The study of the feeding ecology of a given species
generates information about the prey types, most
frequent food items, relative importance of each prey,
inter- and intraspecic relationships and strategies
of food partitioning (Klawinski et al., 1994; Saenz,
1996; Aowphol et al., 2006; Bonglio, Balestrin and
Cappellari, 2006). A high dietary niche breadth is
characteristic for invasive species, which in combination
with adequate ecological conditions, provides a solid
basis for successful colonization of new environments
(Powell, Parmerlee and Rice, 1990; Case, Bolger and
Petren, 1994; Rödder, Solé and Böhme, 2008).
The Tropical House Gecko Hemidactylus mabouia
(Moreau de Jonnés, 1818) (Fig. 1) belongs to the
family Gekkonidae. It is a broadly distributed species
in the tropics (Henderson and Powell, 2009) and has
invaded the West Indies after being introduced from
its native range, like other species of the genus, via
slave ships coming from Africa during the European
colonization (Weiss and Hedges, 2007). This species
is an effective colonizer and nocturnal predator that is
usually associated with human buildings. Hemidactylus
mabouia is well known for using areas around articial
light sources as hunting grounds (Currat, 1980; Powell
and Henderson, 1992; Daniells et al., 2008; Perry et
al., 2008). Population densities can range between
0.04 and 0.21 individuals/ m2 (Howard, Parmerlee and
Powell, 2001). In Cuba, this species was rst reported
from Eastern Cuba (Schwartz and Henderson, 1991).
However, in recent years, H. mabouia has gradually
invaded central and western Cuba, displacing the closely
related invasive species H. angulatus (Díaz, in prep.).
Despite its current colonization in Cuban territory, there
is no information regarding the ecology and especially
the feeding habits of H. mabouia.
Herpetology Notes, volume 6: 11-17 (2013) (published online on 25 January 2013)
Feeding ecology of the Tropical House Gecko
Hemidactylus mabouia (Sauria: Gekkonidae)
during the dry season in Havana, Cuba
Manuel Iturriaga1,* and Rubén Marrero2
1 División de Colecciones Zoológicas. Instituto de Ecología y
Sistemática. Carretera de Varona, Km 3 ½, Capdevila, Boye-
ros, AP 8029, CP 10800, La Habana, Cuba.
2 Facultad de Biología. Universidad de La Habana. Calle 25
# 455 e/ I y J, Vedado, Plaza de la Revolución, La Habana,
*Corresponding author; e-mail: firstname.lastname@example.org
Abstract. The gecko Hemidactylus mabouia is an invasive species and a successful colonizer with a broad distribution in the
tropics. The species is frequently associated with human buildings on which individuals prot from the availability of potential
prey items attracted to articial light sources. The feeding ecology of H. mabouia was studied in an urban area of Havana, Cuba,
in order to evaluate diet composition and potential differences among adult males, adult females and juveniles during the dry
season. The main prey items encountered consisted of non-ying arthropods (cockroaches, spiders and pill bugs). Cockroaches
contributed most to the ingested volume. Snout-vent length did not signicantly differ between adult males and adult females,
whereas signicant differences were recovered in head width among the three sex/age categories. No relationship between body
and head size was found in relation to the number and volume of prey items. All analysed individuals tended to feed on small-sized
prey. Microhabitat selection is suggested to be the main factor that determines diet composition in adult males and juveniles..
Keywords. Feeding ecology, diet composition, dry season, Hemidactylus mabouia, Havana, Cuba.
Figure 1. Male adult of Hemidactylus mabouia from Havana,
Cuba. Photograph by Rubén Marrero.
Manuel Iturriaga & Rubén Marrero
One of the limiting factors that prevents a better
understanding of the problems caused by invasive
species is the lack of information about aspects of their
natural history and ecology (Rocha and Anjos, 2007;
Caicedo-Portilla and Dulcey-Cala, 2011; Hoskin, 2011).
Therefore, this article aims to study the diet composition
of an urban population of H. mabouia from Havana,
Cuba, during the dry season and to determine whether
intraspecic food partitioning in trophic niches exists.
