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Alpine salamander
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== Description == Alpine salamanders are often small in size, and dark brown or black.<ref name="DE MEESTER-2020" /> Members of the subspecies are not wholly black or brown monochrome, but rather have mosaic or spotted patterns.<ref name="DE MEESTER-2020" /> 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="DE MEESTER-2020" /> Distribution of the pigment is dependent on the distribution of certain cells, so may be smooth and even or patchy.<ref name="DE MEESTER-2020" /> Female ''S. atra'' tend to be larger than the males, and can grow up to 151 millimeters, or around 5.9 inches.<ref name="DE MEESTER-2020" /> Males will grow to around 144 millimeters, or 5.6 inches: both measurements include the tail.<ref name="DE MEESTER-2020" /> Males have swollen, visible [[cloacae]], and are more slender than females.<ref name="DE MEESTER-2020" /><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 gland]]s posterior and lateral to their eyes, giving them an elongated head shape.<ref name="DE MEESTER-2020" /> They tend to have between 11 and 13 costal grooves along the 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="DE MEESTER-2020" /> === Coloration === Most alpine salamanders that are either completely black ([[Melanism|melanistic]]) or predominantly black have the dark pigment as a baseline, but the evolution behind this dark coloring has a winding history.<ref name="Burgon-2020">{{Cite journal |last1=Burgon |first1=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=April 2020 |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 |pmid=32159878 |s2cid=212664862 |issn=0962-1083|doi-access=free |hdl=10261/234500 |hdl-access=free }}</ref> Scientists have studied the hypothesis of if the salamanders start completely black, or if they evolved like that over time.<ref name="Bonato-2005">{{Cite journal |last1=Bonato |first1=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 |journal=Italian Journal of Zoology |volume=72 |issue=3 |pages=253–260 |doi=10.1080/11250000509356680 |s2cid=83504324 |issn=1125-0003}}</ref> DNA-evidence traced through maternal lineages suggest the latter: that salamanders evolved their black coloration over time.<ref name="Bonato-2005" /> [[Biological pigmentation]] is determined by presentation of specific color-producing cells, called [[chromatophores]], which absorb and/or reflect [[light]] in a particular way to then appear as a color.<ref name="TREVISAN-1991">{{Cite journal |last1=TREVISAN |first1=PIERLUIGI |last2=PEDERZOLI |first2=AURORA |last3=BAROZZI |first3=GIANCARLO |date=October 1991 |title=Pigmentary System of the Adult Alpine Salamander Salamandra atra atra (Laur., 1768) |journal=Pigment Cell Research |volume=4 |issue=4 |pages=151–157 |doi=10.1111/j.1600-0749.1991.tb00432.x |pmid=1816547 |issn=0893-5785}}</ref><ref>{{Cite journal |last1=PEDERZOLI |first1=AURORA |last2=TREVISAN |first2=PIERLUIGI |date=March 1989 |title=Pigmentary System of the Adult Alpine Salamander Salamandra atra aurorae (Trevisan, 1982) |journal=Pigment Cell Research |volume=3 |issue=2 |pages=80–89 |doi=10.1111/j.1600-0749.1989.tb00266.x |pmid=2385569 |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="TREVISAN-1991" /> while xanthophores produce a yellow pigment,<ref name="Burgon-2020" /> and iridophores are simply light-reflecting.<ref name="Burgon-2020" /> The fully-black phenotype seen in ''S. a. atra'' results from the salamanders' melanophores in the [[dermis]] and [[epidermis]], producing melanin alone.<ref name="Bonato-2005" /><ref name="TREVISAN-1991" /> Xanthophore-iridophore complexes are responsible for production of yellow spotting, which appears bright.<ref name="Bonato-2005" /> In species without yellow patches, it appears that they do not ever develop these cells.<ref name="Bonato-2005" /> 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="TREVISAN-1991" /> The yellow coloring on some alpine salamanders is thought to be an [[aposematic]] strategy to fend off predation.<ref name="Vences-2014">{{Cite journal |last1=Vences |first1=Miguel |last2=Sanchez |first2=Eugenia |last3=Hauswaldt |first3=J. Susanne |last4=Eikelmann |first4=Daniel |last5=Rodríguez |first5=Ariel |last6=Carranza |first6=Salvador |last7=Donaire |first7=David |last8=Gehara |first8=Marcelo |last9=Helfer |first9=Véronique |last10=Lötters |first10=Stefan |last11=Werner |first11=Philine |last12=Schulz |first12=Stefan |last13=Steinfartz |first13=Sebastian |date=2014-04-01 |title=Nuclear and mitochondrial multilocus phylogeny and survey of alkaloid content in true salamanders of the genus Salamandra (Salamandridae) |url=https://www.sciencedirect.com/science/article/pii/S1055790313004442 |journal=Molecular Phylogenetics and Evolution |language=en |volume=73 |pages=208–216 |doi=10.1016/j.ympev.2013.12.009 |pmid=24412216 |issn=1055-7903|url-access=subscription }}</ref> The pure black coloring is also hypothesized to be a form of [[thermoregulation]],<ref name="Vences-2014" /> though it may also be considered a warning to some predators.<ref name="Vences-2014" />
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