Diet Composition in Two Sympatric Amphisbaenian Species

(Amphisbaena ibijara and Leposternon polystegum) from the

Brazilian Cerrado
Author(s): Jerriane O. Gomes, Adriano O. Maciel, João C. L. Costa, and Gilda V.
Andrade
Source: Journal of Herpetology, 43(3):377-384. 2009.
Published By: The Society for the Study of Amphibians and Reptiles
DOI: http://dx.doi.org/10.1670/08-187R1.1
URL: http://www.bioone.org/doi/full/10.1670/08-187R1.1

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Journal of Herpetology, Vol. 43, No. 3, pp. 377–384, 2009
Copyright 2009 Society for the Study of Amphibians and Reptiles

Diet Composition in Two Sympatric Amphisbaenian Species

(Amphisbaena ibijara and Leposternon polystegum) from the
Brazilian Cerrado

JERRIANE O. GOMES,1,2 ADRIANO O. MACIEL,2 JOÃO C. L. COSTA,3 AND GILDA V. ANDRADE

Departamento de Biologia, Universidade Federal do Maranhão, Campus do Bacanga, Av. Portugueses, s/n,
CEP 65080-040, São Luı́s, Maranhão, Brazil

ABSTRACT.—Amphisbaenians are important predators of tropical ecosystems, yet the ecology of most
species is poorly known. We studied the reproduction and diet of two species of amphisbaenian from the
Brazilian Cerrado. Also, we investigated whether snout–vent length is sexually dimorphic and whether
differences existed in the soil depth at which the two species were found. Adults of Leposternon polystegum
were larger than adults of Amphisbaena ibijara. Male L. polystegum were larger than females, but A. ibijara
did not show sexual dimorphism. Amphsibaena ibijara individuals were encountered significantly closer to
the surface than individuals of L. polystegum, although there was some overlap in the soil depth used
between species. Termites and larvae Coleoptera predominated in frequency of occurrence and termites in
number of prey in juveniles and adults of both sexes in A. ibijara. In the diet of juvenile L. polystegum
individuals, there was no dominant prey group. In male L. polystegum, ants and earthworms occurred most
frequently, and termites and ants were most important numerically. In females, earthworms and ants
dominated in frequency of occurrence and in number of individuals, respectively. The mean number of prey
items in stomachs of A. ibijara was high and did not differ either ontogenetically or sexually. In L.
polystegum, the mean number of prey items in stomachs was low. We showed that the diet of A. ibijara at
Urbano Santos did not differ between sexes or size classes, whereas in L. polystegum there is a small shift in
these aspects.

