Alpine salamander: Difference between revisions

Alpine salamander
Conservation status
	; Least Concern (IUCN 3.1)
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Amphibia
Order:	Urodela
Family:	Salamandridae
Genus:	Salamandra
Species:	S. atra
Binomial name
	Salamandra atra; Laurenti, 1768

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The alpine salamander (Salamandra atra) is a black salamander that can be found in the French Alps, and through the mountainous range in Europe^[2] . Their species name, atra, may be derived from the Latin Ater, meaning dull black^[3]. Their life expectancy is at least 10 years. There are four subspecies of the alpine salamander, with varied distribution and physical coloration^[4]. Unlike other salamanders, whose larvae are developed in water, the alpine salamander is a fully terrestrial species. Capture-recapture methods suggest that the species is very stationary^[5]; 12 m (39 ft) was the maximum observed distance traveled by one individual during the summer season. About 120 individuals per hectare were counted in most suitable areas with >2000 individuals/ha also observed, suggesting that this rather cryptic species is quite abundant.^[6]

Description

Alpine salamanders are often small in size, and dark brown or black^[4]. Members of the subspecies are not wholly black or brown monochrome, but rather have mosaic or spotted patterns^[4]. Members of the subspecies Salamandra atra aurorae have bright splotches on its dorsal side and head. The color is often bright yellow, but can range to shades of white or even gray^[4]. Distribution of the pigment is dependent on the distribution of certain cells, so may be smooth and even or patchy^[4].

Female S. atra tend to be larger than the males, and can grow up to 151 millimeters, or around 5.9 inches^[4]. Males will grow to around 144 millimeters, or 5.6 inches: both measurements include the tail^[4]. Males have swollen, visible cloacas, and are more slender than females^[4]^[7]. The salamanders have parotid glands posterior and lateral to their eyes, giving them an elongated head shape^[4]. They tend to have between 11 and 13 costal grooves along theeh sides of their bodies, and a double row of dorsal glands runs down their backs on either side of their spines, down to the tips of their tails^[4].

Coloration

Most alpine salamanders are either completely black (melanistic) or predominantly blackhave black as a baseline, but the evolution behind this dark coloring has a winding history^[8]. Scientists have studied the hypothesis of if the salamanders start completely black, or if they evolved like that over time^[9]. DNA-evidence traced through maternal lineages suggest the latter: that salamanders evolved their black coloration over time^[9].

Pigmentation is determined by presentation of specific color-producing cells, called chromatophores, which absorb andor reflect light in a particular way to then appear as a color to our eyes^[10]^[11]. In S. atra, there are different cells present or activated, which yield different colors: melanophores contribute to the dark coloration by producing the dark pigment melanin^[10], while xanthophores produce a yellow pigment^[8], and iridophores are simply light-reflecting^[8]. The fully-black phenotype seen in S. a. atra results from the salamanders' melanophores in the dermis and epidermis, producing melanin alone^[9]^[10]. Xanthophore-iridophore complexes are responsible for production of yellow spotting, which appears bright^[9]. In species without yellow patches, it appears that they do not ever develop these cells^[9]. In S. a. aurorae or other salamanders with different coloration on different parts of their body, two distinct skin types are present: one that only contains melanophores (black), and one that has melanophores, xanthophores, and iridophores in combination^[10].

Distribution

The alpine salamander is found from the Swiss-French border at the western end of its range, all the way through Austria to the Dinaric Alps at the eastern edge of its territory. This salamander typically lives at altitudes above 700 m (2,300 ft) above sea level, even reaching 2000 meters (6561 ft) of elevation^[4]. The western Alps (in France and Italy) are inhabited by a similar species, Lanza's alpine salamander (Salamandra lanzai), in only one small area[citation needed]. S. atra generally live in forested biomes, particularly deciduous-coniferous mixes^[4]. They also can inhabit meadows or grasslands in the mountains^[4], and tend to do well with a mix of tree types^[12]^[13].

Their range spans several nations, including: Slovenia, Croatia, Bosnia Hersegovina, Montenegro, Kosovo, france, Italy, Austria.

Subspecies

S. a. atra is a fully melanistic (black) subspecies from central, eastern and Dinaric Alps^[14]. This subspecies is the most widespread.
S. a. aurorae, the golden alpine salamander, has golden or yellow spots on its back and primarily lives in a small area in the Venetian Prealps near Asiago, and in the Italian Alps.[citation needed]
S. a. pasubiensis, with fewer yellow spots than S. a. aurorae, lives in a different part of the Venetian Prealps.
S. a. prenjensis lives on Prenj Mountain, part of the Dinaric Alps in Bosnia and Herzegovina. The validity of this subspecies is yet to be confirmed, and some scientists in the field debate if this salamander should be considered its own independent species.^[4]^[15]

Genetic analysis suggests that the Corsican fire salamander (Salamandra corsica) is the closest related species, and the black-yellow coloration is an ancestral feature of alpine salamanders. Proposed colonization from south (Prealps) to Alps was carried out by the fully melanistic (derived feature) S. a. atra after the last retreat of the ice sheets.^[15]

Behavior

Habitat and Environmental Preferences

S. atra tend to live underneath stones or logs, or in rocky crevices in their mountainous habitat^[4]. They also are diurnal, and most active in the day with periods of inactivity, rest, or sleep at night^[13]. They will engage in nocturnal activity on a weather-dependent basis^[16]. Ideal weather for alpine salamanders is rainy or post-rain, at temperatures between 3 and 18C (around 37-64 F)^[13].

Territoriality

They thrive in forest environments that have silver firs and beech trees. Coniferous forests that have high proportions of Norway spruces and European larch trees also provide adequate habitats, even though the salamanders live on the ground floor^[12] . Because the salamanders are completely terrestrial, they have on-land territories that they tend to return to throughout the day and for refuge. They often return to the same sites for much of their lives^[12].Anytime that they leave their sites, they expose themselves to predation and also to the chance of losing their site(). Alpine salamanders are ectothermic, so losing a refuge or shelter could leave them exposed to the elements and be extremely costly, if not fatal, do their lack of an internal thermoregulation mechanism(). This high dependency on a quality nest site supportsthe theory that many terrestrial salamanders, including S. atra, engage in territorial behaviors^[12] .They employ scent-marking techniques to this, and to mark their territories using fecal pellets so they can identify their own shelters^[12]. Scent-marking is an intra-species communication, where chemical signals convey specific messages to other S. atra individuals^[12]. Alpine salamanders can determine if a ound fecal pellet has been left by a member of their same sex and/or species^[12]. Thus, this technique serves a double purpose to warn other salamanders that that particular location has already been claimed^[12]. Females are more likely to return to their home site, while males are more emboldened to enter another male’s territory^[12]. Their fecal pellets allow them to both participate in homing behavior, or returning to their own site, and territoriality, and determine intruders on their territory or invade the spaces of others^[12].

Male-Male Behavior

Males are more likely to engage in chasing other males, as well as actually fighting with each other^[17]. Oftentimes, one male will mount the other, loosely grasp it with his forearms, and start rubbing his head on the other male^[17]. The two males will switch roles, and in one studied interaction, continued like this for seven minutes before parting ways^[17]. In a second documented fight, the behavior was more intense. When one member tried to leave, the other male chased it to re-engage^[17]. This encounter lasted eight minutes. There is video evidence of these behaviors^[17]. Researchers are unsure if it is caused by territoriality, confusion on sexual identificationand mistaken mating, or true combat^[17].

Diet

Male and female alpine salamanders have relatively similar diets^[18]. Some specific organisms they prey on include species such as beetles, snails, millipedes, and spiders^[19], but alpine salamanders display preferences among prey^[18]. S. atra typically consume organisms from the Coloeptera and mollusca taxa^[18]. These taxa are the most crucial component of their diet (). They also tend to eat larger prey since they themselves are larger salamanders^[18]. Though alpine salamanders have definite dietary preferences, they have a substantial amount of variation in their diet^[18]that corresponds to their own optimized physical needs and pre-catching abilities.

Mating:

The alpine salamander engages in a promiscuous mating pattern^[16], meaning that they engage in multiple partner pairings. Males immigrate and travel farther than females do, potentially to find minds and following a scent emitted by females^[16]. Typically, males migrated when they were in the juvenile stage of life^[16]. Female S. atra find and defend their shelters, which is a potential reason that they may stay more local than males^[16].

Gestation and Reproduction

Alpine Salamanders live in a sex ratio of 1:1.^[20] . Mating occurs on land. The male clasps the female at the forelegs, and fertilization is internal. S. atra are categorized as viviparous, meaning that their young are born alive and unlike many other amphibians, do not go through metamorphosis (De meester). They give birth to 2 young, sometimes 3 or 4. New young alpine salamanders may measure as long as 50 mm (2.0 in) at birth, with the mother measuring only 120 mm (4.7 in).