Materials and Methods
The study was carried out from November 2009 to February
2010 at the zoo Jardín Zoológico de La Habana “La Edad de Oro”
(26° 06‘ 41.25“ N, 82° 23‘ 50.18“ W; elevation ca. 39 m SIU)
in Havana, Cuba. During each month, 15 individuals of H. ma-
bouia were captured which resulted in a total of 61 specimens. All
geckos were collected by hand during the period between 18:30
and 22:00 h. Individuals were encountered on tree trunks, on the
ground under rocks and human debris, and on the wall of human
housing, but not under articial lights. Specimens captured were
immediately sacriced and preserved in 70% ethanol. The snout-
vent length (SVL), head length (HL) and head width (HW) were
measured in each individual using a digital calliper (to 0.01 mm).
Sex determination was performed based on secondary sexual cha-
racters, presence of preanal pores in adult males and endolympha-
tic sacs behind of the head in adult females, and nally by gonadal
identication after dissection. The specimens were divided into
three sex/age categories.
Specimens were dissected and their stomachs were removed
for posterior content analyses under a stereomicroscope. Stomach
contents were identied to the taxonomic level of Order. The con-
tents that could not be identied were pooled into a “non-identi-
ed” category. The food items of each specimen were counted
and measured (length and width) using digital calliper for larger
prey and X15 eyepiece micrometre coupled to the binocular ste-
reomicroscope (to 0.014 mm) for smaller prey.
The number of prey was dened as the amount of prey of a
given item in the total diet, and the frequency as the total number
of stomachs in which a given item was found. The volume of
each item (mm3) was estimated by the prolate spheroid volume:
Volume= 4/3π X (L/2) X (W/2)2 where L= length and W= width
(e.g. Colli et al., 2003; Bonglio, Balestrin and Cappellari, 2006).
The importance of each food item consumed (i) was described
by the index of relative importance (Pinkas et al., 1971), which
was calculated as IRIi = %Fi(%Ni+%Vi), where %Fi is the percent
frequency (the number stomachs containing each item i); %Ni is
the percent of the number of i items in all stomachs; and %Vi
is the percent of the volume of items i in all stomachs. Levin´s
standardized niche breadth index (Hurlbert, 1978), B = (1/Σpi
– 1/n-1, was used to determine niche breadth in number and vo-
lume of ingested prey, where pi is the proportion of occurrence of
items i in the diet composition, and n is the total number of prey
taxa. B ranges from 0 to 1; a value of near 1 means that every prey
was used in same proportion, whereas a value near zero means
that only one or a few food items were used in higher proportions
and the rest in lower proportions. Pianka´s index of niche overlap
Figure 2. Linear regression between head length and snout-
vent length (SVL); and between head width and SVL of
juveniles and adult specimens of Hemidactylus mabouia
from zoo Jardín Zoológico de La Habana “La Edad de Oro”,
Havana, Cuba. Solid line represents the regression line and
dashed line represents 95% condence intervals.
Figure 3. Comparisons of head width residual among adult
males, adult females and juveniles of Hemidactylus mabouia
from zoo Jardín Zoológico de La Habana “La Edad de
Oro”, Havana, Cuba. Means with different letters differed
signicantly (Tukey´s HSD post-hoc test, P < 0.05).
Feeding ecology of the Tropical House Gecko Hemidactylus mabouia 13
(Pianka, 1973) was used to determine diet similarity among adult
males, adult females and juveniles: Ojk = ∑PijPik / √(∑Pij
where Pij and Pik is the proportion of use of food item i for the
sex/age categories j and k, respectively.
The data were analysed using Statistica version 8.0 for Windows
and GrasphPad Instat version 3.01 for Windows. The Kolmogo-
rov-Smirnov test and Levene test were calculated to test if the
data were normally distributed and had homogeneous variances,
respectively. A Student-t test was performed to analyse differences
in SVL between adult males and females. A Linear Regression
analysis was used to test linear relationships between HW and HL
with SVL. Calculated residuals from these regressions were used
to compare among adult males, adult females and juveniles using
a one-way ANOVA. Tukey´s HSD test (post-hoc) was applied to
test the differences among particular sex/age categories. This ap-
proach minimizes the effect of body size and keeps the variation
of response variable (Quinn and Keough, 2002). Kruskal-Wallis
ANOVA by ranks for the mean number of consumed prey (the
data did not t a normal distribution) and a one-way ANOVA for
mean volume of ingested prey were implemented to test for dif-
ferences among the three sex/age categories. A Spearman Rank
Correlation was performed to assess the relationship between the
mean number of consumed prey with SVL and HW residuals. A
Linear Regression was computed to test for a linear relationship
between the volume of ingested prey with SVL and HW residuals.
