Lacertid Research Articles

Sexual dimorphism in traits related to locomotion: ontogenetic patterns of variation in Podarcis wall lizards

Sexual dimorphism in body size and shape in animals is normally linked to sexual selection mechanisms that modify the morphological properties of each sex. However, sexual dimorphism of ecologically relevant traits may be amplified by natural selection and result in the ecological segregation of both sexes. In the present study, we investigated patterns of sexual dimorphism of morphological traits relevant for locomotion in two lacertid lizards, Podarcis bocagei and Podarcis carbonelli, aiming to identify ontogenetic sources of variation. We analysed trunk and limb variation in relation to total body size, as well as the covariation of different traits, aiming to shed light on the proximate causation of adult sexual dimorphism. We find that, although immatures are generally monomorphic, adult females have a longer trunk, and adult males have longer fore and hind limbs. Both sexes differ substantially with respect to their growth trajectories and relationships between traits, whereas, in some cases, there are signs of morphological constraints delimiting the observed patterns. Because of the direct connection between limb size/shape and locomotor performance, which is relevant both for habitat use and escape from predators, the observed patterns of sexual dimorphism are expected to translate into ecological differences between both sexes.

Two new types of noose for capturing herps

th July; Revised on 2010, 27 th October; Accepted on 2010, 15 th November. Abstract. We present two new types of noose for capturing herps, which allow adapt- ing the size of the loop to the size of prey rapidly. We use a plastic ring to make the loop: the thread moves perfectly without any impediment or friction, and remains in good conditions for a longer time. The plastic ring can be made during fieldwork. Both new types of noose increase capture effectiveness, as it is possible to noose dif - ferent types of preys with a simple and rapid adaptation of the loop. We have tested this methodology in different species of lacertids, some snakes under rocks, and also some amphibians.

Notes on some Egyptian Lacertidae, including a new subspecies of Mesalina, involving the Seligmann effect

These notes on some Egyptian lacertid lizards contribute to faunal data for the planning of biodiversity conservation. Mesalina bahaeldini curatorum n. subsp. is described from Suez, characterized by coloration and larger (fewer) body scales. Range extensions are documented for Acanthodactylus aegyptius, Mesalina olivieri, and M. rubropunctata. The previously reported occurrence of A. opheodurus in Sinai is doubted. The fact that the M. b. curatorum types were collected in the 1820s demonstrates the importance of museum collections.

Fossil lizards from the locality of Gannat (late Oligocene–early Miocene, France) and a revision of the genus Pseudeumeces (Squamata, Lacertidae)

Pseudeumeces is a distinct lacertid lizard that is known from one species, P. cadurcensis (Filhol). Two species, Ligerosaurus pouiti and “Pseudeumeces” walbeckensis, previously referred to Pseudeumeces, do not belong to this genus. Pseudeumeces cadurcensis is characterized by a heterodont dentition, including large, posterior, crushing teeth on both the maxilla and dentary. Pseudeumeces differs from the Oligocene lacertid lizards Lacerta s.l. filholi and Mediolacerta that have non-expanded, bi- or tricuspid teeth. Dracaenosaurus, an Oligocene lacertid lizard, also possesses crushing, posterior teeth, but its teeth and jaw are much heavier than in those of Pseudeumeces. New specimens from Gannat document cranial elements previously unknown, i.e. frontal and parietal. Pseudeumeces is known with certainty from the early Oligocene (MP25) to the early Miocene (MN1). Notable features of Pseudeumeces (firm lateral contact between the coronoid and the dentary; strongly curved mandibule; tooth morphology and tooth number reduction) corroborate the previous assumption that this lacertid lizard was probably durophagous. Four Oligocene European lacertid lizards form a morphological and chronological series, but the common ancestry of these species has not been definitively proved.