Amphisbaenians are squamate reptiles with
elongate and annulated bodies, reduced eyes,
short tail, and, with the exception of species in
genus Bipes, are limbless animals, being highly
specialized for burrowing. Compared to other
terrestrial groups, we know very little about the
biology of burrowing reptiles (Pough et al.,
2004).
Most ecological studies on amphisbaenians
have analyzed preserved specimens in museum
collections (Colli and Zamboni, 1999; Webb et
al., 2000; Kearney, 2003; see also review in
Andrade et al., 2006). This is not surprising
considering the lack of specific methods for
collecting fossorial animals (Gans, 1969). Fur-
thermore, our knowledge about the ecology and
natural history of amphisbaenians is limited to
short reports based on a few individuals (White
et al., 1992; Cruz-Neto and Abe, 1993; Gil et al.,
1
Corresponding Author. E-mail: jerrianegomes@
yahoo.com.br
2
Present address: Programa de Pos-Graduação em
Zoologia UFPA-MPEG, Museu Paraense Emı́lio
Goeldi/CZO, Caixa Postal 399, CEP 66017-970, Belém,
Pará, Brazil.
3
Present address: Programa de Pos-Graduação em
Ecologia, Unicamp, Instituto de Biologia, Rua Mon-
teiro Lobato, 255, CEP 13083-970, Campinas, São
Paulo, Brazil.
1993; Gomes et al., 2005), with some exceptions
(e.g., Martı́n et al., 1990; López et al., 1991; Vega,
2001; Bernardo-Silva et al., 2006).
Within Brazil, there are six genera in the
family Amphisbaenidae: Amphisbaena, Anops,
Aulura, Bronia, Cercolophia, and Leposternon.
The genus Amphisbaena currently includes ap-
proximately 80 described species of rounded-
head amphisbaenians that are widely distribut-
ed in South America, of which 38 occur in Brazil
(Gans, 2005; SBH, 2005). Amphisbaena ibijara,
recently described by Rodrigues et al. (2003), is
known only from three localities in the north-
eastern Brazilian state of Maranhão: Rosário,
São Luı́s, and Urbano Santos (type locality) (G.
V. Andrade, unpubl. data). Species within
Leposternon are characterized by a head that is
highly modified for burrowing, with a shovel-
shaped snout. Leposternon currently includes
approximately seven described species, six of
which are endemic to Brazil, whereas Leposter-
non microcephalum has a wider distribution in
South America. Leposternon polystegum is re-
stricted to the Brazilian northern and northeast-
ern regions (Porto et al., 2000; Gans, 2005).
Studies examining the feeding ecology of
amphisbaenians have increased in the last
decade (Colli and Zamboni, 1999; Vega, 2001;
Kearney, 2003; Bernardo-Silva et al., 2006). The
378 J. O. GOMES ET AL.
diet of Amphisbaena alba from the central region
of the Brazilian Cerrado consists primarily of
invertebrates such as Coleoptera, ants, spiders,
and termites, although a snake (Liotyphlops sp.,
Anomalepididae), the limb of a lizard (Tropi-
durus sp., Tropiduridae), and a hair, possibly
from a rodent, have also been found in gut
contents, suggesting that this species is a
generalist (Colli and Zamboni, 1999). Leposter-
non microcephalum and Leposternon wuchereri
feed mainly on earthworms, whereas L. poly-
stegum feeds on ants and Coleoptera larvae
(Goeldi, 1902; Barros-Filho and Valverde, 1996;
C. M. Zamprogno, M. G. F. Zamprogno, and I.
Sazima, unpubl. data).
It is important to accumulate information
from local populations to enable future com-
parisons and to provide an understanding of
the ecology of these animals, which may shift
according to geographic and environmental
gradients and conditions (e.g., Pianka, 1970;
Vitt and Colli, 1994; Mesquita and Colli, 2003a).
Sympatric species, although they are subject to
the same spectrum of potential prey, may differ
in their diets because of taxonomic differences,
preferred microhabitat, or body size (e.g.,
Klawinski et al., 1994; Vrcibradic and Rocha,
1996). In addition, prey size that an amphisbae-
nian can capture and ingest is presumably
constrained by morphological specializations