A Chi-square test was used to examine the variation of volume
of ingested prey in diet among the three sex/age categories. The
Volume of ingested prey was pooled in three groups: less 49 mm3,
50-99 mm3 and more 100 mm3.
Among the captured geckos (n=61) during the
sampling period, 48 specimens (15 adult males, 21 adult
females and 12 juveniles) had stomach contents. The
remaining stomachs were empty, representing 21.3% of
the total sample.
No signicant difference was found between SVL
of adult males (51.56 ± 7.20 mm, range = 43.85-59.75
mm; n = 22) and adult females (54.47 ± 10.47 mm;
range = 40.3-60.82 mm; n = 25) (Student-t; t = 0.878;
P = 0.385,). Signicant linear relationships were found
between HW and SVL (R2 = 0.960, F1,59 = 1402.36, P
<0.0001) and HL with SVL (R2 = 0.941, F1,59 = 945.88,
P < 0.0001) (Fig. 2). The one-way ANOVA test revealed
signicant differences in HW residuals among sex/age
categories (F2,58 = 5.272, P = 0.0079, n = 61) where the
adult males showed the higher values (Fig. 3). However,
no differences were found in HL residuals (F2,58 = 2.386,
P = 0.101, n = 61).
The diet was mainly composed of arthropods that
comprised 10 orders of insects, one order of arachnids,
one order of crustaceans and one order of myriapods
(Table 1). In terms of number, the dominant prey in
the diet of H. mabouia were termites (54.55%) for
Table 1. Number (N), volume (in mm3) (V) and frequency of
prey (F) and index of relative importance (IRIi) in the diet
of adult males, adult females and juveniles of Hemidactylus
mabouia from zoo Jardín Zoológico de La Habana “La Edad
de Oro”, Havana, Cuba.
N (%) V (%) F (%) IRI
N (%) V (%) F (%) IRI
N (%) V (%) F (%) IRI
5 (11.36) 42.58 (6.12) 5 ( 21.74) 380.02 7 (14.89) 118.5 (4.93) 7 (18.42) 365.08 4 (13.79) 0.46 (0.08) 4 (16.67) 231.21
2 (4.55) -2 (8.70) -8 (17.02) 167.53 (6.96) 5 (13.16) 315.58 7 (24.14) 60.4 (11.07) 5 (20.83) 733.42
4 (9.09) 466.39 (67.03) 4 (17.39) 1323.73 7 (14.89) 1675.73 (69.65) 6 (15.79) 1334.87 3 (10.34) 386.28 (70.84) 3 (12.5) 1014.75
2 (4.55) 10.36 (1.49) 1 (4.35) 26.27 3 (6.38) 1.64 (0.07) 3 (7.89) 50.89 1 (3.45) -1 (4.17) -
----1 (2.13) -1 (2.63) -2 (6.90) -2 (8.33) -
1 (2.27) 0.80 (0.11) 1 (4.35) 10.35 --------
3 (6.82) 41.82 (6.01) 1 (4.35) 55.81 11 (20.4) 46.43 (1.93) 4 (10.53) 235.13 4 (13.79) 0.28 (0.05) 3 (12.5) 173.00
24 (54.55) 49.69 (7.14) 2 (8.70) 536.70 --------
----2 (4.26) 43.34 (1.80) 2 (5.26) 31.88 2 (6.90) 6.59 (1.21) 2 (8.33) 67.56
1 (2.27) 84.15 (12.09) 1 ( 4.35) 62.47 ----2 (6.90) 26.34 (4.83) 1 (4.17) 48,91
----2 (4.26) 321.95 (13.38) 2 (5.26) 92.79 1 (3.45) 64.94 (11.91) 1 (4.17) 64,05
----4 (8.51) 4.22 (0.18) 3 (7.89) 68.56 1 (3.45) -1 (4.17) -
----1 (2.13) 1.21 (0.05) 1 (2.63) 5.73 ----
----1 (2.13) 25.51 (1.06) 1 (2.63) 8,39 ----
uncountable -3 (13.04) -uncountable -3 (7.89) -----
uncountable -1 (4.35) ---------
2 (4.55) -2 (8.70) -----2 (6.90) -1 (4.17) -
44 (100) 695.79 (100) 46 (100) 2406.06 (100) 29 (100) 545.29 (100)
adult males; ants (20.4%), pill bugs (17.02%), spiders
(14.89%) and cockroaches (14.89%) for adult females
and pill bugs (24.14%) for juveniles. In terms of
volume, the order Blattodea (cockroaches) was the most
important food item, with more than 50% of ingested
volume in each sex/age category. The most frequent prey
items were spiders and cockroaches for adult males and
adult females, as were pill bugs and spiders for juveniles.
