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Alpine salamander

The Alpine salamander (Salamandra atra) is a glossy black, lungless amphibian species renowned for its viviparous reproduction and fully terrestrial life cycle, inhabiting high-altitude humid meadows, woodlands, and rocky crevices above 700 meters in the central, eastern, and Dinaric Alps spanning Austria, Italy, Slovenia, and Croatia.^[1]^[2] This species exhibits a predominantly nocturnal foraging behavior, preying on small invertebrates such as ants, beetles, and earthworms while sheltering diurnally in burrows or under stones to avoid desiccation and predation.^[1]^[3] Unique among most amphibians, S. atra females retain fertilized eggs internally for 2–3 years before giving birth to live, fully metamorphosed juveniles, a reproductive strategy adapted to the cold, aquatic-scarce alpine environment that enhances offspring survival rates.^[4]^[5] The species secretes potent toxic alkaloids from granular glands in its skin, providing chemical defense against predators including birds, mammals, and snakes.^[1] Females typically reproduce every 3–4 years in a promiscuous mating system observed during late spring and summer, with males exhibiting higher dispersal tendencies than philopatric females.^[5] Classified as Least Concern by the IUCN due to its wide distribution and stable populations, S. atra faces localized threats from habitat fragmentation via tourism development, agricultural expansion, and potential climate-driven shifts in suitable elevational ranges, though its adaptability and toxicity confer resilience.^[6]^[7] Certain subspecies, such as the golden S. a. aurorae, remain vulnerable due to restricted ranges and collection pressures, underscoring the need for targeted monitoring amid emerging disease risks like the salamander plague.^[8]^[9]

Taxonomy and phylogeny

Classification and subspecies

The Alpine salamander (Salamandra atra Laurenti, 1768) is classified in the order Urodela, family Salamandridae, subfamily Salamandrinae, and genus Salamandra.^[10]^[1] The species is native to high-altitude regions of the Alps and Dinaric Mountains, with the nominotypical form exhibiting fully melanistic black coloration as an adaptation to alpine environments.^[1] Four subspecies are currently recognized, though the validity of at least one (S. a. prenjensis) has been questioned in some taxonomic assessments due to limited genetic and morphological data.^[1] These subspecies differ primarily in coloration patterns and restricted distributions, reflecting isolation in montane refugia.

S. a. atra: The most widespread subspecies, uniformly black without dorsal markings, occurring from the central Alps (e.g., Switzerland, Austria) eastward to the Dinaric Alps (e.g., Slovenia, Croatia).^[1]^[10]
S. a. aurorae (golden alpine salamander): Distinguished by bright yellow, whitish, greenish, or gray dorsal spots on the head, back, and limbs; confined to a small area (<50 km²) in northeast Italy (Trento-Asiago plateau, 1,300–1,800 m elevation), where it faces high extinction risk from habitat fragmentation.^[1]
S. a. prenjensis: Similar to S. a. atra in dark coloration but potentially distinct in subtle morphological traits; endemic to the Prenj and Čvrsnica mountains near Sarajevo, Bosnia and Herzegovina, with discontinuous distribution from other forms and ongoing debate over its taxonomic status.^[1]^[11]
S. a. pasubiensis: Black with possible minor variations, restricted to the Pasubio massif in the southeastern Italian Prealps.^[10]

Evolutionary history and adaptations

The genus Salamandra diversified in Europe during the late Miocene to Pliocene, with S. atra forming a distinct Alpine-Dinaric clade characterized by terrestrial viviparity and melanism.^[12] Phylogeographic analyses of mitochondrial DNA reveal that S. atra populations experienced divergence influenced by Pleistocene glacial-interglacial cycles, with ancestral lineages retreating to multiple southern refugia, including the Dinaric Alps (e.g., Prenj mountain) and Apennine regions, before post-glacial northward expansion into the Alps.^[13]^[14] Molecular divergence estimates for major mtDNA haplotypes predate the Pleistocene, occurring around 3 million years ago, indicating deep phylogenetic structure predating major Quaternary glaciations.^[15] This history has resulted in genetically differentiated lineages, such as those in the Venetian Prealps and Dinarides, some recognized as subspecies with varying pigmentation (e.g., golden forms in S. a. aurorae).^[16]^[17] Viviparity in S. atra represents an independent evolutionary transition within Salamandra, shifting from ovoviviparity or larviparity in congeners to fully terrestrial parturition of metamorphosed young, likely driven by selective pressures in high-elevation habitats where aquatic larval stages are inviable due to prolonged freezing and short ice-free periods.^[18]^[19] Gestation spans 2–3 years, with intrauterine nourishment via epitheliophagy (embryos consuming uterine tissue), enabling offspring survival in nutrient-poor, cold environments without external water dependence.^[20] Complete melanism, prevalent in nominate S. a. atra, evolved convergently or retentively across lineages, correlating with alpine rock substrates for crypsis and potentially enhanced UV protection or thermoregulation via increased solar heat absorption in low-temperature regimes up to 2,800 m elevation.^[21]^[16] Physiological adaptations include depressed metabolic rates and extended lifespans (up to 20+ years), minimizing energy demands in oligotrophic, high-altitude niches with scarce prey and harsh winters.^[22]^[23]

