on Neotropical Fauna and Environment

2008, iFirst Article, 1–6

ORIGINAL ARTICLE
Food habits of the Broad-snouted Caiman (Caiman latirostris: Crocodylia, Alligatoridae) in
northwestern Uruguay
C. Borteiroa*, F. Gutiérreza, M. Tedrosa and F. Kolencb
a
Departamento de Fisiologı́a, Facultad de Veterinaria, Universidad de la República, Lasplaces 1550, Montevideo 11600,
Uruguay; bCátedra de Bioquı́mica y Bı́ofı́sica, Facultad de Odontologı́a, Universidad de la República, Montevideo, Uruguay
(Received 13 June 2007; accepted 26 September 2008)

The food habits of the Broad-snouted Caiman (Caiman latirostris) were studied in northwestern Uruguay. The
most common prey were insects, the shrimp Pseudopalaemon bouvieri, the snail Pomacea canaliculata, fish and
birds. Spiders, crabs, amphibians, snakes, turtles and mammals were consumed less frequently. Arthropods were
the most frequent prey for juvenile caiman. Fish and snails were consumed by all size classes. The proportion of
stomach contents with invertebrates decreased with increasing caiman size, and that with vertebrates was greater
in the diet of larger caiman. Diet composition and ontogenetic shift are similar to that of other crocodilians
considered as opportunistic generalist predators.
Downloaded By: [Borteiro, C.] At: 20:45 7 November 2008

Se estudiaron los hábitos alimenticios del Yacaré (Caiman latirostris) en el noroeste de Uruguay. Las presas más
comunes fueron insectos, el camarón Pseudopalaemon bouvieri, el caracol Pomacea canaliculata, peces y aves.
Arañas, cangrejos, anfibios, serpientes, tortugas y mamı́feros fueron consumidos menos frecuentemente. Los
artrópodos fueron las presas más frecuentes de juveniles. Peces y aves fueron consumidos por todos los tamaños.
La proporción de contenidos estomacales con invertebrados disminuyó al aumentar el tamaño de los caimanes, y
la de vertebrados fue mayor en los individuos de mayor tamaño. La composición y cambio ontogenético en la
dieta resultan similares a los de crocodilianos considerados predadores generalistas oportunistas.
Keywords: Caiman latirostris; diet; ontogenetic shift; Uruguay

Introduction Alternatively, Magnusson et al. (1987) stated that the

Crocodilians are top predators of many aquatic
ecosystems that prey on a wide variety of taxa
(Pooley 1992). Resource use by different species
varies considerably according to habitat, season, prey
availability, age and size of individuals (Delany &
Abercombie 1986; Hutton 1987; Magnusson et al.
1987; Thorbjarnarson 1993; Delany et al. 1999; Elsey
et al. 2004). An ontogenetic shift in diet is well
documented for some species. Juveniles eat mainly
invertebrates, principally insects, and the relative
importance of vertebrate prey increases with size
(Magnusson et al. 1987; Thorbjarnarson 1993;
Tucker et al. 1996).
It has been proposed that differences in skull and
snout morphology indicate the occurrence of distinct
ecological types of crocodilians (Brochu 2001). For
example, a slender snout relative to cranial length
has been considered an adaptation to piscivory
(Iordansky 1973; Diefenbach 1979; Thorbjarnarson
1990; Pooley 1992). However, dietary studies on the
slender-snouted species Crocodylus johnstoni showed
that trophic niche was not as narrow as expected
(Webb et al. 1982; Lang 1987; Tucker et al. 1996).
wide range of head shape variation in crocodilians is
related to habitat instead of diet: wide flat heads in
swamp-dwelling species and long thin snouts in those
of riverine habitats.
In contrast to longirostrine crocodilians, the
Broad-snouted Caiman (Caiman latirostris Daudin)
exhibits the shortest snout relative to cranial length
among extant species (Medem & Marx 1955). This
fact has led to speculations on possible ecological
implications, such as a diet based on crustaceans and
turtles (Brazaitis 1973) or a decreased ability to prey
on fish (Diefenbach 1979). It was also suggested that
this species feeds mainly on Pomacea snails
(Diefenbach 1979) but the majority of published
reports about the food habits of C. latirostris consist
of qualitative and non-systematic observations (see
Yanosky 1990).
The only previous study of the diet of C.
latirostris is the work of Melo (2002) in southern
Brazil, based on stomach flushing of 44 specimens.
She found that caimans fed mainly on insects, snails,
birds and mammals, and secondarily on crustaceans,
amphibians, reptiles and fish, characterizing C.