According to the Index of Relative Importance (IRIi), the
most important food item consumed was cockroaches
(more than 1000) for three sex/age categories. The pill
bugs were the second most important item in the diet
of juveniles (IRIi = 733.42). For adult males termites
(IRIi = 536) and spiders (IRIi = 380.02) were the second
most important; whereas for adult females were spiders,
pill bugs and ants (IRIi = 365.08, IRIi = 315.58 and IRIi
= 235.13, respectively). The rest of the prey had lower
relative importance indexes (Table 1).
Items unusually found were fragments of vegetal
material (in three stomachs) and shed gecko skins
(one stomach) of adult males. Three non-identied
arthropods were found in two stomachs of adult males
and one of a juvenile.
The mean number of food items consumed by adult
males was 1.92 ± 1.72, by adult females 2.24 ± 1.26
and juveniles 2.42 ± 1.16. There were no signicant
differences among them (Kruskal-Wallis ANOVA by
ranks H2,45 = 3.45; P = 0.178, n = 45). The highest number
of ingested prey was that of an adult male, with 21 food
items (all termites) in its stomach. The mean volume of
ingested prey of adult males ranged between 86.97 ±
146.52 mm3, that of adult females between 169.34 ±
325.17 mm3 and that of juveniles between 68.16 ± 81.11
mm3. The differences among these were not signicant
in mean volume (one-way ANOVA, F2,27 = 0.552; P =
0.582; n = 37). No signicant relationships were found
between the number of food items in each stomach and
body size of geckos (Spearman Rank Correlation, r=
0.009, P = 0.515, n = 47) neither between mean volume
of food items and SVL (Linear Regression, R2 = 0.0014,
F1,28 = 0.039, P = 0.846). Additionally, no signicant
relationships were recovered between number of food
items and HW residuals (Spearman Rank Correlation, r
= -0.2776, P = 0.064, n = 47) nor between mean volume
of ingested prey and HW residuals (Linear Regression,
R2 = 0.119, F1,28 = 3.796, P = 0.062).
Food niche breadth was similar between adult females
and juveniles, but higher in adult males. The trophic
niche overlap in terms of prey frequency was similar
among three sex/age categories, although between adult
female and juveniles this overlap was slightly higher.
The niche breadth and trophic niche overlap in terms
of prey volume were similar (Table 2). No signicant
differences recovered among the three groups (χ2 = 1.40;
Manuel Iturriaga & Rubén Marrero
Niche breadth Trophic niche overlap
Prey Prey Prey frenquency Prey volume
number volume Males Females Juveniles Males Females Juveniles
Males 0.29 0.18 - 0.81 0.85 - 0.95 0.96
Females 0.66 0.11 - 0.90 - 0.99
Juveniles 0.78 0.11 - -
Table 2. Standardized niche breadth and trophic niche overlap indexes based on proportions of prey number and volume, frequency
and volume prey, respectively for adult males, adult females and juveniles of Hemidactylus mabouia from zoo Jardín Zoológico
de La Habana „La Edad de Oro“, Havana, Cuba.
Volume groups (mm3) Adult Males Adult Females Juveniles
≥ 100 1
Total 10 16 11
Table 3. Volume groups of ingested prey by adult males, adult females and juveniles of Hemidactylus mabouia from zoo Jardín
Zoológico de La Habana “La Edad de Oro”, Havana, Cuba.
