Ophidiomyces ophidiicola (Oo) is a fungus primarily affecting the skin of a wide range of snake species. Ophidiomyces ophidiicola has been found in snakes under human care as well as in free-ranging snakes, and the number of documented infections in free-ranging snakes from Europe is increasing. To better understand the distribution of Oo in snakes in Germany, 329 samples were collected from 328 free-ranging snakes from 47 different locations in Germany in 2021-22. Samples included swabs (n=188), shed skins (n=134), and carcasses (n=7) from six different species: 140 dice snakes (Natrix tessellata), 68 European common adders (Vipera berus), 62 Aesculapian snakes (Zamenis longissimus), 29 smooth snakes (Coronella austriaca), 17 whip snakes (Hierophis viridiflavus), and 12 grass snakes (Natrix natrix). Sampled animals were examined for the presence of skin lesions and samples were screened for Oo by real-time PCR (qPCR). Ophidiomyces ophidiicola was detected in 30/328 snakes (9.1%), including dice snakes (23/30, 77%), Aesculapian snakes (5/30, 17%), and grass snakes (2/30, 7%). No differences were found in the likelihood of detection of Oo between individual colubrid species, but colubrid snakes were significantly more likely to be Oo positive than were viperid snakes (Fisher's exact test, P=0.001). Skin lesions were observed in 37/328 snakes (11.3%), of which 8/37 (22%) were Oo positive by qPCR. Significant differences were found in Oo prevalences depending on the presence of skin lesions, and Oo positive snakes were more likely to have skin lesions compared to snakes that were Oo negative (OR=3.37 [95% confidence interval 1.38-8.26%; P<0.05]).

We report the first documented case of interspecific ophiophagy by the naturalised false smooth snake (Macroprotodon mauritanicus), preying on the introduced horseshoe whip snake (Hemorrhois hippocrepis) in Mallorca (Balearic Islands, Spain). The event occurred in a suburban garden and was photographically recorded. While M. mauritanicus is known to consume reptiles, interspecific ophiophagy has not previously been documented. This observation represents a novel trophic interaction between two introduced colubrids. It was likely facilitated by the increasing local abundance of H. hippocrepis and by the ecological constraints of insular systems. Together with a previous report of avian predation on H. hippocrepis in Mallorca, it suggests that both native or naturalised predators may be incorporating this species into their diets. This finding adds new information on the trophic ecology of M. mauritanicus and highlights the dynamic nature of predator-prey interactions in island ecosystems, potentially involving juvenile individuals due to size-related constraints.

Understanding the factors shaping species distributions is essential for predicting their responses to environmental change. The genus Hemorrhois (horseshoe whip snakes) comprises ecologically diverse colubrid snakes found across the Mediterranean Basin, North Africa, the Middle East, and Central Asia. Despite this broad range, their ecological niches and distributional dynamics remain understudied. This study employs ecological niche modeling (ENM) to assess the biogeography, niche differentiation, and potential climate-driven range shifts of H. algirus, H. hippocrepis, H. nummifer, and H. ravergieri under future climate scenarios. Using species occurrence data and bioclimatic variables, I constructed ensemble models to predict suitable habitats, evaluate niche overlap, and quantify potential range changes. Results indicate significant variation in climate-driven distributional responses among species. Hemorrhois algirus is projected to expand across North Africa, whereas H. hippocrepis, H. nummifer, and H. ravergieri may face range contractions under high-emission scenarios. Niche analyses suggest moderate overlap between H. algirus and H. hippocrepis, implying historical and ecological connectivity, while H. nummifer and H. ravergieri display distinct environmental preferences. Climatic and geographic barriers—such as the Sahara Desert, the Dardanelles and Istanbul Straits, the Alps, and the Pyrenees Mountains—play crucial roles in shaping their evolutionary trajectories. Given the increasing threats of climate change and habitat loss, this study underscores the need for conservation strategies prioritizing habitat connectivity, species-specific management, and climate refugia. By integrating ecological and evolutionary perspectives, this research contributes to understanding Mediterranean and Western Palearctic reptile biogeography and their responses to environmental change.