*Corresponding author. Email: borteiro@adinet.com.uy

ISSN 0165-0521 print/ISSN 1744-5140 online
# 2008 Taylor & Francis
DOI: 10.1080/01650520802507572
http://www.informaworld.com
 2 C. Borteiro et al.
latirostris as a generalist predator. Given the scarcity
of data about the trophic ecology of C. latirostris, we
studied its food habits in northwestern Uruguay. Our
objective was to assess whether this extremely blunt-
snouted caiman is a specialized or generalized
predator, and to determine whether an ontogenetic
shift in diet occurs as the caimans grow.
Material and methods
Study area
The study area comprises about 500 km2 of the
Uruguay River basin in the Department of Artigas,
northwestern Uruguay (30u209–30u409S, 57u509–
57u259W). Annual mean temperature in this area is
25uC, and accumulated precipitation is about
1453 mm (available at http://www.meteorologia.com.
uy/estadistica_climat.htm#). The landscape is highly
modified because of agricultural activities, mainly
Downloaded By: [Borteiro, C.] At: 20:45 7 November 2008
rice and sugar cane production.
Surveys and capture techniques
Field surveys were done in impoundments used for
agriculture and cattle ranching over a four-year
period during the austral summer: January to
March 2001, December 2001 to March 2002,
January to February 2003, and January to
February 2004. Caimans were located at night by
spotlighting from a 4.5 m canoe, and captured by a
combination of methods. Small and medium-sized
individuals were caught by hand or with a 4 m
metallic clamp. Adults were noosed.
Dietary analysis
We studied 126 samples obtained by stomach
flushing following the technique described by
Taylor et al. (1978). Caiman snout–vent lengths
(SVL) were measured to the posterior border of the
cloaca. The size range included post-hatchlings to
adults, from 17 to 114 cm of SVL (37–216 cm total
length). After processing, caimans were returned to
their capture location.
Stomach contents were fixed in 10% formalin,
filtered to discard mucus, and examined under a
stereoscopic microscope at 10-fold magnification.
Prey items were grouped in the following categories:
insects, crustaceans, spiders, snails, fish, amphibians,
reptiles, birds and mammals. They were identified to
the lowest taxonomic level possible and classified
either as partially digested (under digestive process),
or as non-digestible prey fragments (remains of
already digested prey) such as hair, feathers, fish
and snake scales, turtle scutes, snail opercula, and
exoskeletons of arthropods.
As the majority of prey were obtained in small
remains it was not possible to accurately determine
prey number, volume or mass. Alternatively, we
analyzed the proportion of stomachs that presented
each food item partially digested. Ontogenetic and
sexual differences were analyzed with the Chi square
test with as many prey categories as possible without
exceeding 20% of expected frequencies (5 (Siegel
1956). In the first case we considered four caiman size
classes (17.8–29.9; 30.0–49.9; 50.0–69.9; >70.0 cm
SVL), and 13 samples obtained in recapture episodes
were excluded to preserve independence of the data.
In the second case, also data obtained from 19 small
individuals that were not confidently sexed were
excluded from the analysis. The statistical analysis of
prey frequency data rely on several assumptions that
according to Magnusson et al. (1987) are: (1) prey of
different categories and volumes remain in the
stomach for similar amounts of time during the
digestive process, regardless of caiman size; (2)
caimans of all size classes had equal access to
different prey types; and (3) prey abundance did not
change greatly over the study period.
To avoid the bias given by the different digestion
rates of prey categories, we separately analyzed the
association of caiman size with each type of prey
(Magnusson et al. 1987). For this analysis, we used
the Spearman r statistic (rs, Siegel 1956), with seven
size classes (17.8–29.9; 30.0–39.9; 40.0–49.9; 50.0–
59.9; 60.0–69.9; 70.0–79.9; >80.0 cm SVL) and the
fraction of individuals in each size class that had
consumed the prey. Prey fragments were excluded
from statistical analysis as they are known to remain
for long periods in crocodilian stomachs (Garnett
1985). Statistical significance was considered when
P,0.05.
Results
Prey diversity
Most stomachs had partially digested prey (122/126,
97%), and most had more than one prey category
(74%). Insects were the most frequent, followed by
freshwater snails, fish, crustaceans, spiders and birds.
Except for fish, fragments were present in a higher
proportion of stomachs than partially digested items.
This was most evident in the case of insects, snails,
spiders, birds, reptiles and mammals (Table 1).
Among identified partially digested insects, the most
common were adult Hemiptera (32%), Orthoptera
(26%), Coleoptera (19%) and immature Odonata
(15%); a few samples had specimens of Diptera (7%)
 Studies on Neotropical Fauna and Environment 3