Female alpine salamanders have uteruses that are composed of a single luminal epithelial cell layer, connective tissue, and smooth muscle^[21].The uterine eggs are large and numerous, but, as a rule, only one fully develops in each uterus. The embryo is nourished on the yolk of the other eggs, which more or less dissolve to form a large mass of nutrient matter. The egg mass can be as long as between 25 mm and 40 mm long^[21]. The embryo passes through three stages^[22]:

The first stage is when they are still enclosed within the egg and living on its own yolk.^[22]
The second stage is when they are free, within the vitelline mass, eating it directly with their mouths.^[21]^[22]
The final stage is where there is no more vitelline mass. The embryo is possessed of long external gills, which serve as an exchange of nutritive fluid through the maternal uterus, these gills functioning in the same way as the chorionic villi of the mammalian egg.^[21]^[22]

Generally, at altitudes of 650-1,000 m above sea level, a pregnancy lasts two years, and at altitudes of 1,400-1,700 m above sea level, the pregnancy lasts around three years, though anything within a 2-4 year range is considered standard^[21]. Alpine salamander embryos are unique in how they are able to take in these nutrients through a long gestation^[21]. A portion of the mother’s uterine wall becomes nourishment after the salamanders have already eaten the unfertilized eggs^[21] (called oophagy^[21], or stage 1 and 2). They ten partake in epitheliophagy, or stage 3, where they ingest these zona trophica cells until birth, and have special tooth-like developments that allow it to do so without detriment to the mother^[21].

Physiology

Glands and Toxins

As mentioned above, alpine salamanders have poison glands. They are known to produce some alkaloid molecules and peptide products, and thus have a mustard-like scent associated with them^[23]. Salamandra bio production is still a developing research area. Salamandarines are a chemical secretion produced by the skin of alpine salamanders, as well as some fire salamanders (De Meester)^[4]^[24]. They are neurotoxins, and are syntehtised via a biochemical pathway independent of dietary intake^[4]. The starting material for this nerve-block is most likely cholesterol, and it is about twice as potent as cyanide^[4] . This pales in comparison to other toxins produced by salamanders, but S. atra do not only use this powerful substance to paralyze prey: they may have antimicrobial properties that protect them against bacterial and fungal infections^[4]^[25]. Salamandorone is another biochemical compound produced by S. atra, and though it is less potent against prey it is the strongest antimicrobial weapon these salamanders have^[4]^[26]^[25].

Immunobiology and Protection

The alpine salamander has been relatively lucky in avoiding infection with amphian chytrid fungus^[25] compared to other amphibian species^[25]^[27]. This dangerous fungal infection, caused by Batrachochytrium dendrobatidis (Bd) has decimated amphibian populations on every continent^[25]. Bd is present in the Alps where alpine salamanders live, but in a study performed in 2012, there were no salamanders who tested positive when swabbed^[25]. This may be because Bd infections are more common in species who spend more of their time in water, and since the alpine salamanders are terrestrial, they are less susceptible^[25]. An alternate hypothesis proposes that S. atra are resistant via their skin microbiome or a produced molecule, thus granting them immunity^[25]^[28]. This theory is untested, but considering how many salamanders excrete biological toxins, plausible^[25]^[28].

Conservation

Alpine salamanders are not resiliant to habitat change, so risks of climate change altering their living spaces is severe^[5]. Though the IUCN reports that alpine salamanders are least vulnerable, their numbers are decreasing^[29]. Additionally, some subspecies of S. atra are in greater danger^[30]. Population numbers are declining in S. atra aurorae, for example. One of the greatest dangers to alpine salamanders overall is commercial deforestation in their habitats^[30]. Machinery like tractors or other forestry tools can compress the soil eliminating some of the small insects that S. atra eat or eliminating potential nooks and burrows for them to use as shelters^[30]. Many scientists propose changes in the lumber industry as an attempt to heal these habitats^[30]. These salamanders may also change their morphology as global temperatures rise^[31]. Amphibians and other organisms that do not internally regulate their body temperature may need adaption mechanisms to remain at ideal physiological temperatures in the face of changing climates^[31]. Other issues like acid rain or precipitation changes could prompt many animals, including alpine salamanders, to be forced into new habitats^[31]. Alpine salamanders play a crucial role in their ecosystems^[32]. There are already animal and ecosystem conservation laws in Europe, but many scholars recommend additional ones to protect the flora and fauna^[33].