P > 0.05) regarding the volume of ingested prey (Table
3). Adult males, adult females and juveniles generally
preferred small sized prey and the volume of consumed
prey was similar among these classes during the dry
The diet of H. mabouia in the study area consisted
essentially of arthropods, coinciding with other
studies on the species (Zamprogno and Teixeira, 1998;
Boniglio, Balestrin and Cappellari, 2006; Rocha and
Anjos, 2007). In general, the data indicated that this
population is generalist and opportunistic. Cockroaches
represented the most important dietary item for all
of the three sex/age categories, followed by pill bugs
for juveniles, termites and spiders for adult males and
spiders, pill bugs and ants for adult females, which
are non-ying arthropods. Capula and Luiselli (1994)
comparatively found that diet of Tarentola mauritanica
and Hemidactylus turcicus in an urban area from Rome,
Italy was primarily composed of terrestrial prey. The
low contribution of ying arthropods in the diet of H.
mabouia, could be explained by the fact that all captures
took place in microhabitats lacking any articial lights.
This approach was performed in order to be able to
evaluate the availability of real prey and eliminate the
bias of consumption of ying prey that are not usually
present in the microhabitats of these geckos. Intraspecic
diet composition of a nocturnal lizard is known to
differ in terms of category of consumed prey between
populations that inhabit urban environments and those
living in natural ones (e.g., Bonglio, Balestrin and
Cappellari, 2006; Rocha and Anjos, 2007). In urban
environments, the ying groups (such as lepidopterans
and dipterans) have a higher number and frequency of
occurrence in the diet of geckos (e.g. Powell, Parmerlee
and Rice, 1990; Klawinski et al., 1994; Saenz, 1996),
because these are attracted by articial light while
sit-and-wait behaviour has been reported to be highly
successful for gecko species (Aowphol et al., 2006).
However, in natural environments the number and
frequency of non-winged groups (such as spiders,
orthopterans, ants, Lepidoptera larvae and termites) is
comparatively higher (e.g., Vitt and Zani, 1997; Vitt,
Zani and Monteiro de Barros, 1997; Zamprogno and
Teixeira, 1998; Colli et al., 2003). The high number of
termites predated by adult males of H. mabouia can be
considered as an unusual result, because the frequency
of this item in their diet composition was low. Probably,
the analysed adult males found a termite nest and they
proted from that opportunity. In arid areas of southern
Africa, termites show a patchy distribution (not only
temporal but also spatial) and are an irregular food
resource fortuitously consumed by Ptenopus garrulus
(Hibbitts et al., 2005). Fragments of vegetal material
and shed gecko skins occasionally found in stomachs of
adult males of H. mabouia were probably ingested by
accidental (the rst) or intentional (the latter).
Compared to previous studies, the proportion of empty
stomachs among the analysed individuals was relatively
high. While this result is in agreement Bonglio, Balestrin
and Cappellari (2006), it differs in respect to other
populations of H. mabouia (Zamprogno and Teixera,
1998; Rocha and Anjos, 2007) and populations of the
congeneric species H. turcicus (Capula and Luiselli,
1994; Saenz, 1996). According to Huey, Pianka and
Vitt (2001), the energetic balance of a particular lizard
population can be estimated by the rate of individuals
with empty stomachs. Nocturnal lizard species tend to
show higher proportions of empty stomachs among their
individuals (more than 20%), when compared to diurnal
species (less than 10%; Huey, Pianka and Vitt, 2001). The
high percentage of empty stomachs can be explained by
two reasons. First, this study was performed during the
dry season. Population densities of arthropods difference
signicantly between the dry season and wet season in
Cuba, where a reproductive explosion and increased
prey availability extraordinarily increases during the
latter (Núñez and Barro, 2003). Second, due to the fact
that most of the captures took place in the early hours of
the night, the geckos were emerging and did not have
enough foraging time to catch prey. Howard, Parmerlee
and Powell (2001) reported that the peak of foraging
activity occurred in late night hours, between 20:00 and
01:00 h, in a population of H. mabouia from Angilla
Island, Lesser Antilles.
Snout-vent length did not signicantly differ between
sexes, which is in agreement with studies of other
populations of the same species (Howard, Parmerlee
and Powell, 2001; Bonglio, Balestrin and Capellari,
2006; Rocha and Anjos, 2007) and populations of H.
turcicus (Klawinski et al., 1994; Saenz and Conner,
1996). A possible explanation for this observation is the
ecological advantage of a larger body size in females
(positive correlation with fecundity), which might
result in parallel increase of body size in both sexes.