for burrowing (Pough et al., 2003).
Despite being one of the world’s hotspots of
biodiversity (Myers et al., 2000), the Brazilian
Cerrado remains poorly known for many
taxonomic groups, including amphisbaenians.
Indeed, almost nothing is known about the
biology of amphisbaenians, except for one study
from Central Brazil (Colli and Zamboni, 1999) and
some biological information provided in the
original description of A. ibijara (Rodrigues et
al., 2003). Therefore, the present study is the first
to provide detailed information about amphis-
baenians in the northeastern Brazilian Cerrado.
Our main objective was to study the feeding
ecologies of A. ibijara and L. polystegum, as well
as to examine ontogenetic and sexual differenc-
es between the species. Furthermore, we inves-
tigated whether sexual dimorphism in snout–
vent length (SVL) exists in the two species, and
whether they occupy different soil depths.
MATERIALS AND METHODS
Study Area.—We carried out this work in a
gallery forest in the Cerrado biome, located at
Fazenda Santo Amaro, municipality of Urbano
Santos (03u129280S, 43u249120W; 41 m elevation),
state of Maranhão, Brazil. Gallery Forest is a
typical vegetation landscape within the cerrado
biome that follows perennial or ephemeral
watercourses, and consists of well-drained soil
(Ratter et al., 1997).
The climate of this region is markedly
seasonal, and is classified as Aw type (Köppen
classification system); 89% of the approximately
1,800 mm annual rainfall occurs from December
to May, and the driest months occur from
August to October (IBGE, 1985). The average
annual temperature varies between 26uC and
27uC, and there is a low thermal variation
between seasons (IBGE, 1985).
Collecting Methods and Analysis.—During five
field trips, which took place in March (three
days), May (three days), July (15 days), Sep-
tember (three days), and October (3 days) of
2004, we searched for amphisbaenians in the
gallery forest, digging with hoes to approxi-
mately 30 cm depth in a 16-ha area, between
0800 and 1730 h on each sampling day. When-
ever possible, we measured the depth at which
we found individuals using a ruler. We collect-
ed and euthanized, with a lethal dose of
anesthetic, all captured amphisbaenians follow-
ing standard approved protocols (Anonymous,
1987). Several variables of collected specimens
were measured using vernier callipers (accura-
cy 0.02 mm), including snout–vent length
(SVL), from the tip of the snout to the posterior
end of the cloaca; tail length (TL), from the
posterior end of the cloaca to the posterior tip of
the tail; head length (HL), from the tip of the
snout to the first annulus; head width (HW), the
maximum width of the head measured in the
temporal region; jaw length (JL), from the tip of
the snout to the posterior end of the jaw; and
jaw width (JW), at the maximum width of the
jaw. We fixed specimens in 10% buffered
formalin solution and preserved them in 70%
ethyl alcohol. We deposited voucher specimems
at Coleção de Herpetologia da Universidade
Federal do Maranhão (HUFMA 239–333), state
of Maranhão, Brazil.
We dissected specimens using a magnifying
glass. We considered individuals as reproduc-
tively active by the presence of enlarged testes
or convoluted epididymides in males and
vitellogenic ovarian follicles or oviductal eggs
in females. We estimated the size at which
sexual maturity is reached for males and
females based on the smallest reproductively
active individual for each sex (see Colli and
Zamboni, 1999).
We removed stomachs and, when possible,
identified their contents to ordinal level and
recorded the frequency of occurrence (number
of stomachs containing the prey category,
divided by the total number of stomachs) and
the number of items of each prey category per
stomach. We estimated the food niche breadth
(B) using the inverse of Simpson’s diversity
 DIET OF AMPHSIBAENA IBIJARA AND LEPOSTERNON POLYSTEGUM 379