Monitoring the performance of wild-born and introduced lizards in a fragmented landscape: Implications for ex situ conservation programmes

Ex situ conservation of animal populations may benefit from captive-breeding programmes, but these are criticised because they are assumed to be difficult, time-consuming and expensive, while they do not guarantee success. However, such assumptions remain untested in most organisms; for example, introductions could be very useful for recovering populations of small-sized species with short generation time, no learned behaviours, and ease to rear in captivity. Here, we document an easy, cheap and successful reintroduction programme of the lacertid lizard Psammodromus algirus. Two captive-bred cohorts (178 juveniles in 2001 and 187 in 2002) were released in four woodland fragments (0.9–5.2 ha) at two localities (B and V); B housed a stable lizard population whereas V apparently lacked a viable population of lizards. We monitored introduced and native lizards during 2002 and 2003, and carried out a corroborative searching in 2006 which confirmed the existence of a lizard population at site V. Introduced lizards had higher activity and dispersed more frequently among woodland fragments than native ones. Survivorship and growth rates were similar for both groups, but introduced juveniles were about 25% larger than native ones, due to both early hatching and better rearing conditions. The whole procedure was easily implemented in our Faculty facilities (mean hatching and hatchling survival rates of 0.90 and 0.87), and cost less than 20,000 € (excluding salaries). Therefore, similar programmes may be of wide application in small animals and of practical importance for species with a meta-population structure living in fragmented landscapes.

HOLASPIS GUENTHERI GLIDES LIKE A FEATHER

Most lacertid lizards are content scurrying in and out of nooks and crannies in walls and between rocks. However, some have opted for an arboreal life style. Holaspis guentheri leap from branch to branch as they scamper through trees in the African forest. There are even anecdotes that the tiny African lizards can glide. But without any obvious adaptations to help them to upgrade a leap to a glide, it wasn't clear whether the reptiles really do take to the air and, if they do, how they remain aloft. Intrigued by all aspects of lacertid locomotion, Bieke Vanhooydonck from the University of Antwerp and her colleagues, Anthony Herrel and Peter Aerts, decided to find out whether H. guentheri really glide. Recruiting undergraduate Greet Meulepas to the team, they began filming dainty H. guentheri lizards,gliding geckos (Ptychozoon kuhli) and landlocked Podarcis muralis lacertid lizards as the animals leapt from a 2 m high platform to see if the African lizards really could glide(p. 2475).Unfortunately, filming the tiny lizards was extremely difficult. Having startled the small animals into leaping off the platform, the team had little control over the animal's direction, and couldn't guarantee that it was parallel to their camera. It was also difficult to capture each trajectory with a single camera and tricky to get the lighting conditions right. But after weeks of persistence the team finally collected enough film, as the lizards leapt, to compare their performances.At first, it didn't look as if the African lizard was gliding any better than the P. muralis. Both animals were able to cover horizontal distances of 0.5 m after leaping from the platform, while the gliding gecko covered distances greater than 1 m, aided by its webbed feet and skin flaps. But when the team compared the lizards' sizes, they noticed that there was a big difference between the two lacertid lizards. The tiny African reptile only weighed 1.5 g, almost 1/3 of the larger lacertid lizard's weight and 1/10 the gliding gecko's mass, so Aerts calculated how far each lizard would travel horizontally if they fell like a stone. This time it was clear that the tiny H. guentheri was travelling 0.2 m further than he would have expected if it were simply jumping off the platform. Holaspis guentheri was definitely delaying its descent and landing more slowly than P. muralis; it was gliding.But how was the tiny lizard able to remain airborne for so long? Maybe the lizard was squashing itself flat while gliding to increase its surface area and generate more lift. But when the team analysed the lizards' trajectories, H. guentheri's shape did not change. And when Aerts calculated the amount of lift each lizard generated as they descended, it was clear that H. guentheri was unable to produce a lift force. The team realised that instead of increasing its surface area to generate lift, H. guentheri is able to glide because it is so light. The lizard's `wing loading' (mass:surface area ratio) was the same as that of the gliding gecko(assisted by skin flaps and webbed feet) so the lizard was able to glide like a feather because it was so light.Curious to find out why H. guentheri is so light, Herrel contacted Renaud Boistel, Paul Tafforeau and Vincent Fernandez at the European Synchrotron Radiation Facility to scan all three lizards' bodies. Visualising the animals' skeletons with X-rays, it was clear that H. guentheri's bones were packed full of air spaces, making the lizard's skeleton feather light for gliding.

Integration of Bayesian molecular clock methods and fossil-based soft bounds reveals early Cenozoic origin of African lacertid lizards.