Table 1. Taxonomic list of prey identified in stomach Table 1. Continued.

contents of Caiman latirostris (n5122), and proportion of
stomachs with prey remains that were partially digested Taxon PD (%) F (%)
(PD, under digestive process) and prey fragments (F).
Values are given only for those taxa for which digestion did Leiuperidae 0.8 –
not prevent identification. Reptiles 0.8 13.1
Chelonia – 1.6
Taxon PD (%) F (%) Squamata
Colubridae 0.8 11.5
Insects 82.0 99.2 Birds 9.8 50.8
Coleoptera 19.0 76.2 Mammals 0.0 6.6
Dytiscidae Unidentified Tetrapoda 12.3 –
Megadytes sp. – – Total invertebrates 91.0 100.0
Hydrophilidae Total vertebrates 46.7 65.6
Dibolocelus sp. – –
Hydrobiomorpha – –
Scarabaeidae – –
Diptera 7.0 – and Ephemeroptera (1%). Snails were represented
Calliphoridae mainly by Pomacea canaliculata, and small specimens
Cochliomyia sp. (larvae) 4.9 0.0 of Biomphalaria straminea were found in two
Muscidae 0.8 –
Nematocera 4.9 –
stomachs. The most common crustacean prey was
Ephemeroptera 1.0 0.0 the small shrimp Pseudopalaemon bouvieri. Only four
Downloaded By: [Borteiro, C.] At: 20:45 7 November 2008