TABLE 1. Summary of biometry (in millimeters) and depth (in centimeters) data for Amphisbaena ibijara and
Leposternon polystegum from a gallery forest in northeastern Brazil. The values are sample size (N), means 6 1
standard deviation, and range.

Amphisbaena ibijara Leposternon polystegum

Variable N Juveniles N Males N Females N Juveniles N Males N Females
SVL 31 122.1 6 25.4 17 210.3 6 23.1 16 200.0 6 17.0 6 177.2 6 9.3 9 384.7 6 58.4 10 276.8 6 66.6
74.3–164.5 166.6–244.1 164.9–227.3 164.3–187.7 295.5–470.1 196.5–419.1
TL 29 13.9 6 3.6 17 21.7 6 7.9 16 21.3 6 7.0 5 9.8 6 0.5 9 17.2 6 2.7 10 13.5 6 2.3
5.0–21.0 17 8.3–31.8 7.4–30.7 9.3–10.6 13.7–21.2 9.7–17.5
HL 31 5.4 6 0.3 16 6.8 6 0.5 16 6.2 6 0.7 5 7.1 6 0.4 9 9.4 6 1.3 9 8.2 6 1.5
4.9–6.0 6.1–7.9 4.0–7.1 6.7–7.7 7.5–11.0 6.5–11.5
JL 31 3.4 6 0.4 16 4.4 6 0.7 16 3.8 6 0.6 5 4.3 6 0.5 9 5.6 6 0.9 10 4.8 6 0.6
2.3–4.5 3.5–6.1 3.2–5.6 3.6–4.8 4.2–7.1 4.0–5.9
HW 31 3.5 6 0.3 16 5.1 6 0.4 16 4.4 6 0.3 5 5.2 6 0.2 9 8.2 6 1.4 10 6.5 6 1.3
2.8–4.3 4.2–5.9 3.5–5.0 5.0–5.5 6.3–9.8 4.9–9.7
JW 31 3.1 6 0.3 16 4.4 6 0.4 16 3.8 6 0.3 5 4.7 6 0.2 9 6.8 6 1.1 10 6.0 6 1.8
2.4–3.8 3.6–5.4 3.0–4.4 4.5–5.0 5.1–8.1 4.7–10.5
DS 21 11.0 6 3.4 12 10.1 6 3.4 11 11.1 6 4.3 4 12.3 6 2.6 6 18.8 6 4.7 2 14.1 6 5.7
6.3–20.7 5.7–16.4 4.5–17.4 9.5–15.5 13.0–24.2 14.1 6 5.7
SVL: snout–vent length; TL: tail length; HL: head length; JL: jaw length; HW: head width; JW: jaw width; DS: depth in the soil

