Fire salamander (Salamandra salamandra sp)
Species: Fire salamander (Salamandra salamandra)
First description: 1758 by Carl von Linné (Salamandra salamandra salamandra), 1789 by Pierre Bonnaterre (Salamandra salamandra terrestris)
Size / Length: average 14-18 cm, max. 24 cm, the ♀ usually grows slightly larger than the ♂.
Weight: average 16-26 g, max. 56 g
Age: 20-50 years (24 years according to AnAge Database of Animal Ageing and Longevity; > 50 years in terrarium according to Böhme 1979 )
Striking black and yellow spot pattern, which serves as a warning and defense against predators. In rare cases with black-orange-red or black-red spot pattern, extremely rare entirely black or albinotic or partially albinotic fire salamanders. The coloration variations originate from at least one anomaly in pigment synthesis. The proportion of red-colored fire salamanders varies from area to area and may be as low as 0%. In common parlance, every 1000th fire salamander exhibits red coloration. In the Stuttgart region, red-spotting is apparently more common (Schweizerbarth 1906 , 1909 , Vogel 1938 , Freytag 1955 ). In Strohgäu (Böblingen district, Baden-Württemberg), there is a fire salamander population with up to 2-3% red-spotted individuals (Rimpp 1992 ). Red-colored fire salamanders are also bred, so a population with exclusively red-colored individuals would be theoretically possible. The spot pattern characteristic of each salamander is not differentiated until about 1.5-2 years after metamorphosis, or from a body length of about 10 cm (Fachbach 1973 ). In younger specimens, the yellow spots are drawn out or apart due to vigorous growth. A documentation shows this change of the color pattern of a young fire salamander within a little more than 5 months (compare Figure 2 / Hillenius 1968 ).
Neotenic or pedomorphic or partially neotenic fire salamanders, which retain their gills in the adult state and do not undergo metamorphosis, have already been documented. As with other caudates, neoteny in fire salamanders is often linked to partial albinism. During investigations of a population in the Wuppertal/Remscheid area (North Rhine-Westphalia) in 1979, a strikingly bright larva was captured and transferred to an aquarium. In the aquarium the larva reached a length of almost 11 cm, until it died in spring 1982 due to severe fungal growth. During the whole time there were no signs of metamorphosis. The conditions of keeping are not the cause, because other larvae in the same tank metamorphosed normally after some time. Since fire salamanders become sexually mature at about 11 to 12 cm in length under field conditions, the present specimen could be described as neoten with reservations. Whether the animal was “stably neoten” can no longer be clarified. Already in the years before 1979 two possibly potentially neotenic individuals were found and captured in the same area. Both individuals died soon after being transferred to the aquarium (Klewen 1982 ). An 8 cm long, fully colored larva with partial neoteny from a shale tunnel in the Hunsrück is also known (Veith 1985 ). Breeding of neotenous fire salamanders has not been successful or is unknown.
The ♀ usually grows slightly larger than the ♂. Adult ♂ with bulging cloaca and ♀ with flat cloaca. During winter, the cloacae of the ♂♂ are often flat as well. The ♂ tend to have a greater proportion of yellow within the same population (Preißler et al. 2019 ).
Subspecies in Germany, Austria and Switzerland
In Germany, Austria and Switzerland the „barred fire salamander“ (Salamandra salamandra terrestris) and the „spotted fire salamander“ (Salamandra salamandra salamandra) can be found. The pattern of the “Spotted Fire Salamander” is irregular and never shows approaches to continuous dorsal stripes. The pattern on the back of the „barred fire salamander“, on the other hand, shows two yellow lines, which can usually be interrupted several times. In both subspecies each individual has an individual color pattern. In the hybrid zones a reliable assignment can only be done by DNA sequencing. Even far away from hybrid zones irritations can occur, as happened in the westernmost areas of the Czech Republic. There the color patterns of some individuals resemble those of „barred fire salamander“ (S. s. terrestris), because they show parallel, continuous bands along the back. However, by DNA sequencing, it could be shown that the individuals are „spotted fire salamander“ (S. s. salamandra) (Brejcha et al. 2021 ).