Hemiptera 31.9 79.5 stomachs had remains of crabs of the genus

Belostomatidae Trichodactylus. A total of 57 stomach contents
Belostoma sp. 31.1 79.5
contained partially digested vertebrate prey, 39 of
Notonectidae
Notonecta sp. 0.8 0.8 which had fish (68%) and 12 of them had birds (21%).
Odonata 15.0 63.1 A small proportion of stomach contents had remains
Corduliidae (larvae) – – of snakes (14, only in one case partially digested),
Libellulidae (larvae) – – hair (eight), turtle scutes (two), and a recently
Orthoptera 26.0 52.5
Romaleidae
ingested amphibian (one). From some individuals
Coryacris angustipennis – – we obtained partially digested viscera and muscle
Acrididae tissue of unidentified tetrapoda that most likely
Marellia remipes – – belonged to birds, as most of these prey remains
Gryllidae 10.7 –
were found along with feathers (11/15). A taxonomic
Snails 59.8 73.0
Architaenioglossa list of prey is summarized in Table 1.
Ampullariidae
Pomacea canaliculata 59.0 73.0
Basommatophora Sexual and ontogenetic variation
Planorbidae No significant differences were found between sexes in
Biomphalaria straminea 1.6 0.0
the proportion of stomachs with partially digested
Crustaceans 25.4 27.9
Decapoda main prey categories: insects, snails, crustaceans,
Palaemonidae spiders, fish, birds and other tetrapoda, x259.7, df 6,
Pseudopalaemon bouvieri 24.6 25.4 P.0.05 (Table 2). However, the proportion of stomach
Trichodactylidae contents with partially digested insects, snails, crusta-
Trichodactylus sp. 0.8 2.5
Spiders 13.1 24.6
ceans and vertebrates (Table 3) differed with caiman
Araneae size (x2544.7, df 12, P,0.005). As caiman size
Lycosidae
Diapontia sp. – –
Fish 32.0 26.2 Table 2. Number of stomach contents of Caiman latirostris
Characiformes with partially digested prey discriminated by sex.
Characidae
Serrasalmus sp. – – Prey Male (n549) Female (n541)
Cyprinodontiformes
Insects 44 29
Poeciliidae
Snails 26 29
Cnesterodon decemmaculatus 4.9 –
Crustaceans 14 10
Perciformes
Spiders 12 3
Cichlidae – –
Fish 14 19
Siluriformes
Birds 4 6
Loricariidae – –
Unidentified Tetrapoda 5 7
Amphibians 0.8 –
 4 C. Borteiro et al.

Table 3. Number of stomach contents of Caiman latirostris with partially digested prey, according to snout–vent length
(SVL).

SVL (cm)

17.8–29.9 (n533) 30.0–49.9 (n538) 50.0–69.9 (n522) >70.0 (n516)

Insects 31 35 15 8
Snails 7 25 20 10
Crustaceans 8 12 6 1
Spiders 8 6 1 0
Vertebrates 9 13 18 19

increased, there was a decrease in the proportion of
stomachs with partially digested insects (rs520.96,
P,0.001) and partially digested spiders (rs520.96,
P,0.001). Partially digested birds (rs50.90, P,0.01)
and other tetrapods (rs50.86, P,0.05) were found
more frequently in larger individuals; they were found
in the stomach content of 33% of caimans >70 cm SVL
Downloaded By: [Borteiro, C.] At: 20:45 7 November 2008
for C. latirostris in northwestern Uruguay were
insects, snails, shrimp, fish and birds (Delany &
Abercombie 1986; Magnusson et al. 1987; Pooley
1992; Thorbjarnarson 1993; Santos et al. 1996; Da
Silveira & Magnusson 1999). The stomach flushing
method likely revealed a true picture of caiman diet
although it is claimed to be biased against bigger and