index (Pianka, 1973): RESULTS

1 Sexual Dimorphism and Size.—There was no
B~ , significant sexual dimorphism in SVL of adult
P
n
pi 2
i~1
where i stands for prey category, n for number
of categories, and p for the proportional
utilization of resource i. We standardized the
food niche breadth values by dividing them by
the number of used prey categories and
calculated the dietary overlap between the two
species using the following equation (Pianka,
1973, 1974):
P
n
(pji :pki )
i~1
Ojk ~ sffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffi
ffi,
P n P n
( 2
p ji : 2
p ki )
i~1 i~1
where Pji and Pki are the proportions of
individuals of the species j and k, respectively,
associated with the resource i. Ojk range from zero
(no overlap) to one (complete overlap). However,
we used overlap only as a similarity index,
making no inferences about the relative abun-
dances of potential prey items in the environ-
ment.
We compared the mean number of prey items
per stomach, the SVL between species and sexes
(sexually mature individuals), and the soil
depths where the amphisbaenians were collect-
ed using a one-way analysis of variance
(ANOVA). We regressed number of prey per
stomach and SVL for each species. Means are
given 6 1 SD.
A. ibijara (F1,31 5 2.1; P 5 0.157), but adult male
L. polystegum were larger than females (F1,17 5
13.9; P 5 0.019; Table 1). In both species, the
smallest reproductively active individuals were
females (Table 1). Adults of L. polystegum were
significantly larger than adults of A. ibijara (F1,50
5 66.5; P , 0.0001; Table 1).
Juveniles of A. ibijara were found during all
five field trips; however, individuals collected
during the dry season (August to October) were
significantly smaller (mean SVL 5 96 mm;
range 5 74.3–117.8 mm; N 5 12) than those
collected during the rainy season (March to
July) (mean SVL 5 138 mm; range 5 106.8–
164.5 mm; N 5 19) (F1,29 5 58.5; P , 0.0001).
Soil Depth.—The two species differed in the
soil depth at which they were found (F1,54 5
15.43; P , 0.0001), with A. ibijara occurring at
shallower depths (10.83 6 3.6 cm; N 5 44) than
L. polystegum (15.90 6 4.79 cm; N 5 12). We
found neither ontogenetic nor sexual (F2,41 5
0.28; P 5 0.75) differences in soil depth for A.
ibijara (Table 1; Fig. 1A). In L. polystegum, males
were found deeper in the soil than females and
juveniles, but the small sample size prevented
us from testing whether these differences were
significant (Table 1; Fig. 1B).
Diet.—Two (3%) and seven (24%) of the
examined stomachs of A. ibijara (N 5 65) and
L. polystegum (N 5 29), respectively, were
empty. In the remaining stomachs, 13 prey
categories were recorded in A. ibijara and 10 in
L. polystegum (Table 2 and 3, respectively).
Ninety percent of all A. ibijara stomachs
380 J. O. GOMES ET AL.
FIG. 1. Relationship between snout–vent length
(SVL) and depth of capture for Amphisbaena ibijara
(A) and Leposternon polystegum (B). Asterisk: males;
circle: females; triangle: juveniles.
contained termites, 23.8% contained Coleoptera
larvae, and 12.7% contained earthworms (Ta-
ble 2). In contrast, 31.8% of L. polystegum
stomachs contained earthworms, followed by
ants (22.7%) and termites (13.6%) (Table 3).
Numerically, 86.2% of all individual prey items
in A. ibijara were termites and 12.8% Coleoptera
larvae, whereas in L. polystegum, 39.3% of
individual prey items were ants, followed by
termites (26.2%), and earthworms (9.5%) (Ta-
ble 2 and 3, respectively).
Molluscs, mites, hymenopterans other than
ants, and scales (possibly from lizards) were
found exclusively in the stomachs of A. ibijara,
whereas Hemiptera only occurred in one
stomach of L. polystegum (Table 2 and 3,
respectively). We considered vegetable debris
as ingested accidentally because of its low
frequency.
The diet of A. ibijara did not differ between
sexes or size classes. Ten different prey items
occurred in the stomachs of juveniles, whereas
in adult stomachs, there were nine different
types of prey (Table 2). Termites and larvae
Coleoptera were found in a higher percentage
of individuals than all other invertebrate orders,
and termites were the most abundant taxa
(Table 2). Larvae Coleoptera were abundant
only in female specimens. In contrast, prey
items varied both ontogenetically and sexually
in L. polystegum individuals (Table 3). In the
stomachs of juveniles, we found three prey
categories (chilopoda, homoptera nymphs, and
nematodes), with similar frequency of occur-
rence, although nematodes were the most
abundant of the three (71.4%) (Table 3). In
males, we found five types of prey items, with
ants (37.5%) and earthworms (25%) dominating
in frequency of occurrence and termites (43.2%)
and ants (40.5%) dominating with respect to the
number of prey. Seven different types of prey
occurred in the stomach contents of females
(Table 3). Earthworms (50%) dominated with
respect to frequency of occurrence, whereas ants
(45%), earthworms, and termites (both with
15%) dominated numerically.
The adjusted food niche for A. ibijara and L.
polystegum adults and juveniles were 0.08 and
0.36 for occurrence and 0.11 and 0.41 for prey
number, respectively. The similarity in food
niche (i.e., niche overlap) of the two species for
occurrence and number of ingested prey were
47.0% and 54.2%, respectively.
We did not find significant relationships
between the number of prey in stomach
contents and SVL for A. ibijara or for L.
polystegum (A. ibijara: r2 5 2.85, P 5 0.097, N 5
60; L. polystegum: r2 5 0.19, P 5 0.67, N 5 15)
(Fig. 2). Also, differences among females, males,
and juveniles for the mean number of prey
items per stomach in A. ibijara were not
significant (females: 93.6 6 132.4, N 5 16; males
80.2 6 98.9, N 5 14; juveniles: 39.6 6 34.9, N 5
30; F2,57 5 2.41; P 5 0.09). In L. polystegum, the
mean number of prey items per stomach was 7.4
6 8.2 (N 5 5) in males, 5.0 6 5.7 (N 5 8) in
females, and 3.5 6 3.5 (N 5 2) in juveniles.
Because of this low sample size, we did not
apply any statistical test, but we do not expect
significant differences because, as in A. ibijara,
we did not find differences between the number
of prey eaten and SVL.
DISCUSSION
Sexual Dimorphism and Size.—López et al.
(1997) suggested that round-headed amphisbae-
 DIET OF AMPHSIBAENA IBIJARA AND LEPOSTERNON POLYSTEGUM 381

TABLE 2. Diet summary for Amphisbaena ibijara from a gallery forest in northeastern Brazil. N 5 Sample size.
The category ‘‘Uap’’ refers to unidentified arthropod parts. FO: frequency of occurrence; NP: number of prey
per stomachs.