The Dahl’s Whip Snake, Platyceps najadum is recorded from Gökçebey district, Zonguldak province (this record is the first case showing a locality of this snake species in this province) and Yenice district, Bartın province (it is the second record except for Eskipazar district) in the western Black Sea region of Türkiye. These results show that the distribution area of the species in the western Black Sea region of the country is wider than known. The morphometric characters and color-pattern features of one dead female (from Yenice) and one female specimen (from Gökçebey) were compared with specimens from the known Turkish localities according to the literature. We observed that the specimens from Yenice and Gökçebey were similar in some characters to the specimens of P. n. dahli reported in the literature.

Abstract Roads are vital for the economic development of countries but they pose major problems for wildlife. The road network in Central America is expanding, yet information about wildlife–vehicle collisions is scarce. We compiled data on vertebrate collisions with vehicles in Guatemala, El Salvador and Honduras, from projects created on the citizen science platform iNaturalist, to provide the first assessment of how these species are affected by roads in northern Central America. Our projects gathered 670 wildlife roadkill records that had been logged by 95 users across the three countries, with 122 species identified. Mammals and reptiles represented 44 and 30% of the records, respectively, with opossums Didelphis spp. and Philander vossi, the common boa Boa constrictor and the neotropical whip snake Masticophis mentovarius being the most frequently reported species (112, 28, 43 and 23 records, respectively). One of the species recorded is categorized as globally Endangered on the IUCN Red List, two as Vulnerable, four as Near Threatened and four have not been evaluated. Forty-six species are listed as Threatened or Endangered nationally. This study is the first roadkill assessment in northern Central America to which both members of the public and specialists contributed, underscoring the value of public engagement and citizen science. We urge further assessment of road impacts on wildlife in this region using standardized methods to identify roadkill rates and hotspots, and the implementation of mitigation measures for existing and planned roads in the region.

Snakes exhibit extraordinary ecological diversity and occupy a broad range of terrestrial, aquatic, and intermediate environments. Despite their widespread capacity for swimming, quantitative assessments of their aquatic locomotion remain scarce. Furthermore, the roles of intraspecific variation in shaping swimming performance and kinematics have been largely overlooked, hindering robust interspecific comparisons. This study systematically investigated intraspecific and interspecific variation in 287 individuals representing seven snake species along a terrestrial to aquatic continuum. Nine conventional swimming traits were quantified, including swimming speed and undulation characteristics such as frequency, wavelength, lateral velocities (head, body and tail) and amplitudes (head, body and tail). Undulation frequency and lateral velocity exhibited strong positive correlations with swimming speed, whereas wavelength showed a strong negative correlation. Semi-aquatic taxa attained the highest swimming speeds, while fully aquatic snakes displayed the lowest speeds. Intraspecific variation in swimming performance and kinematics was moderately reduced in aquatic species. Morphological traits, sex, and reproductive status significantly influenced both speed and kinematic profiles within species. Principal component analyses further revealed distinct kinematic domains among certain species (e.g., no overlap between aspic viper and sea snakes), while others, such as the green whip snake, exhibited broad overlap with all taxa examined. These findings demonstrate that ecological specialization to aquatic habitats does not unilaterally dictate swimming speed or kinematic patterns. Moreover, conventional kinematic parameters alone are insufficient to study the evolutionary trajectories of aquatic locomotion in snakes. Integrating hydrodynamic, endurance capacity, and diving performance will be essential to better understand how natural selection has shaped locomotion in aquatic snakes.

The Western whip snake (Hierophis viridiflavus) is a colubroid snake typically considered aglyphous and non-venomous, although this is currently a subject of debate. Indeed, the maxillary dentition of this species has recently been described as opisthoglyphous due to the presence of enlarged rear maxillary teeth, reportedly characterised by prominent grooves and ridges. Aiming to test the actual presence of these structures and provide a clearer characterisation of the maxillary dentition of the species, we analysed 17 dry-prepared and disarticulated H. viridiflavus specimens using both scanning electron and optical microscopy. The imaging confirmed the presence of posterior, enlarged maxillary teeth displaying a distal carina, highlighted by shallow longitudinal inflections on the disto-labial and disto-lingual tooth surfaces. Nonetheless, the analysis revealed the consistent absence of the meso-labially placed groove typical of opisthoglyphous taxa. Based on the results of our study, the maxillary dentition of the Western whip snake is heterodont, specifically opisthomegadont, and is confirmed to be aglyphous.