(6/18). The percentage of stomachs with partially
digested snails and fish did not change significantly
with caiman size (rs50.18, P50.70, and rs50.57,
P50.18, respectively). Crustaceans were encountered
only in stomach contents of individuals up to 67 cm
SVL, and the percentage tended to decline with
increasing caiman size (rs520.61, P50.15). There
was no significant association of caiman size categories
with the consumption of aquatic and terrestrial prey
that would require different foraging modes (aquatic
insects, shrimp, snails and fishes versus terrestrial
insects, spiders and non-fish vertebrates) (x253.0, df
3, P.0.05).
Other stomach contents
Variable amounts of plant material were present in all
but one stomach. In about 71% of them we identified
rests of the sub-aquatic Ceratophyllum sp., 49% had
grasses (Poaceae), and 18% the floating species Azolla
filiculoides and/or Ricciocarpus natans. Small pieces
of wood were obtained from some individuals, and
gastroliths were found in the stomach of seven
medium to large-sized individuals. Gastric parasites
of the genus Brevimulticaecum (Nematoda) and/or
trematodes were obtained from 41% of stomachs.
Small planorbid snails (,5 mm), Acari, Diptera,
Hymenoptera, copepods, free-living nematodes, and
parasitic Phthiraptera (bird lice) were considered as
secondarily ingested.
Discussion
Prey diversity
Similar to what was reported in other dietary studies
on crocodilians, we found the principal types of prey
recently ingested prey (Jackson et al. 1974; Fitzgerald
1989). So, evidence from stomach flushing supports
the view that the trophic niche of C. latirostris is
broader than previously thought (Brazaitis 1973;
Diefenbach 1979). In fact, food items identified by us
were similar to those reported for this species in
Brazil (Melo 2002) and Uruguay (Orejas-Miranda
1969; Achaval 1980). However, our results did not
confirm the finding that snails (P. canaliculata) are
the most important resource for C. latirostris as
suggested by Achaval (1980) and Diefenbach (1979,
1981) for Rio Grande do Sul (30uS, 50uW). Also, in
contrast to previous studies by Diefenbach (1979)
and Melo (2002) fish were a common prey for C.
latirostris in Uruguay. The occurrence of fish in 32%
of stomachs with partially digested prey in this study
might even be an underestimation of its true
frequency, because of their rapid digestion (Delany
& Abercombie 1986). For the same reason, amphi-
bians may be underrepresented in the stomach
contents analyzed, a finding reported by Melo
(2002) as well. However, many anuran amphibians
are common in the shoreline and shallow areas of
studied agriculture impoundments.
The larvae of the dipteran genus Cochliomyia
found in a few stomachs in this study probably
indicate the ingestion of carrion, as they are usually
found in dead vertebrates. The plant material in
stomach contents is assumed to be secondarily
ingested by crocodilians during foraging (Webb
et al. 1982; Palis 1989), and in our case it indicates
that C. latirostris forages in vegetated areas. The
small pieces of wood could have the same effect as
gastroliths and non-digestible chitinous fragments,
aiding in the mechanical disaggregation of prey

