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Basic and Applied Herpetology 32 (2018) 57-70

Where to live in Lisbon: urban habitat used by the introduced


Italian wall lizard (Podarcis siculus)
Ricardo Ribeiro*, Paulo Sá-Sousa 

ICAAM – Instituto de Ciências Agrárias e Ambientais Mediterrânicas, Universidade de Évora, Pólo da Mitra,
Ap. 94, 7002-554 Évora, Portugal

*Correspondence: E-mail: ricardoavribeiro@gmail.com

Received: 29 March 2018; returned for review: 22 May 2018; accepted 11 September 2018.

Exotic animal invasions constitute a major threat to biodiversity worldwide. Our assessment deter-
mined the core range occupied by the Italian wall lizard (Po d arcis siculus) in the urban area of
Parque das Nações (Lisbon, Portugal), where it was accidentally introduced two decades ago. Des-
pite the apparent current scenario of non-expansion, the alien species interferes with the local dis-
tribution pattern of the native P. virescens, with both lizard species shown to use the available mi-
crohabitats differently. The native P. virescens population displays an overall heterogeneous dis-
tribution in the urban matrix. On the other hand, the occupancy of the exotic species clusters with-
in the original area of introduction (garden in front of the Lisbon Oceanarium), as well as in adja-
cent gardens. There, P. siculus replaced the native P. virescens as the only lizard species present.
Appa-rently, there is no known reason there that prevents the alien lizard from colonizing more
available geographic area and expanding. 

Key words: Invasive species; Po d arcis siculus; Podarcis virescens; spatial segregation.

Invasive species currently represent logical tolerance and plasticity, allied with
one of the major problems in nature con- high propagation capacity (Nevo et al.,
servation (Schulte, 2012). The damage 1972; Capula Ceccarelli, 2003; Crnobr-
caused by invasive reptiles mainly comes nja-Isailovic et al., 2009; Silva-Rocha et
from disruptions and changes in native al., 2014). Podarcis siculus is distributed conti
food webs and ecosystem functions, with -nuously throughout the Italian Peninsula
impacts ranging from predation on sensi- and East Adriatic coast, as well as for big
tive species, intoxication of predators, islands - Sardinia (where recent studies
competition and hybridization with native suggested that it may be allochtonous ins-
species, vectorization of new parasites, as tead of native, Senczuk et al., 2017) and
well as direct and indirect impacts on hu- Sicily - and a multitude of adjacent islets
mans (Kraus, 2009; Fujisaki et al., 2010). (Corti Lo Cascio, 2002). Unlike the re-
Among the European reptile species, the ported pet trade introductions in the USA
Italian wall lizard (Po d arcis siculus, Sau- (Kolbe et al., 2013), it might have been in-
ria: Lacertidae) is considered as both an troduced by ship trade into Corsica, Bale-
opportunistic species and a successful ex- arics and several port areas around the
otic colonizer, characterized by wide eco- Mediterranean Basin e.g. SW France,