BackgroundAlthough current molecular clock methods offer greater flexibility in modelling evolutionary events, calibration of the clock with dates from the fossil record is still problematic for many groups. Here we implement several new approaches in molecular dating to estimate the evolutionary ages of Lacertidae, an Old World family of lizards with a poor fossil record and uncertain phylogeny. Four different models of rate variation are tested in a new program for Bayesian phylogenetic analysis called TreeTime, based on a combination of mitochondrial and nuclear gene sequences. We incorporate paleontological uncertainty into divergence estimates by expressing multiple calibration dates as a range of probabilistic distributions. We also test the reliability of our proposed calibrations by exploring effects of individual priors on posterior estimates.ResultsAccording to the most reliable model, as indicated by Bayes factor comparison, modern lacertids arose shortly after the K/T transition and entered Africa about 45 million years ago, with the majority of their African radiation occurring in the Eocene and Oligocene. Our findings indicate much earlier origins for these clades than previously reported, and we discuss our results in light of paleogeographic trends during the Cenozoic.ConclusionThis study represents the first attempt to estimate evolutionary ages of a specific group of reptiles exhibiting uncertain phylogenetic relationships, molecular rate variation and a poor fossil record. Our results emphasize the sensitivity of molecular divergence dates to fossil calibrations, and support the use of combined molecular data sets and multiple, well-spaced dates from the fossil record as minimum node constraints. The bioinformatics program used here, TreeTime, is publicly available, and we recommend its use for molecular dating of taxa faced with similar challenges.

Concordance between locomotor morphology and foraging mode in lacertid lizards

Foraging behaviors exist along a continuum from highly sedentary, ambush foraging, to more continuous searching, or active foraging. Foraging strategies, or modes, are defined based upon locomotor behaviors (e.g. percent time moving, moves per minute). In lizards, traits correlated with ambush and active foraging have been of interest for some time; however, general patterns of correlated evolution between locomotor morphology and locomotor behavior have only recently begun to be quantified. In this study, variation in hindlimb morphology is investigated in a model group of lizard species that vary between active foraging and more sedentary (or mixed) foraging mode. Canonical variates analysis reveals that the two active foraging species occupy similar regions of the morphospace, while the two more sedentary species occupy different regions. The active foraging species have a narrow pelvis with shorter tibia and femora. The more sedentary species have a wide pelvis, long tibia and femora, and slightly longer metatarsals. Phylogenetic patterns of trait variation were examined through ancestral character state reconstruction and show morphological shifts in concert with foraging mode in these species. The observed shifts in locomotor morphology are discussed in light of published data on sprint speed and endurance in these species. Together, the data show that linking morphological variation to variation in stride length and stride frequency is critical to understanding the evolution of locomotor performance. Much more stride length and frequency data are needed among ambush, mixed, and active foraging species because these parameters, and their morphological components, are likely correlated with variation in food acquisition mode.

Ontogenetically stable dimorphism in a lacertid lizard (Acanthodactylus boskianus) with tests of methodology and comments on life-history

Recent arguments in the literature prompted us to compare methods for assessing sexual dimorphism in body proportions of lacertid lizards, using Acanthodactylus boskianus. Although expressing body-part measurements as proportional to head length was the most effective method, we recommend using trunk length for the baseline as a general method for lizards. We also argue that, when aiming to assess sexual dimorphism in body proportions of lizards, if the context is ecological, all available adults should be included. However, for morphology and taxonomy, small sub-samples of the largest individuals that maximally express their genetic morphological potential should be used. In A. boskianus, the sexual dimorphism of mensural characters in adults was typical: males were larger, with relatively larger head and appendages. However, the ontogeny of this dimorphism was unusual in that the differences existed already in youth and thereafter persisted isometrically. The sexual dimorphism of meristic characters was male-biased in numbers of femoral pores and of caudal vertebrae, and female-biased in numbers of ventral plates along the trunk and of precaudal vertebrae. Size dimorphism may conceivably play a role in sex recognition because two potential visual cues (i.e. size dimorphism and dichromatism) appear to complement each other. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society 2009, 97, 275–288.

Fast radiation of the subfamily Lacertinae (Reptilia: Lacertidae): History or methodical artefact?