Juveniles (N 5 31) Males (N 5 16) Females (N 5 16) All individuals (N 5 63)

Prey types FO (%) FN (%) FO (%) FN (%) FO (%) FN (%) FO (%) FN (%)
Isoptera 30 (96.8) 1187 (96.7) 13 (81.2) 1096 (97.6) 14 (87.5) 1033 (69.0) 57 (90.5) 3316 (86.2)
Coleoptera larvae 6 (19.3) 21 (1.7) 5 (31.2) 16 (1.4) 4 (25.0) 456 (30.5) 15 (23.8) 493 (12.8)
Chilopoda 1 (3.2) 2 (0.2) 1 (6.2) 1 (0.09) 1 (6.2) 1 (0.07) 3 (4.8) 4 (0.1)
Diptera larvae 2 (12.5) 2 (0.1) 2 (3.2) 2 (0.05)
Hymenoptera
Formicidae 1 (6.2) 2 (0.2) 1 (1.6) 2 (0.05)
Others 1 (3.2) 2 (0.2) 1 (1.6) 2 (0.05)
Homoptera
nymph 1 (6.2) 1 (0.09) 1 (1.6) 1 (0.03)
Acarina 2 (6.4) 4 (0.3) 1 (6.2) 2 (0.2) 1 (6.2) 1 (0.07) 4 (6.3) 7 (0.2)
Mollusca 2 (6.4) 2 (0.2) 1 (6.2) 1 (0.07) 3 (4.8) 3 (0.08)
Nematoda 1 (3.2) 1 (0.08) 1 (6.2) 1 (0.09) 1 (6.2) 1 (0.07) 3 (4.8) 3 (0.08)
Oligochaeta 4 (12.9) 4 (0.3) 2 (12.5) 2 (0.2) 2 (12.5) 2 (0.1) 8 (12.7) 8 (0.2)
Scales 1 (3.2) 1 (6.2) 2 (3.2)
Uap 4 (12.9) 4 (0.3) 2 (12.5) 2 (0.2) 6 (9.5) 6 (0.1)
Totals 1227 1123 1497 3847

nians (primitive species) should be sexually males; and in M. anchietae, a shovel-headed

dimorphic in SVL. In contrast, more specialized
amphisbaenians, such as the shoveled-head
species (derived species), should not show
sexual dimorphism with respect to SVL. In fact,
the absence of sexual dimorphism in SVL for A.
ibijara has been reported in other amphisbae-
nians, such as A. alba (Colli and Zamboni, 1999),
Bipes biporus (Kearney, 2003), Blanus cinereus (Gil
et al., 1993), and Monopeltis anchietae (Webb et
al., 2000). In contrast, we found sexual dimor-
phism in SVL of L. polystegum, with males being
larger than females. Webb et al. (2000) also
found that SVL of Zygaspis quadrifrons, a small
amphisbaenian with a rounded snout, is sexu-
ally dimorphic, with females being larger than
species, females have heads significantly wider
than males of similar body size. However,
studies with larger samples and a greater
number of both primitive and derived species
are still lacking, thus preventing definitive
conclusions.
Given that juveniles captured in the dry
season were smaller than those found in the
rainy season, the latter were presumably born at
the end of the dry season or early in the rainy
season. This supports the suggestion of Rodri-
gues et al. (2003) that reproduction in A. ibijara
from Urbano Santos likely occurs during the
dry season. Reproduction during the dry season
is reported for several species from the Cerrado,

TABLE 3. Diet summary for Leposternon polystegum from a gallery forest, northeastern Brazil. N 5 Sample
sizes. The category ‘‘Uap’’ refers to unidentified arthropod parts. FO: frequency of occurrence; NP: number of
prey per stomachs.

Juveniles (N 5 4) Males (N 5 8) Females (N 5 10) All individuals (N 5 22)