DOI: https://dx.doi.org/10.11160/bah.101
RIBEIRO & SÁ-SOUSA

Greece, Turkey, Tunisia or Lybia affected by this P. siculus introduction, al-


(Carretero Silva-Rocha, 2015). though negative impacts are still unknown
Towards Lisbon, some P. siculus indi- (such as expansion, hybridization or extin-
viduals might have been transported ction of native species). Both species under
along with rough materials and ornamen- study coexist in sympatry with other lacer-
tal plants (structures in which they would tids in their native distributions, evidencing
most likely be sheltered in or basking on), ecological segregation (Capula, 1993; Capu-
from their place of origin, during the la et al., 1993; Carretero, 2008; ValdeÓn et
Expo98 event (González de la Vega et al., al., 2010; Grano et al., 2011; Carretero
2001; Silva-Rocha et al., 2012). This appears Silva-Rocha, 2015; Carretero Salvador,
to be a recurrent issue, as similar cases 2016).
were reported by Rivera et al. (2011) and However, the introduction of the Italian
ValdeÓn et al. (2010), after the observation of wall lizard in other areas has been shown to
several P. siculus individuals in olive trees, be detrimental to autochthonous lacertids,
imported from Calabria (Italy) to Catalo- especially in micro-insular environments,
nia (Spain). with evidence of competitive spatial coex-
In urban environments, which are istence or exclusion (Nevo et al., 1972; Cap-
typically characterized by spatially and ula, 2002; Capula Ceccarelli, 2003; Pod-
temporally heterogeneous thermal mos- nar et al., 2005; ValdeÓn et al., 2010; Mateo
aics, ecological interactions with poten- et al., 2011; Carretero Silva-Rocha,
tial competitors and predators may also 2015).
influence patterns of occurrence and abun- Based on previous personal reports
dance of individuals of a given species from Paulo Sá-Sousa and bibliography
(Kolbe et al., 2016). In those environments, it from Loureiro et al. (2008), this study’s main
has been observed that sympatric lacertids objectives consisted in determining the cur-
(namely Pod arcis muralis and P. siculus) rent range area occupied by the introduced
are spatially organized according to each P. siculus and assessing the degree of spatial
species specific ecological needs, and there- segregation exerted by the exotic popula-
fore not limited to interspecific interactions tion upon the autochthonous species. This
(Capula et al., 1993). Lizard species often tend required the knowledge of the current spa-
to be associated with specific microhabitat tial distribution of both species in Lisbon, as
patches, with minimized interspecific com- well as each species microhabitat prefer-
petition through their spatial niche distri- ences, according to the available space. 
bution (Smith Ballinger, 2001; Rugiero
Luiselli, 2007; Vitt Caldwell, 2014).
Materials and methods
However, there may be cases where the The study area covered the location
microhabitat occupied by a certain species where P. siculus was first introduced
is restricted by the presence of another and consequently reproduced (González
(Arnold, 1987). That could be the case with de la Vega et al., 2001), within the so-called
autochthonous Podarcis virescens (Geniez Parque das Nações, a leisure and residential
et al., 2014; Dias et al., 2016), which might be area of Lisbon (38°45ʹ43.7ʺN 9°05ʹ39.0ʺW,

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HABITAT OCCUPANCY BY PODARCIS SICULUS IN LISBON

Figure 1: Total prospected area limited in red; Pod arcis siculus introduction spot highlighted by a
black square.

Fig. 1). For lizard sampling one large poly- were favourable weather conditions for
gon was delimited to address the possible lizard thermoregulation). However, some
expansion and establishment of new po- sectors took longer to cover in cases of
pulations towards adjacent spots. The high occurrence of observations. All lizard
sampling area was subsequently divided observations (each corresponding to one
in 16 plots or sectors with approximate individual) consisted of visual encounters
dimensions, to facilitate the visual encoun- (Lambert, 1984; Eekhout, 2010), where the
ter of lizards. Although these plots consis- following data was recorded: GPS point of
ted of urban areas, there are contrasting location, species (P. siculus or P. vi-
differences between them, ranging from rescens), behaviour activity, perch type, height
those that show predominance of artificial and slope, distance to vegetation cover,
elements, to those that hold some garden refuge type and distance to refuge, along
spots with lawn or a more rustic appear- with the substrate texture, number of
ance, usually consisting of an eclectic se- trees, shrubs, non-woody plants and dead
lection of plant species from various re- branches/trunks, found within a four me-
gions of the world (Pedrosa, 2013). ters radius surrounding each observation
Due to statistical and data indepen- point (the dissertation from which this
dence reasons, the sectors were sampled article was adapted covered a wider range
following a random order. Each sector was of variables, including sex and size class,
surveyed by walking at a slow, steady which were considered negligible for the
pace, during one sunny day (when there study´s purpose, primarily focused on

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RIBEIRO & SÁ-SOUSA

Figure 2: Spatial distribution


of both species (A - total
area; B - focus in the pre-
sence of Po d arcis siculus).
Red symbols – introduced
Podarcis siculus; Green sym-
bols - autochthonous Po -
darcis virescens.

habitat use and microspatial distribution, 0.05) (Strode Brokaw, 2015). The chi-
and therefore discarded from this paper). squared test was used for comparison be-
The first round of sampling covered all (16) tween species, regarding the data collected
sectors, providing a general acquaintance of on the constituent elements of the micro-
both the study area and its lizard occupan- habitats used by both species, where the
cy. Based on this previous data assessment, formulated null hypothesis was that “there
the second and third rounds covered only 8 are no differences between the use given by
of the 16 sectors, excluding those where P. the invasive and autochthonous species to
siculus was clearly absent in the first round. the elements present in the microhabitat”.
Thus, the first round took place between For this test’s application we used raw data,
April 25th, 2016 and July 3rd, 2016; the sec- that is, simple counts (observations), ins-
ond began on September 8th, 2016 and en- tead of percentages or proportions (Strode
ded on September 21st, 2016; the third lasted Brokaw, 2015), presented in table, which
from September 24th, 2016 to October 11th, included the observed and expected values
2016. for several categories, within each con-
Further statistical analyses were per- sidered parameter (Dytham , 2011). Since
formed using the SigmaPlot® 13.0, Statisti- multiple tests were performed simultane-
ca 13® and Microsoft Excel 2016 softwares, ously, further statistical corrections were
tested for a confidence level of 95% (α = made using the Holm-Bonferroni method,

