Description
Salamandra salamandra is a large, conspicuous salamander that grows up to 250 mm in body length, sometimes almost 300 mm, and has conspicuous parotoid glands. The tail is cylindrical and shorter than body with head.

In life, the conspicuous parotoid glands behind eyes are pigmented. Typically, the dorsal and lateral skin is black, with large yellow to orange spots and/or bands. Yellow pattern varies among subspecies (see variation below), and with individual variation so that it is not entirely reliably for subspecies identification. The belly skin is black or brownish.

Salamandra salamandra is widely distributed with geographically specific coloration variation. As a result, 12 - 17 subspecies have been proposed, with the majority of the variation occurring in the Iberian Peninsula (Burgon et al. 2021). The subspecies with the most phylogenetic support are: S. s. almanzoris, S. s. bejarae, S. s. bernardezi, S. s. crespoi, S. s. fastuosa, S. s. gigliolii, S. s. "hispanica", S. s. longirostris, and S. s. morenica, with S. s. gallaica, S. s. salamandra, S. s. terrestris and S. s. werneri being paraphyletic (see Comments section for more information). Salamandra s. almanzoris is found at Sierra de Gredos and Sierra de Guadarrama in central Spain, and is small with a reduced yellow dorsal stripe on a back dorsum. Salamandra s. bejarae is found in central and central­ eastern Meseta areas in Spain in dry highlands, and is small with irregular yellow dorsal spots on a black dorsum, a pointed snout, and a short, high tail. Salamandra s. bernardezi can be found in the Asturia, western Cantabria and north­eastern Galicia regions of Spain, and is small with a mostly yellow dorsum with a black stripe. However, the dorsum may also be dirty grey-­yellow, grey, brown, orange-­brown or olive­green. This subspecies may also be mistaken for S. s. fastuosa. Salamandra s. crespoi is found between Serra de Monchique in southern Portugal to to the Serra de Grandola in the north and east to the Spanish border. They are large with many small yellow spots on a black dorsum, long tail, long limbs, long fingers, and small paratoids. Salamandra s. fastuosa is found in the in the western and central Pyrenees and has a range overlap with S. s. bernardezi at eastern Asturia and western Cantabria, however the former is larger in size. Salamandra s. fastuosa has two bright yellow dorsolateral strips. Salamandra s. gigliolii is found in Italy and is long with extensive yellow coloration, almost masking the black background. They also have red throats and ventrums. Salamandra s. "hispanica" is found in the eastern Pyrenees. Salamandra s. longirostris is found in Cadiz and Malaga of southern Spanish, and is large with lemon-yellow, comma or horseshoe-shaped spots on a black dorsum, four large yellow spots on the head, and two yellow spots above the corners of the mouth. Salamandra s. morenica is found in the Sierra Morena, from the Portuguese boarder to Murcia, Spain. The species is large with red coloration and small yellow spots on a black dorsum. Among the paraphyletic subspecies Salamandra s. gallica is found in Northern Portugal, and is large with yellow, horseshoe or comma-shaped spots on a black background. Salamandra s. salamandra is the nominate species and can be found ranging from central Italy north throughout central Europe and the Balkan Peninsula. It has a stocky body with variable, irregular yellow spots that sometimes merge into bands. Salamandra s. terrestris is found in France and in the Pyrenees region. They are moderately sized with two either continuous our discontinuous yellow dorsolateral stripes. They can also be found with spots instead of stripes or with red coloration. Lastly, S. s. werneri are found in southern and central Greece, and are large with irregular small spots, sometime forming longitudinal rows, that can have red centers (Sparreboom 2014, Burgon et al. 2021).

Females are generally larger than males and possess relatively shorter extremities and tail. Male's cloaca much more swollen than female's cloaca.

Distribution and Habitat

Country distribution from AmphibiaWeb's database: Albania, Austria, Belgium, Bosnia and Herzegovina, Bulgaria, Croatia, Czech Republic, France, Germany, Greece, Hungary, Italy, Luxembourg, Macedonia, the Former Yugoslav Republic of, Montenegro, Netherlands, Poland, Portugal, Romania, Serbia, Slovakia, Slovenia, Spain, Switzerland, Turkey, Ukraine

	View distribution map in BerkeleyMapper.
	View Bd and Bsal data (3172 records).

The species is distributed from the Iberian Peninsula to Iran and from North Africa to North Germany. The genus consists of variable forms, the taxonomy of which has not been revised as yet. Some of the former subspecies of Salamandra salamandra are now recognized as separate species and the separation of more species can be expected. Populations of S. s. salamandra from Turkey are genetically closely related to the S. s. infraimmaculata group. The species inhabits mainly deciduous and mixed, sometimes conifer forests. Populations inhabiting anthropogenic landscapes and unforested habitats can be considered, as a rule, as relicts of formerly forest dwellers. The spotted coloration of this salamander seems to play two roles: cryptic, when the spots on black background allow the animal to hide on the forest floor, where there are alternate spots of sun and shadow, and aposematic, where bright yellow spots indicate poisonous skin secretions.

