The reproductive ecology of exotic Trachemys scripta elegans in an

invaded area of southern Europe

NATIVIDAD PEREZ-SANTIGOSA, CARMEN DIAZ-PANIAGUA* and JUDITH HIDALGO-VILA

Estación Biológica de Doñana-CSIC, P.O. Box 1056, 41080 Sevilla, Spain

ABSTRACT
1. The main reproductive parameters of exotic Trachemys scripta elegans in established populations from two
ponds in southern Spain are described.
2. Females were found to reproduce at 5 years of age, although sexually mature individuals of 4 and 3 years of
age were also detected. Annual reproductive frequency was estimated to be 80–86% of reproductive females.
Mean clutch size was 11.5 eggs and was not correlated with maternal body size.
3. The number of follicles of different size in ovaries suggested that four or more clutches could be laid
per season. Eggs were laid from April to June (inclusive) mainly during morning hours, when females were easily
detected during nesting trips. 79.5% of eggs were fertile.
4. Reproductive parameters of this species in southern Spain were of similar or even higher values than in
native areas, thus placing them among chelonian species of earlier maturity.
5. Although chelonians might never be considered explosive breeders, the number of T. s. elegans individuals
could surpass that of native aquatic chelonians in southern Spain, where T. s. elegans reach maturity earlier, are
more fecund and their eggs are more fertile.
Copyright # 2008 John Wiley & Sons, Ltd.

KEY WORDS: exotic turtles; reproduction; life history traits; population establishment

INTRODUCTION Nowadays, free-living individuals are reported in countries on

the five continents of the world (Newberry, 1984; Gasperetti
The introduction of exotic species poses a serious threat to et al., 1993; Luiselli et al., 1997; Chen and Lue, 1998; Brinjsøe,
aquatic ecosystems. Trachemys scripta elegans, an intentionally 2001; Cadi et al., 2004; Feldman, 2007; Perry et al., 2007).
introduced species, is considered to be among the most While in some countries successful reproduction is considered
common reptile pets traded worldwide (Salzberg, 1995, 1998; unviable (e.g. in New Zealand: Feldman, 2007), in many
Lowe et al., 2000; Telecky, 2001; Reed and Gibbons, 2003). In others, such as Mediterranean countries, these turtles find
recent decades millions of hatchlings of this species, a popular favourable conditions for field incubation of eggs. Successful
pet, have been exported from USA farms to many countries. reproduction has been reported in several countries outside of

