Podarcis Sicula Research Articles

Getting ahead: exploitative competition by an invasive lizard

Biological invasions are a contemporary global threat because invasive species can have substantial negative economic and ecological impacts. Invasive species can outcompete native species through two main mechanisms: interference competition (direct, negative interactions like aggression) and/or exploitative competition (indirect, negative interactions resulting from species using the same, limited resources like food). The invasive Italian wall lizard (Podarcis siculus) was introduced into Lisbon, Portugal, 20 years ago, and is believed to be locally displacing the native green Iberian wall lizard (Podarcis virescens). We experimentally tested for competition between these two lizard species by establishing heterospecific (one pair of each species) and conspecific (two pairs of the same species; control) treatments in enclosures containing a high- and a low-quality refuge. Lizards were fed from food dishes every other day. We tested if species showed interference (aggressive behaviour, stealing food and shelter exclusion) or exploitative competition (tolerance between species but differences in food consumption efficiency). We found evidence for exploitative competition: the invasive species arrived first at food stations, consumed more food and gained more weight than the native species. We suggest that exploitative competition may, in part, explain the observed displacement of P. virescens from contact areas with the invasive P. siculus. Deciphering the competitive mechanisms between invasive and native species is vital for understanding the invasion process. To become successful invaders, alien species must often outcompete native species they encounter in a new location. But how can an alien species outcompete a resident with a long evolutionary history in an environment in which the resident is expected to be better adapted? We studied an invasive and native congeneric pair of sympatric lizards to understand how they interact and potentially compete in a controlled environment. The invasive Italian wall lizard and a native congener were very tolerant of each other; however, the invasive species was first to arrive at food, ate more and grew faster, suggesting exploitative competition. This contrasts with previous studies in other introduced locations where the Italian wall lizard was more aggressive towards native lizards, suggestive of interference competition. Our results help explain why the Italian wall lizard is so successful and suggests it may compete in different ways, possibly in response to local environmental conditions and which species it may be competing with, but with the same outcome: the displacement of native species.

Rapid Dietary Shift in Podarcis siculus Resulted in Localized Changes in Gut Function.

AbstractNatural dietary shifts offer the opportunity to address the nutritional physiological characters required to thrive on a particular diet. Here, we studied the nutritional physiology of Podarcis siculus, with populations on Pod Mrčaru, Croatia, that have become omnivorous and morphologically distinct (including the development of valves in the hindgut) from their insectivorous source population on Pod Kopište. We compared gut structure and function between the two island populations of this lizard species and contrasted them with an insectivorous mainland out-group population in Zagreb. On the basis of the adaptive modulation hypothesis, we predicted changes in gut size and structure, digestive enzyme activities, microbial fermentation products (short-chain fatty acids [SCFAs]), and plant material digestibility concomitant with this dietary change. The Pod Mrčaru population had heavier guts than the mainland population, but there were no other differences in gut structure. Most of the enzymatic differences we detected were between the island populations and the out-group population. The Pod Mrčaru lizards had higher amylase and trehalase activities in their hindguts compared with the Pod Kopište population, and the Pod Kopište lizards had greater SCFA concentrations in their hindguts than the omnivorous Pod Mrčaru population. Interestingly, the differences between the Pod Mrčaru and Pod Kopište populations are primarily localized to the hindgut and are likely influenced by microbial communities and a higher food intake by the Pod Mrčaru lizards. Although subtle, the changes in hindgut digestive physiology impact the digestibility of plant material in adult lizards-Pod Mrčaru lizards had higher digestibility of herbivorous and omnivorous diets fed over several weeks in the laboratory than did their source population.

Spermatogenesis and regulatory factors in the wall lizard Podarcis sicula

Spermatogenesis is an extraordinarily complex process, regulated by several factors, which leads to the differentiation of spermatogonia into spermatozoa. Among vertebrates, several reports have been focused on the lizard Podarcis sicula, a seasonal breeder and a good model for the study of reproductive processes. The goal of this review is to resume all the available data about systemic and above all local control factors involved in the control of P. sicula testicular activity. During the seasonal reproductive cycle, the variation of the expression levels of these factors determines significant variations that induce the activation or blocking of spermatogenesis. The data supplied in this review, in addition to analyze the current literature regarding the main actors of Podarcis sicula spermatogenesis, will hopefully provide a basic model that can be used for further studies on the intratesticular interaction between molecular factors that control spermatogenesis.

