Reproductive tactics of Ameiva ameiva (Lacertilia: Teiidae) in a seasonally fluctuating

tropical habitat

LAURIE
J. VITT'
Savannah River Ecology Laboratory, Drawer E, Aiken, SC, U.S.A. 29801
Received April 26, 1982
Can. J. Zool. Downloaded from www.nrcresearchpress.com by University of Saskatchewan on 11/23/12

VITT,L. J. 1982. Reproductive tactics of Ameiva ameiva (Lacertilia: Teiidae) in a seasonally fluctuating tropical habitat. Can. J.
ZOO^. 60: 31 13-3120.
The reproductive ecology of Ameiva ameiva was studied for 12 months in a caatinga habitat of northeast Brazil. Even though
rainfall was seasonal, the female reproductive cycle was not associated with this seasonality. Females reproduced year-round,
with peak reproductive periods during August-October and January-February. Clutch size ranged from one to nine and was
correlated to female size but egg size was constant. Males showed evidence of reproductive activity throughout the year. Fat body
mass of males and females varied greatly among individuals. There was no association between fat storage and wet-dry
seasonality. In females, fat body mass tended to decrease during peak reproductive periods. Most striking was the observation
that 97.8% of all adult Ameiva possessed enlarged fat bodies, suggesting that resource periods low enough to affect reproduction
did not exist during 1977- 1978. The reproductive tactics of Ameiva were similar to those of other tropical macroteiids, regardless
of their distribution, but very different than reproductive tactics of sympatric iguanid lizards. Compared with iguanid lizards,
resources may be less limiting for macroteiids because their widely foraging behavior for prey acquisition may allow them to find
rich patches of resources which would be unavailable to habitat specific sit-and-wait foragers, like most iguanid lizards.

VITT,L. J. 1982. Reproductive tactics of Ameiva ameiva (Lacertilia: Teiidae) in a seasonally fluctuating tropical habitat. Can. J.
For personal use only.

Zool. 60: 31 13-3120.

L'Ccologie de la reproduction d'Ameiva ameiva a CtC CtudiCe durant une pCriode de 12 mois, dans un habitat de
caatinga a i nord-est du Brksil. Bien que la p6riode de precipitations soit iaisonnikre, le cycle de reproduction de
la femelle n'est pas associC aux saisons. Les femelles se reproduisent durant toute I'annCe, mais on peut reconnaitre deux
pkriodes d'activitk reproductrice plus intenses, d'aoiit a octobre et en janvier-fkvrier. Le nombre d'oeufs dans une portCe va de
1 a 9 selon la taille de la femelle, mais la taille des oeufs est constante. Les miles manifestent des signes d'activitC reproductrice
durant toute I'annCe. La masse des corps gras varie beaucoup d'un individu a l'autre, chez les miles comme chez les femelles. La
mise en rCserve des graisses ne suit pas les changements de saisons. Chez les femelles, la masse des corps gras tend a diminuer
durant les pCriodes intenses d'activitk reproductrice. Durant la pCriode d'Ctude, 97,8% des adultes d'Ameiva possidaient des
corps gras de taille considkrable; il semble donc qu'il n'y ait pas eu, en 1977- 1978, de pCriode oh les ressources ont diminuC au
point d'affecter la reproduction. Les tactiques reproductrices d'Ameiva ressemblent a celles d'autres macrotCidCs tropicaux,
quelle que soit leur rkpartition, mais diffkrent considkrablement des stratigies reproductrices des lCzards iguanidks sympatrides.
Les ressources semblent jouer un r81e limitant moins important pour les macrotCiidCs que pour les iguanidks, puisque le
comportement de recherche de nourriture sur de grandes surfaces permet aux macrotCiidCs de trouver des sources abondantes de
proies en certains endroits, sources inaccessibles a la plupart des iguanidCs confinks a un habitat particulier et dont la stratCgie de
chasse est d'attendre que des proies se prksentent.