*Correspondence to: C. Diaz-Paniagua, Estacion Biologica de Donana-CSIC, P.O. Box 1056, 41080 Sevilla, Spain. E-mail: [email protected]
its native range, such as in France (Cadi et al., 2004), Italy sample of 100 females from El Acebuche and 99 females from
(Ferri and Soccini, 2003), Taiwan (Chen and Lue, 1998), and El Portil were included in this study. From 2002–2004, 53
in a wide number of localities within Spain (Brinjsøe, 2001; females were captured by hand while migrating for nesting,
Pleguezuelos, 2004). Frequent release of pet turtles by their and 60 females and juveniles were trapped with baited
owners coupled with their considerable reproductive output submerged traps. In 2005, 86 turtles were captured with
favour the establishment of naturalized populations. However, basking traps, which notably increased the efficiency of exotic
established populations in countries where they are non-native turtle trapping (Perez-Santigosa et al., 2006b). The hour (in
have only been reported in Taiwan (Chen and Lue, 1998), the GMT) at which nesting females were found was recorded. All
Caribbean islands (Perry et al., 2007) and in Spain (Perez- individuals were euthanized with thiopental sodium injection
Santigosa et al., 2006a), although the actual distribution of (Tiobarbital, Braun Medical), and their straight carapace
reproductive populations has not previously been studied in length (SCL) and plastron length (PL) were measured with
detail. callipers ( ± 1 mm), and body mass ( ± 1 g) was recorded with
In Spain, the introduction of exotic turtles is considered to an electronic balance. The ovaries and oviducts from 34
threaten the status of the two native species of aquatic turtles females from El Acebuche and 31 from El Portil were
(Emys orbicularis and Mauremys leprosa), which are at present extracted, weighed and preserved for later examination.
listed as vulnerable, with declining populations (Pleguezuelos Body mass of females with shelled eggs in oviducts was
et al., 2004). Recent studies have demonstrated the influence of recalculated by subtracting clutch mass.
competition of this species with the European pond turtle After a preliminary inspection of external characters of
Emys orbicularis, the most widely distributed aquatic individuals, they were classified as juveniles or adults.
chelonian in Europe, at present considered to be a Dissected young females with ovaries with developing
threatened species (Cadi and Joly, 2003, 2004). In Spain, follicles, eggs, or distended oviducts indicative of recent
free-living individuals of T. s. elegans were first reported in oviposition were considered adults. Shelled eggs borne in
1996 (Garcıa-Parıs and Martın, 1997), and the species now females were counted and their length, width and mass
occurs throughout the country, including the Canary and measured. Clutch mass was estimated as the sum of masses
Balearic Islands (Pleguezuelos, 2004). of all eggs, and relative clutch mass (RCM) was calculated as
This paper is based on the analyses of exotic turtles the ratio of clutch mass to body mass of females. Follicles in
removed from two established populations in southern oviducts were classified in four groups, according to diameter
Spain. The aim of this study was to describe their main (I) 57 mm (their number were not included in results), (II) 7–
reproductive characteristics, information basic to the analysis 13 mm, (III) 14–20 mm; (IV) >21 mm; following Moll and
of population trends and demography, to enable a comparison Legler (1971). The number of clutches that females lay
with the dynamics of native turtle species. Reproductive per season was estimated from the number of follicles of
parameters, such as age at maturity and fecundity, may be various sizes and eggs (after Moll and Legler, 1971). Age of
used to evaluate the ability of turtles to establish wild individuals was estimated by skeletochronology, counting the
populations in natural habitats. Other detailed information, number of annuli in an 18 m section of the femur, after tinction
such as the description of the nesting season, may be of special with haematoxylin (see details of this technique in Castanet
interest for improving programmes to eradicate these exotic and Smirina (1990)). Total number of annuli was considered
turtles. equivalent to the age of females. Annual frequency of
reproduction was estimated from data on females captured
with aquatic traps in 2004 (18 in El Acebuche, and 21 in El
Portil), not from data on nesting females, as the latter data
METHODS could have led to an overestimation of frequency.
Females captured in 2004 and 2005 were inspected by
This study was carried out in two coastal lagoons, El Acebuche inguinal palpation to detect oviductal eggs and seven gravid
and El Portil (in the province of Huelva, south-western Spain). females from El Acebuche and two from El Portil were
Wild populations of T. s. elegans have established themselves induced to oviposit by injection of oxytocin (Ewert and
in these lagoons after successful reproduction of pet turtles Legler, 1978). Eggs were also obtained from 14 nests (of
released by owners around 1996–1998. A detailed description recent oviposition) from El Portil and two from El Acebuche.
of these populations was reported in Perez-Santigosa et al. In order to estimate the proportion of fertile eggs, 227 eggs
(2006a). from 25 different clutches were incubated in moistened
From 2002–2006, a programme for removing exotic turtles vermiculite at a constant temperature of 28.58C. These eggs,
was carried out in both localities. More than 250 adult turtles used in different experiments, were not uniformly subject to
were removed from El Acebuche and 400 from El Portil. A identical moisture conditions, and so the values obtained for