Proximate and ultimate drivers of variation in bite force in the insular lizardsPodarcis melisellensisandPodarcis sicula

AbstractBite force is a key performance trait in lizards because biting is involved in many ecologically relevant tasks, including foraging, fighting and mating. Several factors have been suggested to impact bite force in lizards, such as head morphology (proximate factors), or diet, intraspecific competition and habitat characteristics (ultimate factors). However, these have been generally investigated separately and mostly at the interspecific level. Here we tested which factors drive variation in bite force at the population level and to what extent. Our study includes 20 populations of two closely related lacertid species, Podarcis melisellensis and Podarcis sicula, which inhabit islands in the Adriatic. We found that lizards with more forceful bites have relatively wider and taller heads, and consume more hard prey and plant material. Island isolation correlates with bite force, probably by driving resource availability. Bite force is only poorly explained by proxies of intraspecific competition. The linear distance from a large island and the proportion of difficult-to-reduce food items consumed are the ultimate factors that explain most of the variation in bite force. Our findings suggest that the way in which morphological variation affects bite force is species-specific, probably reflecting the different selective pressures operating on the two species.

Differences in cautiousness between mainland and island Podarcis siculus populations are paralleled by differences in brain noradrenaline/adrenaline concentrations

Adaptive behavior is shaped by the type and intensity of selection pressures coming from the environment, such as predation risk and resource availability, and can be modulated by individual's neuroendocrine profile involving steroid hormones and the brain-stem monoaminergic circuits projecting to forebrain structures. Boldness when faced with a predator and exploration/activity when confronted with a new environment reflect the degree of cautiousness and/or “risk-taking” of an individual. In this study we have explored to which extent two populations of Podarcis siculus occupying different ecological niches: mainland (ML) and an islet (ISL) differ in the level of cautiousness and whether these differences are paralleled by differences in their monoaminergic profiles. Boldness was tested in the field as antipredator behavior, while novel space and object explorations were tested in a laboratory setting in an open field apparatus. Finally, serotonin, dopamine, noradrenaline (NA) and adrenaline (ADR) concentrations were measured in whole brain samples by ELISA. Lizards from ML population spent significantly more time hiding after a predator encounter in the field, displayed lower intensity of novel space exploration in a laboratory setting, and contained significantly higher whole-brain concentrations of NA and ADR than their ISL counterparts. Parallelism between the level of risk-taking behavior and concentrations of neurotransmitters mediating alertness and reaction to stress suggests that the differing environmental factors on ML and ISL may have shaped the degree of cautiousness in the residing lizard populations by affecting the activity of NA/ADR neural circuits.

First record of an Italian Wall Lizard (<i>Podarcis siculus</i>) in British Columbia, Canada

Italian Wall Lizard (Podarcis siculus) is known from scattered locations in the United States, including Orcas Island, Washington, where a population went unreported for about a decade, but was confirmed in 2017. On 19 June 2019, a single adult female P. siculus was caught in an industrial area along the northern arm of the Fraser River in Vancouver, British Columbia; this represents the first record of this species on the loose in Canada. No other P. siculus were sighted in the Vancouver area. We assume this Canadian P. siculus was a lone stow-away and had not deposited eggs. We could not determine how it entered the province.

Effects of alkylphenols mixture on the adrenal gland of the lizard Podarcis sicula

Alkylphenols (AP) are widespread environmental compounds belonging to the large family of substances known as Endocrine Disrupting Chemicals (EDCs). The present study was carried out to assess the effects of Octylphenol (OP) alone and in combination with Nonylphenol (NP) on the hypothalamus-pituitary-adrenal gland (HPA) axis of the lizard Podarcis sicula. Lizards are good bioindicators due to their features such as wide distribution, large population and good sensitivity to contaminants. Results obtained showed a time and dose-dependent stimulation of the HPA together with a high variation of both catecholamine plasma levels and greater vascularization and hypertrophy of steroidogenic cord of adrenal gland after both OP and OP + NP treatments. Interestingly, the OP + NP mixture treatment has provoked a state of stress of the adrenal gland which in fact appeared to be characterized by the presence of a marked macrophage infiltration which can be seen especially close to the connective capsule surrounding the gland. This macrophage infiltration could be an evidence of a particularly pronounced inflammatory state to indicate, probably, an animal’s response to a non-physiological situation.

Urban invaders are not bold risk-takers: a study of 3 invasive lizards in Southern California.