The Iberian Peninsula hosts a unique diversity of fire salamander (Salamandra salamandra) lineages (up to nine described subspecies). In northern Spain, the subspecies S. s. bernardezi represents a distinct lineage (closely related to a northern Spanish lineage, S. s. fastuosa, and the Italian S. s. gigliolii), with remarkable phenotypic diversity across its small range. The subspecies demonstrates the largest variety of colour patterns observed in any Salamandra species, ranging from entirely bright yellow, over the more typical black with yellow stripes, to entirely black-brown animals. Salamandra s. bernardezi is well known for its stunning reproductive shift, evolving from larviparity (females give birth to aquatic larvae) to pueriparity (females deliver fully terrestrial juveniles). Since all taxa within the genus Salamandra are considered highly vulnerable to Bsal chytridiomycosis, the small range of the genetic entities of S. s. bernardezi puts them at high risk of massive population declines and possibly extinction if Bsal would invade (Guillermo Velo-Antón, pers.com.).

Life History, Abundance, Activity, and Special Behaviors
Over a large part of its range, S. salamandra seems to be not a rare species, but its abundance declines in many regions. In unwooded areas the species is generally rarer than in forests.

Females are active in daytime during the breeding period; afterwards adults are active at twilight, spending the day under logs, snags, stones, rodent burrows and holes. During rainy weather salamanders regularly leave their hiding places by day. The appearance of active salamanders on the land surface in day time indicates the approach of rain. Hibernation, typically in groups, occurs in the northern part of the range, whereas in the south (e.g., in Israel) activity ceases during hot summer period. Similarly, in central Europe reproduction occurs between spring and autumn, whereas at the south of the range it is confined to winter. Mating takes place on land, and male-male combats for a female often takes place. The species is typically viviparous, and the female releases the young into water, usually shallow brooks. The number of larvae per female, as well as their stage at birth time, varies among subspecies. Salamandra salamandra bernardezi and, sometimes, S. s. fastuosa give birth to completely metamorphosed young salamanders. Larval development takes several months, but in many cases they overwinter and finish their metamorphosis in the next year. Most larvae occur in fishless parts of brooks, which is caused by fish predation. As a rule, larvae start active feeding just after birth. Age changes in diet during ontogeny are minor and related mainly to the use of larger prey. Larvae consume primarily upon rheophilous invertebrates: Gammaridae, larval Ephemeroptera, Diptera, etc. In semi-flowing waters, typical limnophilous preys (e.g., Diaptomidae) are included in their diet. Adults do not consume the small preys that are eaten by juveniles: Acarina, Geophylomorpha, and Collembola. However, they eat large Mollusca, Myriapoda (Oniscomorpha, Polydesmida and Juliformia), Coleoptera, etc.

Trends and Threats
In historical perspective, the range seemed to be constricted, mainly due to deforestation. In some places (e.g., in Ukrainian Carpathians) declines of populations take place due to anthropogenic influences.

Although the IUCN status has not been updated, since 2010, there has been a precipitous decline in the species bringing it close to extinction by 2013. A captive breeding problem resulted in 49% of the captive animals dying of, then, unknown causes. It was later determined that a newly identified chytrid fungus was infecting the species, causing anorexia, apathy, and ataxia then death within 7 - 27 days (depending on method of exposure). The new fungus was named Batrachochytrium salamandrivorans and is not known to infect frogs (Martel et al. 2013).

Relation to Humans
Habitat destruction, pollution, and collecting for commercial purposes (mainly pet trade) are the main threats for the populations. Destruction of forests and overcollecting cause the declines of some populations.

Possible reasons for amphibian decline

General habitat alteration and loss
Habitat modification from deforestation, or logging related activities
Long-distance pesticides, toxins, and pollutants
Disease
Intentional mortality (over-harvesting, pet trade or collecting)

Comments
The phylogenetic relationships in the family Salamandridae have a long history of confusion. Recent analyses (Bayesian Inference, Maximum Likelihood, Network Analysis, and Principal Components Analysis) on 4905 RAD-loci (nDNA) support the existence of six species: S. algira, S. atra, S. corsica, S. infraimmaculata, S. lanzai, and S. salamandra. These analyses indicate that S. salamandra is most closely related to S. algira and together they are most closely related to the clade formed by S. atra, S. corsica and S. lanzai (Burgon et al. 2021).