Physical description

Morphology and size variation

The Alpine salamander (Salamandra atra) possesses a robust, elongated body with short, well-developed limbs suited to navigating rocky alpine terrains. The head is slightly elongated and broader than long, featuring prominent kidney-shaped parotoid glands posterior to the eyes, which serve as defensive structures containing toxic alkaloids. The skin is smooth, moist, and typically glossy black, lacking scales or osteoderms common in some other salamanders. Flanks bear 11–13 distinct costal grooves, and the tail comprises approximately 40–50% of total length, ending in a short, bluntly rounded fin that originates near the pelvic girdle.^[1]^[24] Adult body size ranges from 80 to 150 mm in total length, with notable sexual dimorphism: females attain a maximum of 151 mm, exceeding males at 144 mm, reflecting differences in reproductive investment and growth rates.^[1] Intraspecific size variation is evident across geographic ranges and elevations; higher-altitude populations, such as those in the southern Alps, exhibit reduced adult sizes and earlier maturity, potentially as an adaptation to shorter active seasons and resource limitations.^[25] Subspecies display morphometric distinctions, including variations in head dimensions, limb proportions, and overall robustness; for example, analyses of 14 traits from central European and Dinaric populations confirm significant differences supporting taxonomic separation.^[26] These patterns align with environmental gradients, where body size correlates inversely with elevation in some lineages, though genetic and phenotypic plasticity both contribute.^[27]

Coloration and melanism

The Alpine salamander (Salamandra atra) displays a predominantly melanistic coloration, characterized by a uniform black appearance resulting from high concentrations of melanin pigments in the skin.^[21] This fully melanistic form is typical of the nominate subspecies S. a. atra, which inhabits central, eastern, and Dinaric Alpine regions.^[21] Coloration varies across subspecies, with some exhibiting yellow or golden patches on the dorsal surface, head, and limbs. The subspecies S. a. aurorae is invariably yellow-patched, while S. a. pasubiensis shows intermediate variability, ranging from limited yellow patches to complete melanism.^[21] These yellow markings, when present, contrast sharply with the black ground color and are thought to represent a derived aposematic pattern in ancestral lineages, though reduced or absent in higher-altitude, fully melanistic forms.^[21] Phylogenetic reconstruction based on mitochondrial DNA reveals that the yellow-patched condition is the plesiomorphic (ancestral) state within S. atra, with evolutionary transitions to melanism occurring through progressive reduction in yellow patch size.^[21] This shift to full melanism in S. a. atra likely predates the last glacial period, potentially driven by genetic drift or natural selection associated with Pleistocene climatic oscillations, though the precise adaptive mechanisms remain unclear.^[21] Full melanism has arisen independently in related lineages, such as Salamandra lanzai, underscoring convergent evolution in color loss within alpine environments.^[21] The black phenotype may confer crypsis against dark rocky substrates in high-elevation habitats, contrasting with the warning function of yellow-black patterns in lowland fire salamanders.^[28]

Distribution and habitat

Geographic range

The Alpine salamander (Salamandra atra) inhabits high-elevation regions across central and southern Europe, with its core distribution spanning the Alps from southeastern France through Switzerland, Austria, southern Bavaria in Germany, and northern and northeastern Italy to Slovenia.^[10] Isolated populations extend southward into the Dinaric Alps and adjacent highlands of Croatia, Bosnia and Herzegovina, and Montenegro.^[10] This range reflects adaptation to montane environments, with the species absent from lower valleys and plains.^[1] Subspecies exhibit varying degrees of geographic restriction within this overall distribution; for instance, the nominate subspecies S. a. atra occupies much of the central and eastern Alps, while S. a. aurorae is confined to a small area (less than 50 km²) between Trento and Asiago in northeastern Italy.^[1] Other subspecies, such as S. a. prenjensis, are limited to specific massifs like Mount Prenj in Bosnia and Herzegovina.^[10] These localized distributions highlight the species' patchy occurrence, influenced by historical glaciation and habitat fragmentation.^[29]