[Traduit par le journal]

Introduction Lacher 1981). Reproductive tactics are also known to

During the past 15 years, a diversity of reproductive vary among geographically separated populations of the
tactics among species of lizards has been identified same species (Dunham 1981; Parker and Pianka 1975,
(Fitch 1970; Tinkle 1969; Tinkle et al. 1970). This 1976; Pianka 1970; Tinkle and Ballinger 1972) as well
diversity is reflected among lizard species in general, as among years within the same population (Ballinger
among species in the same taxon (Barbault 1976; and Congdon 1981; Dunham 1981; Martin 1977; Tinkle
Gorman and Licht 1974; Licht and Gorman 1970; 1967; Vinegar 1975; Vitt et al. 1978). Resource availa-
Sexton et al. 1971; and others), and among species of bility or its correlates (rainfall, temperature) are usually
different taxa occurring at the same locality (Barbault identified as determinants of variability in such repro-
1976; Dixon and Soini 1975; Duellman 1978; Inger and ductive characteristics as clutch size, clutch frequency,
Greenberg 1966; Vitt and Goldberg 1983; Vitt and age at maturity, and even egg size under certain
circumstances (Derickson 1976; Ferguson and Bohlen
'present address: Biology Department, University of Cali- 1978; Ferguson et al. 1980; Nussbaum 1981).
fornia, Los Angeles, CA, U.S.A. 90024. Even though there has been a recent interest in
0008-4301 /82/123113-08$01 .OO/O
01982 National Research Council of Canada/Conseil national de recherches du Canada
 31 14 CAN. J . ZOOL. VOL. 60, 1982
reproduction of tropical lizards (see Sherbrooke (1975)
and Duellman (1978) for review), very little is known
concerning reproductive tactics of tropical lizards in
certain environments. The Caatinga is a semiarid region
of northeast Brazil which harbors a saurofauna including
seven gekkonids, four iguanids, five teiids, one scincid,
one anguid, and one amphisbaenid (Vanzolini et al.
1980).
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In this paper, I focus on the male and female repro-
ductive and fat body cycle of the macroteiid lizard
Ameiva ameiva. This wide-ranging lizard occurs from
northern Argentina east of the Andes to Panama and in
many areas is very abundant. It has been studied in
Amazonian Ecuador (Duellman 1978; Simmons 1975)
and the Iquitos region of Peru (Dixon and Soini 1975).
Other species of Ameiva have been studied in Costa Rica
(Hirth 1963; Smith 1968). Utilizing data collected over a
12-month period, I investigated the following questions:
(i) Is the reproductive or fat cycle correlated to season-
ality in rainfall? (ii) Does clutch size vary with female
size? (iii) Is egg size associated with female size,
season, or is it constant? (iv) Is Ameiva reproduction at
For personal use only.
this locality the same as at other localities? and (v) Is
seasonality in the reproductive pattern of Ameiva similar
to that of sympatric lizards?
Methods
Study area and climate
This study was carried out near Exu, Pernambuco, a small
town near the geographical center of "zonas das caatingas," a
vast semiarid region of northeastern Brazil (Reis 1976). The
region is of particular interest with respect to studies on lizard
reproductive tactics in that it is the largest dry region entirely
contained within the tropics, it is surrounded by mesic
habitats, and it contains a saurofauna exhibiting taxonomic and
ecological diversity.
Most Ameiva sampled were taken from Low Caatinga
(Caatinga Baixa) habitats. Low Caatinga communities contain
xerophytic trees ranging from 3 to 5 m, occasional emergents
up to 8m, and large cacti including Cereus jamacura,
Cephalocereus gounellei, and Zehntherella squamulosa
(Mares et al. 1981). A variety of Euphorbiacea and Legumi-
nosae are also common. The thorn forest habitat is interrupted
by occasional granitic extrusions (Lajeiros) which provide
refugia for many reptiles, amphibians, and mammals (see for
example, Lacher 1981; Mares et al. 1981; Vanzolini et al.