References

^ Franco Andreone, Mathieu Denoël, Claude Miaud, Benedikt Schmidt, Paul Edgar, Milan Vogrin, Jelka Crnobrnja Isailovic, Rastko Ajtic, Claudia Corti, Idriz Haxhiu (2009). "Salamandra atra". IUCN Red List of Threatened Species. 2009: e.T19843A9023725. doi:10.2305/IUCN.UK.2009.RLTS.T19843A9023725.en. Retrieved 13 November 2021.{{cite journal}}: CS1 maint: multiple names: authors list (link)
^ "Salamandre noire unique - La Salamandre". Archived from the original on 2020-07-02.
^ "ater - WordSense Dictionary". www.wordsense.eu. Retrieved 2022-11-14.
^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j ^k ^l ^m ⁿ ^o ^p ^q ^r ^s ^t DE MEESTER, Gilles; ŠUNJE, Emina; PRINSEN, Els; VERBRUGGEN, Erik; VAN DAMME, Raoul (2020-10-13). "Toxin variation among salamander populations: discussing potential causes and future directions". Integrative Zoology. 16 (3): 336–353. doi:10.1111/1749-4877.12492. ISSN 1749-4877.
^ ^a ^b Basile, Marco; Romano, Antonio; Costa, Andrea; Posillico, Mario; Scinti Roger, Daniele; Crisci, Aldo; Raimondi, Ranieri; Altea, Tiziana; Garfì, Vittorio; Santopuoli, Giovanni; Marchetti, Marco; Salvidio, Sebastiano; De Cinti, Bruno; Matteucci, Giorgio (2017-09-12). "Seasonality and microhabitat selection in a forest-dwelling salamander". The Science of Nature. 104 (9–10). doi:10.1007/s00114-017-1500-6. ISSN 0028-1042.
^ Bonato, Fracasso. Movements, distribution pattern and density in a population of Salamandra atra aurorae (Caudata: Salamandridae). Amphibia-Reptilia 2003, 24, 251-260.
^ "Salamandra atra". AmphibiaWeb. 6 December 2018.
^ ^a ^b ^c Burgon, James D.; Vieites, David R.; Jacobs, Arne; Weidt, Stefan K.; Gunter, Helen M.; Steinfartz, Sebastian; Burgess, Karl; Mable, Barbara K.; Elmer, Kathryn R. (2020-04). "Functional colour genes and signals of selection in colour‐polymorphic salamanders". Molecular Ecology. 29 (7): 1284–1299. doi:10.1111/mec.15411. ISSN 0962-1083. {{cite journal}}: Check date values in: |date= (help)
^ ^a ^b ^c ^d ^e Bonato, Lucio; Steinfartz, Sebastian (2005-01-01). "Evolution of the melanistic colour in the Alpine salamander Salamandra atra as revealed by a new subspecies from the Venetian Prealps". Italian Journal of Zoology. 72 (3): 253–260. doi:10.1080/11250000509356680. ISSN 1125-0003.
^ ^a ^b ^c ^d TREVISAN, PIERLUIGI; PEDERZOLI, AURORA; BAROZZI, GIANCARLO (1991-10). "Pigmentary System of the Adult Alpine Salamander Salamandra atra atra (Laur., 1768)". Pigment Cell Research. 4 (4): 151–157. doi:10.1111/j.1600-0749.1991.tb00432.x. ISSN 0893-5785. {{cite journal}}: Check date values in: |date= (help)
^ PEDERZOLI, AURORA; TREVISAN, PIERLUIGI (1989-03). "Pigmentary System of the Adult Alpine Salamander Salamandra atra aurorae (Trevisan, 1982)". Pigment Cell Research. 3 (2): 80–89. doi:10.1111/j.1600-0749.1989.tb00266.x. ISSN 0893-5785. {{cite journal}}: Check date values in: |date= (help)
^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j Gautier, Patrick G.; Miaud, Claude (2003-01-01). "Faecal pellets used as an economic territorial marker in two terrestrial alpine salamanders". Écoscience. 10 (2): 134–139. doi:10.1080/11956860.2003.11682759. ISSN 1195-6860.
^ ^a ^b ^c Romano, Antonio; Anderle, Matteo; Forti, Alessandro; Partel, Piergiovanni; Pedrini, Paolo (2018-12-31). "Population density, sex ratio and body size in a population of Salamandra atra atra on the Dolomites". Acta Herpetologica: 195–199 Pages. doi:10.13128/ACTA_HERPETOL-22592.
^ "Salamandre noir". Karch.Ch (Swiss information center for amphibians and reptiles). Retrieved 29 June 2020.
^ ^a ^b Bonato & Steinfartz. Evolution of the melanistic color in the Alpine salamander Salamandra atra as revealed by a new subspecies from the Venetian prealps. Italian Journal of Zoology 2001, 72, 253-260.
^ ^a ^b ^c ^d ^e Helfer, V.; Broquet, T.; Fumagalli, L. (2012-10). "Sex-specific estimates of dispersal show female philopatry and male dispersal in a promiscuous amphibian, the alpine salamander ( Salamandra atra )". Molecular Ecology. 21 (19): 4706–4720. doi:10.1111/j.1365-294X.2012.05742.x. {{cite journal}}: Check date values in: |date= (help)
^ ^a ^b ^c ^d ^e ^f Di Nicola, Matteo Ricardo (November 2022). "Male-male interactions in Alpine salamanders, Salamandra atra atra Laurenti, 1768, with an overview of the main cases reported for the whole genus Salamandra Garsault, 1764". Herpetology Notes. Volume 15: 601–604. {{cite journal}}: |volume= has extra text (help)
^ ^a ^b ^c ^d ^e Roner, Luca; Costa, Andrea; Pedrini, Paolo; Matteucci, Giorgio; Leonardi, Stefano; Romano, Antonio (2020-05). "A Midsummer Night's Diet: Snapshot on Trophic Strategy of the Alpine Salamander, Salamandra atra". Diversity. 12 (5): 202. doi:10.3390/d12050202. ISSN 1424-2818. {{cite journal}}: Check date values in: |date= (help)CS1 maint: unflagged free DOI (link)
^ Šunje, Emina; Courant, Julien; Vesnić, Adi; Koren, Toni; Lukić Bilela, Lada; Van Damme, Raoul (2022-09-20). "Patterns of variation in dietary composition among four populations of Alpine salamanders (Salamandra atra prenjensis)". Amphibia-Reptilia: 1–15. doi:10.1163/15685381-bja10100. ISSN 0173-5373.
^ Body size, population structure and fecundity traits of Salamandra atra atra (Amphibia, Urodela, Salamandridae) population from the northeastern Italian Alps. Vol. 68. Luiselli, Andreone, Capizzi, Anibaldi: Italian Journal of Zoology. 2001. pp. 125–130.
^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ Guex, G. -D.; Chen, P. S. (1986-12). "Epitheliophagy: Intrauterine cell nourishment in the viviparous alpine salamander,Salamandra atra (Laur.)". Experientia. 42 (11–12): 1205–1218. doi:10.1007/bf01946392. ISSN 0014-4754. {{cite journal}}: Check date values in: |date= (help)
^ ^a ^b ^c ^d Chisholm, Hugh, ed. (1911). "Salamander" . Encyclopædia Britannica. Vol. 1 (11th ed.). Cambridge University Press.
^ Lüddecke, Tim; Schulz, Stefan; Steinfartz, Sebastian; Vences, Miguel (2018-09-04). "A salamander's toxic arsenal: review of skin poison diversity and function in true salamanders, genus Salamandra". The Science of Nature. 105 (9–10). doi:10.1007/s00114-018-1579-4. ISSN 0028-1042.
^ De Meester, Gilles; Šunje, Emina; Prinsen, Els; Verbruggen, Erik; Van Damme, Raoul (2021-05). "Toxin variation among salamander populations: discussing potential causes and future directions". Integrative Zoology. 16 (3): 336–353. doi:10.1111/1749-4877.12492. ISSN 1749-4877. {{cite journal}}: Check date values in: |date= (help)
^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ R, Lötters, S Kielgast, J Sztatecsny, M Wagner, N Schulte, U Werner, P Rödder, D Dambach, J Reissner, T Hochkirch, A Schmidt, B (2012-04-30). Absence of infection with the amphibian chytrid fungus in the terrestrial Alpine salamander, Salamandra atra. Deutsche Gesellschaft für Herpetologie und Terrarienkunde (DGHT). OCLC 1030045649.{{cite book}}: CS1 maint: multiple names: authors list (link)
^ Spitzen - van der Sluijs, Annemarieke; Stegen, Gwij; Bogaerts, Sergé; Canessa, Stefano; Steinfartz, Sebastian; Janssen, Nico; Bosman, Wilbert; Pasmans, Frank; Martel, An (2018-02-28). "Post-epizootic salamander persistence in a disease-free refugium suggests poor dispersal ability of Batrachochytrium salamandrivorans". Scientific Reports. 8 (1). doi:10.1038/s41598-018-22225-9. ISSN 2045-2322.
^ Feldmeier, Stephan; Schefczyk, Lukas; Wagner, Norman; Heinemann, Günther; Veith, Michael; Lötters, Stefan (2016-10-31). "Exploring the Distribution of the Spreading Lethal Salamander Chytrid Fungus in Its Invasive Range in Europe – A Macroecological Approach". PLOS ONE. 11 (10): e0165682. doi:10.1371/journal.pone.0165682. ISSN 1932-6203. PMC 5087956. PMID 27798698.{{cite journal}}: CS1 maint: PMC format (link) CS1 maint: unflagged free DOI (link)
^ ^a ^b Woodhams, D. C.; Rollins-Smith, L. A.; Alford, R. A.; Simon, M. A.; Harris, R. N. (2007-11). "Innate immune defenses of amphibian skin: antimicrobial peptides and more". Animal Conservation. 10 (4): 425–428. doi:10.1111/j.1469-1795.2007.00150.x. ISSN 1367-9430. {{cite journal}}: Check date values in: |date= (help)
^ IUCN (7 July 2021). "The IUCN Red List of Threatened Species". ” IUCN Red List of Threatened Species.
^ ^a ^b ^c ^d Romano, Antonio; Costa, Andrea; Salvidio, Sebastiano; Menegon, Michele; Garollo, Elena; Tabarelli de Fatis, Karol; Miserocchi, Danio; Matteucci, Giorgio; Pedrini, Paolo (2018-09). "Forest management and conservation of an elusive amphibian in the Alps: Habitat selection by the Golden Alpine Salamander reveals the importance of fine woody debris". Forest Ecology and Management. 424: 338–344. doi:10.1016/j.foreco.2018.04.052. ISSN 0378-1127. {{cite journal}}: Check date values in: |date= (help)
^ ^a ^b ^c Ficetola, Gentile Francesco; Colleoni, Emiliano; Renaud, Julien; Scali, Stefano; Padoa-Schioppa, Emilio; Thuiller, Wilfried (2016-06). "Morphological variation in salamanders and their potential response to climate change". Global Change Biology. 22 (6): 2013–2024. doi:10.1111/gcb.13255. PMC 4972144. PMID 26910389. {{cite journal}}: Check date values in: |date= (help)CS1 maint: PMC format (link)
^ Reinthaler-Lottermoser, Ursula; et al. (8 October 2018). "A new approach for surveying the Alpine Salamander (Salamandra atra) in Austria". Acta Herpetologic: 249–253. {{cite journal}}: Explicit use of et al. in: |last= (help)
^ Scalera, Riccardo (2004-07-01). "The legal framework for the protection of amphibians and reptiles in Italy: an annotated overview of the provisions at the international, European community, national and regional level". Italian Journal of Zoology. 71 (sup002): 21–32. doi:10.1080/11250003.2004.9525560. ISSN 1125-0003.

External links

Media related to Alpine salamander at Wikimedia Commons
Data related to Salamandra atra at Wikispecies
European Field Herping Community. "Picture gallery of S. a. atra, S. a. aurorae and S. a.pasubiensis subspecies". Archived from the original on 2015-09-24. Retrieved 2011-02-20.
Emina Šunje. "Salamandra atra prenjensis". Archived from the original on 2021-04-27. Retrieved 2011-02-20. (in Bosnian)

[iucn_status_13_November_2021-1] Franco Andreone, Mathieu Denoël, Claude Miaud, Benedikt Schmidt, Paul Edgar, Milan Vogrin, Jelka Crnobrnja Isailovic, Rastko Ajtic, Claudia Corti, Idriz Haxhiu (2009). "Salamandra atra". IUCN Red List of Threatened Species. 2009: e.T19843A9023725. doi:10.2305/IUCN.UK.2009.RLTS.T19843A9023725.en. Retrieved 13 November 2021.{{cite journal}}: CS1 maint: multiple names: authors list (link)

[2] "Salamandre noire unique - La Salamandre". Archived from the original on 2020-07-02.

[3] "ater - WordSense Dictionary". www.wordsense.eu. Retrieved 2022-11-14.

[:0-4] ^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j ^k ^l ^m ⁿ ^o ^p ^q ^r ^s ^t DE MEESTER, Gilles; ŠUNJE, Emina; PRINSEN, Els; VERBRUGGEN, Erik; VAN DAMME, Raoul (2020-10-13). "Toxin variation among salamander populations: discussing potential causes and future directions". Integrative Zoology. 16 (3): 336–353. doi:10.1111/1749-4877.12492. ISSN 1749-4877.

[:1-5] Basile, Marco; Romano, Antonio; Costa, Andrea; Posillico, Mario; Scinti Roger, Daniele; Crisci, Aldo; Raimondi, Ranieri; Altea, Tiziana; Garfì, Vittorio; Santopuoli, Giovanni; Marchetti, Marco; Salvidio, Sebastiano; De Cinti, Bruno; Matteucci, Giorgio (2017-09-12). "Seasonality and microhabitat selection in a forest-dwelling salamander". The Science of Nature. 104 (9–10). doi:10.1007/s00114-017-1500-6. ISSN 0028-1042.