Habitat requirements and microhabitats

The Alpine salamander (Salamandra atra) requires cool, moist habitats characteristic of subalpine and alpine zones, with a typical elevational range of 600 to 2400 meters above sea level, though extremes extend from 430 to 2800 meters.^[30] These salamanders thrive in environments providing high humidity and stable, low temperatures, often in meadows, woodlands, and forested landscapes where precipitation and shade prevent desiccation.^[30] Habitat suitability is influenced by landscape elements such as forests and alpine pastures above 1000 meters, with slower adaptation to changes suggesting persistence in marginally suitable areas.^[31] Microhabitats selected by S. atra emphasize refugia that buffer against aridity and temperature fluctuations, including the upper soil layer, under stones or logs, and within rocky crevices.^[32] In forested settings, individuals show preference for areas dominated by silver fir (Abies alba), European beech (Fagus sylvatica), and grassy clearings over dense thickets, facilitating access to epigean (surface) activity zones.^[32] Soil moisture and pH play critical roles, with salamanders favoring damp substrates that support their fully terrestrial, lungless respiration and prevent dehydration, as evidenced by comparative studies on related plethodontid species adapted to similar conditions.^[22] These microhabitat choices underscore the species' limited dispersal and vulnerability to fragmentation from drying trends or intensive land use.^[30]

Behavior and ecology

Territoriality and movement patterns

Alpine salamanders (Salamandra atra) display territorial behavior primarily through chemical signaling and occasional aggressive encounters. Individuals deposit faecal pellets under cover objects within their home ranges, serving as low-cost scent markers to advertise presence and deter intruders, as observed in populations at high altitudes in France.^[33] Male-male interactions often involve agonistic displays such as chasing, mounting attempts, and chin rubbing, which may function in territorial defense or mate competition, with reports from subspecies like S. a. atra and S. a. aurorae indicating these behaviors occur during nocturnal activity periods.^[34] ^[35] Movement patterns are characterized by limited dispersal and high site fidelity in adults, who maintain home ranges typically spanning a few hundred square meters, adapting to stable alpine microhabitats with rock crevices and forest litter.^[30] Adults exhibit nocturnal foraging excursions covering 4–22 meters per night, rarely venturing far from shelters, which supports territorial maintenance in resource-scarce environments. Juveniles, in contrast, show greater mobility, dispersing primarily on the ground surface to establish new territories, while ontogenetic shifts lead adults to restrict movements to confined areas.^[36] Dispersal exhibits sexual dimorphism, with genetic analyses revealing female philopatry—remaining near natal sites—and higher male dispersal rates, likely driven by promiscuous mating strategies that favor mate-searching in males across fragmented habitats.^[5] This pattern contributes to gene flow primarily through males, as evidenced by assignment tests in Swiss populations, though overall population connectivity remains low due to the species' alpine endemism and topographic barriers.

Foraging and diet

The Alpine salamander (Salamandra atra) is a nocturnal forager, active primarily during humid nights to exploit available terrestrial invertebrates while minimizing desiccation risk in its high-elevation habitat. Stomach content analyses indicate low feeding rates, with an average of 2.78–3.98 prey items per individual across studies, suggesting selective or infrequent foraging rather than continuous consumption. This pattern aligns with a sit-and-wait or opportunistic ambush strategy, where salamanders target slow-moving or accessible prey in moist microhabitats such as under rocks, logs, and leaf litter.^[37]^[3] The diet consists predominantly of invertebrates, reflecting a generalist trophic strategy at the population level but with notable selectivity for specific taxa that offer higher nutritional value or ease of capture, such as those with larger size, lower mobility, or softer exoskeletons. In S. a. atra, dominant prey include dipteran larvae (38 items recorded in one sample), mollusks (19 items), chilopods (17 items), coleopterans (13 items), and arachnids (12 items), comprising 139 analyzable items from 45 individuals sampled in midsummer. Subspecies like S. a. aurorae show similar composition, emphasizing myriapods, non-ant hymenopterans, gastropods, and coleopterans, with 199 prey items across 53 individuals positively selected over more abundant environmental options like collembolans and ants. DNA metabarcoding of stomach flushes has further revealed an exceptionally diverse diet, underscoring breadth beyond traditional morphological identification.^[37]^[3]^[38] Prey selection indices demonstrate positive bias toward nocturnal-available taxa like chilopods and coleopterans, while rejecting ubiquitous but less profitable items such as formicids and collembolans, despite their higher environmental abundance. High inter-individual diet variation (e.g., modularity Q=0.47, variation E=0.86–0.87) indicates potential individual specialization within the generalist framework, possibly driven by microhabitat differences or experience-based learning. Dietary composition varies modestly among populations and subspecies, but consistently excludes vertebrates or plant matter, confirming an exclusively carnivorous, invertebrate-based niche adapted to alpine prey scarcity.^[37]^[3]^[39]