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HABITAT OCCUPANCY BY PODARCIS SICULUS IN LISBON

Table  1: Sampled sector maximums for both The native P. virescens population
species in each sector. The three sectors with shows an overall heterogeneous distribu-
occurrence of both species highlighted in grey. tion in the urban matrix. On the other hand,
the occupancy of the invasive species only
Sector  Max. P. virescens  Max. P. siculus  clusters within the original area of intro-
duction (garden in front of the Lisbon
1  26  1 
Oceanarium), as well as in adjacent gardens
2  9  26  (Fig. 2a). A single individual of P. siculus
3  22  96  was also o-bserved in a previously
4  41  0  unknown location (a garden spot placed
5  13  0  some hundreds of meters away from the
6  7  0  main area), where P. virescens often occurs
7  4  0  (Fig. 2b).
8  8  0 
9  27  0  Degree of spatial segregation between
10  24  0  Podarcis siculus and Podarcis virescens
11  2  0  The Chi-squared tests of independence
12  10  0  support the main differences found in the
13/14  33  0  spatial variables tested between P. siculus
15  9  0  and P. virescens, as explained in the follo-
16  16  0  wing sections concerning habitat, through
Sub‐total  251  123  the analysis of the graphics and tables (P <
0.001 for nearly all values, Table 2), regar-
ding the behaviour shown by the lizards
during observation, the elements present in
narrowing the possibility of generating a the microhabitat and its use by both spe-
statistically significant result (type I error) cies. The only exception was the parameter
(Holm, 1979). To enlighten these intricate “Distance to Refuge”, which was similar for
multiple results, a Multiple Correspon- both species (P = 0.223, Table 2). The Holm-
dence Analysis test was performed compre- Bonferroni correction method confirmed
hending all parameters, to visualize the the obtained results, whereas the previous-
pattern of relationships between the micro- ly mentioned parameter remained as the
habitat features and their usage by both only to not reject the null hypothesis.
species (Abdi Valentin, 2007). The Multiple Correspondence Analysis
Results test plots for both species (Fig. 6) showed
that, despite overlapping for the most part,
Lizard occupancy
P. siculus only occupies part of the spatial
A total of 699 observations were regis- niche of P. virescens at the multivariate le-
tered, 476 (68%) belonging to the native P. vel, which could provide some answers
virescens and 223 (32%) to the introduced P. regarding the possible event of future ex-
siculus, present in only three sectors (Table 1). pansion and its impacts on P. virescens.

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RIBEIRO & SÁ-SOUSA

Table 2: Results of the chi-Squared statistical tests, used for comparison between species. Signifi-
cant P-values highlighted in bold. Adjusted Holm-Bonferroni P-values on the right.

Holm‐Bonferroni  P‐
Tested Parameters  χ2  df  P‐value 
values 
Behaviour  19.401  3  P < 0.001  P = 0.01 

Perch type  81.089  3  P < 0.001  P = 0.005 


Perch height  51.731  3  P < 0.001  P = 0.03 
Perch slope  22.236  2  P < 0.001  P = 0.01 

Refuge type  98.363  6  P < 0.001  P = 0.01 

Distance to refuge  4.379  3  P = 0.223  P = 0.01 

Number of Trees  34.2  1  P < 0.001  P = 0.02 

Number of Shrubs  43.719  2  P < 0.001  P = 0.01 


Number of non-woody plants  12.945  2  P = 0.002  P = 0.05 

Number of dead branches/trunks  33.585  3  P < 0.001  P = 0.004 

Substrate texture  62.558  2  P < 0.001  P = 0.01 


Distance to vegetation cover  21.584  2  P < 0.001  P = 0.01 

Among the variables which contributed but were also present in areas with 1 to 5
the most for the relative inertia values, (23% and 39%) shrubs. Both species used
were the Refuge and Perch Type Rock, as areas with a moderate presence of dead
well as Vertical Perch Slope, and Artificial wooden matter (up to 30 branches/trunks/
Perch Type and Substrate Texture. etc.), counting for 71% of P. siculus and
56% of P. virescens observations (Fig. 3d).
Habitat use
Substrate texture (Fig. 3e) enhances P. sicu-
Both species seem to occur in larger lus as having occupation for earthy open spaces
numbers (84% of the P. siculus total and (79%), while P. virescens appears to be a
97% of P. virescens) in areas with few or more generalist species, though often
no trees (0 to 2) (Fig. 3a), with the autoch- found in earthy substrate (50%), dividing
thonous species being less dependent on the remaining half of observations almost
the existence of arboreal and non-woody equally between artificial and mixed
plants cover on site (Fig. 3c - the congeners grounds.
showed a marked preference for areas Micro‐habitat features
without non-woody plants, 73% of P. sicu- The most used perch for both lacertids
lus and 80% P. virescens). Similarly, 58% of (Fig. 4a) appears to be the ground (more
P. siculus and 56% of P. virescens individuals evident for P. siculus, 81%, than for P. vi-
appeared in spots without shrubs (Fig. 3b) rescens, 53%), whereas P. virescens also choos-