Copyright # 2008 John Wiley & Sons, Ltd. Aquatic Conserv: Mar. Freshw. Ecosyst. 18: 1302 – 1310 (2008)
DOI: 10.1002/aqc
hatching rate were not included in this study. Fertility rate Reproductive frequency
was estimated as the number of eggs which hatched or
contained embryos as a proportion of the total number of Considering only those dissected females not captured in their
incubated eggs. All those eggs which did not hatch were nesting migrations, 12 out of 15 (80%) females captured in El
opened and inspected to determine the presence or absence of Acebuche had enlarged or preovulatory follicles, eggs or
embryos. distended oviducts, suggesting recent oviposition. The
Comparison of variables from females, eggs and hatchlings remaining three females did not show signs of oviposition
of the two study sites were tested with ANOVA, using locality nor follicle development, although they were sexually mature.
as a grouping factor. ANOVA with month as a grouping In El Portil lagoon, 18 out of 21 females examined (85.7%)
factor was also used to analyse ovarian mass variation, showed signs of reproduction.
including data from all dissected females from both
localities. The relationship among clutch size and SCL of The annual reproductive cycle
females was analysed with Pearson correlation coefficients.
The description of the main reproductive characteristics is Ovaries differed significantly in mass among individuals
indicated by mean ± standard deviation (sd). captured in different months (F6,51=0.587, P=0.0001).
Females with heaviest ovaries were found in April, and a
gradual decrease of ovarian mass was recorded in the
following months until July. In those months most females
had follicles of the three largest size-classes, and eggs (Table 2).
RESULTS
Lowest ovarian masses were recorded during summer months
(July–September), when a period of quiescence is considered to
Body size and age at maturity occur, as reported for Trachemys venusta from Panama (Moll
Adult females from both localities did not differ in SCL, PL
and body mass (Table 1). The youngest adult female was 3
years old, although only 32% of females of this age appeared
sexually mature, with mature ovaries. Also 64% of 4-year-
olds and 92% of 5-year-old females were mature, as were all
older females. Eggs or recent oviposition were only detected
in females 5 years or older. Because these are young
populations, no individual older than 13 years was
captured during the study period (Figure 1). The smallest
adult female was 170.7 mm SCL (Table 1), while the smallest
egg-bearing female was 184.9 mm SCL, 179 mm PL and Figure 1. Age estimated by annuli counted in sections of humeri in
864 g. adult and subadult female Trachemys scripta elegans.

Table 1. Mean, standard deviation and range (min-max) of straight carapace length (SCL), plastron length (PL) and body mass (excluding clutch
mass) of females; clutch size, length, width and mass of eggs; relative clutch mass (RCM=clutch mass/body mass); and straight carapace length,
plastron length and mass of hatchlings. Results of ANOVA between values of the two populations studied are also indicated.

Acebuche Portil ANOVA

n x̄ std dev min-max n x̄ std dev min-max

Female SCL (mm) 99 212.06 15.87 170.7–257.8 95 209.90 18.48 171.0–248.3 F1,192=0.765, P=0.383
Female PL (mm) 99 201.06 14.42 159.5–236.0 95 196.59 17.19 159.5–232.1 F1,192=3.851, P=0.051
Female mass (without eggs) (g) 100 1419.50 321.00 718.0–2700.0 96 1397.26 356.58 797.0–2204.0 F1,194=0.211, P=0.647
Clutch size (eggs) 29 11.34 2.45 6–15 22 11.73 2.07 9–16 F1,49=0.338, P=0.563
Egg length (mm) 26 33.48 2.03 29.45–39.10 20 34.12 3.02 28.26–39.53 F1,44=0.732, P=0.397
Egg width (mm) 26 21.81 0.82 20.35–23.47 20 21.89 1.39 18.80–24.87 F1,44=0.056, P=0.815
Egg mass (g) 26 9.43 1.14 6.88–11.80 20 9.74 1.85 7.04–14.32 F1,44=0.463, P=0.500
RCM (%) 25 7.15 1.70 3.85–10.10 20 7.30 1.99 3.91–11.31 F1,42=0.052, P=0.821
Hatchling SCL (mm) 8 28.383 1.82 24.35–30.55 10 29.16 1.75 26.47–31.60 F1,16=0.851, P=0.370
Hatchling PL (mm) 8 27.15 2.15 22.0–28.5 10 26.92 2.01 24.25–29.79 F1,16=0.056, P=0.816
Hatchling mass (g) 8 6.71 0.98 4.50–7.48 10 6.81 1.21 4.88–8.64 F1,16=0.034, P=0.856