[6] Bonato, Fracasso. Movements, distribution pattern and density in a population of Salamandra atra aurorae (Caudata: Salamandridae). Amphibia-Reptilia 2003, 24, 251-260.

[7] "Salamandra atra". AmphibiaWeb. 6 December 2018.

[:2-8] Burgon, James D.; Vieites, David R.; Jacobs, Arne; Weidt, Stefan K.; Gunter, Helen M.; Steinfartz, Sebastian; Burgess, Karl; Mable, Barbara K.; Elmer, Kathryn R. (2020-04). "Functional colour genes and signals of selection in colour‐polymorphic salamanders". Molecular Ecology. 29 (7): 1284–1299. doi:10.1111/mec.15411. ISSN 0962-1083. {{cite journal}}: Check date values in: |date= (help)

[:3-9] Bonato, Lucio; Steinfartz, Sebastian (2005-01-01). "Evolution of the melanistic colour in the Alpine salamander Salamandra atra as revealed by a new subspecies from the Venetian Prealps". Italian Journal of Zoology. 72 (3): 253–260. doi:10.1080/11250000509356680. ISSN 1125-0003.

[:4-10] TREVISAN, PIERLUIGI; PEDERZOLI, AURORA; BAROZZI, GIANCARLO (1991-10). "Pigmentary System of the Adult Alpine Salamander Salamandra atra atra (Laur., 1768)". Pigment Cell Research. 4 (4): 151–157. doi:10.1111/j.1600-0749.1991.tb00432.x. ISSN 0893-5785. {{cite journal}}: Check date values in: |date= (help)

[11] PEDERZOLI, AURORA; TREVISAN, PIERLUIGI (1989-03). "Pigmentary System of the Adult Alpine Salamander Salamandra atra aurorae (Trevisan, 1982)". Pigment Cell Research. 3 (2): 80–89. doi:10.1111/j.1600-0749.1989.tb00266.x. ISSN 0893-5785. {{cite journal}}: Check date values in: |date= (help)

[:5-12] ^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j Gautier, Patrick G.; Miaud, Claude (2003-01-01). "Faecal pellets used as an economic territorial marker in two terrestrial alpine salamanders". Écoscience. 10 (2): 134–139. doi:10.1080/11956860.2003.11682759. ISSN 1195-6860.

[:6-13] Romano, Antonio; Anderle, Matteo; Forti, Alessandro; Partel, Piergiovanni; Pedrini, Paolo (2018-12-31). "Population density, sex ratio and body size in a population of Salamandra atra atra on the Dolomites". Acta Herpetologica: 195–199 Pages. doi:10.13128/ACTA_HERPETOL-22592.

[14] "Salamandre noir". Karch.Ch (Swiss information center for amphibians and reptiles). Retrieved 29 June 2020.

[bonato2001-15] Bonato & Steinfartz. Evolution of the melanistic color in the Alpine salamander Salamandra atra as revealed by a new subspecies from the Venetian prealps. Italian Journal of Zoology 2001, 72, 253-260.

[:7-16] Helfer, V.; Broquet, T.; Fumagalli, L. (2012-10). "Sex-specific estimates of dispersal show female philopatry and male dispersal in a promiscuous amphibian, the alpine salamander ( Salamandra atra )". Molecular Ecology. 21 (19): 4706–4720. doi:10.1111/j.1365-294X.2012.05742.x. {{cite journal}}: Check date values in: |date= (help)

[:8-17] ^ ^a ^b ^c ^d ^e ^f Di Nicola, Matteo Ricardo (November 2022). "Male-male interactions in Alpine salamanders, Salamandra atra atra Laurenti, 1768, with an overview of the main cases reported for the whole genus Salamandra Garsault, 1764". Herpetology Notes. Volume 15: 601–604. {{cite journal}}: |volume= has extra text (help)

[:9-18] Roner, Luca; Costa, Andrea; Pedrini, Paolo; Matteucci, Giorgio; Leonardi, Stefano; Romano, Antonio (2020-05). "A Midsummer Night's Diet: Snapshot on Trophic Strategy of the Alpine Salamander, Salamandra atra". Diversity. 12 (5): 202. doi:10.3390/d12050202. ISSN 1424-2818. {{cite journal}}: Check date values in: |date= (help)CS1 maint: unflagged free DOI (link)

[19] Šunje, Emina; Courant, Julien; Vesnić, Adi; Koren, Toni; Lukić Bilela, Lada; Van Damme, Raoul (2022-09-20). "Patterns of variation in dietary composition among four populations of Alpine salamanders (Salamandra atra prenjensis)". Amphibia-Reptilia: 1–15. doi:10.1163/15685381-bja10100. ISSN 0173-5373.

[20] Body size, population structure and fecundity traits of Salamandra atra atra (Amphibia, Urodela, Salamandridae) population from the northeastern Italian Alps. Vol. 68. Luiselli, Andreone, Capizzi, Anibaldi: Italian Journal of Zoology. 2001. pp. 125–130.

[:10-21] ^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ Guex, G. -D.; Chen, P. S. (1986-12). "Epitheliophagy: Intrauterine cell nourishment in the viviparous alpine salamander,Salamandra atra (Laur.)". Experientia. 42 (11–12): 1205–1218. doi:10.1007/bf01946392. ISSN 0014-4754. {{cite journal}}: Check date values in: |date= (help)

[:11-22] Chisholm, Hugh, ed. (1911). "Salamander" . Encyclopædia Britannica. Vol. 1 (11th ed.). Cambridge University Press.

[23] Lüddecke, Tim; Schulz, Stefan; Steinfartz, Sebastian; Vences, Miguel (2018-09-04). "A salamander's toxic arsenal: review of skin poison diversity and function in true salamanders, genus Salamandra". The Science of Nature. 105 (9–10). doi:10.1007/s00114-018-1579-4. ISSN 0028-1042.

[24] De Meester, Gilles; Šunje, Emina; Prinsen, Els; Verbruggen, Erik; Van Damme, Raoul (2021-05). "Toxin variation among salamander populations: discussing potential causes and future directions". Integrative Zoology. 16 (3): 336–353. doi:10.1111/1749-4877.12492. ISSN 1749-4877. {{cite journal}}: Check date values in: |date= (help)

[:12-25] ^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ R, Lötters, S Kielgast, J Sztatecsny, M Wagner, N Schulte, U Werner, P Rödder, D Dambach, J Reissner, T Hochkirch, A Schmidt, B (2012-04-30). Absence of infection with the amphibian chytrid fungus in the terrestrial Alpine salamander, Salamandra atra. Deutsche Gesellschaft für Herpetologie und Terrarienkunde (DGHT). OCLC 1030045649.{{cite book}}: CS1 maint: multiple names: authors list (link)

[26] Spitzen - van der Sluijs, Annemarieke; Stegen, Gwij; Bogaerts, Sergé; Canessa, Stefano; Steinfartz, Sebastian; Janssen, Nico; Bosman, Wilbert; Pasmans, Frank; Martel, An (2018-02-28). "Post-epizootic salamander persistence in a disease-free refugium suggests poor dispersal ability of Batrachochytrium salamandrivorans". Scientific Reports. 8 (1). doi:10.1038/s41598-018-22225-9. ISSN 2045-2322.

[27] Feldmeier, Stephan; Schefczyk, Lukas; Wagner, Norman; Heinemann, Günther; Veith, Michael; Lötters, Stefan (2016-10-31). "Exploring the Distribution of the Spreading Lethal Salamander Chytrid Fungus in Its Invasive Range in Europe – A Macroecological Approach". PLOS ONE. 11 (10): e0165682. doi:10.1371/journal.pone.0165682. ISSN 1932-6203. PMC 5087956. PMID 27798698.{{cite journal}}: CS1 maint: PMC format (link) CS1 maint: unflagged free DOI (link)

[:13-28] Woodhams, D. C.; Rollins-Smith, L. A.; Alford, R. A.; Simon, M. A.; Harris, R. N. (2007-11). "Innate immune defenses of amphibian skin: antimicrobial peptides and more". Animal Conservation. 10 (4): 425–428. doi:10.1111/j.1469-1795.2007.00150.x. ISSN 1367-9430. {{cite journal}}: Check date values in: |date= (help)

[29] IUCN (7 July 2021). "The IUCN Red List of Threatened Species". ” IUCN Red List of Threatened Species.