Predation and defensive strategies

The Alpine salamander (Salamandra atra) faces predation primarily from avian species such as tawny owls (Strix aluco), mammalian carnivores including stone martens (Martes foina) and red foxes (Vulpes vulpes), and reptiles like the common European viper (Vipera berus), though successful attacks are infrequent due to the species' high-altitude distribution and defensive adaptations.^[40] Predation pressure from snakes appears to exert stronger selective influence on toxin profiles compared to mammals or birds, as evidenced by comparative analyses of steroidal alkaloid (SAM) composition across populations.^[40] The principal defensive mechanism involves cutaneous toxins secreted from parotoid and granular glands, comprising samandarines and related steroidal alkaloids that act as neurotoxins, deterring predators through unpalatability and potential lethality upon ingestion.^[41] These compounds provide dual protection against predation and microbial infections, with population-level variation in toxin profiles—such as higher SAM concentrations in areas with elevated snake presence—suggesting adaptive responses to local ecological pressures rather than uniform aposematic signaling, given the species' uniform black melanism.^[42] Unlike the congeneric fire salamander (S. salamandra), S. atra lacks documented projectile spraying of secretions, relying instead on passive release during handling or attack.^[40] Behavioral defenses include crypsis facilitated by dark coloration blending with alpine rock crevices and nocturnal activity patterns that minimize encounters, supplemented by occasional tail-powered jumps for evasion when threatened.^[43] Tail autotomy, a common urodeles trait, allows escape from grasping predators, though regeneration in S. atra is slow and energetically costly in its oligotrophic habitat.^[44] Overall, the integration of chemical and cryptic strategies renders S. atra highly resistant to predation, contributing to its persistence in predator-scarce montane ecosystems.^[41]

Reproduction

Mating systems and interactions

The Alpine salamander (Salamandra atra) exhibits a promiscuous mating system characterized by multiple partners for both sexes, with internal fertilization occurring terrestrially without reliance on aquatic environments.^[5] Mating activity peaks in late spring to summer, specifically May to June at lower altitudes (around 650 m above sea level) and June to July at higher elevations (around 1700 m), though observations can extend to the end of the active season.^[45] Courtship involves the male grasping the female in a ventral amplexus, followed by the deposition of one or more gelatinous spermatophores on the substrate, which the female collects using her cloaca to achieve sperm transfer—a behavior unique among salamandrids to the genus Salamandra.^[46]^[47] Male-male interactions during the mating period can include aggressive encounters, as documented in the golden Alpine salamander subspecies (S. a. aurorae), where males engage in prolonged physical contests involving attempts to mount each other and chin-rubbing, lasting up to four minutes.^[48] Such aggression likely serves to establish dominance for access to females in this low-density, territorial species. Females demonstrate philopatry, remaining near natal sites, while males exhibit greater dispersal, potentially facilitating gene flow in the promiscuous system.^[5] Reproduction is infrequent, with females typically breeding every three to four years due to the energetic demands of viviparity.^[5]

Gestation, viviparity, and offspring development

The Alpine salamander (Salamandra atra) exhibits pueriparity, a form of viviparity in which females give birth to fully metamorphosed, terrestrial juveniles rather than larvae or eggs.^[1] This reproductive mode involves internal fertilization followed by extended intrauterine development, with typically one embryo developing per oviduct, resulting in an average litter size of two offspring.^[8] Rarely, litters of three or four have been documented, though such cases may reflect developmental anomalies rather than normative fecundity.^[49] Gestation duration varies with elevation and climatic conditions, lasting 2 years at altitudes of 650–1000 m and extending to 3 years at 1400–1700 m, with records up to 4 years in higher, colder environments.^[1]^[46]^[50] Mating occurs primarily in late spring or early summer, after which sperm storage enables fertilization; females typically mate every 2–4 years, aligning with the protracted gestation cycle.^[45] Embryonic development proceeds in two nutritional phases post-hatching within the uterus: initial oophagy, where embryos consume unfertilized eggs or siblings, followed by epitheliophagy, involving ingestion of maternal uterine epithelial cells for sustained matrotrophic nourishment beyond yolk reserves.^[51]^[20] This process ensures complete metamorphosis in utero, bypassing an aquatic larval stage and adapting offspring for immediate terrestrial independence. At birth, juveniles measure 40–50 mm in total length, with no significant correlation between maternal body size and offspring dimensions observed in field studies.^[1]^[46] Birth occurs on land, often in moist microhabitats, supporting the species' fully terrestrial lifecycle.^[8]