(Garnett 1985). The detection of parasites in a high
percentage of individuals is a common feature in
crocodilian dietary studies (Magnusson 1985; Delany
& Abercombie 1986; Thorbjarnarson 1993).
Ontogenetic variation in the diet of C. latirostris
The ontogenetic shift from a diet composed pre-
dominantly of small invertebrates to one dominated
by vertebrates agrees with the results obtained by
Melo (2002) and appears to be a generalized pattern
among crocodilians (Webb et al. 1982; Delany &
Abercombie 1986; Hutton 1987; Magnusson et al.
1987; Delany 1990; Thorbjarnarson 1993; Tucker
et al. 1996; Da Silveira & Magnusson 1999; Delany
et al. 1999). The greater relative importance of
vertebrates for larger individuals was due to the
feeding on tetrapods, mostly birds. For example,
water birds from the families Rallidae and Ardeidae
Downloaded By: [Borteiro, C.] At: 20:45 7 November 2008
were abundant at the study sites. Thus, although we
did not estimate prey volume or mass it was evident
that as caimans grow they consumed larger prey.
Nevertheless, large C. latirostris (.70 cm SVL) also
continued to eat small invertebrates such as insects
and snails. Small invertebrates were also frequent in
the diet of adult Paleosuchus trigonatus, P. palpebro-
sus and Osteolaemus tetraspis osborni (Ouboter 1996;
Riley & Huchzermeyer 2000), which are similar in
size to adult C. latirostris. Since the fraction of small
invertebrates that are secondarily ingested by croco-
dilians is almost neglible in dietary studies (Webb et
al. 1982; Tucker et al. 1996), as crocodilians grow
they do not just shift to larger animals but appear to
increase the upper size limit of potential prey and the
frequency of large items while continuing to consume
some small prey (Tucker et al. 1996; Da Silveira &
Magnusson 1999).
Specialist or generalist predator?
Food habits of C. latirostris and its ontogenetic shift
are similar to those of other crocodilians considered as
generalist opportunistic predators. We found no
evidence that its peculiar cranial morphology has led
to specialization in diet, and our observations support
the hypothesis that crocodilian skull morphology is
related to habitat (Magnusson et al. 1987). The broad
and strong head of C. latirostris would allow the
capture of different prey in vegetated habitats by
smashing through the vegetation, which is presumably
an adaptation to living in swamps (Magnusson 1998).
In fact, prey spectrum of C. latirostris (both aquatic
and terrestrial) and the secondary ingestion of diverse
plant material suggest the display of various foraging
modes in lentic and vegetated habitats.
Studies on Neotropical Fauna and Environment 5
Acknowledgements
The authors wish to thank S. Ripoll and M. Silva for
their invaluable help during field work, and
Cooperativa Azucarea del Litoral Norte Uruguayo
(CALNU), landowners and landworkers of Bella
Unión for their hospitality. Marı́a C. Machado, J.C.
Borteiro and H. Etcheverry gave logistic support, G.
Speranza, S. Borteiro, M. Letamendı́a, F. Gagliardi,
M. Borteiro, J.M. Venzal, L.A. Chaves and G. de
Souza assisted in the field. The suggestions made by
W. Magnusson, A. Zillikens and two anonymous
referees greatly improved the manuscript. We are
grateful to C. Piña, L.M. Verdade, L. Lavarello, A.
Ferreira, F. Scarabino and A. Larriera for their
comments, to L.M. Verdade, D. Baldo, A.L. Aquino-
Ortiz, G.J.W. Webb and A. Corrent for sharing
bibliography, and to O. Castro, E. Morelli, E. Lorier,
A. Verdi and M. Simó for the identification of
invertebrates. This work was partially funded by
CIDEC, Facultad de Veterinaria, and CSIC,
Universidad de la República. Caimans were captured
under authorization of División Fauna-MGAP,
Uruguay.
References
Achaval F. 1980. El Yacaré. Bol Mus Nat Hist Nat Montev.
2(29):5–6.
Brazaitis P. 1973. The identification of living crocodilians.
Zoologica (N Y). 58:59–101.
Brochu CA. 2001. Crocodylian snouts in space and time:
phylogenetic approaches toward adaptive radiation. Am Zool.
41:564–585.
Da Silveira R, Magnusson WE. 1999. Diets of Spectacled and
Black Caiman in the Anavilhanas Archipielago, Central
Amazonia, Brazil. J Herpetol. 33(2):181–192.
Delany MF. 1990. Late summer diet of juvenile American
Alligators. J Herpetol. 24(4):418–421.
Delany MF, Abercombie CL. 1986. American Alligator food