1980; Vitt 1980, 1981). Only one lizard, Platynotus semitae-
niatus, is restricted primarily to Caatinga (see Vanzolini
1976), the remaining lizard fauna resulting from colonization
by species from the Amazon Basin, Atlantic Forest, and the
remaining open formations (Cerrado and Chaco) which extend
from northeastern Brazil to Paraguay and Argentina (Vanzo-
lini 1974, 1976). Ameiva ameiva is most often associated with
open areas between the lajeiros and is uncommon where
vegetation is dense.
The Caatinga climate is considered semiarid with annual
rainfall varying from less than 400 to just over 1000 mm.
Rainfall is highly unpredictable in timing and intensity and the
region is susceptible to extreme droughts or flooding (Eicit
1968). During 1977- 1978, monthly rainfall (recorded approx-
imately 50 km from Exu at Ouricuri, Sudene, Divisiio de
Hydrologia) was similar to long-term averages (Vitt and
Goldberg 1983). In typical years, there is a dry season
extending from May until November, considering months
receiving less than 50 mm rainfall to be dry. During 1977-
1978, the dry season began in June and persisted until nearly
the end of November (Vitt and Goldberg 1983).
Thermally, the Caatinga can be considered equable, with
monthly temperatures fluctuating very little. The mean differ-
ence between maximum and minimum daily temperatures
calculated over 2-week periods throughout the year was 10.19
2 1.40C (SD). There was no significant temperature differ-
ence between the wet and dry seasons (Vitt and Goldberg
1983).
Field and laboratory methods
One hundred and ninety-four male and 167 female Ameiva
ameiva were sampled for reproductive analysis over a 12-
month period extending from March 1977 through February
1978. Lizards were collected by shooting with a pellet gun, by
hand, and by digging them from burrows. Lizards killed in the
field were immediately placed on ice until return to the
laboratory. All lizards brought to the laboratory alive were
killed with Nembutal within 1 h after collection, a variety of
measurements were taken, and lizards were fixed in 10%
formalin. Measurements on fresh animals included snout-vent
length (SVL) and total body mass. Following fixation, lizards
were autopsied. For males, length of each testis was recorded
and observations on condition of testes and epididymides were
recorded. For females, the number of corpora lutea, vitello-
genic follicles, and (or) oviductal eggs was recorded. Diameter
of vitellogenic follicles and width and length of oviductal eggs
were recorded. Testes with attached epididymides, fat bodies,
and oviductal eggs were removed and transferred to fresh
Formalin. Formalin-fixed testes, fat bodies, and individual
eggs were rolled dry on paper towels, and weighed to 0.001 g.
There was no apparent change in size or shape of testes or fat
bodies due to Formalin fixation and thus their masses are
considered reasonable estimates of fresh mass. Oviductal
eggs, however, appeared to swell in fixative (see also Martin
1978)and thus my wet mass estimates may be high. Eggs were
then soaked for 48 h in water and freeze-dried. Dry mass of
eggs should not be subject to variation due to Formalin fixation
or variation attributable to hydration state of the eggs following
ovulation.
To gain the maximum amount of information from the data
set, females were divided into five categories: (1)containing
oviductal eggs, (2) containing enlarged vitellogenic follicles
and oviductal eggs or corpora lutea, (3) containing corpora
lutea only, (4) containing enlarged vitellogenic follicles only,
and (5)nonreproductive, i.e., containing no evidence of recent
reproductive activity. Females in any of the first three
categories were considered "reproductive" because their re-
productive state indicates that they would deposit eggs in a
short time (categories 1 and 2) or they had recently deposited
eggs (category 3 ) .
For estimation of clutch size, I used only counts of enlarged
vitellogenic follicles (>3.0 mm) or oviductal eggs. Number of
corpora lutea was also recorded but used only in comparing
 VITT
clutch size within an individual. Only the simultaneous
occurrence of enlarged vitellogenic follicles and oviductal
eggs or corpora lutea was considered adequate evidence of
individuals producing more than one clutch. Minimum size
(SVL) at sexual maturity for females was based on the smallest
female containing oviductal eggs or enlarged vitellogenic
follicles. Minimum size at sexual maturity for males was based
on presence of enlarged testes and convoluted epididymides.
