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Map of the introduced populations of Podarcis sicula analysed. (A) populations in Iberian Peninsula, United Kingdom and Switzerland , (B) populations in United States, (C) populations in Greece and Turkey.
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Biological invasions are a challenge to conservation and constitute a threat to biodiversity worldwide. The Italian wall lizard Podarcis sicula has been widely introduced, and seems capable of adapting to most of the regions where it is established and to impact on native biota. Here we construct a phylogenetic framework to assess the origin of the...
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Citations
... Introduction of P. siculus in different areas was indicated to be disturbing to autochthonous lacertids, particularly in micro-insular habitats (Nevo et al., 1972;Capula, 2002;Capula and Ceccarelli, 2003;Podnar et al., 2005;Valdeón et al., 2010;Mateo et al., 2011;Carretero and Silva-Rocha, 2015;Ribeiro and Sá-Sousa, 2018). Podarcis siculus is thought as both an opportunistic species and an accomplished exotic colonizer, having high ecological tolerance, associated with superior distribution ability (Nevo et al., 1972;Capula and Ceccarelli, 2003;Isailovic et al., 2009;Silva-Rocha et al., 2014;Ribeiro and Sá-Sousa, 2018). ...
The present study provides a new locality record (Hıdırlık Street in Safranbolu district of Karabük province) of Podarcis siculus from the western Black Sea region of Türkiye. Our findings show that the distribution of this species in Türkiye may cover more areas than known. It is possible that the species, which has not been reported to be distributed anywhere between Bolu and Samsun provinces so far, is also likely to be found in the areas between Karabük and Samsun provinces. Pholidolial and morphometric characteristics of the Safranbolu specimens were compared to those of the specimens reported in the literature from other parts of Türkiye. It was concluded that the samples from the Hıdırlık population were similar to the P. siculus samples reported in the literature.
... Lizards of the genus Podarcis have been introduced and successfully established in various locations around the globe (Hedeen 1984;Podnar et al. 2005;Heym et al. 2013;Silva-Rocha et al. 2014;Michaelides et al. 2015;Ribeiro & Sá-Sousa 2018). In addition to these repeated invasions, almost all species in the genus are color polymorphic (Brock et al. 2022a), making Podarcis an ideal study system to understand the role of discrete color polymorphisms in the context of invasion. ...
Many species exhibit color polymorphisms which have distinct physiological and behavioral characteristics. However, the consistency of morph trait covariation patterns across species, time, and ecological contexts remains unclear. This trait covariation is especially relevant in the context of invasion biology and urban adaptation. Specifically, physiological traits pertaining to energy maintenance are crucial to fitness, given their immediate ties to individual reproduction, growth, and population establishment. We investigated the physiological traits of Podarcis muralis , a versatile color polymorphic species that thrives in urban environments (including invasive populations in Ohio, USA). We measured five physiological traits (plasma corticosterone and triglycerides, hematocrit, body condition, and field body temperature), which compose an integrated multivariate phenotype. We then tested variation among co‐occurring color morphs in the context of establishment in an urban environment. We found that the traits describing physiological status and strategy shifted across the active season in a morph‐dependent manner—the white and yellow morphs exhibited clearly different multivariate physiological phenotypes, characterized primarily by differences in plasma corticosterone. This suggests that morphs have different strategies in physiological regulation, the flexibility of which is crucial to urban adaptation. The white‐yellow morph exhibited an intermediate phenotype, suggesting an intermediary energy maintenance strategy. Orange morphs also exhibited distinct phenotypes, but the low prevalence of this morph in our study populations precludes clear interpretation. Our work provides insight into how differences among stable polymorphisms exist across axes of the phenotype and how this variation may aid in establishment within novel environments.