Prey types FO (%) FN (%) FO (%) FN (%) FO (%) FN (%) FO (%) FN (%)
Isoptera 1 (12.5) 16 (43.2) 2 (20.0) 6 (15.0) 3 (13.6) 22 (26.2)
Coleoptera larvae 2 (20.0) 3 (7.5) 2 (9.1) 3 (3.6)
Chilopoda 1 (25.0) 1 (14.3) 1 (12.5) 1 (2.7) 2 (9.1) 2 (2.4)
Diptera larvae 2 (20.0) 3 (7.5) 2 (9.1) 3 (3.6)
Hymenoptera
Formicidae 3 (37.5) 15 (40.5) 2 (20.0) 18 (45.0) 5 (22.7) 33 (39.3)
Homoptera nymph 1 (25.0) 1 (14.3) 1 (4.5) 1 (1.2)
Hemiptera 1 (10.0) 1 (2.5) 1 (4.5) 1 (1.2)
Nematoda 1 (25.0) 5 (71.4) 1 (4.5) 5 (6.0)
Oligochaeta 2 (25.0) 2 (5.4) 5 (50.0) 6 (15.0) 7 (31.8) 8 (9.5)
Uap 3 (37.5) 3 (8.1) 3 (30.0) 3 (7.5) 6 (27.3) 6 (7.1)
Totals 7 37 40 84
382 J. O. GOMES ET AL.
FIG. 2. Relationship between prey totals and
snout–vent length (SVL) for Amphisbaena ibijara (A)
and Leposternon polystegum (B). Asterisk: males; circle:
females; triangle: juveniles.
such as A. alba (Colli and Zamboni, 1999), and
the lizards Cnemidophorus ocellifer (Mesquita and
Colli, 2003b) and Gymnodactylus geckoides amar-
ali (Colli et al., 2003). Several hypotheses have
been proposed to explain the seasonality of
reproduction in the tropics (see Mesquita and
Colli, 2003b; Brown and Shine, 2006). Colli et al.
(2003) suggested that, in G. g. amarali, a species
that ingests a large number of termites (81.46%
of total prey volume), reproduction may occur
during the dry season because of the increase in
foraging activity of termites during this period
(Pinheiro et al., 2002). The same may be true for
A. ibijara, whose adults and juveniles ingest
large numbers of termites (86.2% of total
number of prey). In addition, eggs might be
flooded if they were laid during the wet season
and, thus, increase the probability of egg
mortality (Brown and Shine, 2006).
Depth.—The difference in microhabitat use,
with L. polystegum found in deeper soil layers,
was expected because this species has more
sophisticated adaptations for fossorialism
(Pough et al., 2004) than does A. ibijara.
Differences in diet composition between these
two species may be, in part, a consequence of
the different depths they occupy in the soil,
although foraging depths probably overlap
between the two species.
Diet.—Like Webb et al. (2000) and Kearney
(2003), we found a high proportion of individ-
uals with food items in their stomachs, both in
A. ibijara (97%) and in L. polystegum (76%),
indicating that these amphisbaenians probably
feed frequently (see Avery, 1973). We found a
total of 14 prey types in the two amphisbaenian
species. Other studies with cogeneric species
showed that these animals use a variety of
similar taxa in other localities (Webb et al.,
2000). At Urbano Santos, A. ibijara and L.
polystegum, although overlapping in the soil
depth where they were found, differed in the
types of prey that were ingested. The spectrum
of prey consumed for the two species was
narrow, with termites and Coleoptera larvae
dominating in A. ibijara, whereas in L. poly-
stegum there was a shift in diet, both ontogenet-
ically and sexually. In L. polystegum juveniles,
prey items were smaller than those ingested by
adults, and there was no dominant prey taxon.
In contrast, ants (occurrence) and termites
(number of prey) dominated in the stomachs
of males, and earthworms (occurrence) and ants
(number of prey) dominated in the stomachs of
females. These observed shifts in the diet
composition of L. polystegum may be caused by
morphological constraints, which may prevent
the smallest individuals from ingesting the
largest prey. However, additional studies are
needed to confirm whether this trend holds
true. Overall, the prey types consumed by L.
polystegum at Urbano Santos did not differ from
the few previous reports about the diet of this
species in other localities (see Barros-Filho and
Valverde, 1996). Although the value of the
adjusted niche was lower in A. ibijara than in
L. polystegum, both juveniles (nine categories)
and adults (13 categories) of A. ibijara ingested
more prey categories than juveniles (three
categories) and adults (eight categories) of L.
polystegum. This supports the suggestion of
Gans (1968) that amphisbaenians with modified
shovel-shaped heads are less efficient predators