Physiology

Skin glands and chemical defenses

The skin of the Salamandra atra features granular glands distributed across the body, with prominent parotoid glands—clusters of these granular glands situated posterodorsally to the eyes—serving as primary sites for toxin production and storage, alongside mucus glands that facilitate secretion and maintain dermal integrity.^[52]^[1] These granular glands synthesize steroidal alkaloids collectively termed samandarines, including samandarine as the dominant compound and up to nine identified derivatives such as samandarone, which are released upon mechanical stimulation or stress.^[53]^[52] The parotoid glands of an individual S. atra contain approximately 5 mg of samandarine, a lower quantity compared to the 20 mg in the related fire salamander (S. salamandra), reflecting adaptations to its high-altitude, low-predation environment.^[52] When threatened, the salamander orients toward the predator and adjusts its posture—often lowering the head—to emphasize the parotoid glands, triggering expulsion of viscous, milky secretions that adhere to attackers and cause mucous membrane irritation, neurotoxic effects, or lethality upon ingestion, thereby deterring predation.^[54] These alkaloids also possess antimicrobial properties, inhibiting bacterial and fungal growth on the skin and potentially conferring resistance to pathogens like Batrachochytrium dendrobatidis.^[55] Geographic variation in toxin profiles occurs among S. atra populations, with differences in samandarine composition and poison gland sizes observed across sites in the Dinaric Alps, though total toxin quantities do not differ significantly; such variability may arise from genetic or dietary factors rather than predation pressure or infection risk, influencing localized defense efficacy.^[56] Empirical assays confirm that heavier individuals produce higher absolute amounts of these alkaloids, correlating with larger parotoid glands (F<sub>1,124</sub>=62.92, p<0.001).^[40]

Immunological adaptations and disease resistance

The Alpine salamander (Salamandra atra) demonstrates resistance to chytridiomycosis caused by Batrachochytrium dendrobatidis (Bd), with no infections detected in surveys of 92 individuals across 11 populations in the Alps.^[57] This pathogen, which disrupts amphibian skin electrolyte balance leading to cardiac arrest, relies on aquatic zoospores for transmission, and S. atra's fully terrestrial, viviparous life cycle—lacking free-living larval stages and involving minimal water immersion—severely limits exposure risk.^[57] While innate skin defenses, such as antimicrobial peptides common in salamandrids, could provide supplementary protection, empirical confirmation specific to S. atra remains absent. Surveys for Batrachochytrium salamandrivorans (Bsal), the causative agent of salamander chytridiomycosis or "salamander plague," have similarly yielded no positive detections in 818 S. atra and closely related S. lanzai individuals from 40 Alpine sites between 2017 and 2022.^[9] Experimental infections indicate high susceptibility within the Salamandra genus, with rapid mortality from skin erosion and systemic invasion, yet wild populations persist uninfected, likely due to geographic isolation at high elevations (above 700 m) where cooler, drier microhabitats inhibit fungal viability and dispersal.^[9] ^[1] Habitat suitability models predict potential Bsal establishment under climate warming, underscoring vulnerability despite current absence.^[9] Mortality events in vulnerable subspecies, such as S. atra prenjensis in the Dinarides, show no association with major amphibian pathogens including ranaviruses, Bd, or Bsal, as qPCR and histopathology on deceased specimens tested negative.^[58] Ranaviruses, which cause hemorrhagic disease with high lethality in other urodeles, appear undetected in S. atra monitoring efforts.^[59] This pattern suggests effective barriers to pathogen incursion, potentially bolstered by low population densities and limited inter-population gene flow that curtail epidemic spread.^[30] Specific immunological adaptations in S. atra are underexplored, but as a poikilotherm adapted to subzero temperatures and hypoxia at altitudes exceeding 2,000 m, its innate immune components—such as antimicrobial peptides and lysozymes in skin mucus—likely function efficiently under cold stress, mirroring patterns in related salamandrids that mount cellular and humoral responses to allografts.^[60] Viviparity may facilitate maternal antibody transfer to offspring, enhancing neonatal resistance during prolonged uterine gestation (up to 3 years), though direct evidence is lacking.^[1] Overall, pathogen exclusion stems more from ecological and behavioral traits than documented immune novelty, with ongoing threats from pathogen spillover necessitating vigilant surveillance.^[9]