Lacertinae is one of the three lacertid lizard subfamilies with a geographical distribution confined to the Palaearctic. Several past attempts to reconstruct its phylogeny resulted in unresolved bush-like topologies. We address the question of whether the lack of resolution is due to insufficient data or whether this lack reflects a rapid succession of speciation events. We analyzed four partial and one complete gene sequences from mitochondrial and nuclear genomes, totalling roughly 3600 bp. We included 29 species representing all 19 genera suggested in recent revision of Lacertinae [Arnold, E.N., Arribas, O., Carranza, S., 2007. Systematics of the palaearctic and oriental lizard tribe Lacertini (Squamata: Lacertidae: Lacertinae), with descriptions of eight new genera. Zootaxa 1430, 1–86]. The resulting phylogeny, first, corroborates monophyly at the genus level for the suggested genera, as well as the finding that Atlantolacerta andreanskyi, until recently part of Lacertinae, belongs to the subfamily Eremiadinae. Second, we find that increasing the sequence length and combining multiple nuclear and mitochondrial sequences did not resolve the polytomy, suggesting that the inferred topology indicates a multiple cladogenesis within a short geological period, rather than a methodical artefact. Divergence time estimates, based on previous estimates of several node ages, range from 13.9 to 14.9 million years for the radiation event, however with very broad confidence interval. To associate the radiation with a narrower geological time we consider palaeogeographic and palaeoclimatic data, assuming that the Lacertinae probably evolved in Central Europe and W Asia after the collision of Africa and Eurasia. We suggest that this radiation may date to the late Langhian (ca. 14–13.5 million years) when geological events caused abrupt changes in regional water–land distribution and climate, offering a window of distinct conditions.

Pheromone Communication in Amphibians and Reptiles

This selective review considers herpetological papers that feature the use of chemical cues, particularly pheromones involved in reproductive interactions between potential mates. Primary examples include garter snake females that attract males, lacertid lizards and the effects of their femoral gland secretions, aquatic male newts that chemically attract females, and terrestrial salamander males that chemically persuade a female to mate. Each case study spans a number of research approaches (molecular, biochemical, behavioral) and is related to sensory processing and the physiological effects of pheromone delivery. These and related studies show that natural pheromones can be identified, validated with behavioral tests, and incorporated in research on vomeronasal functional response.

Complete mitochondrial genomes of three lizard species and the systematic position of the Lacertidae (Squamata)

Complete mitochondrial (mt) genomes were sequenced from representatives of three lacertid lizards: Podarcis siculus, Podarcis muralis and Phoenicolacerta kulzeri. In all three genomes the arrangement of the 22 tRNAs, the two rRNAs and the 13 protein-coding genes conforms to the common vertebrate arrangement. The phylogenetic position of Lacertidae within the order Squamata was determined through sequence analyses based on large sections of complete mt genomes. The number of nucleotide sites used for tree construction was 9234 when outgroup taxa were included, and 10 499 when only Squamata were compared. The phylogenetic analyses confirmed the sister group relationship between Lacertidae and Amphisbaenia as previously proposed on the basis of molecular data. Additionally, Bayesian analysis revealed a well supported clade comprising (Gekkonidae (Lacertidae + Amphisbaenia)), which is not in accordance with the traditional morphological view and most of the previous molecular studies. It confirms, however, the close relationship between Gekkonidae and Amphisbaenia as revealed in a recent study based on complete mt genomes from a smaller number of taxa. Intra- and intergeneric sequence comparisons of six commonly used marker genes showed rather high levels of divergence within the Lacertidae. In the intrageneric comparison the control region proved to be considerably more conserved than the protein coding genes.

Can scent-mediated female mate preference explain an abrupt mtDNA cline in Lacerta schreiberi?

AbstractDivergence in female mate preferences can strongly influence the structure and dynamics of hybrid zones. We examined the potential role of female mate preferences in maintaining an abrupt west-east mtDNA cline between two deeply divergent genetic lineages of Lacerta schreiberi, a lizard endemic to the Iberian Peninsula. The lineages are largely morphologically cryptic but with respect to the mtDNA cline, western males tend to be less parasitized, in better body condition and more intensely coloured than eastern males, a pattern that cannot be explained by environmental variation alone. The lineages may also differ in unmeasured aspects such as physiology, behaviour and olfactory signals, which may influence mate choice. As female mate attraction has been found to vary with olfactory cues in lacertid lizards, we experimentally tested whether females were differentially attracted to femoral pore secretions of males from the two genetic backgrounds. Females did not prefer scents of 'higher quality' western males, nor did they prefer the scents of males belonging to their own genetic background. This suggests the abrupt mtDNA cline is unlikely to be explained by assortative mating of matrilineages that distinguish male genetic background based on scent.

