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Reproductive Biology of Tantilla Melanocephala (Linnaeus, 1758) (Serpentes,

Colubridae) from Eastern Amazonia, Brazil

Article in Journal of herpetology · September 2009

DOI: 10.1670/0022-1511(2006)40[553:RBOTML]2.0.CO;2

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Journal of Herpetology, Vol. 40, No. 4, pp. 556–559, 2006
Copyright 2006 Society for the Study of Amphibians and Reptiles

Reproductive Biology of Tantilla melanocephala (Linnaeus, 1758) (Serpentes,

Colubridae) from Eastern Amazonia, Brazil
MARIA CRISTINA DOS SANTOS-COSTA,1,2 ANA L. C. PRUDENTE,3 AND MARCOS DI-BERNARDO4
1
Centro de Ciências Biológicas, Universidade Federal do Pará, 66075-110 Belém, Pará, Brazil
3
Laboratório de Herpetologia, Departamento de Zoologia, Museu Paraense Emı́lio Goeldi, 66040-170 Belém, Pará, Brazil
4
Laboratório de Herpetologia, Museu de Ciências e Tecnologia and Faculdade de Biociências, Pontifı́cia Universidade
Católica do Rio Grande do Sul, 90619-90 Porto Alegre, Rio Grande do Sul, Brazil

ABSTRACT.—The reproductive biology of Tantilla melanocephala was studied in eastern Amazonia, Pará,
Brazil. Males and females attained sexual maturity at approximately twice the length (snout–vent length 5
SVL) of newborn snakes. Mature males had a larger tail length and more subcaudal scales than mature
females, whereas females were larger (SVL) and had more ventral scales than males. Total length did not
differ between mature males and females. Reproduction was not seasonal. There was no correlation between
the number of vitellogenic follicles and oviductal eggs. The number of eggs was positively correlated with
the female SVL. The data presented here indicate that the eastern Amazonia population differs from
populations in southeastern Brazil in important morphological and reproductive aspects, including the mean
number of eggs produced (mean 5 1.53), which was smaller in eastern Amazonian populations.

The genus Tantilla comprises 53 species of small
oviparous and semifossorial snakes, ranging from the
United States to southern South America (Peters and
Orejas-Miranda, 1970; Wilson, 1987; Sawaya and
Sazima, 2003). Tantilla melanocephala occurs from
Guatemala throughout Central America into South
America east of the Andes, southeastern Peru, Bolivia,
northern Argentina and Uruguay; it also occurs on
Trinidad and Tobago (Wilson and Mena, 1980). In
Brazil, it occurs in Amazon rain forest, Caatinga,
Cerrado, and Savannas (Cunha and Nascimento, 1978;
Vanzolini et al., 1980; Strüssmann, 2000; Lema, 2002).
In Brazilian Amazonia, specimens are found in open
fields as well as in forest litter, inside termite nests and
rotting trunks, and under rocks or fallen trunks, in
habitats including primary and secondary forests
(Cunha and Nascimento, 1978; Martins and Oliveira,
1998).
Marques and Puorto (1998) recorded seasonal
reproduction and low fecundity for T. melanocephala
in open areas of southeastern Brazil, a pattern also
found in the neartic species Tantilla rubra and Tantilla
coronata (Easterla, 1975; Aldridge and Semlitsch, 1992).
In the southeastern Brazilian population of T. melano-
cephala, vitellogenesis and ovulation occurred only
during the rainy season (September to January), and
recruitment period was short (Marques and Puorto,
1998). Duellman (1978) mentioned the scarcity of data
concerning the extent of the reproductive period in
Amazonian snakes and suspected the existence of
several patterns (from restricted to unrestricted),
though the only known consistent data at that time
(for six species) indicated continuous reproductive
cycles.
Despite the wide geographic distribution of T.
melanocephala, data on natural history are available
only for the southeastern Brazilian population. We
present data on reproductive parameters (e.g., re-
productive cycle, fecundity, size, and age at matura-
tion) of a population of T. melanocephala from eastern
Amazonia, Brazil, a region geographically distant and
both climatically and physionomically different from
the one in which T. melanocephala was studied by
Marques and Puorto (1998).
MATERIALS AND METHODS
We examined 255 specimens of T. melanocephala
from State of Pará, eastern Amazonia, Brazil, captured
between 1960 and 2002. All specimens are deposited
in the herpetological collection of Museu Paraense
Emı́lio Goeldi (MPEG), Belém, Brazil. The climate
type in the study region is Am (Köeppen classifica-
tion): humid tropical with excessive rainfall during
some months (December, January, February and
March) reaching 2500–3000 mm per year, relative
humidity higher than 80%, and mean temperature
ranging from 25–31uC throughout the year (Cunha
and Nascimento, 1978).
We recorded the following data for each specimen
(length in millimeters): snout–vent length (SVL), tail
length (TL), total length (TOL), number of ventral (VS)
and subcaudal (SC) scales, length of the largest
vitellogenic follicle, and condition of the deferent
ducts. Sex was determined by dissection and analysis
of gonads. Males were recognized as mature when
they presented opaque and convoluted deferent ducts
(Shine, 1977) and females when they had an SVL equal
to or larger than the smallest female with ovarian
follicles . 5 mm (Marques and Puorto, 1998), oviduc-
tal eggs, or flaccid oviducts (Shine, 1988). The smallest
immature specimens presenting a visible umbilical
scar were considered newborns. Means are given with
the respective standard deviation.