Conservation and threats

Current status and population dynamics

The Alpine salamander (Salamandra atra) is assessed as Least Concern on the IUCN Red List, reflecting its relatively wide distribution across high-altitude regions of the central and eastern Alps, including Austria, Switzerland, Italy, Slovenia, and parts of Germany, France, and Liechtenstein, typically above 700 meters elevation.^[1] However, the global population trend is decreasing, driven by habitat fragmentation, climate-induced shifts in suitable microhabitats, and potential vulnerability to emerging pathogens, though no widespread declines have been documented in core populations. Subspecies such as S. a. aurorae in Italy have been considered highly endangered due to restricted ranges and low densities. Population dynamics are characterized by slow growth and low fecundity, with females producing 2–50 live young after 2–3 years of gestation, leading to limited recruitment and sensitivity to mortality events.^[1] Monitoring in the Alps indicates stable abundances in protected forested and rocky habitats, but isolated peripheral populations, such as those in the Dinaric Alps or Montenegro's Mt. Orjen, exhibit genetic bottlenecks and heightened extinction risk from stochastic events.^[61] Recent surveys confirm absence of the fungal pathogen Batrachochytrium salamandrivorans (Bsal) in Alpine populations as of 2024, averting immediate mass die-offs observed elsewhere in Europe, though modeling predicts up to 30% of urodelan species at high local extinction risk from Bsal by 2030.^[62] ^[9] Climate change poses a long-term threat, with projections indicating reduced climatic suitability by 2070 due to warming and altered precipitation, potentially contracting habitats to suboptimal higher elevations and exacerbating isolation.^[7] Quantitative population estimates remain scarce, as the species' nocturnal and crevice-dwelling habits hinder comprehensive censuses, but localized studies report densities of 0.1–1 individual per square meter in optimal sites.^[22] Conservation efforts emphasize habitat connectivity and disease surveillance to mitigate these pressures, with legal protections in countries like Austria and Switzerland supporting persistence.^[1]

Anthropogenic impacts and mitigation

Habitat alteration from infrastructure development, including road construction and ski slope preparation, poses localized threats to Salamandra atra populations by fragmenting rocky and forested refuges essential for their terrestrial lifecycle.^[22]^[63] Ground leveling and clearcutting associated with these activities disrupt microhabitats, reducing shelter availability and increasing exposure to desiccation and predators.^[22] In the Venetian Alps, deforestation for logging has acutely endangered the subspecies S. a. aurorae, prompting calls for immediate forest protection in areas like Altopiano dei Sette Comuni.^[64] Road mortality affects certain subpopulations, particularly in Switzerland, where vehicle traffic intersects migration or foraging paths, leading to direct fatalities.^[8]^[65] Forestry practices contribute to habitat loss through fragmentation, though mature forests with retained woody debris and riparian buffers support higher salamander densities by preserving moisture and cover.^[66] Climate change, driven by anthropogenic greenhouse gas emissions, is projected to render much of the current high-altitude range unsuitable by 2070, shifting suitable niches upslope and compressing habitats against topographic limits.^[7] Emerging pathogens like Batrachochytrium salamandrivorans (Bsal), potentially spread via human-mediated trade in amphibians, represent a further risk, though peripheral populations remain uninfected as of 2025 assessments.^[9]^[67] Mitigation efforts emphasize legal safeguards and habitat management. S. atra is protected under Appendix II of the Bern Convention, with subspecies S. a. aurorae additionally listed on Annex II of the EU Habitats Directive, mandating conservation measures like habitat restoration and development restrictions.^[65] National laws in Austria and Switzerland prohibit collection and enforce habitat protections.^[1] Forestry guidelines promote selective logging to maintain structural complexity, including retention of refuges and avoidance of clearcuts in core ranges.^[66] Targeted interventions, such as road underpasses or fencing in high-mortality zones, could reduce vehicle impacts, though implementation remains limited.^[8] For climate-vulnerable areas, predictive modeling informs priority conservation zones, prioritizing refugia with stable microclimates while minimizing overlap with co-occurring species.^[7] Public awareness campaigns advocate reduced tourism disturbance and pollution controls to curb indirect effects like altered hydrology from uphill development.^[43] Despite these, ongoing monitoring is essential given the species' overall Least Concern status but localized declines.^[1]