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Immature males had very small testes and the epididymides
were tiny translucent structures. In addition, the heads of
mature males were large relative to the body as compared with
immature males.
Standard parametric statistical tests were used when the
assumptions of the tests were not seriously violated. Appropri-
ate nonparametric tests were used when the assumptions of the
parametric test could not be met. A probability level of 0.05
was considered adequate for falsification of null hypotheses.
Unless otherwise stated, means are presented as i+ standard
error (SE) .
Results
Body size and apparent sexual dimorphism in size
Ameiva ameiva in northeastern Brazil range from 40
to 176 mm SVL (Fig. 1). Males reach sexual maturity at
For personal use only.
99mm SVL and females reach sexual maturity at
102mm SVL. Comparisons of SVL and body mass
between reproductively mature males and females re-
vealed significant differences with males larger. The
means of SVL and body mass for reproductively mature
males were 141.4 + 1.6 (N = 172) mm and 90.7 +
3.0 g. Means of SVL and body mass for reproductively
mature females were 128.8 2 1.1 (N = 144) mm and
61.0 + 1.5 g. Both mean SVL and mean body mass
differed significantly between sexes (SVL: Mann-
Whitney U = 177250, p < 0.0001; body mass:
Mann-Whitney U = 17 3 11.5, p < 0.0001). Analysis
of covariance with log SVL as the covariate revealed
significant differences in slopes (F1,354= 10.22, p =
0.0015) and intercepts (F1,355= 8.66, p = 0.035) of the
log body mass - log SVL relationships between sexes
with males attaining significantly heavier mass at a
given SVL than females. The latter analysis included all
Ameiva sampled.
Female reproduction
The seasonal distribution of reproductive categories
in Ameiva females appears in Fig. 2. The greatest
proportions of reproductive females occurred during
June, July, and August, even though there were repro-
ductive females during every month. The greatest
proportions of nonreproductive females were observed
from September to February.
During March-July and again from December to
February, size of vitellogenic follicles varied consider-
ably among females (Fig. 3). Follicles 1 3.0 mm con-
tained yolk material rendering them easily distin-
guishable from the translucent and small (<3.0mm)
31 15
I Females I
40 60 80 100 120 140 160 160
S N O U T - V E N T L E N G T H (mm)
FIG. 1. Body size distribution of male and female Ameiva
ameiva . Arrows indicate minimum size at sexual maturity.
/ 1 a n d 0vlductaIEggb o r
/
C a r p o r o Lutao
-0vlductal Eggs
M ' A ' M ' J ' J ' A ' s ' o ' N ' D ' J ~
MONTH
FIG. 2. Seasonal distribution of reproductive categories in
Ameiva ameiva from northeast Brazil. Monthly sample size is
indicated across the top of the figure.
nonvitellogenic follicles. The wide variation in size of
vitellogenic follicles within any given month suggests
that at least some females were close to ovulation during
each of these months. The time period during which few
females had vitellogenic follicles large enough to
suggest that ovulation was near (>10 mm), corres-
ponded to the time period when many females contained
oviductal eggs (August-October) and was followed by
a period during which no females sampled contained
oviductal eggs (November-January) . Relative clutch
mass (clutch massltotal mass) for 25 females averaged
0.159 2 0.007.
Clutch and egg characteristics
Clutch size ranged from one to nine in Ameiva with
larger females producing larger clutches (Fig. 4). Mean
clutch size based on oviductal egg counts was 5.7 2
0.29 (N = 26) and mean clutch size based on counts of
vitellogenic follicles was 5.6 + 0.19 (N = 80). The
models for the relationships of number of oviductal eggs
to SVL (OVI = -9.46 + 0.114 SVL) and number of
vitellogenic follicles to SVL (FOLL = -7.47 + 0.099
SVL) were both significant (OVI-SVL; F, ,24 = 19.7, p
= 0.0002: FOLL-SVL; F1,9= 51.2, p < 0.001) and
SVL explained 45.1 and 39.3% of the variance (r2),
 31 16 CAN. J. ZOOL. VOL. 60, 1982
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p 20

1 3 5 7 9
Clutch Size
I I 1 I 1 I I
110 120 130 140 150 160
Snout-Vent L e n g t h (mm)
MONTH
FIG. 4. The relationship between clutch size and snout-
FIG. 3. Seasonal distribution of vitellogenic follicle diam- vent length in Ameiva ameiva based on counts of oviductal
eters in Ameiva ameiva. The line at the top of the figure eggs and vitellogenic follicles. The size of dot indicates
indicates the percent of the total monthly sample containing number of data points at a given x, y coordinate. The inset
oviductal eggs or both oviductal eggs and vitellogenic follicles shows frequency of clutch sizes based on counts of oviductal
(see Fig. 2 for monthly sample sizes). eggs (shaded) and vitellogenic follicles (open).