Biological invasions threaten biodiversity worldwide, and therefore, understanding the traits of successful invaders could mitigate their spread. Many commonly invasive species do well in disturbed habitats, such as urban environments, and their abilities to effectively respond to disturbances could contribute to their invasiveness. Yet, there are noninvasive species that also do well in disturbed habitats. The question remains whether urban invaders behave differently in urban environments than noninvaders, which could suggest an “urban-exploiting” phenotype. In Southern California, the co-occurrence of invasive Italian wall lizards Podarcis siculus, brown anoles Anolis sagrei, and green anoles A. carolinensis, and native western fence lizards Sceloporus occidentalis offers an opportunity to test whether invasives exhibit consistent differences in risk-taking within human-altered habitats compared with a native species. We predicted that invasive lizards would exhibit more bold behavior by having shorter flight-initiation distances (FIDs) and by being found farther from a refuge (behaviors that would presumably maximize foraging in low-risk environments). Invasive populations had similar or longer FIDs, but were consistently found at distances closer to a refuge. Collectively, invasive lizards in urban habitats were not bolder than a native species. Reliance on nearby refuges might help species successfully invade urban habitats, and if a general pattern, may pose an added challenge in detecting or eliminating them.

Do lizards (Podarcis siculus) react to whip snake (Hierophis viridiflavus) scents? A comparative test on odour stimuli recognition

Abstract We tested the ability in the ruin lizard (Podarcis siculus) to discriminate between odour of a predator (the whip snake Hierophis viridiflavus) and those from harmless sources. We analysed two lizard populations: one (PP) predated by snakes and another (PNP) where no snakes occur. We tested the rate of tongue flick directed to cotton-tips impregnated with odours, and the rate of tongue flick, the time spent in immobility, escaping and exploring the terrarium containing a diffuse odour. We used the smell of the snake as dangerous stimulus, water as blank control, and cologne as complex control. Both populations did not discriminate snake cue from complex odour in both the experiments. PP individuals were more active and prone to analysing and exploring the stimuli and the environment than PNP lizards. In PP, the higher interest towards cologne and snake scents could be triggered just by their complexity that requires more tongue flicks to be ‘analysed’, with no apparent adaptive anti-predatory value. Overall, we observed a total responsiveness and activity pattern higher in the PP respect to PNP.

First molecular identification of the tapeworm Mesocestoides litteratus from an Italian wall lizard (Podarcis siculus)

The cyclophyllidean cestodes of the genus Mesocestoides are parasites infecting a variety of carnivorous vertebrates. The complete life cycle of these tapeworms is not clear yet, but some reptiles, amphibians, birds and micro-mammals have been indicated as intermediate hosts. Although Mesocestoides spp. are considered endemic in Europe, infections by the metacestode form, the tetrathyridium, are reported less commonly than are infections by adult worms. In the present study, a peritoneal infection by Mesocestoides tetrathyridia in a wall lizard (Podarcis siculus) collected in central Italy is described. The phylogenetic position of the metacestodes was assessed using the barcode region COI and species delimitation methods, i.e. Automatic Barcode Gap Discovery (ABGD) and Bayesian implementation of the Poisson Tree Processes model (bPTP). COI sequence analysis assigned the isolates from the wall lizard to Mesocestoides litteratus, as confirmed by species delimitation and phylogenetic analyses. M. litteratus has been observed in Europe mainly in the definitive hosts (dogs, cats and foxes), and never identified in the Mediterranean region. This study represents the first evidence of parasitic infection by tetrathyridia of M. litteratus in P. siculus in Italy.

Immunolocalization of aromatase P450 in the epididymis of Podarcis sicula and Rattus rattus.

The goal of this study was to evaluate P450 aromatase localization in the epididymis of two different vertebrates: the lizard Podarcis sicula, a seasonal breeder, and Rattus rattus, a continuous breeder. P450 aromatase is a key enzyme involved in the local control of spermatogenesis and steroidogenesis and we proved for the first time that this enzyme is represented in the epididymis of both P. sicula and R. rattus. In details, P450 aromatase was well represented in epithelial and myoid cells and in the connective tissue of P. sicula epididymis during the reproductive period; instead, during autumnal resumption this enzyme was absent in the connective tissue. During the non-reproductive period, P450 aromatase was localized only in myoid cells of P. sicula epididymis, whereas in R. rattus it was localized both in myoid cells and connective tissue. Our findings, the first on the epididymis aromatase localization in the vertebrates, suggest a possible role of P450 aromatase in the control of male genital tract function, particularly in sperm maturation.