Follicle ablation increases offspring size in a lizard with a low clutch frequency

AbstractStudies of lizards and snakes have shown that an experimental reduction of offspring number sometimes, but not always, results in an increase in offspring size. We applied the “follicle ablation” technique to an oviparous lacertid lizard (Eremias argus) to test the hypothesis that offspring size can be easily altered by manipulating clutch size in species with a low clutch frequency. Our manipulation of clutch size had the effect of inducing variation in egg size in the first post-surgical clutch, with follicle-ablated females producing fewer larger eggs than did controls. Follicle-ablated females produced a second post-surgical clutch as normally as did controls. The proportional amount of resources allocated to reproduction did not shift seasonally in E. argus, but females normally switched from producing a larger number of smaller eggs early in the breeding season to a smaller number of larger eggs later in the season. Females used in this study never produce more than two clutches per breeding season. Therefore, our data validate the hypothesis tested. Our data also provide an inference that maximization of reproductive success could be achieved in females of E. argus by diverting a larger enough, rather than an extraordinarily high, fraction of the available energy to individual offspring in single reproductive episodes.

Different habitats, different pressures? Analysis of escape behaviour and ectoparasite load in Podarcis sicula (Lacertidae) populations in different agricultural habitats

AbstractHuman agricultural activities can deeply alter the environment thus provoking major impacts on a variety of organisms. Agricultural habitats however can be very different from one another in terms of habitat structure and management intensity, presenting varying pressures and/or benefits for different species. Agro-ecosystems can have opposing effects on reptiles and in some circumstances the presence of a species can even been enhanced by agricultural practices. We focused our study on Podarcis sicula, a relatively widespread lacertid lizard commonly present in agro-environments in Italy. We examined escape behaviour, caudal autotomy rates and ectoparasite load (tick infestation) in populations living in two different land uses, olive tree plantations and vineyards. All three aspects seemed to be deeply influenced by habitat structure. Predation pressure, as evaluated by tail break frequency, was lower in olive tree plantations, the most structurally complex habitats. In this type of habitat lizard escape behaviour was characterised by a clear preference for olive trees as refuges: individuals ran farther distances on average to reach the trees and hid inside them for a relatively long time. In vineyards, on the contrary, a less clear escape strategy was observed, showing a use of more temporary refuges. Also tick (Ixodes ricinus) infestation differed among land uses, being higher in olive tree plantations, probably in relation to vegetation cover features. Differences were found also between managements (with a higher tick load in traditional cultivations) and sexes, with males being more parasitized.

Karyotype, chromosome structure, reproductive modalities of three Southern Eurasian populations of the common lacertid lizard, Zootoca vivipara (Jacquin, 1787)

According to a hypothesis of the evolution of viviparity the lacertid lizard Zootoca vivipara, rare relict oviparous populations of the species might occur in south - ern-eastern part of its distribution area. Such a hypothesis has been verified by compar - ing the karyotype, chromosome structure, and reproductive modality of three popula - tions of south-eastern part of Russia, including Altai and neighbouring regions, where small territories remained isolated during the Pleistocene cooling and where Pleistoce - nic fossils of Z. vivipara have been found. The chromosomal study was carried out by conventional staining method and banding methods, namely C-banding and sequential staining of C-banding+ fluorochromes, CMA 3 and DAPI. All studied females displayed viviparous reproductive modality and showed a karyotype of 2N = 35 acrocentric chro - mosomes, with a Z1Z2W sex chromosome system. Chromosome W was subtelocentric . No inter-population variability on karyotype and heterochromatin distribution and composition was observed. From the obtained data the three studied south-eastern Russian viviparous populations belong to the Russian viviparous form of Z. v. vivipara.

Fine-scale phylogeographic structure of Borrelia lusitaniae revealed by multilocus sequence typing.