2
Corresponding Author. Present address: Universi-
dade Federal do Pará, Centro de Ciências Biológicas,
Rua Augusto Corrêa, 01, 66075-110 Belém, Pará,
Brazil. E-mail: mcsc@ufpa.br
RESULTS
The smallest mature male had an SVL of 181 mm,
and the smallest mature female had an SVL of
189 mm (Table 1). Sixty-two percent of males with
SVL between 191 and 210 mm and 95% of males with

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 SHORTER COMMUNICATIONS 557

TABLE 1. Morphology of mature females and males of Tantilla melanocephala from State of Pará, eastern
Amazonia, Brazil, with mean and standard deviation (6 SD). The amplitude of variation and the number of
specimens studied for each character (N) are within parenthesis. SVL 5 snout–vent length; TL 5 tail length; TOL
5 total length; VS 5 ventral scales number; SC 5 subcaudal scales number. Asterisks indicate significant
differences between females and males (see text).

Mature females Mature males

*SVL (mm) 229.8 6 17.0 (189–266; N 5 81) 217.7 6 20.9 (181–296; N 5 65)
*TL (mm) 61.6 6 7.74 (44–89; N 5 59) 70.2 6 8.66 (55–95; N 5 43)
TOL (mm) 291 6 22.3 (249–355; N 5 60) 285 6 25.2 (242–350; N 5 43)
*VS (mm) 145.8 6 3.7 (135–156; N 5 75) 136.7 6 3.5 (126–147; N 5 98)
*SC (mm) 49.4 6 4.37 (39–59 N 5 75) 55.0 6 4.58 (42–69; N 5 98)

SVL between 211 and 230 mm were mature; above
230 mm SVL, 100% of the males were mature.
The SVL of mature females was larger than the SVL
of mature males (t144 5 3.81; P , 0.05; Table 1).
Mature males have TL larger than mature females (U
5 531, P ,, 0.05, Table 1). Total length was not
significantly different between mature females and
males (U 5 1099; P 5 0.21). The number of ventral
scales was higher in females (U 5 266; P ,, 0.05),
and the number of subcaudal scales was higher in
males (t171 5 8,14; P , 0.01; Table 1, Fig. 1).
Vitellogenic follicles were found throughout the
year; eggs were recorded from March to November in
13 (26%) of the 50 mature females studied. Newborns
were recorded in February, July, September, and
October (N 5 4; SVL 5 94–112 mm).
The number of oviductal eggs varied from 1–3
(mean 5 1.53 6 0.63; N 5 15) and did not differ (U 5
139; P .. 0.05; N 5 34) from the number of
vitellogenic follicles (mean 5 1.52 + 0.51; range 5 1–
2; N 5 19). There was a positive significant correlation
between SVL and the number of eggs (r 5 0.54; P ,
0.05; N 5 15).
DISCUSSION
Male T. melanocephala from eastern Amazonia reach
sexual maturity at a smaller size (SVL) than females.
The same is true for T. melanocephala in southeastern
Brazil (Marques and Puorto, 1998) (and this is the
pattern for snakes in general; see Parker and Plum-
mer, 1987). The data made available by Marques and
Puorto (1998) indicate that SVL in mature males from
southeastern Brazil population is larger than in the
eastern Amazonia population. In both populations,
males and females become mature when they reach
about twice the SVL of newborns. In the population of
T. melanocephala from southeastern Brazil, the age at
maturity was estimated as 10 months for males and
20 months for females, based on the seasonal distri-
bution of size classes of specimens of both sexes
captured along all the months of several years
(Marques and Puorto, 1998). In eastern Amazonia,

FIG. 1. Correlation between the number of ventral and subcaudal scales in females (solid circles) and males
(open circles) of Tantilla melanocephala from State of Pará, eastern Amazonia, Brazil.