Spring and Fall recrudescence in Podarcis siculus ovaries: A role for progesterone

In the lacertid Podarcis siculus the reproductive cycle is typically biphasic, with alternate recrudescence (Spring and Fall) and resting (Summer and Winter) phases. This study aimed to shed some light on the role exerted by progesterone during the two recrudescence periods; to this purpose, exogenous progesterone was administered intraperitoneally and the effects on oogonial proliferation, oocyte recruitment, and follicle cells apoptosis were determined. The presence and distribution of progesterone receptors was also investigated by immunohistochemistry and western blotting. Results indicate that progesterone would play different roles and follow different route of action in the two recrudescence periods thus confirming the complexity of the mechanisms controlling oogenesis in this species of vertebrate.

Lizard colour plasticity tracks background seasonal changes.

ABSTRACTEnvironmental heterogeneity on a spatial and temporal scale fosters an organism's capacity to plastically alter coloration. Predation risk might favour the evolution of phenotypic plasticity in colour patterns, as individuals who change colour throughout the year may be able to improve their fitness. Here we explored the change in dorsal pigmentation of the Italian wall lizard (Podarcis siculus campestris) at three time points (March, July, October) during a period of activity in a Mediterranean natural area in southern Italy. Following a preliminary investigation conducted in 2018, during 2019 we captured 135 lizards and took a picture of their ventral scales to check for possible recapture over the sessions. Lizard dorsal pictures were collected in the field with the support of a reference chart to quantitatively estimate chromatic variables (hue, saturation and value). At the same time, pictures of the environmental background were collected. Our findings suggest that lizards are capable of altering dorsal coloration during seasonal change. They vary from green at the onset of spring, to brownish in the middle of summer and to a greyish colour in October. This modification closely followed environmental background colour variation and enhanced lizard crypsis during each season.

Podarcis siculus latastei (Bedriaga, 1879) of the Western Pontine Islands (Italy) raised to the species rank, and a brief taxonomic overview of Podarcis lizards

In recent years, great attention has been paid to many Podarcis species for which the observed intra-specific variability often revealed species complexes still characterized by an unresolved relationship. When compared to other species, P. siculus underwent fewer revisions and the number of species hidden within this taxon may have been, therefore, underestimated. However, recent studies based on genetic and morphological data highlighted a marked differentiation of the populations inhabiting the Western Pontine Archipelago. In the present work we used published genetic data (three mitochondrial and three nuclear gene fragments) from 25 Podarcis species to provide a multilocus phylogeny of the genus in order to understand the degree of differentiation of the Western Pontine populations. In addition, we analyzed new morphometric traits (scale counts) of 151 specimens from the main islands of the Pontine Archipelago. The phylogenetic analysis revealed five principal Podarcis groups with biogeographic consistency. The genetic distinctiveness of the Podarcis populations of the Western Pontine Islands is similar or even more ancient than those observed in numerous other pairs of Podarcis sister species. In the light of these evidences we raise the Western Pontine lizards to specific rank; thus they should be referred to as Podarcis latastei.

Fixation of genetic variation and optimization of gene expression: The speed of evolution in isolated lizard populations undergoing Reverse Island Syndrome.

The ecological theory of island biogeography suggests that mainland populations should be more genetically divergent from those on large and distant islands rather than from those on small and close islets. Some island populations do not evolve in a linear way, but the process of divergence occurs more rapidly because they undergo a series of phenotypic changes, jointly known as the Island Syndrome. A special case is Reversed Island Syndrome (RIS), in which populations show drastic phenotypic changes both in body shape, skin colouration, age of sexual maturity, aggressiveness, and food intake rates. The populations showing the RIS were observed on islets nearby mainland and recently raised, and for this they are useful models to study the occurrence of rapid evolutionary change. We investigated the timing and mode of evolution of lizard populations adapted through selection on small islets. For our analyses, we used an ad hoc model system of three populations: wild-type lizards from the mainland and insular lizards from a big island (Capri, Italy), both Podarcis siculus siculus not affected by the syndrome, and a lizard population from islet (Scopolo) undergoing the RIS (called P. s. coerulea because of their melanism). The split time of the big (Capri) and small (Scopolo) islands was determined using geological events, like sea-level rises. To infer molecular evolution, we compared five complete mitochondrial genomes for each population to reconstruct the phylogeography and estimate the divergence time between island and mainland lizards. We found a lower mitochondrial mutation rate in Scopolo lizards despite the phenotypic changes achieved in approximately 8,000 years. Furthermore, transcriptome analyses showed significant differential gene expression between islet and mainland lizard populations, suggesting the key role of plasticity in these unpredictable environments.

Studies on Gastrointestinal Helminth of Three Lacertid Lizard Species, Podarcis Muralis, Podarcis Siculus and Ophisops Elegans (Sauria: Lacertidae) from Bursa, North-Western Turkey.