Borrelia lusitaniae is an Old World species of the Lyme borreliosis (LB) group of tick-borne spirochetes and prevails mainly in countries around the Mediterranean Basin. Lizards of the family Lacertidae have been identified as reservoir hosts of B. lusitaniae. These reptiles are highly structured geographically, indicating limited migration. In order to examine whether host geographic structure shapes the evolution and epidemiology of B. lusitaniae, we analyzed the phylogeographic population structure of this tick-borne bacterium using a recently developed multilocus sequence typing (MLST) scheme based on chromosomal housekeeping genes. A total of 2,099 questing nymphal and adult Ixodes ricinus ticks were collected in two climatically different regions of Portugal, being ∼130 km apart. All ticks were screened for spirochetes by direct PCR. Attempts to isolate strains yielded 16 cultures of B. lusitaniae in total. Uncontaminated cultures as well as infected ticks were included in this study. The results using MLST show that the regional B. lusitaniae populations constitute genetically distinct populations. In contrast, no clear phylogeographic signals were detected in sequences of the commonly used molecular markers ospA and ospC. The pronounced population structure of B. lusitaniae over a short geographic distance as captured by MLST of the housekeeping genes suggests that the migration rates of B. lusitaniae are rather low, most likely because the distribution of mediterranean lizard populations is highly parapatric. The study underlines the importance of vertebrate hosts in the geographic spread of tick-borne microparasites.

“Meiotic abnormality in dominant genic male sterile Brassica napus”

We have isolated and characterised sequences of a SINE family specific for squamate reptiles from a genome of lacertid lizard that we called Squam1. Copies are 360-390 bp in length and share a significant similarity with tRNA gene sequence on its 5'-end. This family was also detected by us in DNA of representatives of varanids, iguanids (anolis), gekkonids, and snakes. No signs of it were found in DNA of mammals, birds, amphibians, and crocodiles. Detailed analysis of primary structure of the retroposons obtained by us from genomic libraries or GenBank sequences was carried out. Most taxa possess 2-3 subfamilies of the SINE in their genomes with specific diagnostic features in their primary structure. Individual variability of copies in different families is about 85% and is just slightly lower on the genera level. Comparison of consensus sequences on family level reveals a high degree of structural similarity with a number of specific apomorphic features which makes it a useful marker of phylogeny for this group of reptiles. Snakes do not show specific affinity to varanids when compared to other lizards, as it was suggested earlier.

Seasonal Shifts in Reproductive Investment of Female Northern Grass Lizards (Takydromus septentrionalis) from a Field Population on Beiji Island, China

Understanding seasonal variation in reproductive investment is crucial to explore life-history strategies of a given species. We measured the timing of reproduction and the reproductive output of recently captured females to test whether the conclusions on seasonal shifts of reproductive investment made in the laboratory can apply to field populations in a lacertid lizard Takydromus septentrionalis. Female T. septentrionalis started producing eggs in April and ceased reproduction in August. During the midreproductive season (May to July), more than 60% of field-captured females were gravid. The females produced more eggs and thus had heavier total egg mass in May than in July, although egg mass and hatchling size did not vary seasonally. The seasonal variation in clutch size did not induce seasonal shifts in trade-offs between clutch size and egg size. In general, these results are largely consistent with data collected in the laboratory and support the conclusion of previous studies in this species that seasonal shifts in reproductive output are determined ultimately by natural selection. However, Relative Clutch Mass (RCM) of recently captured females was greater than that of females in captivity, which suggests that RCMs could be affected noticeably by proximate factors. Therefore, when explaining the evolutionary strategies of an organism based on intraspecific comparisons of RCMs, investigators should measure them in a common-garden environment so as to control the potential influence of environmental factors.

Two new species of Podarcis (Squamata; Lacertidae) from Greece

Recently, several works have focused on the lacertid lizards of the genus Podarcis, revealing cases of hidden diversity and paraphyly, and offering evidence that suggests the revision of the extant taxonomical arrangements within the genus. Hidden diversity and paraphyly have been shown to exist in the relationships between the Balkan species P. peloponnesiaca and P. erhardii as well. Here we couple a molecular (mtDNA) dataset with a corresponding morphological one, consisting of morphometric and pholidotic characters, to check for concordance between the two. Phylogenetic analyses reinforced previous suggestions for paraphyly of P. erhardii with respect to P. peloponnesiaca. We found the variation of certain pholidotic characters concordant with the relationships inferred from partial mtDNA sequences, whereas morphometric characters were not. The latter is possibly due to greater influence of morphometric characters by environmental factors. To avoid the observed paraphyly we proceed with the description of the populations from Crete and the islet of Pori, until now designated as P. erhardii, as separate taxa at the species level.