Burgon et al. (2021) also examined the genetic relationship between subspecies. The subspecies of S. salamandra have long been disputed with 12 - 17 potential subspecies, however, understanding their evolutionary history is vital to conservation management as the species as a whole is threatened by the chytrid fungus, Batrachochytrium salamandrivorans. Burgon et al.'s results found a major split with two northern Spanish and one Italian subspecies, S. s. bernardezi, S. s. fastuosa and S. s. gigliolii respectively, forming one clade, while all other subspecies formed another. The analyses supported the monophyly of S. s. bernardezi, S. s. fastuosa and S. s. gigliolii, but merged S. s. alfredschmidti into S. s. bernardezi. Of the remaining subspecies, the monophyly of S. s. almanzoris, S. s. bejarae, S. s. crespoi, S. s. longirostris, S. s. morenica, and the disputed S. s. hispanica were supported. Salamandra s. almanozoris was placed basally in this clade. Salamandra s. gallaica, S. s. salamandra, S. s. terrestria, and S. s. werneri were all found to be paraphyletic with S. s. molleri merged into S. s. gallaica, and S. s. beschkovi and S. s. carpathica merged into S. s. salamandra. The analyses also indicate some introgression between subspecies, but did not identify where introgression zones are located. The study also lacked samples from France, which may complicate these findings. However, these questions are being investigated.

This is the first species from which Batrachochytrium salamandrivorans (Bsal) infections have been identified (Martel et al. 2013).

This species was featured as News of the Week on 24 June 2019:

Although Fire Salamanders are assumed to be an aposematic species, where bright colors are viewed by potential predators as warning of toxicity, the costs associated with the honesty of their signaling and predator learning behavior has not been quantified. Preißler et al. (2019) tested whether the alkaloid content matched yellow coloration in a highly variable Fire Salamander population in Solling, Germany, (Salamandra salamandra terrestris) to determine if the species had an honest signal of toxicity. They found that the amount of yellow patterning did not correlate with the amount of toxins in individuals, as would be expected for a true aposematic species. The authors also found that the population was sexually dichromatic, with males being more yellow than females. Although these findings could be explained by sexual selection, statistical analysis of color variation indicated that site specific female choice was not a factor. These findings indicate that other, yet to be verified, biological processes are playing a role in the yellow coloration of Fire Salamanders, and that the toxins are produced by conserved bioactivity (Written by Ann T. Chang).

This species was featured as News of the Week on 17 August 2020:

The pathogenic amphibian fungus known as Bsal (Batrachochytrium salamandrivorans) may be the most potent amphibian disease and poses extreme risk to natural populations, especially in salamanders. First detected in Fire Salamanders (Salamandra salamandra) in extreme southeastern Netherlands and adjacent Belgium and reported in 2013, it has spread to western Germany (with new reports from Bavaria), where it is having devastating effects. An entire issue of the journal Salamandra (2020, vol 56, issue 3, open access and available as PDF) is devoted to Bsal research centered in Germany. Salamander populations have essentially disappeared from the northern Eiffel region and are threatened in the southern Eiffel and Ruhr regions. Bsal has been present in Germany for at least 16 years and has been found in laboratory populations of the Common Frog, Rana temporaria, and field populations of the Great Crested Newt, Triturus cristatus. It is known to infect salamandrid species from southeast Asia, which appear to have been the source of the European outbreaks via pet trade importation. The goal in highlighting this important set of papers as stated by the editors "must go beyond documenting declines towards understanding spatio-temporal disease dynamics and the factors influencing the spread and impact of Bsal in different situations." In light of the seriousness of the Bsal threat in Germany, the authors' common goal is a national Bsal Action Plan, which would be of great importance for the international community of amphibian biologists and for the public (Written by David B. Wake).

This species was featured as News of the Week on 6 September 2021:

The chytrid fungus Batrachochytrium salamandrivorans (Bsal) has recently emerged in Europe, where it is devastating salamander and newt populations. While the disease mainly affected northwestern Europe, it recently caused mass mortality in northern Spain. The Iberian peninsula is a hotspot of salamanders and newts, and in a joint effort between Spanish, Belgian and Australian scientists, Bosch et al. (2021) assessed the risk of Bsal for this salamander-rich region. Fortunately, Bsal appears to be limited to a single outbreak site at present. However, we identified six conservation units at high risk and priority areas as targets for the most urgent disease surveillance and biosecurity measures. The study stresses the importance of region specific conservation strategies, coupling efficient wildlife disease surveillance to the ability to rapidly and drastically respond, but equally points to the importance of "clean trade" (absence of pathogens from the amphibian trade), amphibian population monitoring and closing gaps of knowledge regarding salamander diversity. (Written by Frank Pasmans)

References

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Species Account Citation: AmphibiaWeb 2021 Salamandra salamandra: Fire Salamander <https://amphibiaweb.org/species/4284> University of California, Berkeley, CA, USA. Accessed Jan 27, 2022.

Citation: AmphibiaWeb. 2022. <https://amphibiaweb.org> University of California, Berkeley, CA, USA. Accessed 27 Jan 2022.

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