Copyright # 2008 John Wiley & Sons, Ltd. Aquatic Conserv: Mar. Freshw. Ecosyst. 18: 1302 – 1310 (2008)
DOI: 10.1002/aqc
Table 2. Monthly variation of the percentage of females with follicles of different size groups, or with oviductal eggs; and of the mean number and
range of follicles or eggs
Month n Class II Class III Class IV Eggs
April 6 % females 100 100 83.3 66.7
mean number of follicles 16.2 (15–19) 23.5 (12–41) 10.0/11.8(5–19) 11.0 (8–13)
May 11 % females 90.9 100.0 18.2 54.6
mean number of follicles 15.5 (12–22) 25.8 (12–42) 3.9/17.0 (15–19) 11.8 (10–13)
June 11 % females 81.8 100.0 36.4 90.9
mean number of follicles 11.8 (11–23) 18.3 (10–24) 5.5/13.5 (5–17) 11.4 (9–14)
July 18 % females 77.8 55.6 5.6 50.0
mean number of follicles 15.1(11–51) 16.8 (7–26) 0.5/7.0 (7–7) 11.0 (8–15)
August 11 % females 100 18.2 0 0
mean number of follicles 25.5 (7–46) 25.5 (15–36) 0 0
September 4 % females 75.0 25.0 0 0
mean number of follicles 37 (35–39) 8.0 0 0
November 2 % females 100 100 0 0
mean number of follicles 47.5 (42–53) 16.0 (11–21) 0 0

had finished, it was assumed that these follicles were not

necessarily related to clutches produced in the current year,
but might correspond to clutches of the following year.
Thus, as only a proportion of follicles in class III and all of
class IV are involved in annual fecundity, the number of
clutches was approximated by the sum of the number of
Class III and IV follicles and of eggs divided by the number
of oviductal eggs of the last clutch when detected (or by
mean clutch size, when females had already oviposited). In
April and May, females seemed to bear at least three
clutches, with eggs and follicles of Groups III and IV. The
large number of enlarged follicles in group III and IV of
some females suggests that they could lay four or even more
clutches per year (Table 2).

Figure 2. Monthly variation in mass of ovarian follicles measured in Clutch size and egg characteristics
female Trachemys scripta elegans from established wild population in
southern Spain. Mean clutch size was 11.5 ± 2.31 eggs, and did not differ
between both localities (F1,49=0.338, P=0.563) nor among the
and Legler, 1971). The oviposition period was April–July. different months of the nesting season (F3,43=1.450,
From August–September, females did not bear eggs, nor large P=0.242). The lowest mean value of clutch size was
follicles (class IV), but mainly contained small follicles (Class found in April and the largest in May (Figure 3). SCL and
II); a small proportion of females also contained a small body mass of females were not correlated with clutch
number of preovulatory follicles (Class III). In November, an size (SCL: r=0.113, P=0.432; Body mass: r=0.046,
increase in mass of the ovaries was detected, corresponding to P=0.751). Average values of egg length, width and mass
the phase of follicle enlargement, just before the winter are given in Table 1, using all values obtained from nests
inactivity period when all females had follicles of Class I and and oviducts, as significant differences among measurements
II only (Figure 2, Table 2). were not observed for eggs obtained with different techniques.
These egg variables did not differ between females from both
localities (Table 1), and neither did they differ in relation
Clutch number
to clutch size. Clutch mass constituted on average 7.2 ± 1.8%
Because a wide variation in the number of Class II follicles of female mass, and RCM did not differ between localities
was recorded every month, even after the oviposition period (Table 1).