respectively, in the clutch size estimates. Intercepts and
slopes were significantly different from zero in both
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models (all p values 5 0.0 1) but not from each other ( p

> 0.05). On the average, one egg is added for each
28mm-increase in SVL. Mean monthly clutch size
varied from 4 to 6.5 based on oviductal egg counts and
2.9 and 7.0 based on vitellogenic follicle counts. Based
on the Duncan's multiple range test, all monthly mean
oviductal egg counts were the same. Only 1 month,
January, was identified as differing from other months in
the mean count of vitellogenic follicles. The mean
clutch size in January was only 2.9. A paired t-test
revealed no significant difference (t = 2.70, df = 17, p
> 0.05) between follicle number and oviductal egg or
corpora lutea number in females simultaneously con-
taining both (two females with damaged ovaries were
not used in this analysis).
Wet mass of 147 preserved eggs from 26 females
averaged 1.9783 + 0.038 1 g and there was no correla-
tion (F1,22 = 1.07, p = 0.31) between mean egg wet
mass (preserved) per female and SVL. Dry egg mass
MONTH
averaged 0.7874 2 0.0063 g and the mean egg dry mass
per female was not correlated to SVL ( F ,22 = 2.85, p = FIG. 5. Seasonal cycle of testes mass (TM) and fat body
0.11). Water content of preseved eggs averaged 59.22 mass (FBM) for male Ameiva ameiva. Sample size is indicated
2 1.10%. Based on the Duncan's multiple range test, all above each figure.
monthly means of preserved egg wet and dry mass were
the same. Thus egg size does not vary with female size seasonality in testes size. Testes were largest near the
or season in Ameiva ameiva. end of the wet season and into the beginning of the dry
season (April-June, Fig. 5) with a rapid decline during
Male reproductive cycle the dry season. Virtually all males greater than 99 mm
An analysis of covariance with SVL as the covariate SVL exhibited highly convoluted epididymides and thus
revealed that testes mass was associated with SVL (r2 = appeared capable of reproducing year-round. Continu-
0.86, F23,146 = 39.0, p < 0.0001) and thus least square ous breeding capability by males is further suggested by
means were calculated on a monthly basis to examine the year-round availability of reproductive females.

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MONTH
FIG. 6. Seasonal cycle of fat body mass (FBM) in female
Ameiva ameiva (lower graph). Sample size appears above each
For personal use only.
figure. The upper graph ciepicts rainfall calculated on a 2-week
basis. The number above each bar indicates number of days
receiving greater than 1 mm rain and T indicates that only a
trace of rain was recorded.
Male and female fat body cycles
Analysis of covariance with SVL as the covariate was
used to adjust means of monthly fat body mass (FBM)
because of a significant effect of body size on FBM
(males: r2 = 0.28, F23,147 = 2.5, p = 0.0005; females:
r5 = 0.63, F23,119 = 8.7, p < 0.0001). Fat body mass in
both sexes was highly variable among individuals (Figs.
5 and 6). In males (Fig. 5), FBM tended to decrease
when testes mass increased (May-June) and tended to
increase when testes mass decreased (September-Oc-
tober). There was no clear association between FBM
and tropical wet-dry seasonality. In females, FBM
tended to increase slightly at the peak of the dry season
and into the first part of the wet season (December-Feb-
ruary), a time period just following peak reproductive
investment by females (Fig. 2). Thus it appears that
reproductive investment and fat storage are inversely
related.