... Accumulating evidence is strongly identifying the "plant nursery trade" (commerce in live plants for ornamental purposes) as one of the main forms of reptile introduction into new territories 4 , since these animals often use plants and trees for refuge and thermoregulation e.g., 72,73 . This kind of human-mediated trade in the Mediterranean has been responsible, for instance, for the introduction of the Italian wall lizard, Podarcis siculus e.g., 8,74 , the brahminy blind snake, Indotyphlops braminus 75 , and the colubrid snakes Hemorrhois hippocrepis, Malpolon monspessulanus, and Zamenis scalaris 53 , outside their native geographic ranges. Curiously, many of these introductions result from the trade of old olive trees transported from Italy [76][77][78] . ...
In this study, we aimed to understand the contemporary and ancient colonization routes of the Moorish gecko, Tarentola mauritanica, using simple sequence repeats. By analyzing the genetic diversity of populations in different regions, we found that Morocco is the genetic diversity hotspot for the species, followed by the Iberian Peninsula. However, historical gene flow estimates identified the Iberian Peninsula, not Morocco, as the primary contributor of colonizing individuals, along with continental Italy to a lesser extent. Currently, mainland Italy is the main source of introduced individuals, likely due to the plant nursery trade. The study suggests that human-facilitated introductions from various geographical origins, with numerous regions colonized through continental Italy during two distinct periods, are responsible for the recurrent entry of individuals belonging to the European lineage of T. mauritanica into the Mediterranean and Macaronesia. These findings can inform better monitoring surveys and conservation programs by identifying putative current colonization routes of alien species.
... These previous findings coincide with the present work, as it has been observed that the moorish gecko population inhabiting the island of Mallorca is able to recognize and respond to the novel predator H. hippocrepis only after sixteen years of its introduction. Rapid responses to changes in the environment of some lizard species are thought to be related to a great adaptability and capacity of colonizing new territories (Vervust et al., 2007;Silva-Rocha et al., 2014;Mencía et al., 2017), which could be applied to T. mauritanica, since its wide distribution would suggest a considerable capacity to inhabit new regions. ...
Stressful situations induce an increase in the production of reactive oxygen species (ROS) which can lead to molecular damage and alteration of cell function. The introduction of new potential predators induces physiological stress in native fauna. However, behavioral responses have been reported in preys, demonstrating an induction of the defenses against alien species. Behavioral and antioxidant enzyme responses in the moorish gecko, Tarentola mauritanica, against the invasive predator horseshoe whip snake (Hemorrhois hippocrepis) were assessed. Behavior was recorded and a tissue sample from the tail was collected after placing the gecko in a terrarium with previous absence or presence of the snake in ‘Control’ and ‘H. hippocrepis’ groups, respectively. Fifteen behavioral variables were examined, including tongue flick (TF) and locomotion patterns. Antioxidant enzyme activities -catalase (CAT), superoxide dismutase (SOD), glutathione peroxidase (GPx), glutathione reductase (GR)-, and the levels of reduced (GSH) and oxidized glutathione (GSSG), glutathione/glutathione disulfide ratio (GSH/GSSG) and malondialdehyde (MDA) concentrations were measured in the tissue sampled. Geckos exposed to the snake's odor showed a higher number of TF, longer amounts of time remaining motionless or moving in slow motion and they spent less time on the ground in comparison to the ‘Control’ group. The presence of the snake produced a significant increase in the activities of CAT, SOD and GR and a decrease in the GSH/GSSG ratio in T. mauritanica individuals exposed to the snake's scent. Thus, both behavioral responses and oxidative stress biomarkers clearly showed that T. mauritanica is able to recognize H. hippocrepis as a potential predator, despite being a recently introduced snake at the Balearic Islands.
... A total of 277 sequences from the Italian Peninsula, Corsica and Sardinia (Senczuk et al., 2017), accession numbers: KY064841-KY065117 were downloaded. Additionally, 41 published sequences from other introduced populations in Eurasia: 33 sequences from the Iberian Peninsula and Menorca (Silva- Rocha et al., 2012;Garin-Barrio et al., 2020), accession numbers: JX072938-JX072960, MW192534-MW192543; seven sequences from Turkey, Greece, and United Kingdom (Silva- Rocha et al., 2014), accession numbers: KP036396-KP036402. The samples from Switzerland (Schulte and Gebhart, 2011) were not included in the analysis because they were not available in GenBank but the locations were added to the map according to Silva-Rocha et al. (2014). ...