Copyright # 2008 John Wiley & Sons, Ltd. Aquatic Conserv: Mar. Freshw. Ecosyst. 18: 1302 – 1310 (2008)
DOI: 10.1002/aqc
Hatchlings (mean=30.91 ± 2.70, F1,18=6.41, P=0.021) than hatchlings
from laboratory-incubated eggs. These differences may be
Mean values of SCL, PL and body mass of hatchlings are explained because individuals were measured in the laboratory
shown in Table 1. These values corresponded to individuals immediately after hatching, when their shell could still be
hatched from eggs incubated in the laboratory, and were not curved due to their position inside the egg, while hatchlings
significantly different from eggs from both localities. However, incubated in the field but not yet emerged from the nest were
data recorded from six hatchlings from two nests naturally extracted from the nest approximately 1 month after hatching.
incubated in the field did not reveal differences in mass Hatchling SCL and mass were not correlated with maternal
(mean=7.11 ± 2.36 g), but they had longer SCL characteristics, but they were positively correlated with
(mean=32.24 ± 3.20 mm, F1,18=5.98, P=0.025) and PL egg length (regg length×hatchling mass ¼ 0:591; P=0.008;
regg length×hatchling length ¼ 0:593; P=0.007) and mass
(regg mass×hatchling mass ¼ 0:607; P=0.006; regg mass×hatchling length
¼ 0:458; P=0.048).

Nesting
Females were found migrating for nesting or coming back to
the lagoon from early April to 23 July. Except for one female,
which was found nesting in the evening in June (17:00), all
females nested from 7:30 to 14:00 (Figure 4). However, a
variation in the time at which females nested was observed
among the different months of the nesting season
Figure 3. Clutch size of female turtles captured in different months of (F3,26=3.182, P=0.041). In April females were found nesting
the nesting season. from 09:30 to 14:30, while later in the season nesting was

Figure 4. Number of Trachemys scripta elegans females captured on land in their nesting migrations, and monthly variation of the mean (maximum
and minimum) hour at which females were found nesting throughout the nesting season. An exceptional female nesting at 17 h in June is not shown.

Copyright # 2008 John Wiley & Sons, Ltd. Aquatic Conserv: Mar. Freshw. Ecosyst. 18: 1302 – 1310 (2008)
DOI: 10.1002/aqc
mainly concentrated earlier in the morning, before the warmest Morin, 1988; Congdon and Gibbons, 1990). The earlier
periods of the day in summer months (Figure 4). maturity ages of female T. s. elegans, although detrimental
to adult longevity, may benefit the initial establishment of
Egg fertility these exotic species populations, as it reduces the time elapsed
before incorporation of new individuals. In fact, the time
On average, 79.5% of the eggs of each clutch were fertile, with required for establishment of these populations in the wild is at
no significant differences between localities (F1,2350.001, least 3–4 years, the minimum time that the first individuals
p=0.997). All eggs (100%) were fertile in 50% of the born in the wild need to become reproductive. Compared with
clutches obtained from El Acebuche and 33.3% from El other species of chelonians, maturing from 4 to 25 years of age
Portil. The minimum fertility rate corresponded to one clutch (see review in Iverson, 1992), T. s. elegans females in southern
from El Acebuche, in which only one of the three eggs Spain reach maturity at a notably early age.
incubated was fertile.