Discussion
In many tropical lizards, a majority of female repro-
duction occurs during the wet season (Andrews and
Rand 1974; Duellman 1978; Fitch 1973; Fleming and
Hooker 1975; Licht and Gorman 1970; Sexton et al.
1971; and others). Some species, however, have repro-
ductive cycles which do not correspond clearly to
wet-dry seasonality (Barbault 1976; Gorman and Licht
1974; Vitt 1983). Ameiva in Caatinga of northeast Brazil
reproduce year-round, but the greatest intensity of
reproduction occurs during the peak of the dry season
(June-August) and again during the early wet season
(January-February) . Most likely, resource availability
is the proximate factor dictating intensity of reproduc-
tion in continuously breeding tropical lizards.
In the absence of data on resource availability, cycles
in fat bodies (a measure of lipid levels; Dessauer 1955;
Gorman and Licht 1974) can provide indirect evidence
of resource availability, at least in respect to its effect on
reproduction. Figures 5 and 6 revealed that males and
females had fat bodies during every month of the year.
Indeed, only 7 of 316 (2.2%) adult Ameiva lacked
enlarged fat bodies, suggesting that resource levels were
not only adequate for reproduction, but adequate for
storage above and beyond that needed for reproduction.
A correspondence of reproductive intensity to resource
levels or rainfall, which is associated with resource
abundance in seasonal tropical environments (Janzen
and Schoener 1968), might only be observed during
extended droughts, when individual lizards are not
capable of accumulating energy year-round for repro-
duction.
An alternate possibility is that the rate of egg laying
has a seasonal component. Fat bodies of females were
largest when the frequency of reproductive females was
reduced (September-February ; Figs. 2 and 5), sugges-
ting a reduced egg-laying rate during this period
allowing greater fat storage. Coincidentally, male testes
were reduced in size, suggesting a reduction in repro-
ductive activity. Thus, although reproduction is clearly
continuous in caatinga Ameiva, there appear to be
seasonal fluctuations in the intensity of reproduction or
the rate of egg laying. A major limitation of these data is
that there is no way to determine whether individual
females reproduce year-round (although many produce
at least two clutches) or whether individual females are
somewhat seasonal, but females in the population are
not synchronous.
Clutch size, based on oviductal egg or vitellogenic
follicle counts was shown to increase with body size
(and presumably age). Because females can produce
more offspring by increasing body size, selection should
favor rapid growth and large body size in females.
Trivers (1972) suggested that female body size should
tend to be large in species which can vary clutch size
(considering clutch size a correlate of reproductive
success) with body size. In Anolis garmani, a species
which does not vary clutch size with body size (clutch
size is consistently one), reproductive success as mea-
sured by number of copulations, was highest in larger
females, presumably because they could produce
clutches faster than smaller females (Trivers 1972).
In the absence of competition among males for
females, females should be larger than males if repro-
ductive success is associated with size. However, males
 31 18 CAN. J. ZOOL. VOL. 60, 1982
tend to be larger than females in Ameiva, suggesting the
possibility of competition among males. On several
occasions I observed premating , mating, or aggressive
behavior in Ameiva. Pertinent to sexual selection,
chasing and fighting among male teiids is not uncom-
mon (Stamps 1977). Intrasexual aggression has been
reported, for example, in Ameiva quadrilineata (Hirth
1963) and a number of other teiids (Stamps 1977).
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Reproductive success in males may be associated with
size and this perhaps explains the apparent size dimor-
phism in Ameiva ameiva. An alternate hypothesis is that
reproduction is riskier for females than males and that
apparent sexual dimorphism in size is simply a reflec-
tion of higher mortality rates on larger sized females.