... Additionally, 41 published sequences from other introduced populations in Eurasia: 33 sequences from the Iberian Peninsula and Menorca (Silva- Rocha et al., 2012;Garin-Barrio et al., 2020), accession numbers: JX072938-JX072960, MW192534-MW192543; seven sequences from Turkey, Greece, and United Kingdom (Silva- Rocha et al., 2014), accession numbers: KP036396-KP036402. The samples from Switzerland (Schulte and Gebhart, 2011) were not included in the analysis because they were not available in GenBank but the locations were added to the map according to Silva-Rocha et al. (2014). One sequence from Podarcis melisellensis from GenBank was used as an outgroup (accession number AY185057), following Silva- Rocha et al., (2014). ...
... The samples from Switzerland (Schulte and Gebhart, 2011) were not included in the analysis because they were not available in GenBank but the locations were added to the map according to Silva-Rocha et al. (2014). One sequence from Podarcis melisellensis from GenBank was used as an outgroup (accession number AY185057), following Silva- Rocha et al., (2014). Sequences were edited using Geneious Prime v.2020.1 (https://www.geneious.com). ...
In recent decades, many reptile species have been introduced outside their native ranges, either accidentally through the transportation of goods and materials (e.g., plants, construction materials), but also intentionally through the pet trade. As a paradigmatic example, the Italian wall lizard, Podarcis siculus, native to the Italian Peninsula, Sicily and the north Adriatic coast, has been introduced in several nearby islands since historical times (Corsica, Sardinia, Menorca). Besides these regions, scattered populations were later reported from the Iberian Peninsula, France, Swit-zerland, Turkey, Greece, the United Kingdom and North America. Here, we provide molecular evidence regarding the introduction and origin of P. siculus in six new populations outside its native range: Romania (Bucharest and Alba Iulia), inland Croatia (Zagreb and Karlovac), Italy (Lampedusa Island) and Azerbaijan (Baku). Phylogenetic analysis suggests that the Alba Iulia (Romania) population originated from a single clade (Tuscany), while the population from Azerbaijan is admixed including two distinct clades, one similar to those found in Sicily and the other present across the Tuscany clade. Samples from Bucharest also have admixed origins in Tuscany and the Adriatic clades. Less surprisingly, samples from Zagreb and Karlovac are included in the Adriatic clade while those from Lampedusa originated from Sicily. Overall, our results further demonstrate that P. siculus is able to establish outside of its native range even under different climatic conditions, not particularly from specific clades or source areas. Also, for the first time in this species, our results indicate that repeated human introductions promote lineage admixture and enhance their invasive potential.
... These previous findings coincide with the present work, as it has been observed that the moorish gecko population inhabiting the island of Mallorca is able to recognize and respond to the novel predator H. hippocrepis only after sixteen years of its introduction. Rapid responses to changes in the environment of some lizard species are thought to be related to a great adaptability and capacity of colonizing new territories (Vervust et al., 2007;Silva-Rocha et al., 2014;Mencía et al., 2017), which could be applied to T. mauritanica, since its wide distribution would suggest a considerable capacity to inhabit new regions. ...
... siculus) successfully imitate conspecifics and members of a local congeneric species. This ability might be linked to the species' opportunistic nature (Capula and Aloise 2011;Mačát et al. 2015), ecological flexibility (Vervust et al. 2010;Kapsalas et al. 2016;Wehrle et al. 2020) and invasive success (Silva-Rocha et al. 2014). 'Opportunistically' foraging species are more likely to exhibit social learning (Klopfer 1959(Klopfer , 1961. ...