SummaryA total of 80 specimens of three species of lacertid Podarcis muralis (39), Podarcis siculus (18) and Ophisops elegans (23) from Bursa were examined for helminths. One species of Digenea, Plagiorchis elegans, 1 species of Cestoda, Mesocestoides sp. (tetrathyridium); and 3 species of Nematoda, Skrjabinodon medinae, Spauligodon saxicolae and Skrjabinelazia hoffmanni were found. The helminths reported in this study are generalist helminths that infect a number of lizards.

HSP70 localization in Podarcissiculus embryos under natural thermal regime and following a non-lethal cold shock

The Heat Shock Proteins (HSPs) are a superfamily of molecular chaperones that maintain cellular homeostasis under stress. HSP70 represents the major stress-inducible family member, often activated in response to changes in thermal ranges of organisms, and therefore playing an important role enhancing thermal tolerance limits in ectothermic animals. The present study aimed to investigate the presence and the localization of HSP70 through the development of Podarcis siculus, an oviparous lizard inhabiting temperate Mediterranean regions, showing a limited potential to tolerate thermal changes during embryogenesis. Immunohistochemical analysis demonstrated that HSP70 protein is constitutively present in early embryonic stages, abundantly distributed in eye, in encephalic domains (predominantly in ventricular areas and in grey matter), in grey matter of spinal cord, in lung, gut mucosa, hepatic cords and kidney tubules. Interestingly, a severe drop in incubation temperature (5°C for 3 days) does not induce enhancements in HSP70 levels nor changes in tissues localization. These results suggest that the HSP70 found in P. siculus embryos represents a non-inducible, constitutive molecular chaperone that should be better called Heat Shock Cognate 70 (HSC70); the presence of stress-induced members of the HSP family in P. siculus has yet to be proven.

Diet variability among insular populations of Podarcis lizards reveals diverse strategies to face resource-limited environments.

Access to resources is a dynamic and multicausal process that determines the success and survival of a population. It is therefore often challenging to disentangle the factors affecting ecological traits like diet. Insular habitats provide a good opportunity to study how variation in diet originates, in particular in populations of mesopredators such as lizards. Indeed, high levels of population density associated with low food abundance and low predation are selection pressures typically observed on islands. In the present study, the diet of eighteen insular populations of two closely related species of lacertid lizards (Podarcis sicula and Podarcis melisellensis) was assessed. Our results reveal that despite dietary variability among populations, diet taxonomic diversity is not impacted by island area. In contrast, however, diet disparity metrics, based on the variability in the physical (hardness) and behavioral (evasiveness) properties of ingested food items, are correlated with island size. These findings suggest that an increase in intraspecific competition for access to resources may induce shifts in functional components of the diet. Additionally, the two species differed in the relation between diet disparity and island area suggesting that different strategies exist to deal with low food abundance in these two species. Finally, sexual dimorphism in diet and head dimensions is not greater on smaller islands, in contrast to our predictions.

Lizards and the city: A community study of Lacertidae and Gekkonidae from an archaeological park in Rome

The study of animal communities is mainly concerned with how the groupings of species are distributed in nature and the ways in which the members of these communities assemble and interact with each other. Pianka suggested that animals partition environmental resources in three basic ways: temporally, spatially, and trophically. Such differences in activities separate the niches, reduce competition, and presumably allow the coexistence of a variety of species in the communities. The present work compares the spatial, trophic and temporal niches of four reptile species, two geckos (Tarentola mauritanica and Hemidactylus turcicus) and two lizards (Podarcis muralis and Podarcis siculus) living in a same roman-age archeological park situated area inside Rome metropolitan area (central Italy), and sharing the same two-dimensional habitat (the roman aqueduct walls). The results showed an overall overlap among species higher than expected by chance for all considered ecological dimensions except for the spatial resource related to the vertical position on the wall. Surprisingly, despite living in a two-dimension habitat would expect an increase of interspecific interaction rate, no partition was observed between species with the most convergent ecological requirements. Although our hypotheses about lizard communities structuring and interactions in the urban habitat require further investigation, we think that our lizard community is based more on the ecological needs of each species rather than on species’ interactions. The observed spatial segregation between diurnal geckos and lizards would not support alone the interaction hypothesis. We speculated that the coexistence of ecologically overlapping species at the study area is allowed by a non-limiting availability of resources (above all food) that would result in the reduction of heterospecific competitive interactions and an overall wide overlap of resource use.

Extreme necrosis in an Italian Wall Lizard (Podarcis sicula)

Extreme necrosis in an Italian Wall Lizard (Podarcis sicula)