Reproductive frequency
DISCUSSION Turtles are considered organisms of moderate investment in
reproduction (Wilbur and Morin, 1988; Congdon and
Female size and age at maturity Gibbons, 1990), and females of many species do not
reproduce every year (see references in Frazer et al., 1990).
Several studies have analysed the size and age of sexual
In their original range, 50–70% of female T. scripta reproduce
maturity in female Trachemys in North America, where a wide
annually in some localities (Congdon and Tinkle, 1982), and
variation is observed among populations, species and
27.2–47.1% in others (Cagle, 1950; Gibbons and Greene,
subspecies (Cagle, 1950; Moll and Legler, 1971; Gibbons
1990). In contrast, in southern Spain the reproductive
et al., 1981, 1982; Gibbons and Greene, 1990; Mitchell and
frequency of exotic female T. scripta was notably larger (80–
Pague, 1990; Moll and Moll, 1990; Vogt, 1990; Close and
85% reproductive females per year), thus increasing the
Seigel, 1997; Tucker and Warner, 1999; Aresco, 2004).
reproductive potential of newly established populations
In Trachemys scripta, sexual maturity is influenced mainly
relative to populations in their original range. This difference
by body size rather than age of females (Cagle, 1950; Moll,
may be due to the younger age of females in the exotic
1979; Wilbur and Morin, 1988; Gibbons and Greene, 1990).
populations studied. Wild populations in North America are
Females mature with a plastron length of 160–170 mm, and
composed of individuals of a wider age structure, and it is
observed differences in growth rate imply differences in the age
probable that reproductive frequency may be affected by the
of sexual maturity. Thus, in very productive habitats, turtles
age of females, or even by senescence of older individuals
reached mature size at a younger age (4 years old, Congdon
(Frazer et al., 1990). In fact, the oldest female captured in
and Gibbons, 1983) than in other localities, where females
Spain was 13 years old, while in native US populations,
matured at 6–8 years old (Congdon and Gibbons, 1983;
females are reported to reach at least 31 years (Frazer et al.,
Gibbons et al., 1981; Frazer et al., 1990; Mitchell and Prague,
1990).
1990). These data correspond to the subspecies T. s. scripta,
while for the subspecies T. s. elegans, there is no precise
information about the age and size at maturity, although Nesting period and clutch number
reproductive females were reported from 167 mm PL (Tucker
and Moll, 1997). North American female T. scripta nest from mid-April to early
The data from the present study showed that, as an exotic August (Cagle, 1950; Congdon and Gibbons, 1983; Jackson,
invader, T. s. elegans in southern Spain may reach maturity at 1988; Aresco, 2004), although the subspecies T. s. elegans is
159.5 mm PL and at 3 years old, although at this age only a reported to nest only in May and June (Aresco, 2004). In the
small percentage of females were mature, while most 5-year- present study area, the nesting period of this exotic species was
old females were mature. This early maturity suggests that in 4 months, similar to the longer period in the original range of
these localities, exotic turtles have a similar, or an even higher, T. scripta. This long period is favoured by the climatic
growth rate than in the most productive habitats of the native character of the study area, in which other reptiles such as the
range of the species. The delayed age of maturity of chelonian terrestrial chelonian Testudo graeca (Dıaz-Paniagua et al.,
species, attributable mainly to a greater investment in survival 1996) have a similar nesting season. In contrast, the two native
than in reproduction (Congdon and Gibbons, 1990; Iverson, aquatic turtles of the same study area concentrate their nesting
1992), is characteristic of a life-history strategy with low in only two months, mainly June and July (Keller and Busack,
reproductive rates and low population increase (Wilbur and 2001; Roques et al., 2006).

Copyright # 2008 John Wiley & Sons, Ltd. Aquatic Conserv: Mar. Freshw. Ecosyst. 18: 1302 – 1310 (2008)
DOI: 10.1002/aqc
 In North America, females mainly nest on warm days after to fertilize all clutches. However, although the proportion of
rains. In southern Spain, a gradual increase in temperature and males in relation to females was found to be low in one of the
notable decrease in rainfall occurs throughout the nesting two populations studied (Perez-Santigosa et al., 2006a),
period, so that the last two months are dry and hot. A similar 4 fertility rates were high, and all clutches inspected had fertile
month nesting period was reported for these exotic aquatic eggs. The low proportion of males could be compensated for
turtles in France, although delayed to May–August (Cadi by sperm storage, which has been reported to occur in this
et al., 2004). However, in other European localities, such as species (Gist and Congdon, 1998). Fertility rates were even
central Italy, the viability of reproduction has been questioned higher in clutches of T. s. elegans than in native aquatic turtle
(Luiselli et al., 1997). species, which may also use stored sperm; however, fertility in
native clutches fertilized by stored sperm has been reported to
Clutch number decrease (Roques et al., 2006).