In many animals, decision-making is influenced by social learning, i.e. the acquisition of insights through the observation of other individuals’ behaviours. In cases where such socially obtained information conflicts with personally acquired knowledge, animals must weigh up one form of information against the other. Previous studies have found that individual animals differ consistently in how they trade-off socially versus personally acquired knowledge, but why this is so remains poorly understood. Here, we investigate whether an animal’s cognitive profile affects its decision to use either prior personal or new, conflicting social information, using the Italian wall lizard (Podarcis siculus) as our model species. We trained lizards to associate one of two colour cues with food, and subsequently allowed them to observe a conspecific trained for the opposite colour. After social demonstrations, lizards overall tended to use the ‘fake’, non-rewarding social information, but some individuals were more likely to do so than others. Lizards that showed faster spatial learning were more likely to copy social information even in the presence of reliable previous knowledge. Our study highlights the existence of significant inter-individual variation in social learning in a lizard, possibly mirroring variation in cognitive abilities.
Significance statement
Animals often use social information in daily decision-making. Whenever knowledge obtained through personal experience conflicts with observations of decisions made by conspecifics, individuals must weigh these two types of information against each other. Individuals tend to differ in whether they prioritize social versus private information, but why this is so, remains poorly understood. Using lizards, we tested whether an animal’s cognitive profile affects its decision to use either prior personal or new, conflicting social information. We found considerable variation among individuals in social information use and tendency to rely on personal or public knowledge. Specifically, faster spatial learners inclined to copy social information, even in the presence of reliable previous knowledge, suggesting that intrinsic, cognitive aspects influence how individuals trade-off public and private information.
... An introduced population of P. siculus was discovered in Athens (Greece) in March 2014 at the southern coastline of the city, likely corresponding to recent introduction (Adamopoulou 2015). Molecular analyses revealed that the new population originated most probably from the Adriatic region (Silva-Rocha et al. 2014). At the time of the first discovery, the population comprised around 60 adults and plenty of subadults (Adamopoulou 2015) that quickly increased, thus threatening the several Greek endemic lacertid lizards which might come in contact with the invader in the future (Kapsalas et al. 2016). ...
... It is important to mention that the site where the species occurs is very close to the largest port of Greece and hence may act as an invasion hub (Lacarella et al. 2020). Secondary introductions have already been reported for P. siculus (Silva-Rocha et al. 2014), thus highlighting the importance of keeping the population of Athens restricted, since the port of Piraeus is less than 10 km from the area of introduction. To represent the worst scenario, we considered an eventual human-mediated invasion in Milos island, putting the endemic Milos wall lizard under threat. ...
... In other areas, higher neophilia has also been reported (Damas-Moreira et al. 2019). Such traits are likely behind the repeated invasion success of P. siculus in Southern Europe (Silva-Rocha et al. 2014;Adamopoulou 2015;Damas-Moreira et al. 2019 and America (Kolbe et al. 2013). Our results reinforce previous research suggesting that the Italian wall lizard exhibits a series of behavioral and physiological traits that favor survival in new environments, even in the presence of native congeneric species Vanhooydonck et al. 2000;Downes and Bauwens 2002;Vervust et al. 2010;Damas-Moreira et al. 2019Nikolic et al. 2019). ...
Though biological invasions constitute one of the biggest threats for global biodiversity, our understanding of the mechanisms that enable invasive species to outperform native species is still limited, especially, in terms of behavior. Most available studies have examined behavioral traits which favor invasive species on the later stages of invasion, however, our knowledge on earlier stages, namely, when alien species face novel environments and must exploit new resources, remains obscure. Here, we focus on one crucial behavioral trait, finding food. The Italian wall lizard (Podarcis siculus) has been widely introduced and established viable populations in S. Europe and N. America. We examined whether P. siculus has enhanced exploratory behavior and abilities to find food compared to two native congeneric species with which it may come in contact in the near future, an insular endemic (P. milensis) and a widely distributed lizard (P. erhardii). We performed a Y-maze experiment, in which we varied arm markings in a standard way to prevent learning. Podarcis siculus was more efficient than its congenerics in finding and consuming food. This exploitative superiority was persistent, more frequent and repetitive. Interesting behavioral differences were also detected within the native species. Some P. milensis individuals showed no interest in exploring the maze, while few P. erhardii individuals remained rather indifferent to food even after detecting it. Our results suggest that the invasive P. siculus displays behavioral traits that could provide better opportunities for survival in the new environment and thus facilitate establishment even in the presence of congenerics. This provides further support to the idea that behavior plays a crucial role in animal invasions.