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Salamandra - an overview | ScienceDirect Topics
A total of 10–11 costal grooves. Tail equal in length to SVL or slightly smaller. Tail fin short, originating anterior to the pelvic girdle, with a bluntly ...<|separator|>
[25]
Variations in age, size at maturity and gestation duration among two ...
Feb 28, 2006 · Variations in age, size at maturity and gestation duration among two neighbouring populations of the alpine salamander (Salamandra lanzai). C.
[26]
Morphometric characteristics of Alpine salamanders: A support for ...
Aug 5, 2025 · The present study investigates 14 morphological traits of Alpine salamanders originating from Austria and from Bosnia and Herzegovina (B&H).Missing: sources | Show results with:sources
[27]
Morphological variation in salamanders and their potential response ...
Across 33 species of salamanders from Europe and Asia, the variation of body size was strongly related to the number of vertebrae (PGLS: F1,31 = 33.7, B ± SE = ...
[28]
Alpine salamander (Salamandra atra) - Picture Nature
The color of alpine salamander is deep black throughout, providing excellent camouflage in its rocky habitat. The species doesn't have any unique markings ...Missing: coloration | Show results with:coloration
[29]
Distribution map of Salamandra atra (dark gray area, based on ...
1). The species is mostly found between 800 and 2000 m a.s.l., on alpine rocky meadows and pastures, in or at the edges of open deciduous, mixed or coniferous ...
[30]
Conservation Genetics of an Isolated Alpine Salamander Population
The Alpine salamander (Salamandra atra) is a mountain-dwelling species with limited dispersal and a preference for cool, moist habitats.
[31]
[PDF] Ecological requirements of the Alpine Salamander Salamandra atra
These landscape patterns consist of the variables forest, alpine pasture (above 1000 m. a. s. l., grazing only seasonal, no mowing), agricultural area (below ...Missing: microhabitats | Show results with:microhabitats
[32]
(PDF) Epigean habitat of a population of Salamandra atra aurorae
Adults were found in the upper layer of the soil more frequently in areas with Abies alba, Fagus sylvatica and grassy clearings than in thicker forest areas ...Missing: microhabitats | Show results with:microhabitats
[33]
Faecal pellets used as an economic territorial marker in two ... - jstor
a population at 2,300 m a.s.l. (44° 42' n, 7° 03' e; France). The alpine salamander, Salamandra atra, is also a viviparous species, endemic to the alpine arc.
[34]
Male-male interactions in Alpine Salamanders, Salamandra atra ...
Aug 5, 2022 · ... territoriality by. salamanders and a comparison with the territorial behavior of ... Alpine salamander Salamandra atra aurorae (Caudata, ...
[35]
First record of a male-male aggressive interaction in the golden ...
Apr 20, 2023 · We describe an aggressive interaction between two males which fought for four minutes, trying to go on top of each other and rubbing their chin ...
[36]
Movements, distribution pattern and density in a population of ...
Salamanders changed their behaviour with growth: while juveniles usually dispersed on the ground, adults showed a strong fidelity to a limited area, a few ...
[37]
A Midsummer Night's Diet: Snapshot on Trophic Strategy of the ...
May 17, 2020 · Small number of preys per stomach could indicate a dimensional selectivity; for a large species such as Salamandra atra, foraging on few but ...
[38]
DNA metabarcoding analysis of stomach flushing contents reveals ...
Sep 30, 2025 · ... feeding behavior and diet preferences ... A midsummer night's diet: snapshot on trophic strategy of the Alpine salamander Salamandra atra.
[39]
(PDF) Trophic ecology of alpine salamander, Salamandra atra atra ...
Sep 16, 2019 · Considering that Salamandra atra is a typical nocturnal species, we collected food availability (with pitfalls traps) separately for diurnal and ...
[40]
[PDF] Toxin variation among salamander populations - UA-repository.
There are several reasons why predation by snakes is likely a stronger selective pressure for. 261. SAM composition in S. atra than predation by mammals or ...<|separator|>
[41]
Toxin variation among salamander populations: discussing potential ...
Amphibians produce defensive chemicals which provide protection against both predators and infections. Within species, populations can differ considerably ...
[42]
Toxin variation among salamander populations: discussing potential ...
Sep 23, 2020 · Amphibians produce defensive chemicals which provide protection against both predators and infections. Within species, populations can ...
[43]
salamandraatra.org
The alpine salamander, scientifically known as Salamandra atra, stands as an enchanting example of high-altitude adaptation. Resilient and distinctive ...
[44]
Anti-Predator Mechanisms in Salamanders – Kari Yoshida - UO Blogs
Salamanders use toxic skin secretions, warning colors, immobility, biting, tail autotomy, flipping, warning postures, tail lashing, and vocalization as anti- ...<|control11|><|separator|>
[45]