Anatomical analyses reveal the presence of two rear- fanged models among European colubroids, categorized by different types of venom glands and rear maxillary fangs and differences in the features of the maxillary bones. The first model, opisthoglyphous, is characterized by the presence of purely serous venom glands, from which the secretions produced flow in the grooved posterior maxillary fangs. The posterior fangs are separated from the anterior ones by an alveolar diastema and a deviation of the maxillary bone. The second model, blade-fanged, has a venom gland composed of seromucous cells, anatomically positioned in a more recessed position compared to the opisthoglyphous model. In this model the posterior maxillary fangs are enlarged but have distal and mesial keels instead of grooves, but are not separated from the anterior teeth by diastemas or deviations of the maxillary bone. The European group of colubroids, previously composed almost exclusively of snakes considered “aglyphous,” must be reconsidered in light of a system that consists mostly of opisthoglyphous snakes (predominantly the large group of Whip snakes, in addition to some species of Colubridae and Lamprophiidae), “aglyphous” snakes (exclusively Colubridae, including Ratsnakes and Dwarf snakes), and blade-fanged snakes (Natricidae, including Grass snakes and Water snakes). This reassessment is justified and supported by a comparative analysis of various European and non-European species, which confirms that “aglyphous” colubrids are residual forms of a rear-fanged model.

S nakes of the western Palearctic display a broad spectrum of phenotypic variation, sometimes mimicking other distinct species (Jablonski et al., 2017; 2023).This resemblance can lead to misidentification by local people (Beddek, 2017;Bakhouche & Escoriza, 2017).Hemorrhois hippocrepis (Linnaeus, 1758) has a predominantly west-Mediterranean distribution, spanning North Africa, the Iberian Peninsula, and the islands of Ibiza, Formentera, Sardinia, Pantelleria and Zembra where it was introduced (Mateo et al., 2011;Faraone et al., 2020).The dorsal colouration of H. hippocrepis is quite variable across its geographic range, where its ground colour can have diverse hues.A form of partial melanism, where the overall dorsal colour is blackish except for small light dots that surround the elements of the pattern, is fairly common in this species, especially in some of its eastern populations (Bruno & Hotz, 1976;Cattaneo, 1985).This dark morph appears to be purely ontogenetic, as it is exclusively present in adults, whereas juveniles are typically light coloured (Fig. 1) (Cattaneo, 1985;Feriche, 2017).Partially or totally melanistic juveniles are quite rare in H. hippocrepis (Garcia-Mars et al., 2015), as is the case in juveniles and adults in congeneric species (Tuniyev et al., 1997;Jablonski & Soran, 2023).Here, we describe for the first time a melanistic juvenile H. hippocrepis from its African geographic range.As part of an investigation into people's perception of snakes, we recovered the individuals killed by the local population in Algiers region.After examination, we identified 13 individuals of H. hippocrepis and six Natrix maura.In the same context, on January 2024, we found a black, small snake at the USTHB university campus in the locality of Bab Ezzouar (36 4247 N, 3 1045 E).The snake was collected for further examination, due to its unusual colouration.The observation took place in a habitat that was formerly a marshland but has since been heavily urbanised.Upon further morphological examination, the snake was determined as H. hippocrepis.The specimen had a snout-vent length of 298.1 mm and tail length of 76.9 mm.Consistent with our specific determination (Feriche, 2017) the specimen had the following head scalation characters (right side): 10 supralabials, 8 infralabials, 1 preocular, 2 postoculars, 3 suboculars, 1 loreal, 2 nasals, 3 anterior temporals and 3 posterior temporals.Based on its size and according to