... Habitat loss, urbanization, wildfires, overgrazing, and inbreeding have all been proposed as potential driving factors [18,19,[24][25][26][27][28]. It has been suggested that historical declines of P. raffonei might be related with the expansion of the Italian wall lizard P. siculus [18,19], which has been introduced across the whole Mediterranean basin and adjacent areas, as well as in the UK and the US and Canada, by human activities [29,30]. The competition with this introduced lizard is thus a possible driver of the decline of P. raffonei [28]. ...
The Aeolian wall lizard Podarcis raffonei is an island endemic that survives only on three tiny islets, and on the Capo Grosso peninsula of the Vulcano island, thus is among the European vertebrates with the smallest range and one of the most threatened by extinction. This species is declining due to competition and hybridization with the non-native lizard Podarcis siculus , but a regular monitoring program is lacking. Here we assessed the size and status of the Capo Grosso population of P . raffonei on Vulcano. In September 2015 we captured 30 individuals showing the typical brown phenotype of P . raffonei , while one single male showed a green phenotype, apparently intermediate between P . raffonei and the non-native Podarcis siculus . In May 2017, only 47% of 131 individuals showed the typical brown phenotype ( P . raffonei -like) and 53% showed the green phenotype ( P . siculus -like). Based on N -mixture models and removal sampling the estimated size of the Capo Grosso population was of 800–1300 individuals in 2017, being similar to 2015; available data suggest that the total range of the species could be as small as 2 ha. The frequency of individuals with the typical P . raffonei phenotype dramatically dropped between two samplings with a parallel increase of individuals displaying the green phenotype. Observation on outdoor captive-bred individuals demonstrates plasticity for colouration in P . raffonei individuals from Capo Grosso, with several individuals showing the typical brown pattern in September 2017 and a green pattern in March 2021. Non-exclusive hypotheses, including hybridization with P . siculus and plasticity in colour pattern of P . raffonei , are discussed to explain the phenotypic shifts of the P . raffonei population of Capo Grosso. While genomic evidence is required to reach conclusions and investigate eventual hybridization, it is urgent to undertake a programme for the monitoring and management of this lizard.
... An additional aspect that we should consider is that not even a single mitochondrial haplotype belonging to the continental lineages of P. siculus has been found on the western Pontine Archipelago. Since that P. siculus has colonized in recent times by long distance passive transport several islands of the Mediterranean Sea (Silva-Rocha et al., 2014;Santos et al., 2019), including for example the Tremiti Islands and the Eastern Pontine (Senczuk et al., 2017), it seems very strange to us that this has not happened for the Western Pontine. Although there are signs of sporadic introgression of some nuclear haplotypes (Senczuk et al., 2018b), the absence of mitochondrial haplotypes indicates some type of "blocking" of invasive lizards belonging to the mainland. ...
Based on genetic and morphological evidence, Senczuk et al. (2019) formally raised the Podarcis populations from the Western Pontine Islands, previously classified as several subspecies of P. siculus, to species rank, i.e. Podarcis latastei (Bedriaga, 1879). This taxonomic change was not accepted in the checklist of the European herpetofauna by Speybroeck et al. (2020), recently published on Amphibia-Reptilia. In this note we respond to the reasons given by Speybroeck and colleagues and support the validity of Podarcis latastei as an endemic Italian species.