8 Reproductive cycle of Salamandra atra. Mating peaks in May and ...
Mating peaks in May and June at 650 m a.s.l. and in June and July at 1700 m a.s.l. Matings can be observed, however, until the end of the season. Fertilization ...Missing: life | Show results with:life
[46]
[PDF] The Alpine Salamander - Zobodat
Here, we will focus on the entire distribution of S. atra to learn more about its evolutionary history, and highlight different lineages in a perspective of ...
[47]
[PDF] Ethology, 74, 133–145 (1987) - Stevan J. Arnold
the male usually deposited three or four spermatophores in a courtship encounter. Salamandra atra: The courtships of S. atra and S. salamandra are very.
[48]
[PDF] First record of a male-male aggressive interaction in the golden ...
Apr 20, 2023 · It can now be confirmed that aggressive male-male interactions are present in the golden Alpine salamander. Considering the rarity of in-.
[49]
(PDF) Body size, population structure and fecundity traits of a ...
... Salamandra atra atra (Amphibia, Urodela ... gestation, were named 'pregnant females ... 4 mm). Pregnancy and offspring parameters. Throughout the fiel ...
[50]
Salamandra atra Laurenti, 1768 - Alpensalamander. | Request PDF
Salamandra atra Laurenti, 1768 - Alpensalamander. January 2004 ... pregnancy every 2-4 years (Vilter & Vilter, 1960). Gestation duration of up to four years ...
[51]
intrauterine cell nourishment in the viviparous alpine salamander ...
The intrauterine nourishment of the viviparous alpine salamander, Salamandra atra, has been demonstrated to consist of two phases: oophagy - after hatching ...Missing: offspring scientific
[52]
Salamandra - an overview | ScienceDirect Topics
The toxicity is presumably due to very potent local anesthetic activity. Remarkably, the fire salamander is sensitive to the toxic effects of samandarine.Missing: strategies | Show results with:strategies
[53]
Nuclear and mitochondrial multilocus phylogeny and survey of ...
The major toxic compounds in the skin secretions of S. salamandra and S. atra are steroidal alkaloids called samandarines, of which nine derivatives have so far ...
[54]
(PDF) A salamander's toxic arsenal: review of skin poison diversity ...
Salamandra skin toxins do not only represent a potent antipredator defence but may also have antimicrobial effects.
[55]
A salamander's toxic arsenal: review of skin poison diversity and ...
Sep 4, 2018 · Salamandra skin toxins do not only represent a potent antipredator defence but may also have antimicrobial effects.<|separator|>
[56]
https://onlinelibrary.wiley.com/doi/full/10.1111/1749-4877.12492
[57]
(PDF) Absence of infection with the amphibian chytrid fungus in the ...
Aug 6, 2025 · Resistance to Chytridiomycosis in European Plethodontid Salamanders of the Genus Speleomantes ... North America and the neotropics harbor nearly ...
[58]
(PDF) Recorded mortality in the vulnerable Alpine salamander ...
Feb 19, 2018 · Recorded mortality in the vulnerable Alpine salamander, Salamandra atra prenjensis (Amphibia: Caudata), is not associated with the presence of ...
[59]
Emina Sunje - Establishment of the Regional Monitoring Program for ...
Feb 27, 2018 · The alpine salamander (Salamandra atra) has a continuous ... Ranavirus. It is crucial to understand if the origin of the virus comes ...
[60]
Salamandra salamandra - an overview | ScienceDirect Topics
Gestation is shorter in S. salamandra, which gives birth to larval salamanders. The large size of these offspring indicates that considerable nutrient ...
[61]
Another Isolated Relic Population of the Alpine Salamander ...
We provide information on an isolated locality of Salamandra atra on Mt. Orjen in southern Montenegro that fills in a large gap in the species distribution ...<|separator|>
[62]
Alpine salamanders at risk? The current status of an emerging ...
May 17, 2024 · Members of the genus Salamandra are highly susceptible to Bsal leading to the lethal disease. Moreover, ecological modelling has shown that the ...
[63]
About the Large Alpine Salamander in France
It is often confused with the similar Salamandra atra ... Threats are principally human activities, tourism, road building and other infrastructure projects.
[64]
Salamandra atra aurorae in immediate danger due to deforestation!
Jul 15, 2020 · An urgent call has been made to protect the forests in the Venetian Alps – more concretely the area of the Altopiano dei Sette Comuni.<|control11|><|separator|>
[65]
[PDF] Salamandra atra
Conservation Measures: Salamandra atra is listed on Appendix II of the Berne Convention and. Salamandra atra aurorae is listed on Annex II of the EU Natural.
[66]
Forest management and conservation of an elusive amphibian in ...
Sep 15, 2018 · The critically endangered Salamandra lanzai from the Alps may be severely exposed to future climate change effects and its suitable climate ...<|control11|><|separator|>
[67]
Absence of Bsal in Peripheral Populations of the Golden Alpine ...
Jan 8, 2025 · Batrachochytrium salamandrivorans (Bsal) poses a serious threat to Urodeles, causing mass mortality in populations naive to the pathogen.