Melanism, the process of heavier melanin deposition, can interact with climate variation at both micro and macro scales, ultimately influencing color evolution in organisms. While the ecological processes regulating melanin production in relation to climate have been extensively studied, intraspecific variations of melanism are seldom considered. Such scientific gap hampers our understanding of how species adapt to rapidly changing climates. For example, dark coloration may lead to higher heat absorption and be advantageous in cool climates, but also in hot environments as a UV or antimicrobial protection mechanism. To disentangle such opposing predictions, here we examined the effect of climate on shaping melanism variation in 150 barred grass snakes (Natrix helvetica) and 383 green whip snakes (Hierophis viridiflavus) across Italy. By utilizing melanistic morphs (charcoal and picturata in N. helvetica, charcoal and abundistic in H. viridiflavus) and compiling observations from 2002 to 2021, we predicted that charcoal morphs in H. viridiflavus would optimize heat absorption in cold environments, while offering protection from excessive UV radiation in N. helvetica within warm habitats; whereas picturata and abundistic morphs would thrive in humid environments, which naturally have a denser vegetation and wetter substrates producing darker ambient light, thus providing concealment advantages. While picturata and abundistic morphs did not align with our initial humidity expectations, the charcoal morph in N. helvetica is associated with UV environments, suggesting protection mechanisms against damaging solar radiation. H. viridiflavus is associated with high precipitations, which might offer antimicrobial protection. Overall, our results provide insights into the correlations between melanin-based color morphs and climate variables in snake populations. While suggestive of potential adaptive responses, future research should delve deeper into the underlying mechanisms regulating this relationship.

During two visits to the northeastern part of the Greek island of Lefkada, one in October 2022 and another in June 2023, 14 species of herpetofauna were recorded (two amphibian species, twelve reptile species), including one previously unrecorded species, the European cat snake Telescopus fallax. In addition, live specimens of Marginated tortoise Testudo marginata were found and location data for Dahl’s whip snake Platyceps najadum has been recorded. The observation localities of four species, Testudo hermanni, Testudo marginata, Platyceps najadum and Telescopus fallax are shown in an overview map.

Although extensive research has been conducted on snake venoms, the effects of bites inflicted by non-front-fanged colubroid snakes remain incompletely understood, particularly for species of uncertain medical relevance. The Western Whip Snake (Hierophis viridiflavus) is a colubrid snake typically classified as non-venomous and harmless to humans. Nevertheless, old works reporting the presence of Duvernoy's glands in this species raise questions regarding its presumed lack of venom. This report presents the first case of a prolonged bite from a wild Western Whip Snake (Hierophis viridiflavus carbonarius) that occurred in Italy, and provides a detailed account of the resulting effects. The primary symptom experienced by the bitten subject was painless, marked local oedema, which subsided within 24h after the bite. The clinical manifestations observed in the current study suggest that Hierophis viridiflavus could have the potential to inflict bites that lead to mild local effects consistent with envenoming.

The established methodology in detection and long-term reptile monitoring programmes has so far focused on visual encounter surveys (VES), a method that can yield very high capture rates for snakes, but, even when carried out in favourable habitats, can also be very limited. Therefore, one passive sampling method was tested: fibrocement slabs. A total of 81 slabs were placed in nine sampling stations in the southern part of Irún (Basque Autonomous Community, Spain). During the two years of sampling (2020–2021) a total of 79 reptile records were obtained, of which 33 were records of the European whip snake corresponding to 28 individuals. There was at least one reptile record in each transect and 75% of the reptile species present in the study area were detected. However, we did not identify any trend over time that indicated an increase in the number of individuals captured during the second year. All in, the data obtained have provided positive results; therefore, it could prove to be a very useful methodology for reptile studies, in particular for ophidians. Keywords: European whip snake, conservation, methodology, artificial refuge, capture, detection, Basque Autonomous Community, Spain

Abstract The skin of snakes acts as the first interface between these reptiles and the environment they live in. It plays an important role in the ecology of these organisms, and specialised structures, such as tactile corpuscles, are crucial in the sensing of a wide variety of stimuli. This study, by examining the presence, localisation and colocalisation of serotonin, substance P, PGP 9.5 and tubulin, wants to demonstrate the sensory role that these structures play in the ecology of the Green whip snake Hierophis viridiflavus (Lacépède, 1789). This study presents the first evaluation of nerve and immune molecules (TLR2) in the tactile corpuscles of H. viridiflavius ventral skin, indicating a potential defence role of these structures. These findings provide valuable information on the ecology of snakes, their defence systems, and their evolutionary morphostructural adaptations. Additionally, the identification of these corpuscles could provide valuable support for taxonomic recognition. Furthermore, this study expands the knowledge about stimuli sensing in snakes.