[:14-30] Romano, Antonio; Costa, Andrea; Salvidio, Sebastiano; Menegon, Michele; Garollo, Elena; Tabarelli de Fatis, Karol; Miserocchi, Danio; Matteucci, Giorgio; Pedrini, Paolo (2018-09). "Forest management and conservation of an elusive amphibian in the Alps: Habitat selection by the Golden Alpine Salamander reveals the importance of fine woody debris". Forest Ecology and Management. 424: 338–344. doi:10.1016/j.foreco.2018.04.052. ISSN 0378-1127. {{cite journal}}: Check date values in: |date= (help)

[:15-31] Ficetola, Gentile Francesco; Colleoni, Emiliano; Renaud, Julien; Scali, Stefano; Padoa-Schioppa, Emilio; Thuiller, Wilfried (2016-06). "Morphological variation in salamanders and their potential response to climate change". Global Change Biology. 22 (6): 2013–2024. doi:10.1111/gcb.13255. PMC 4972144. PMID 26910389. {{cite journal}}: Check date values in: |date= (help)CS1 maint: PMC format (link)

[32] Reinthaler-Lottermoser, Ursula; et al. (8 October 2018). "A new approach for surveying the Alpine Salamander (Salamandra atra) in Austria". Acta Herpetologic: 249–253. {{cite journal}}: Explicit use of et al. in: |last= (help)

[33] Scalera, Riccardo (2004-07-01). "The legal framework for the protection of amphibians and reptiles in Italy: an annotated overview of the provisions at the international, European community, national and regional level". Italian Journal of Zoology. 71 (sup002): 21–32. doi:10.1080/11250003.2004.9525560. ISSN 1125-0003.