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HABITAT OCCUPANCY BY PODARCIS SICULUS IN LISBON

Figure  4: Results concerning each species use


of the available micro-habitat: Perch Type (A),
Figure  3: Results concerning micro-habitat Height (B), Slope (C) and Distance to Vegeta-
elements: number of Trees (A), Shrubs (B), tion Cover (D).
Non-woody Plants (C), dead Branches/Trunks
(D) and Substrate Texture (E).

es perches located in artificial stru-ctures (90% for P. siculus and 75% for P. vi-
(34%). The rocky perches were the least rescens) of vegetation cover (herbaceous,
chosen, by both species. Concerning to the shrubs, trees – Fig. 4d) near to the perch
perch height, soil level was the most ob- (up to 0.5 m) was detected.
served (Fig. 4b), with P. siculus showing The behaviour activity of lizards (Fig.
ground dwelling attributes (82%), similar 5a) shows that both species were often
to slightly more than half of the P. vi- seen basking (57% for P. siculus and 64%
rescens individuals. Of the latter, 21% are still for P. virescens), but also active (18 to
distributed at a reduced height (from 0 to 20%), suggesting a similar daily tasks
0.25 m), with only 9% of the individuals pattern. When disturbed, the lizards often
found at heights above 0.75 m. The slope seek the available vegetation cover as ref-
assumed by each lizard at perch site (Fig. uge (Fig. 5b), which was evident (50%) for
4c) was mostly horizontal for both species P. siculus, while the native lizard chose be-
(84% of P. siculus and 70% of P. virescens). tween ve-getation cover and artificial
The “Vertical” posture was the second structures (26% and 32%, respectively), the
most observed in P. virescens and the least latter one which the exotic species appar-
observed in P. siculus. High dependency ently tends to avoid. Approximately 22%

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RIBEIRO & SÁ-SOUSA

Figure 5: Results concerning each spe-


cies use of the available micro-habitat:
Behaviour (A), Refuge Type (B) and
Distance to Refuge (C).

of the observations for both species did servations. There, this species individuals
not seek refuge, remaining in the perch show the behaviour and ecological
without mo-ving, or escaping without patterns typical of their own species
seeking shelter in the nearby elements. (Corti Lo Cascio, 2002). However, the
About 40% of P. siculus and 47% of P. vi- obtained data suggests that local spatial
rescens prefer to carry out their daily tasks segregation was most probably due to
near (up to 0.25 m) to a possible refuge competitive exclusion, as P. siculus re-
(Fig. 5c), with the number of observations placed P. virescens as the only lizard spe-
decreasing as the distance to refuge in- cies present at the original introduction
creases.  site. In fact, P. siculus has a larger body
size than P. virescens, which is probably
Discussion
an advantage in competitive encounters
Although there were found some si- since, according to Downes Bauwens
gnificant differences in the use of several (2002), larger species tend to be dominant
categories of habitat tested between both over smaller ones. This appears to have
lizard species, that is not the major evi- favoured the exotic wall lizard to the detri-
dence. Indeed, all observations in Parque ment of the native species, since in Lisbon
das Nações (Lisbon) point to a scenario of the species are shown to be sympatric but
absence of expansion by the alien species never syntopic.
P. siculus, as previously suggested by Lourei- Sympatric species, when in direct com-
ro et al. (2008) and Sá-Sousaʹs personal ob- petition, often spatially separate them-

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HABITAT OCCUPANCY BY PODARCIS SICULUS IN LISBON

Figure  6: Multiple Corre-


spondence Analysis Scatter-
plot map, comprehending
all surveyed parameters for
both species. Black symbols
represent introduced P.
siculus; purple, open symbols
represent autochthonous P.
virescens.