than species with rounded heads (e.g., genus
Amphisbaena).
 DIET OF AMPHSIBAENA IBIJARA AND LEPOSTERNON POLYSTEGUM
The absence of a significant relationship
between the number of prey items and SVL, as
observed in A. ibijara and L. polystegum, was also
reported for Amphisbaena mertensii and Cercolo-
phia roberti (Cruz-Neto and Abe, 1993), Amphis-
baena gonavensis (White et al., 1992), B. cinereus
(López et al., 1991) and other burrowing reptiles
(Henle, 1989; White et al., 1992). This may reflect
morphological specializations for burrowing
that presumably restrict the size of prey that
can be subdued and swallowed (Gans, 1968;
Andrews et al., 1987), indicating that adults and
juveniles apparently feed on food items of
similar size, although juveniles feed more
frequently than adults.
Termites are a frequent item in the diet of
amphisbaenians and lizards from the Cerrado
(e. g., Colli and Zamboni, 1999; Colli et al., 2003;
Mesquita and Colli, 2003b; this study). This may
be a consequence of the elevated abundance of
these arthropods in this biome (Colli et al.,
2003). However, there are no data on the
availability of potential prey in the environment
to evaluate whether these animals prefer ter-
mites or whether they consume prey according
to relative abundance.
Two specimens of A. ibijara, a juvenile and an
adult female, had scales in their stomachs,
possibly of lizards, suggesting that these ani-
mals may be predators of reptiles, which has
been observed for A. alba (Colli and Zamboni,
1999). Kearney (2003) indicated that, although
the literature suggests that amphisbaenians
exhibit morphological specializations for feed-
ing on vertebrates, few empirical data exist to
support its occurrence in the wild. One speci-
men of A. gonavensis contained ‘‘vertebrate
connective tissue remains’’ (see Cusumano
and Powell, 1991), and in A. alba, the limb of a
lizard and a snake have been found in stomach
contents (see Colli and Zamboni, 1999). More-
over, Sazima and Haddad (1992) found mam-
mal fur in the stomach of A. alba. Therefore, the
presence of scales in the stomach of A. ibijara (a
relatively small-bodied species) may be consid-
ered additional evidence of vertebrate predation
by amphisbaenians, even though accidental
ingestion cannot be discounted.
Kearney (2003), studying the diet composition
of 203 individuals of B. biporus, found the
remains of both subterranean and surface-
dwelling animals in stomachs; all individuals
found during the day were underground, but at
night individuals were found both below and
above ground (N 5 79). She concluded that B.
biporus may feed on the surface of the soil at
night. This same behavior appears to occur in A.
ibijara and L. polystegum, which also consumed
prey that live above and below ground. Al-
though all individuals of A. ibijara collected
383
during the day were underground, a previous
study in the same area (L. D. Cruz and J. O.
Gomes, unpubl. data) that employed pitfall
traps that were checked twice daily (at dusk
and dawn) found specimens of A. ibijara only
after sunrise, suggesting that this species is
active on the surface during the night. Con-
versely, we found a specimen of L. polystegum
swimming in a pond during the day, and one of
us (GVA) found a specimen of Aulura anomala
feeding during the day on the surface of the soil
underneath a stone at other locality. Therefore,
it is seems possible that these amphisbaenians
forage both during the day and night.
Acknowledgments.—We thank T. C. S. Avila-
Pires and T. Mott for the valuable comments on
earlier drafts and S. Peters for English review.
We are grateful to Walt Meshaka, Geoffrey R.
Smith, and to two anonymous reviewers for
useful comments on the manuscript. Many
thanks are extended to M. Mendonça for help
in the identification of some arthropods and to
Falcão for assistance with the fieldwork. The
Comercial e Agrı́cola Paineiras Ltda. partially
supported this research. P. L. Peloso helped in
the configuration of Figure 1. Permits for
collecting amphisbaenians was granted by
IBAMA (12 and 13/05; Process 02012.001879/
2004-35 – NUFAU/IBAMA).
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