Subsequently, both sexes reach their maximum size
by varying niche- and habitat constraints (Saenz and
Conner, 1996). Alternatively, Colli et al. (2003) stated
that a larger size of females in Gymnodactylus geckoides
Feeding ecology of the Tropical House Gecko Hemidactylus mabouia 15
amarali could be associated with increased clutch and
offspring size. The HW was shown to be being broader
in adult males. Similar results were recovered by Saenz
and Conner (1996) and Colli et al. (2003). Head-size
dimorphism is a common trait in squamate reptiles that
may be inuenced by ecological segregation and sexual
selection. The rst explains intersexual differences as
a mechanism to reduce competition for food resources
both at intra- and interspecic levels (Klawinski et al.,
1994; Gifford, Powell and Steiner, 2000; Dame and
Petren, 2006); whereas the second invokes intrasexual
competition among males for access to females (Colli et
al., 2003; Dame and Petren, 2006).
The high dietary composition overlap between adults
of H. mabouia can be explained by the high availability
of prey (cockroaches and spiders) in the study area,
while both adult sexes utilize similar microhabitats.
High availability of prey either in urban or natural
environments minimizes intraspecic competition of
food resources (e.g., Colli et al., 2003; Hibbitt et al.,
2005; Rocha and Anjos, 2007). The adult females also
showed dietary overlap with juveniles. Therefore, adult
females seem occupy the widest trophic niche breadth
of the three age/sex classes. Possibly, this is related to
guarantee energy for reproduction (Vitt and Caldwell,
2009). During the study period, seven adult females
captured had oviductal eggs and four other individuals
showed vitellogenic ovaries. The main difference in diet
observed among the three sex/age categories represented
the consumption of ground-dwelling prey (pill bugs) by
During this study, surface-dwelling juveniles were
commonly observed. Microhabitat selection therefore
seems to be a major factor in determining diet
composition. A similar use of microhabitat implicates
higher dietary overlap (Rocha and Anjos, 2007) but
different habitat associations can lead to consumption of
different prey items (Saenz, 1996; Howard, Parmerlee
and Powell, 2001). Possibly, the high frequency of
juveniles encountered on or near the ground is driven
by avoidance behaviour in relation to aggression by
adult males. Several examples have been described
concerning juvenile predation by geckos, due to which
juveniles tend to be in or near the ground to avoid
such behaviour by adults (e.g., Gifford, Powell and
Steiner, 2000; Howard, Parmerlee and Powell, 2001;
Bonglio, Balestrin and Cappellari, 2006). Summing
up, the diet of H. mabouia in an urban area of Havana
is composed of arthropods, mainly non-ying groups
(cockroaches, spiders and pill bugs). Adult females
show diet overlap with adult males and juveniles.
No signicant relationships were recovered between
body size and head size, in relation to the number and
volume of food items. There is a structural partitioning
between juveniles and adult males that determines the
consumption of ground-dwelling- prey by the former
and avoids predation by the latter.
Acknowledgements. We thank Javier Torres, Addison Díaz and
Javier Fernández Forrellat for their help during the eld work. We
thank Elier Fonseca from Facultad de Biología, Universidad de
La Habana, Cuba for identication of invertebrates in of stomach
contents. We thank Nayla García and Jans Morffe from Instituto
de Ecología y Sistemática, Havana, Cuba for by providing the
eyepiece micrometer and other microscope equipments. Senior
author thanks José Rances Caicedo-Portilla from Universidad
Nacional de Colombia, Bogotá D.C., Colombia for the useful
bibliographies offered so kindly and Vicente Berovides from
Facultad de Biología, Universidad de La Habana, Cuba for his
advices in statistical analysis. We thank Jans Morffe and Pedro
P. Herrera from the Instituto de Ecología y Sistemática, Havana,
Cuba; Ansel Fong from BIOECO, Santiago de Cuba, Cuba, José
Rances Caicedo-Portilla from Universidad Nacional de Colombia,
Bogotá D.C., Colombia and Van Wallach from Museum
of Comparative Zoology, Harvard University, Cambridge,
Massachusetts, USA; for their critical revision of previous drafts
of the manuscript.
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Accepted by Wouter Beukema
Feeding ecology of the Tropical House Gecko Hemidactylus mabouia 17