Presumably, egg or offspring size is optimized in
animals (Smith and Fretwell 1974; Brockleman 1975)
and the lack of association of egg size in Ameiva with
any measure of parental size or season adds support to
this hypothesis. Recently, offspring size of certain
female lizards has been shown to vary in association
with seasonality in resource levels (Derickson 1976;
Ferguson et al. 1980; Nussbaum 1981). The low value
for relative clutch mass (RCM) reported here (0.159) is
For personal use only.
similar to that reported for other macroteiids and
supports models predicting low RCM in widely foraging
species for ecological and energetic reasons (Vitt and
Congdon 1978; Vitt and Price 1982).
It is difficult to make accurate geographic compari-
sons among Ameiva populations. Ameiva ameiva re-
production has only been studied in Amazonian Ecuador
(Duellman 1978; Simmons 1975) and Peru (Dixon and
Soini 1975). Simmons ( 1975) considered the reproduc-
tive season to extend from late May through early
December, although his data were incomplete from
January to March. The observations of Dixon and Soini
(1975) indicate that Ameiva ameiva breed continuously
in the Iquitos region of Peru. Hirth (1963) suggested
nearly continuous egg laying by Ameiva quadrilineata
in Costa Rica. Other tropical macroteiids including
Cnemidophorus ocellifer (Vitt 1983), Cnemidophorus
deppii and Cnemidophorus lemniscatus (Fitch 1970)
apparently breed continuously.
Lizards of various taxa which are sympatric with
Ameiva ameiva in northeast Brazil exhibit a diversity of
reproductive tactics. The iguanids Tropidurus torquatus
and Platynotus semitaeniatus reproduce primarily dur-
ing the dry season and early wet season (Vitt and
Goldberg 1983), whereas another iguanid, Polychrus
acutirostris, reproduces at the beginning of the wet
season over a 3-month period (Vitt and Lacher 1981).
Thus it appears that Ameiva ameiva reproduction in
northeast Brazil is more like reproduction in other
tropical macroteiids regardless of geographic location
than it is like sympatric species of different taxa. A quite
different situation exists among sympatric skinks in
Africa. Barbault (1976) found that Mabuya buttneri
reproduced in the late wet season, M . maculilabris
reproduced in the dry season, and Panaspis nimbaensis
reproduced in the wet season. Thus tropical iguanids and
skinks exhibit a diversity of reproductive tactics within
and among localities in respect to seasonality, whereas
macroteiids do not. The capability of macroteiids to
reproduce continuously may be an adaptation allowing
them to take advantage of high resource habitats that
may be unpredictable in time and space. The high
mobility of these widely foraging lizards may make it
possible for them to find high resource patches that
would be unavailable to sit-and-wait predators such as
most iguanid lizards. The latter tend to be habitat
specialists (Vitt and Price 1982) and thus many patches
are unavailable to them. The specific habitat patches
used by sit-and-wait foragers may have seasonal re-
source variation which affects timing of reproduction.
It is clear that a great deal remains to be learned about
reproductive tactics of tropical lizards. Sorting out
phylogenetic , historical, and ecological factors contri-
buting to the suite of reproductive characteristics that
renders individuals of a species successful in a particular
environment remains to be one of the overriding
challenges to evolutionary ecologists.
Acknowledgments
T. E. Lacher, K. E. Streilein, and M. Willig provided
occasional assistance during the field portion of this
study. The people of Exu, Pernambuco, were particu-
larly cooperative and provided a setting which fostered a
great amount of research. I thank Chico Ventura for
allowing me access to the relatively undisturbed Fa-
zenda Batente, where much of the fieldwork was carried
out. Dr. Paulo E. Vanzolini, Museo de Zoologia da
Universidade de S i o Paulo, provided the impetus for
this and other studies in addition to keeping up the flow
of much-needed supplies. Aristides Leio and the people
at the Brazilian Academy of Sciences were of unmea-
surable help. Comments of two anonymous reviewers
aided greatly in the final revision. Financial support was
provided by the people and Federal government of
Brazil in support of the project: Estudos Ecologicos no
Nordeste Semi-arido do Brasil, undertaken by the
Academia Brasileira de Cikncias. The final stages of this
research and manuscript preparation were conducted
under contract EY-76-C-09-0819 between the United
States Department of Energy and the University of
Georgia.
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