habits in Northcentral Florida. J Wildl Manag. 50(2):348–353.
Delany MF, Linda SB, Moore CT. 1999. Diet and condition of
American Alligators in 4 Florida lakes. In: Proceedings of the
21st Annual Conference of the Southeast Association of fish and
wildlife Agencies. p. 220–222.
Diefenbach OC. 1979. Ampullarid gastropod-staple food of
Caiman latirostris? Copeia. 1979(1):162–163.
Diefenbach OC. 1981. Regurgitation is normal in Crocodylia.
Ciênc Cult. 33(1):82–83.
Elsey RM, Trosclair PL, Linscomb JT. 2004. The American
Alligator as a predator of Mottled Ducks. Southeast Nat.
3(3):381–390.
Fitzgerald LA. 1989. An evaluation of stomach flushing techniques
for crocodilians. J Herpetol. 23(2):170–172.
Garnett ST. 1985. The consequences of slow chitin digestion on
crocodilian diet analyses. J Herpetol. 19:303–304.
Hutton J. 1987. Growth and feeding ecology of the Nile Crocodile
Crocodylus niloticus at Ngezi, Zimbabwe. J Anim Ecol. 56:25–38.
Iordansky NN. 1973. The skull of the Crocodilia. In: Gans S,
Parsons TS, editors. Biology of the Reptilia. IV Morphology.
London: Academic Press. p. 201–262.
 6 C. Borteiro et al.
Jackson JF, Campbell HW, Campbell KE. 1974. The feeding
habits of crocodilians. Validity of the evidence from stomach
contents. J Herpetol. 8(4):378–381.
Lang JW. 1987. Crocodilian behaviour: implications for manage-
ment. In: Webb JW, Manolis SC, Whitehead PJ, editors. Wildlife
management: crocodiles and alligators. Sydney (Australia):
Surrey Beatty & Sons and The Conservation Commission of
the Northern Territory. p. 273–294.
Magnusson WE. 1985. Habitat selection, parasites and injuries in
amazonian crocodilians. Amazoniana. 9(2):193–204.
Magnusson WE. 1998. Crocodiles and alligators. In: Cogger HG,
Zweifel RG, editors. Encyclopedia of reptiles and amphibians.
San Diego (CA): Academic Press. p. 224–233.
Magnusson WE, Vieira da Silva E, Lima AP. 1987. Diets of
amazonian crocodilians. J Herpetol. 21(2):85–95.
Medem F, Marx H. 1955. An artificial key to the New-World
species of crocodilians. Copeia. 1955(1):1–2.
Melo MTQ. 2002. Dieta do Caiman latirostris no sul do Brasil. In:
Verdade LM, Larriera A, editors. Conservação e manejo de
jacarés e crocodilos da América Latina. II. Piracicaba (Brazil):
C.N. Editoria. p. 116–125.
Orejas-Miranda BR. 1969. Reptiles. In: Klappenbach MA, Orejas-
Miranda BR, editors. Anfibios y reptiles. Montevideo
Downloaded By: [Borteiro, C.] At: 20:45 7 November 2008
(Uruguay): Nuestra Tierra 11. p. 41–68.
Ouboter PE. 1996. Ecological studies on crocodilians in Suri-
name. Niche segregation and competition in three preda-
tors. Amsterdam (Netherlands): SPB Academic Publishing.
139 p.
Palis JG. 1989. Alligator mississippiensis (American Alligator).
Foraging behavior. Herp Rev. 20(3):69.
Pooley AC. 1992. Dieta y hábitos alimentarios. In: Ross CA,
Garnett S, editors. Cocodrilos y caimanes. Barcelona (Spain):
Encuentro Editorial. p. 76–91.
Riley J, Huchzermeyer FW. 2000. Diet and lung parasites of
swamp forest Dwarf Crocodiles (Osteolaemus tetraspis osborni)
in the northern Congo Republic. Copeia. 2000(2):582–586.
Santos SA, Stoll M, Silva M, Campos Z, Magnusson WE, Mourão
G. 1996. Diets of Caiman crocodilus yacare from different
habitats in the Brazilian Pantanal. Herpetol J. 6:111–117.
Siegel S. 1956. Nonparametric statistics for the behavioral sciences.
New York: McGraw-Hill. 312 p.
Taylor JA, Webb GJW, Magnusson WE. 1978. Methods of
obtaining stomach contents from live crocodilians (Reptilia,
Crocodilidae). J Herpetol. 12(3):415–417.
Thorbjarnarson JB. 1990. Notes on the feeding behavior of the
Gharial (Gavialis gangeticus) under semi-natural conditions. J
Herpetol. 24(1):99–100.
Thorbjarnarson JB. 1993. Diet of the Spectacled Caiman (Caiman
crocodilus) in the Central Venezuelan Llanos. Herpetologica.
49(1):108–117.
Tucker AD, Limpus CJ, McCallum HI, McDonald KR. 1996.
Ontogenetic dietary partitioning by Crocodylus johnstoni during
the dry season. Copeia. 1996(4):978–988.
Webb GJW, Manolis SC, Buckworth R. 1982. Crocodylus johnstoni
in the McKinlay River Area, N. T. I. Variation in the diet, and a
new method of assessing the relative importance of prey. Aust J
Zool. 30:877–899.
Yanosky AA. 1990. Histoire naturelle du Caiman à museau large
(Caiman latirostris), un Alligatoriné mal connu (1). Rev Fr
Aquariol Herpetol. 17(1):19–31.