Reptiles, including snakes, can be asymptomatically infected with multiple pathogen microorganisms, including Salmonella spp., which is considered an important concern for public and animal health. Small and uninhabited isles are quite ecologically different from mainland and represent interesting fields of study, to discover unexpected biological and microbiological aspects of their wild inhabitants. This work reports the presence of the very rare Salmonella enterica serovar Yopougon, isolated in a carcass of a native wild snake (Hierophis viridiflavus) from an Italian uninhabited island of Mediterranean Sea, Montecristo. To our knowledge, S. enterica serovar Yopougon was previously isolated only once 34 years earlier in Ivory Coast, from a human fecal sample. In the present study, we present the genomic characterization of the new isolate, the phylogenetic comparison with the previously isolated S. enterica serovar Yopougon strain of human origin and with other sequences available in public databases. In addition, an extensive review of available data in the literature and from our case history is provided. Our finding represents an example of the ability of some pathogens to travel for very long distances within their hosts and then to infect others, even from different taxa.

Spatial and temporal patterns of road mortality in the Caspian whip snake (Dolichophis caspius Gmelin 1758) in Romania

Abstract Speciation is the fundamental process that underlies the diversification of species. Different steps of speciation, such as reproductive isolation or ecological niche separation, can evolve in different orders and over different time scales. The two subspecies of the green whip snake, Hierophis viridiflavus viridiflavus and Hierophis viridiflavus carbonarius, sometimes considered as separate species, are ideal to study speciation because they diverged in isolated glacial refugia. So far, little is known about whether these two subspecies are reproductively isolated from each other. Furthermore, it is unknown whether the common melanism of the subspecies H. v. carbonarius might be genetically encoded. We quantified gene flow along two transects across the contact zone of the two subspecies using double-digest restriction site-associated DNA. Our analyses implicated the existence of two well-defined clades that experience substantial gene flow along a contact zone that is ~300 km wide. We detected a clear association between the genetic assignment and the quantitative extent of melanism within the contact zone, which is likely to be genetically polygenic. Given the level of gene flow, it seems that strong postzygotic reproductive barriers have not evolved. We conclude that the two subspecies represent evolutionarily significant units for conservation and suggest treating them taxonomically as subspecies.

Phylogeographic and paleoclimatic modelling studies have been combined to infer the role of Pleistocene climatic oscillations as drivers of the genetic structure and distribution of Mediterranean taxa. For the European whip snake, Hierophis viridiflavus, previous studies based on paleoclimatic modelling have depicted a low reliability in the pattern of past climatic suitability across the central Mediterranean Basin, which barely fits the species' genetic structure. In this study, we combined phylogeographic and paleoclimatic modelling tools to improve our understanding of the biogeographic history of H. viridiflavus, particularly extending the sampling and phylogeographic inferences to previously under-sampled regions. Phylogeographic analyses recovered two major clades that diverged at the beginning of the Pleistocene and had diversified in different ways by the late Pleistocene: the east clade (composed of three subclades) and the west clade (with no further structure). Paleoclimatic models highlighted the temperate character of H. viridiflavus, indicating range contractions during both the last inter-glacial and last glacial maximum periods. Range expansions from southern-located climatic refugia likely occurred in the Bølling-Allerød and Middle Holocene periods, which are supported by signals of demographic growth in the west clade and South-East-North subclade. Overall, this work improves our understanding of the historical biogeography of H. viridiflavus, providing further insights into the evolutionary processes that occurred in the Mediterranean Basin hotspot.

Seventeen predation attempts on four Hoopoe Upupa epops nests in an heterogenous agricultural Mediterranean area of central Italy were recorded during breeding seasons 2020-21. Camera traps identified 7 different predators’ species, mainly Western whip snake Hierophis viridiflavus and Black rat Rattus rattus. Uropygial secretions and hissing calls from chicks and incubating female inside the breeding cavity always saved the clutches, while no active defense was performed by parents if outside the nest.

Further evidence of ophiophagy, including cannibalism, by the western whip snake Hierophis viridiflavus in Italy