selves due to differences in the structural artificial substrates. It shows, to some ex-
niche (e.g. diameter and height of the cho- tent, dependence on the existence of some
sen perch) and, in some cases, in the mi- type of vegetation cover (herbaceous,
croclimatic environment (eg. open vs. shrubs, trees) relatively close to the perch,
shaded areas) or by displacement of indi- being mainly found at soil level. Despite
viduals through both competition and anti its observable proximity to vegetation co-
-predatorial/survival behaviour (Downes ver, the microhabitats used by P. siculus
Bauwens, 2002; Žagar et al., 2015). Syntop- tend to be poor in tree and shrub cover,
ic lizard populations were shown to alter with most of the individuals observed also
their predation risk by using specific habi- showing preference for areas with little to
tats based on the availability of refu-ges none dead wooden matter on the ground.
that provide protection against shared It was also observed, in this species, ten-
predators, which act as competition medi- dency to avoid artificial structures as pos-
ators, indirectly inducing asymmetrical sible shelter, preferring the available vege-
competition and shifts in segregation tation cover for this purpose - and thus
patterns (Smith Ballinger, 2001; Žagar perhaps limiting its distribution in this
et al., 2015). The results in Lisbon suggest that predominantly urban area.
both species use the available microhabi- It is tricky to explain the present lack of
tats differently. expansion of P. siculus in Lisbon, especial-
The exotic species P. siculus preferen- ly since this species often spreads into ur-
tially uses microhabitats having a mostly ban areas in native Italy, where it is sym-
earthy substrate texture, avoiding fully patric with P. m uralis: P. siculus prefers the

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RIBEIRO & SÁ-SOUSA

driest places, while P. muralis the moisten little or no shelter to these P. siculus indi-
ones (Corti Lo Cascio, 2002). Perhaps it viduals, they might attempt to expand. If
is still a question of a short time lapse (two landscape work is carried out in the adja-
decades) after the presumed date of intro- cent areas and gardens, it may possibly
duction (not yet the sufficient time and promote the dispersal of P. siculus, by al-
effort required to efficiently use the newly tering their previously unfavourable con-
occupied habitat - Stamps, 1995), the possi- ditions and resources, since dispersal be-
bility of the exotic species larger body size haviour seems to be in some aspects influ-
being a disadvantage at this particular in- enced by external factors, such as intraspe-
troduction site (making this lizard more cific competition and habitat quality
prone to predation or less capable of dea- (Vignoli et al., 2012). This brings us to the
ling with the introduction site’s condi- single individual found off the core area of
tions), and/or lack of a reproductive effe- introduction, which may occur most likely
ctive high enough to promote recruitment due to the translocation of plants or trees
of young lizards towards new adjacent between gardens of Parque das Nações, an
spots (more individuals = more potential action witnessed during the sampling
dispersers - Vignoli et al., 2012). rounds. Most of these movements are usu-
This diminished reproductive effective ally carried out in the winter, due to the
might also be due to high predation pres- reduced plant activity, which facilitates
sure, as several individuals of Tarento la the success of the transplant, but also fa-
mauritanica were observed in the P. siculus vours exotic reptile species introductions,
place of occurrence and nearby sites, a often hibernating inside available cavities
species which may consume individuals of in trees or plants, so that these movements
the latter and other species of lacertids, do not cause the individual to escape from
both immature and adult (Pellitteri-Rosa the refuge (ValdeÓn et al., 2010). Despite
et al., 2015). This population growth control the contribution of such activities and reg-
may be carried out by P. siculus adult in- ular maintenance work to introductions,
dividuals as well, in cases of cannibalism they can also rake the soil and diminish
(such as those previously described by the available vegetation cover, destroy
Ouboter, 1981, Burke Mercurio, 2002, lizard eggs buried in the ground or sand,
Cattaneo, 2005, Capula Aloise, 2011 reduce the availability of perches for ther-
and Grano et al., 2011), or by cats, dogs or moregulation and disturb the existing mi-
seagulls present in the area (Burke Ner, croclimate at the place of introduction –
2005). Other lacertid found inside the yet again providing a plausible argument
study area limits was Psam m o d ro m us al- for the absence of expansion. This cluste-
girus, which could spark interactions with yet ring could be due to high predation pres-
unknown consequences in case of future sure, short time span since introduction,
contact with the exotic population. competition with autochthonous species,
Although the introduction area in lack of available favourable resources and/
Parque das Nações is surrounded by artifi- or microhabitat degradation – or synergy
cial structures that apparently provide of all the previous factors. If expansion is

66
HABITAT OCCUPANCY BY PODARCIS SICULUS IN LISBON

to occur, data suggests that it possibly to Leonor Melo for the additional review
would not exclude the autochthonous spe- of the article. Miguel A. Carretero and one
cies entirely, since P. siculus seems to not anonymous reviewer provided construc-
occupy all the available habitats in the tive comments and suggestions on the
Parque das Nações area, with a reduced manuscript. 
usage of the more artificial and/or rocky
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