Constraints of abdominal cavity influence clutch size in

chelonians, but females may increase their annual fecundity
by producing multiple clutches per year (Moll, 1979; Congdon Comparison with native turtles and implications for
and Gibbons, 1985). Species with longer nesting seasons conservation
increase the possibility of laying more clutches in relation to
other species having a shorter nesting period. Species laying Trachemys s. elegans is considered at present to be among the
multiple clutches have longer reproductive periods than species 100 most invasive species in the world (Lowe et al., 2000). Low
laying a unique annual clutch (Moll, 1979). Female Trachemys population increase and moderate reproduction characteristic
venusta lay up to six clutches in Panama (Moll and Legler, of chelonians imply that this is not an explosive species capable
1971) and four clutches in Mexico (Vogt, 1990), in nesting of rapid colonization of feral habitats. Among chelonians,
seasons of 4–5 months. North American female T. s. elegans however, T. s. elegans is one of the species with earliest
may lay at least three different clutches annually, with maturity and greater fecundity than most other species, which
internesting periods of 14–22.5 days (Tucker, 2001). favour population establishment in new habitats. In
Although information on successive clutches from females in competition with native chelonians, invasive T. s. elegans
southern Spain was not obtained, the number of different could reach a larger number of individuals in optimal
groups of follicles and eggs suggest that at least up to four environments, with favourable availability of resources. In
clutches annually may be produced. Considering the inter- comparison with native aquatic turtles from the localities
nesting interval described for this subspecies, a maximum studied in southern Spain (M. leprosa and E. orbicularis),
number of five to eight clutches could be expected in the four exotic turtles reach maturity at an earlier age, are more fecund
months of the nesting season in this area. and fertile (Table 3), and would be favoured by their larger
body size in agonistic encounters for resources. In fact, T. s.
elegans has been reported to affect survival of E. orbicularis,
Fertility rates
competing for basking sites (Cadi and Joly, 2004) while the
In a newly established population, the sex ratio could be displacement of individuals of M. leprosa has also been
unequal, and the number of available males may be insufficient detected in the study areas (unpublished data). Thus, the

Table 3. Comparison of the main reproductive characteristics of Trachemys scripta elegans and of the two native aquatic turtles from southern Spain
Trachemys s. elegans Mauremys leprosa Emys orbicularis
a
Female SCL (mm) 211.0 (170.7–257.85) 176 (131.5– 228) 142.6 (129–167)a
Female mass (g) 1408.4 (718–2700) 743.7 (329–1650)a 526.6 (340–820)a
Nesting season April–July May–Junea June–Julyb
Clutch number >=4 2a 2–3a
Clutch size 11.5 (6–16) 6.4 (3–13)a 6.4 (3–10)a
Age at maturity (years) 3–4–5 7a Approx. 5e
Size at maturity (mm) 170–171 120–140a,d 129.8a
Fertility 79.5% 78.27%c 76.44%b
a
Keller (1997).
b
Roques et al. (2006).
c
Authors’ unpublished data based on incubation of 76 eggs (from 14 clutches) at constant 288C.
d
Perez et al. (1979).
e
Estimated from the age of females calculated in relation to the size (SCL) at maturity, after Keller (1997).

Copyright # 2008 John Wiley & Sons, Ltd. Aquatic Conserv: Mar. Freshw. Ecosyst. 18: 1302 – 1310 (2008)
DOI: 10.1002/aqc
introduction of this exotic species constitutes a major threat to introduced slider turtle (Trachemys scripta elegans) in the
the conservation of these two native species. South of France. Aquatic Conservation: Marine and
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reproductive characteristics of T. s. elegans provides basic, scripta troostii (Holbrook). Ecological Monographs 20: 31–
54.
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Copyright # 2008 John Wiley & Sons, Ltd. Aquatic Conserv: Mar. Freshw. Ecosyst. 18: 1302 – 1310 (2008)
DOI: 10.1002/aqc