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@@ Line 13: / Line 13: @@
 [[File: Salamandra atra_accouplement_MHNT.jpg|thumb|''Salamandra atra'' - [[MHNT]]]]
-The '''alpine salamander''' (''Salamandra atra'') is a shiny black [[salamander]] found in the Alps from the Swiss-French border at the western end of its range  <ref>{{cite web| url = https://www.salamandre.org/article/salamandre-noire-unique/ |url-status=dead |archive-url=https://web.archive.org/web/20200702202548/https://www.salamandre.org/article/salamandre-noire-unique/ |archive-date=2020-07-02 |title=Salamandre noire unique - La Salamandre}}</ref> through Austria to the [[Dinaric Alps]] at the eastern end of its range, at altitudes above {{convert|700|m|abbr=on}}. The [[western Alps]] (in France and Italy) are inhabited by a similar species, [[Lanza's alpine salamander]] (''Salamandra lanzai''), in only one small area. No differences in length are reported between the sexes ({{convert|9|-|14|cm|abbr=on}}) and the sex ratio is 1:1.<ref>{{cite book|location=Luiselli, Andreone, [[Capizzi]], Anibaldi|title= Body size, population structure and fecundity traits of Salamandra atra atra (Amphibia, Urodela, Salamandridae) population from the northeastern Italian Alps|publisher=Italian Journal of Zoology|year=2001|volume=68|pages=125–130}}</ref> Their life expectancy is at least 10 years. Unlike other salamanders, whose larvae are developed in water, the alpine salamander is a fully terrestrial species. Capture-recapture methods suggest that the species is very stationary; {{convert|12|m|abbr=on}} was the maximum observed distance travelled by one individual during the summer season. About 120 individuals per [[hectare]] were counted in most suitable areas with >2000 individuals/ha also observed, suggesting that this rather cryptic species is quite abundant.<ref>Bonato, Fracasso. Movements, distribution pattern and density in a population of Salamandra atra aurorae (Caudata: Salamandridae). Amphibia-Reptilia 2003, 24, 251-260.</ref>
+The '''alpine salamander''' (Salamandra atra) is a black '''[[salamander]]''' that can be found in the [[French Alps|French Alps,]] and through the mountainous range in Europe<ref>{{cite web| url = https://www.salamandre.org/article/salamandre-noire-unique/ |url-status=dead |archive-url=https://web.archive.org/web/20200702202548/https://www.salamandre.org/article/salamandre-noire-unique/ |archive-date=2020-07-02 |title=Salamandre noire unique - La Salamandre}}</ref> . Their species name, ''atra'', may be derived from the Latin ''Ater'', meaning dull black<ref>{{Cite web |title=ater - WordSense Dictionary |url=https://www.wordsense.eu/ater/ |access-date=2022-11-14 |website=www.wordsense.eu |language=en}}</ref>. Their life expectancy is at least 10 years. There are four subspecies of the alpine salamander, with varied distribution and physical coloration<ref name=":0">{{Cite journal |last=DE MEESTER |first=Gilles |last2=ŠUNJE |first2=Emina |last3=PRINSEN |first3=Els |last4=VERBRUGGEN |first4=Erik |last5=VAN DAMME |first5=Raoul |date=2020-10-13 |title=Toxin variation among salamander populations: discussing potential causes and future directions |url=http://dx.doi.org/10.1111/1749-4877.12492 |journal=Integrative Zoology |volume=16 |issue=3 |pages=336–353 |doi=10.1111/1749-4877.12492 |issn=1749-4877}}</ref>. Unlike other salamanders, whose larvae are developed in water, the alpine salamander is a fully terrestrial species. Capture-recapture methods suggest that the species is very stationary<ref name=":1">{{Cite journal |last=Basile |first=Marco |last2=Romano |first2=Antonio |last3=Costa |first3=Andrea |last4=Posillico |first4=Mario |last5=Scinti Roger |first5=Daniele |last6=Crisci |first6=Aldo |last7=Raimondi |first7=Ranieri |last8=Altea |first8=Tiziana |last9=Garfì |first9=Vittorio |last10=Santopuoli |first10=Giovanni |last11=Marchetti |first11=Marco |last12=Salvidio |first12=Sebastiano |last13=De Cinti |first13=Bruno |last14=Matteucci |first14=Giorgio |date=2017-09-12 |title=Seasonality and microhabitat selection in a forest-dwelling salamander |url=http://dx.doi.org/10.1007/s00114-017-1500-6 |journal=The Science of Nature |volume=104 |issue=9-10 |doi=10.1007/s00114-017-1500-6 |issn=0028-1042}}</ref>; 12 m (39 ft) was the maximum observed distance traveled by one individual during the summer season. About 120 individuals per hectare were counted in most suitable areas with >2000 individuals/ha also observed, suggesting that this rather cryptic species is quite abundant.<ref>Bonato, Fracasso. Movements, distribution pattern and density in a population of Salamandra atra aurorae (Caudata: Salamandridae). Amphibia-Reptilia 2003, 24, 251-260.</ref>
 == Description ==
+Alpine salamanders are often small in size, and dark brown or black<ref name=":0" />. Members of the subspecies are not wholly black or brown monochrome, but rather have mosaic or spotted patterns<ref name=":0" />. Members of the subspecies ''Salamandra atra aurorae'' have bright splotches on its dorsal side and head. The color is often bright yellow, but can range to shades of white or even gray<ref name=":0" />. Distribution of the pigment is dependent on the distribution of certain cells, so may be smooth and even or patchy<ref name=":0" />.
-Total length up to 144&nbsp;mm for males, up to 151&nbsp;mm for females. Slightly elongated head, with two large kidney-shaped paratoid glands. The body has 11-13 costal grooves on either side. The tail is square in cross-section. One double row of poison glands runs down the center of the back. One single row of poison glands runs along either side of the body onto the tail. Usually black or brown-black, but the subspecies ''Salamandra atra aurorae'' has a bright coloration on the head, back, and dorsal side of the extremities. This coloration can consist of continuous patches or be spotted or blotched. It can vary in color from whitish or yellow to greenish or gray. Males have a slightly more pronounced cloaca than females.<ref>{{Cite web|url=http://amphibiaweb.org:8000/cgi/amphib_query?where-genus=Salamandra&where-species=atra|title=Salamandra atra|date=6 December 2018|website=AmphibiaWeb}}</ref>
+Female S. atra tend to be larger than the males, and can grow up to 151 millimeters, or around 5.9 inches<ref name=":0" />. Males will grow to around 144 millimeters, or 5.6 inches: both measurements include the tail<ref name=":0" />. Males have swollen, visible cloacas, and are more slender than females<ref name=":0" /><ref>{{Cite web |date=6 December 2018 |title=Salamandra atra |url=http://amphibiaweb.org:8000/cgi/amphib_query?where-genus=Salamandra&where-species=atra |website=AmphibiaWeb}}</ref>. The salamanders have parotid glands posterior and lateral to their eyes, giving them an elongated head shape<ref name=":0" />. They tend to have between 11 and 13 costal grooves along theeh sides of their bodies, and a double row of dorsal glands runs down their backs on either side of their spines, down to the tips of their tails<ref name=":0" />.
+=== Coloration ===
+Most alpine salamanders are either completely black (melanistic) or predominantly blackhave black as a baseline, but the evolution behind this dark coloring has a winding history<ref name=":2">{{Cite journal |last=Burgon |first=James D. |last2=Vieites |first2=David R. |last3=Jacobs |first3=Arne |last4=Weidt |first4=Stefan K. |last5=Gunter |first5=Helen M. |last6=Steinfartz |first6=Sebastian |last7=Burgess |first7=Karl |last8=Mable |first8=Barbara K. |last9=Elmer |first9=Kathryn R. |date=2020-04 |title=Functional colour genes and signals of selection in colour‐polymorphic salamanders |url=https://onlinelibrary.wiley.com/doi/10.1111/mec.15411 |journal=Molecular Ecology |language=en |volume=29 |issue=7 |pages=1284–1299 |doi=10.1111/mec.15411 |issn=0962-1083}}</ref>. Scientists have studied the hypothesis of if the salamanders start completely black, or if they evolved like that over time<ref name=":3">{{Cite journal |last=Bonato |first=Lucio |last2=Steinfartz |first2=Sebastian |date=2005-01-01 |title=Evolution of the melanistic colour in the Alpine salamander Salamandra atra as revealed by a new subspecies from the Venetian Prealps |url=https://doi.org/10.1080/11250000509356680 |journal=Italian Journal of Zoology |volume=72 |issue=3 |pages=253–260 |doi=10.1080/11250000509356680 |issn=1125-0003}}</ref>. DNA-evidence traced through maternal lineages suggest the latter: that salamanders evolved their black coloration over time<ref name=":3" />.
+Pigmentation is determined by presentation of specific color-producing cells, called chromatophores, which absorb andor reflect light in a particular way to then appear as a color to our eyes<ref name=":4">{{Cite journal |last=TREVISAN |first=PIERLUIGI |last2=PEDERZOLI |first2=AURORA |last3=BAROZZI |first3=GIANCARLO |date=1991-10 |title=Pigmentary System of the Adult Alpine Salamander Salamandra atra atra (Laur., 1768) |url=http://dx.doi.org/10.1111/j.1600-0749.1991.tb00432.x |journal=Pigment Cell Research |volume=4 |issue=4 |pages=151–157 |doi=10.1111/j.1600-0749.1991.tb00432.x |issn=0893-5785}}</ref><ref>{{Cite journal |last=PEDERZOLI |first=AURORA |last2=TREVISAN |first2=PIERLUIGI |date=1989-03 |title=Pigmentary System of the Adult Alpine Salamander Salamandra atra aurorae (Trevisan, 1982) |url=http://dx.doi.org/10.1111/j.1600-0749.1989.tb00266.x |journal=Pigment Cell Research |volume=3 |issue=2 |pages=80–89 |doi=10.1111/j.1600-0749.1989.tb00266.x |issn=0893-5785}}</ref>. In ''S. atra'', there are different cells present or activated, which yield different colors: melanophores contribute to the dark coloration by producing the dark pigment melanin<ref name=":4" />, while xanthophores produce a yellow pigment<ref name=":2" />, and iridophores are simply light-reflecting<ref name=":2" />. The fully-black phenotype seen in ''S. a. atra'' results from the salamanders' melanophores in the dermis and epidermis, producing melanin alone<ref name=":3" /><ref name=":4" />. Xanthophore-iridophore complexes are responsible for production of yellow spotting, which appears bright<ref name=":3" />. In species without yellow patches, it appears that they do not ever develop these cells<ref name=":3" />. In ''S. a. aurorae'' or other salamanders with different coloration on different parts of their body, two distinct skin types are present: one that only contains melanophores (black), and one that has melanophores, xanthophores, and iridophores in combination<ref name=":4" />.
+==Distribution==
+The alpine salamander is found from the Swiss-French border at the western end of its range, all the way through Austria to the [[Dinaric Alps]] at the eastern edge of its territory. This salamander typically lives at altitudes above 700 m (2,300 ft) above sea level, even reaching 2000 meters (6561 ft) of elevation<ref name=":0" />. The [[western Alps]] (in France and Italy) are inhabited by a similar species, [[Lanza's alpine salamander]] (Salamandra lanzai), in only one small area[citation needed]. ''S. atra'' generally live in forested biomes, particularly deciduous-coniferous mixes<ref name=":0" />. They also can inhabit meadows or grasslands in the mountains<ref name=":0" />, and tend to do well with a mix of tree types<ref name=":5">{{Cite journal |last=Gautier |first=Patrick G. |last2=Miaud |first2=Claude |date=2003-01-01 |title=Faecal pellets used as an economic territorial marker in two terrestrial alpine salamanders |url=https://doi.org/10.1080/11956860.2003.11682759 |journal=Écoscience |volume=10 |issue=2 |pages=134–139 |doi=10.1080/11956860.2003.11682759 |issn=1195-6860}}</ref><ref name=":6">{{Cite journal |last=Romano |first=Antonio |last2=Anderle |first2=Matteo |last3=Forti |first3=Alessandro |last4=Partel |first4=Piergiovanni |last5=Pedrini |first5=Paolo |date=2018-12-31 |title=Population density, sex ratio and body size in a population of Salamandra atra atra on the Dolomites |url=https://oaj.fupress.net/index.php/ah/article/view/1840 |journal=Acta Herpetologica |language=en |pages=195–199 Pages |doi=10.13128/ACTA_HERPETOL-22592}}</ref>.