Journal of Herpetology hpet-40-04-18.3d 23/9/06 00:43:27 557 Cust # 212-05N

558 SHORTER COMMUNICATIONS
recruitment is aseasonal (see below), which prevents
confident conclusions concerning the age at maturity
(specimens with similar body size were found in
several months of the year). In both eastern Amazonia
and southeastern Brazil (Marques and Puorto, 1998),
sexual dimorphism in SVL of T. melanocephala is
similar to that which is found in other species of the
tribe Sonorini (Shine, 1994), with the exception of
some species of the genus Sonora, which present male
combat behavior, and have male SVLs larger than or
equal to conspecific females (Shine, 1994). Males of T.
melanocephala present SVL consistently smaller than in
females, which could be an indication of absence of
combat related to mating.
The fecundity of T. melanocephala in eastern Ama-
zonia is low, as in the population in southeastern
Brazil (Marques and Puorto, 1998) and other species of
Tantilla (Easterla, 1975; Aldridge and Semlitsch, 1992).
The low fecundity may be related to the small size of
these snakes (fecundity usually varies as a direct rate
to body size; Vitt and Vangilder, 1983; Shine, 1993;
Marques, 2002; Pizzatto and Marques, 2002), to the
low energy cost required (Lillywhite, 1987), and to the
semifossorial habit (Cadle and Greene, 1993; Webb et
al., 2000, 2001; Balestrin and Di-Bernardo, 2005).
Fecundity is lower in eastern Amazonia than in
southeastern Brazil (1.53 vs. 1.95 eggs per female,
respectively), probably because the SVL of mature
females in the Amazonian population is smaller. Fitch
(1985) found positive correlation between fecundity
(clutch size) and latitude for 60% of the snake species
he studied. The known data of T. melanocephala
corroborate Fitch’s prediction (Fitch, 1985).
The absence of a significant difference between
number of vitellogenic follicles and number of eggs
indicates a conversion rate of vitellogenic follicle into
egg of about 100%. Similarly high conversion rates
were recorded for other species of oviparous (e.g.,
Oxyrhopus rhombifer, G. F. Maschio, pers. obs.) and
viviparous (e.g., Helicops infrataeniatus, Aguiar and Di-
Bernardo, 2005) Neotropical snakes, which allows the
use of the number of vitellogenic follicles as a fecun-
dity parameter. The use of this parameter as an
indicator of fecundity is particularly informative
because in most snake samples studied the number
of females containing eggs is relatively small (Al-
dridge and Semlitsch, 1992; Goldberg, 2001; Webb et
al., 2001; Hartmann et al. 2002).
Recent studies (e.g., Martins and Oliveira, 1998)
corroborated Duellman’s (1978) idea and pointed out
several species from central Amazonia with seasonal
reproductive cycles. In eastern Amazonia, the record
of newborns of T. melanocephala in February, July,
September, and October, and females with secondary
vitellogenic follicles or eggs in all months, indicate
a continuous reproductive cycle. However, this
species presents seasonal reproduction in southeast-
ern Brazil (Marques and Puorto, 1998), where the
climate is subtropical. Fitch (1985) pointed out that
96% of the snakes he studied had some kind of
geographic variation of reproductive patterns. Our
results and those of Marques and Puorto (1998) show
that the reproductive cycle of T. melanocephala shows
geographic variation. Further studies could contribute
with relevant information concerning local adapta-
tions in different environmental conditions, helping to
Journal of Herpetology hpet-40-04-18.3d 23/9/06 00:43:31
identify which characteristics could be selected as
a function of specific ecological conditions.
Acknowledgments.—We thank M. Hoogmoed and T.
C. Avila-Pires for criticism and suggestions on the
manuscript, and A. P. Maciel, A. B. Bonaldo, and W.
Wosiacki for improving the English. The senior author
thanks the staff of the Museu Paraense Emı́lio Goeldi
for facilities and for permission to use the laboratory.
MD is partially financed by the Conselho Nacional de
Desenvolvimento Cientı́fico e Tecnológico (Process
CNPq 300.164/00-9).
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Accepted: 1 August 2006.
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