+Their range spans several nations, including: Slovenia, Croatia, Bosnia Hersegovina, Montenegro, Kosovo, france, Italy, Austria.
 ==Subspecies==
-*''S. a. atra'' is a fully melanistic (black) subspecies from central, eastern and Dinaric Alps.<ref>{{cite web |title=Salamandre noir |url=http://www.karch.ch/karch/home/amphibien/amphibienarten-der-schweiz/alpensalamander.html |website=Karch.Ch (Swiss information center for amphibians and reptiles) |accessdate=29 June 2020}}</ref>
+*''S. a. atra'' is a fully melanistic (black) subspecies from central, eastern and Dinaric Alps<ref>{{cite web |title=Salamandre noir |url=http://www.karch.ch/karch/home/amphibien/amphibienarten-der-schweiz/alpensalamander.html |website=Karch.Ch (Swiss information center for amphibians and reptiles) |accessdate=29 June 2020}}</ref>. This subspecies is the most widespread.
-*''[[Salamandra atra aurorae|S. a. aurorae]]'', the golden alpine salamander, has golden or yellow spots on its back and lives in a small area in the [[Venetian prealps|Venetian Prealps]] near [[Asiago]].
+*''[[Salamandra atra aurorae|S. a. aurorae]]'', the golden alpine salamander, has golden or yellow spots on its back and primarily lives in a small area in the [[Venetian prealps|Venetian Prealps]] near [[Asiago]], and in the Italian Alps.[citation needed]
-*''[[Salamandra atra pasubiensis|S. a. pasubiensis]]'', with less yellow spots than ''S. a. aurorae'', lives in a different part of the [[Venetian Prealps]].
+*''[[Salamandra atra pasubiensis|S. a. pasubiensis]]'', with fewer yellow spots than ''S. a. aurorae'', lives in a different part of the [[Venetian Prealps]].
-*''S. a. prenjensis'' lives on [[Prenj]] Mountain, part of the Dinaric Alps in [[Bosnia and Herzegovina]]. The validity of this subspecies is yet to be confirmed.<ref name=bonato2001>Bonato & Steinfartz. Evolution of the melanistic color in the Alpine salamander ''Salamandra atra'' as revealed by a new subspecies from the Venetian prealps. Italian Journal of Zoology 2001, 72, 253-260.</ref>
+*''S. a. prenjensis'' lives on [[Prenj]] Mountain, part of the Dinaric Alps in [[Bosnia and Herzegovina]]. The validity of this subspecies is yet to be confirmed, and some scientists in the field debate if this salamander should be considered its own independent species.<ref name=":0" /><ref name=bonato2001>Bonato & Steinfartz. Evolution of the melanistic color in the Alpine salamander ''Salamandra atra'' as revealed by a new subspecies from the Venetian prealps. Italian Journal of Zoology 2001, 72, 253-260.</ref>
 Genetic analysis suggests that the [[Corsican fire salamander]] (''Salamandra corsica'') is the closest related species, and the black-yellow coloration is an ancestral feature of alpine salamanders. Proposed colonization from south (Prealps) to Alps was carried out by the fully melanistic (derived feature) ''S. a. atra'' after the last retreat of the ice sheets.<ref name=bonato2001/>
@@ Line 28: / Line 40: @@
 [[File:Salamandra atra aurorae TREVISAN, 1982.jpg|thumb|''S. a. aurorae'' (Golden alpine salamander)]]
-==Reproduction==
+==Behavior==
-Mating occurs on land, the male clasping the female at the forelegs, and fertilization is internal. ''S. atra'' is a [[viviparity|viviparous]] [[amphibian]], giving birth to two live young, or rarely three or four. They may measure as long as {{convert|50|mm|abbr=on}} at birth, with the female measuring only {{convert|120|mm|abbr=on}}. The uterine [[egg (biology)|eggs]] are large and numerous, but, as a rule, only one fully develops in each uterus, the embryo being nourished on the yolk of the other eggs, which more or less dissolve to form a large mass of nutrient matter. The embryo passes through three stages:<ref>{{Cite EB1911|wstitle=Salamander|volume=1}}</ref>
+=== Habitat and Environmental Preferences ===
-#still enclosed within the egg and living on its own yolk
+S. atra tend to live underneath stones or logs, or in rocky crevices in their mountainous habitat<ref name=":0" />. They also are diurnal, and most active in the day with periods of inactivity, rest, or sleep at night<ref name=":6" />. They will engage in nocturnal activity on a weather-dependent basis<ref name=":7">{{Cite journal |last=Helfer |first=V. |last2=Broquet |first2=T. |last3=Fumagalli |first3=L. |date=2012-10 |title=Sex-specific estimates of dispersal show female philopatry and male dispersal in a promiscuous amphibian, the alpine salamander ( Salamandra atra ) |url=https://onlinelibrary.wiley.com/doi/10.1111/j.1365-294X.2012.05742.x |journal=Molecular Ecology |language=en |volume=21 |issue=19 |pages=4706–4720 |doi=10.1111/j.1365-294X.2012.05742.x}}</ref>. Ideal weather for alpine salamanders is rainy or post-rain, at temperatures between 3 and 18C (around 37-64 F)<ref name=":6" />.
-#free, within the [[wiktionary:vitelline|vitelline]] mass, which is directly swallowed by the mouth
-#with no more vitelline mass, the embryo is possessed of long external gills, which serve as an exchange of nutritive fluid through the maternal uterus, these gills functioning in the same way as the [[chorionic villi]] of the mammalian egg
+=== Territoriality ===
+They thrive in forest environments that have silver firs and beech trees. Coniferous forests that have high proportions of Norway spruces and European larch trees also provide adequate habitats, even though the salamanders live on the ground floor<ref name=":5" /> . Because the salamanders are completely terrestrial, they have on-land territories that they tend to return to throughout the day and for refuge. They often return to the same sites for much of their lives<ref name=":5" />.Anytime that they leave their sites, they expose themselves to predation and also to the chance of losing their site(). Alpine salamanders are ectothermic, so losing a refuge or shelter could leave them exposed to the elements and be extremely costly, if not fatal, do their lack of an internal thermoregulation mechanism().  This high dependency on a quality nest site supportsthe theory that many terrestrial salamanders, including S''. atra'', engage in territorial behaviors<ref name=":5" /> .They employ scent-marking techniques to this, and to mark their territories using fecal pellets so they can identify their own shelters<ref name=":5" />. Scent-marking is an intra-species communication, where chemical signals convey specific messages to other S. atra individuals<ref name=":5" />. Alpine salamanders can determine if a ound fecal pellet has been left by a member of their same sex and/or species<ref name=":5" />. Thus, this technique serves a double purpose to warn other salamanders that that particular location has already been claimed<ref name=":5" />. Females are more likely to return to their home site, while males are more emboldened to enter another male’s territory<ref name=":5" />. Their fecal pellets allow them to both participate in homing behavior, or returning to their own site, and territoriality, and determine intruders on their territory or invade the spaces of others<ref name=":5" />.
+=== Male-Male Behavior ===
+Males are more likely to engage in chasing other males, as well as actually fighting with each other<ref name=":8">{{Cite journal |last=Di Nicola |first=Matteo Ricardo |date=November 2022 |title=Male-male interactions in Alpine salamanders, Salamandra atra atra Laurenti, 1768, with an overview of the main cases reported for the whole genus Salamandra Garsault, 1764. |journal=Herpetology Notes |volume=Volume 15 |pages=601-604}}</ref>. Oftentimes, one male will mount the other, loosely grasp it with his forearms, and start rubbing his head on the other male<ref name=":8" />. The two males will switch roles, and in one studied interaction, continued like this for seven minutes before parting ways<ref name=":8" />. In a second documented fight, the behavior was more intense. When one member tried to leave, the other male chased it to re-engage<ref name=":8" />. This encounter lasted eight minutes. There is video evidence of these behaviors<ref name=":8" />. Researchers are unsure if it is caused by territoriality, confusion on sexual identificationand mistaken mating, or true combat<ref name=":8" />.
+=== Diet ===
+Male and female alpine salamanders have relatively similar diets<ref name=":9">{{Cite journal |last=Roner |first=Luca |last2=Costa |first2=Andrea |last3=Pedrini |first3=Paolo |last4=Matteucci |first4=Giorgio |last5=Leonardi |first5=Stefano |last6=Romano |first6=Antonio |date=2020-05 |title=A Midsummer Night’s Diet: Snapshot on Trophic Strategy of the Alpine Salamander, Salamandra atra |url=https://www.mdpi.com/1424-2818/12/5/202 |journal=Diversity |language=en |volume=12 |issue=5 |pages=202 |doi=10.3390/d12050202 |issn=1424-2818}}</ref>. Some specific organisms they prey on include species such as beetles, snails, millipedes, and spiders<ref>{{Cite journal |last=Šunje |first=Emina |last2=Courant |first2=Julien |last3=Vesnić |first3=Adi |last4=Koren |first4=Toni |last5=Lukić Bilela |first5=Lada |last6=Van Damme |first6=Raoul |date=2022-09-20 |title=Patterns of variation in dietary composition among four populations of Alpine salamanders (Salamandra atra prenjensis) |url=http://dx.doi.org/10.1163/15685381-bja10100 |journal=Amphibia-Reptilia |pages=1–15 |doi=10.1163/15685381-bja10100 |issn=0173-5373}}</ref>, but alpine salamanders display preferences among prey<ref name=":9" />. S. atra typically consume organisms from the Coloeptera and mollusca taxa<ref name=":9" />. These taxa are the most crucial component of their diet (). They also tend to eat larger prey since they themselves are larger salamanders<ref name=":9" />. Though alpine salamanders have definite dietary preferences, they have a substantial amount of variation in their diet<ref name=":9" />that corresponds to their own optimized physical needs and pre-catching abilities.
+=== Mating: ===
+The alpine salamander engages in a promiscuous mating pattern<ref name=":7" />, meaning that they engage in multiple partner pairings. Males immigrate and travel farther than females do, potentially to find minds and following a scent emitted by females<ref name=":7" />. Typically, males migrated when they were in the juvenile stage of life<ref name=":7" />. Female S. atra find and defend their shelters, which is a potential reason that they may stay more local than males<ref name=":7" />.
+== Gestation and Reproduction ==
+Alpine Salamanders live in a sex ratio of 1:1.<ref>{{cite book |title=Body size, population structure and fecundity traits of Salamandra atra atra (Amphibia, Urodela, Salamandridae) population from the northeastern Italian Alps |publisher=Italian Journal of Zoology |year=2001 |volume=68 |location=Luiselli, Andreone, [[Capizzi]], Anibaldi |pages=125–130}}</ref> . Mating occurs on land. The male clasps the female at the forelegs, and fertilization is internal. ''S. atra'' are categorized as viviparous, meaning that their young are born alive and unlike many other amphibians, do not go through metamorphosis (De meester). They give birth to 2 young, sometimes 3 or 4. New young alpine salamanders may measure as long as 50 mm (2.0 in) at birth, with the mother measuring only 120 mm (4.7 in).
+Female alpine salamanders have uteruses that are composed of a single luminal epithelial cell layer, connective tissue, and smooth muscle<ref name=":10">{{Cite journal |last=Guex |first=G. -D. |last2=Chen |first2=P. S. |date=1986-12 |title=Epitheliophagy: Intrauterine cell nourishment in the viviparous alpine salamander,Salamandra atra (Laur.) |url=http://dx.doi.org/10.1007/bf01946392 |journal=Experientia |volume=42 |issue=11-12 |pages=1205–1218 |doi=10.1007/bf01946392 |issn=0014-4754}}</ref>.The uterine eggs are large and numerous, but, as a rule, only one fully develops in each uterus. The embryo is nourished on the yolk of the other eggs, which more or less dissolve to form a large mass of nutrient matter. The egg mass can be as long as between 25 mm and 40 mm long<ref name=":10" />. The embryo passes through three stages<ref name=":11">{{Cite EB1911|wstitle=Salamander|volume=1}}</ref>:
+# The first stage is when they are still enclosed within the egg and living on its own yolk.<ref name=":11" />
+# The second stage is when they are free, within the vitelline mass, eating it directly with their mouths.<ref name=":10" /><ref name=":11" />
+# The final stage is where there is no more vitelline mass. The embryo is possessed of long external gills, which serve as an exchange of nutritive fluid through the maternal uterus, these gills functioning in the same way as the chorionic villi of the mammalian egg.<ref name=":10" /><ref name=":11" />
+Generally, at altitudes of 650-1,000 m above sea level, a pregnancy lasts two years, and at altitudes of 1,400-1,700 m above sea level, the pregnancy lasts around three years, though anything within a 2-4 year range is considered standard<ref name=":10" />. Alpine salamander embryos are unique in how they are able to take in these nutrients through a long gestation<ref name=":10" />. A portion of the mother’s uterine wall becomes nourishment after the salamanders have already eaten the unfertilized eggs<ref name=":10" /> (called oophagy<ref name=":10" />, or stage 1 and 2). They ten partake in epitheliophagy, or stage 3, where they ingest these zona trophica cells until birth, and have special tooth-like developments that allow it to do so without detriment to the mother<ref name=":10" />.
+== Physiology ==
+=== Glands and Toxins ===
+As mentioned above, alpine salamanders have poison glands. They are known to produce some alkaloid molecules and peptide products, and thus have a mustard-like scent associated with them<ref>{{Cite journal |last=Lüddecke |first=Tim |last2=Schulz |first2=Stefan |last3=Steinfartz |first3=Sebastian |last4=Vences |first4=Miguel |date=2018-09-04 |title=A salamander’s toxic arsenal: review of skin poison diversity and function in true salamanders, genus Salamandra |url=http://dx.doi.org/10.1007/s00114-018-1579-4 |journal=The Science of Nature |volume=105 |issue=9-10 |doi=10.1007/s00114-018-1579-4 |issn=0028-1042}}</ref>. Salamandra bio production is still a developing research area. Salamandarines are a chemical secretion produced by the skin of alpine salamanders, as well as some fire salamanders (De Meester)<ref name=":0" /><ref>{{Cite journal |last=De Meester |first=Gilles |last2=Šunje |first2=Emina |last3=Prinsen |first3=Els |last4=Verbruggen |first4=Erik |last5=Van Damme |first5=Raoul |date=2021-05 |title=Toxin variation among salamander populations: discussing potential causes and future directions |url=https://onlinelibrary.wiley.com/doi/10.1111/1749-4877.12492 |journal=Integrative Zoology |language=en |volume=16 |issue=3 |pages=336–353 |doi=10.1111/1749-4877.12492 |issn=1749-4877}}</ref>. They are neurotoxins, and are syntehtised via a biochemical pathway independent of dietary intake<ref name=":0" />. The starting material for this nerve-block is most likely cholesterol, and it is about twice as potent as cyanide<ref name=":0" /> . This pales in comparison to other toxins produced by salamanders, but ''S. atra'' do not only use this powerful substance to paralyze prey: they may have antimicrobial properties that protect them against bacterial and fungal infections<ref name=":0" /><ref name=":12">{{Cite book |last=R |first=Lötters, S Kielgast, J Sztatecsny, M Wagner, N Schulte, U Werner, P Rödder, D Dambach, J Reissner, T Hochkirch, A Schmidt, B |url=http://worldcat.org/oclc/1030045649 |title=Absence of infection with the amphibian chytrid fungus in the terrestrial Alpine salamander, Salamandra atra |date=2012-04-30 |publisher=Deutsche Gesellschaft für Herpetologie und Terrarienkunde (DGHT) |oclc=1030045649}}</ref>. Salamandorone is another biochemical compound produced by S. atra, and though it is less potent against prey it is the strongest antimicrobial weapon these salamanders have<ref name=":0" /><ref>{{Cite journal |last=Spitzen - van der Sluijs |first=Annemarieke |last2=Stegen |first2=Gwij |last3=Bogaerts |first3=Sergé |last4=Canessa |first4=Stefano |last5=Steinfartz |first5=Sebastian |last6=Janssen |first6=Nico |last7=Bosman |first7=Wilbert |last8=Pasmans |first8=Frank |last9=Martel |first9=An |date=2018-02-28 |title=Post-epizootic salamander persistence in a disease-free refugium suggests poor dispersal ability of Batrachochytrium salamandrivorans |url=http://dx.doi.org/10.1038/s41598-018-22225-9 |journal=Scientific Reports |volume=8 |issue=1 |doi=10.1038/s41598-018-22225-9 |issn=2045-2322}}</ref><ref name=":12" />.
+=== Immunobiology and Protection ===
+The alpine salamander has been relatively lucky in avoiding infection with amphian chytrid fungus<ref name=":12" /> compared to other amphibian species<ref name=":12" /><ref>{{Cite journal |last=Feldmeier |first=Stephan |last2=Schefczyk |first2=Lukas |last3=Wagner |first3=Norman |last4=Heinemann |first4=Günther |last5=Veith |first5=Michael |last6=Lötters |first6=Stefan |date=2016-10-31 |title=Exploring the Distribution of the Spreading Lethal Salamander Chytrid Fungus in Its Invasive Range in Europe – A Macroecological Approach |url=https://journals.plos.org/plosone/article?id=10.1371/journal.pone.0165682 |journal=PLOS ONE |language=en |volume=11 |issue=10 |pages=e0165682 |doi=10.1371/journal.pone.0165682 |issn=1932-6203 |pmc=PMC5087956 |pmid=27798698}}</ref>. This dangerous fungal infection, caused by Batrachochytrium dendrobatidis (Bd) has decimated amphibian populations on every continent<ref name=":12" />. Bd is present in the Alps where alpine salamanders live, but in a study performed in 2012, there were no salamanders who tested positive when swabbed<ref name=":12" />. This may be because Bd infections are more common in species who spend more of their time in water, and since the alpine salamanders are terrestrial, they are less susceptible<ref name=":12" />. An alternate hypothesis proposes that S. atra are resistant via their skin microbiome or a produced molecule, thus granting them immunity<ref name=":12" /><ref name=":13">{{Cite journal |last=Woodhams |first=D. C. |last2=Rollins-Smith |first2=L. A. |last3=Alford |first3=R. A. |last4=Simon |first4=M. A. |last5=Harris |first5=R. N. |date=2007-11 |title=Innate immune defenses of amphibian skin: antimicrobial peptides and more |url=https://onlinelibrary.wiley.com/doi/10.1111/j.1469-1795.2007.00150.x |journal=Animal Conservation |language=en |volume=10 |issue=4 |pages=425–428 |doi=10.1111/j.1469-1795.2007.00150.x |issn=1367-9430}}</ref>. This theory is untested, but considering how many salamanders excrete biological toxins, plausible<ref name=":12" /><ref name=":13" />.
+== Conservation ==
-Generally, at altitudes of 650-1,000 m above sea level, a pregnancy lasts two years, and at altitudes of 1,400-1,700 m, the pregnancy lasts around three years (1,140 days).
+Alpine salamanders are not resiliant to habitat change, so risks of climate change altering their living spaces is severe<ref name=":1" />. Though the IUCN reports that alpine salamanders are least vulnerable, their numbers are decreasing<ref>{{Cite web |last=IUCN |date=7 July 2021 |title=The IUCN Red List of Threatened Species. |url=https://www.iucnredlist.org/species/19843/89706038 |website=” IUCN Red List of Threatened Species}}</ref>. Additionally, some subspecies of ''S. atra'' are in greater danger<ref name=":14">{{Cite journal |last=Romano |first=Antonio |last2=Costa |first2=Andrea |last3=Salvidio |first3=Sebastiano |last4=Menegon |first4=Michele |last5=Garollo |first5=Elena |last6=Tabarelli de Fatis |first6=Karol |last7=Miserocchi |first7=Danio |last8=Matteucci |first8=Giorgio |last9=Pedrini |first9=Paolo |date=2018-09 |title=Forest management and conservation of an elusive amphibian in the Alps: Habitat selection by the Golden Alpine Salamander reveals the importance of fine woody debris |url=http://dx.doi.org/10.1016/j.foreco.2018.04.052 |journal=Forest Ecology and Management |volume=424 |pages=338–344 |doi=10.1016/j.foreco.2018.04.052 |issn=0378-1127}}</ref>. Population numbers are declining in S. atra aurorae, for example. One of the greatest dangers to alpine salamanders overall is commercial deforestation in their habitats<ref name=":14" />. Machinery like tractors or other forestry tools can compress the soil eliminating some of the small insects that ''S. atra'' eat or eliminating potential nooks and burrows for them to use as shelters<ref name=":14" />. Many scientists propose changes in the lumber industry as an attempt to heal these habitats<ref name=":14" />. These salamanders may also change their morphology as global temperatures rise<ref name=":15">{{Cite journal |last=Ficetola |first=Gentile Francesco |last2=Colleoni |first2=Emiliano |last3=Renaud |first3=Julien |last4=Scali |first4=Stefano |last5=Padoa-Schioppa |first5=Emilio |last6=Thuiller |first6=Wilfried |date=2016-06 |title=Morphological variation in salamanders and their potential response to climate change |url=https://onlinelibrary.wiley.com/doi/10.1111/gcb.13255 |journal=Global Change Biology |language=en |volume=22 |issue=6 |pages=2013–2024 |doi=10.1111/gcb.13255 |pmc=PMC4972144 |pmid=26910389}}</ref>. Amphibians and other organisms that do not internally regulate their body temperature may need adaption mechanisms to remain at ideal physiological temperatures in the face of changing climates<ref name=":15" />. Other issues like acid rain or precipitation changes could prompt many animals, including alpine salamanders, to be forced into new habitats<ref name=":15" />. Alpine salamanders play a crucial role in their ecosystems<ref>{{Cite journal |last=Reinthaler-Lottermoser, et al. |first=Ursula |date=8 October 2018 |title=A new approach for surveying the Alpine Salamander (Salamandra atra) in Austria |journal=Acta Herpetologic |pages=249-253}}</ref>. There are already animal and ecosystem conservation laws in Europe, but many scholars recommend additional ones to protect the flora and fauna<ref>{{Cite journal |last=Scalera |first=Riccardo |date=2004-07-01 |title=The legal framework for the protection of amphibians and reptiles in Italy: an annotated overview of the provisions at the international, European community, national and regional level |url=https://doi.org/10.1080/11250003.2004.9525560 |journal=Italian Journal of Zoology |volume=71 |issue=sup002 |pages=21–32 |doi=10.1080/11250003.2004.9525560 |issn=1125-0003}}</ref>.
 ==References==