Subspecies in general
There are a total of 13 subspecies, most of which are found on the Iberian Peninsula. S. infraimmaculata, S. algira, and S. corsica were also previously listed as subspecies, but are now all considered to be separate species. Based on phylogenetic analyses, they are thought to have split off 5 to 13 million years ago (Steinfartz et al. 2001 ). Each of these salamander species inhabits a specific geographic area, with the exception of S. salamandra. S. salamandra occurs throughout Europe from Spain to Greece, suggesting that it was the only species to recolonize central Europe after the last ice age. It is believed that the two subspecies S. s. terrestris and S. s. salamandra were able to persist in climatically milder areas of the Iberian Peninsula and the Balkans during the ice age and from there recolonized central Europe with the end of the last ice age (Steinfartz et al. 2001 ). Both subspecies now show an overlapping distribution zone in Central Europe. To test the assumption of recolonization, alleles (gene variants) of individuals from different regions were compared. Alleles from western Germany and eastern Germany were compared with those from southern France and Greece. The comparisons show that the identical allele from western Germany is found in southern France and that the identical allele from eastern Germany is found in Greece (Steinfartz et al. 2001 ).
High genetic similarity between two geographically distant subspecies
Comparison of mitochondrial DNA shows that the “Italian fire salamander” S. s. gigliolii from southern Italy and the “Oviedo fire salamander” S. s. bernardezi from northern Spain are more closely related to each other than to the other subspecies. The high genetic similarity is surprising because both subspecies occur separately and far away from each other. S. s. gigliolii and S. s. bernardezi have both maintained their genetic identity in their territories, although they are not separated by geographic barriers from very closely related subspecies. These populations are thought to be remnants of a large homogeneous population that colonized central Europe during an earlier interglacial period about 500,000 years ago. The animals from these populations were apparently unsuccessful in a later recolonization of central Europe (Steinfartz et al. 2001 ).
Overview of sister species and subspecies
Figure 3 (Burgon et al. 2021 ) presents a classification of fire salamander species and subspecies based on an extensive phylogenetic analysis. Five species (S. infraimmaculata, S. atra, S. lanzai, S. corsica, S. algira und S. salamandra) are distinguished, with the species S. atra and S. lanzai being alpine salamanders. Within the species S. salamandra, 13 subspecies are listed, divided into two main clades: one clade containing the Apennine subspecies S. s. gigliolii within the Iberian S. s. bernardezi / fastuosa, and a second clade containing all other Iberian, Central and Eastern European subspecies. Salamandra s. longirostris, sometimes considered a distinct species, has been classified as a subspecies within S. salamandra. S. s. alfredschmidti is not listed here at all, as it is no longer considered a subspecies of its own but is considered S. s. bernardezi (Beukema et al. 2016 ).
Usually in hilly and mountainous areas up to over 1000 m a.s.l. In moist deciduous forests with clear, cold and oxygen-rich waters, usually streams. In exceptional cases, populations are also found at low altitudes. A very low elevation population is found in Stârmina Hill in Romania at an elevation of 78 m (Covaciu et al. 2017 ). In Germany, there is a fire salamander population in Hasbruch (Oldenburg County, Lower Saxony) at elevations starting at 28 m. Other low-altitude populations are known from Nodrhein-Westfalen in Münster 60 m. a.s.l. and in Ostbevern in Warendorf County 54 m. a.s.l.. The more southern the mountain ranges the higher areas are colonized. In the Jura, heights up to 1000 m, in the Massif Central up to 1550 m, in the Alps up to 1800 m and in the Pyrenees up to 2350 m are reached. In warm regions flat areas are hardly or not colonized. In floodplain forests the fire salamander is not found, possibly because of its poor swimming abilities. In pure coniferous forests, the fire salamander is rare or completely absent.
Living on the ground, during the day and in dry conditions hidden in foliage, under stones, dead wood, tree stumps and in rocks. Fire salamanders are active only when humidity is high, which is often at night when the sky is overcast. Accordingly, the vast majority of fire salamanders are active at night during rainy weather. After a long dry period, Fire Salamanders also show up during the day when it rains, although daytime activity can be observed especially in the months of October and November, and rather rarely in the summer. On warm winter days, the fire salamanders can be observed even in the middle of winter if there is sufficient humidity. Fire salamanders are most active at temperatures between 8 and 18.6° C (Strübing 1954 ), at humidity above 85%, and at low light levels below 10 lux (Klewen 1985 , Seifert 1991 , Thiesmeier et al. 2004 ). The range of action is rather small. Documented fire salamanders traveled 35 to 350 meters at night (⌀ 127 m) and returned to their daytime hiding places in 79 of 84 cases (Klewen 1985 ). The same daytime roost is consequently usually visited again. Orientation is visual, with little need for light. Like the adults, the larvae are also quite site-frequent. However, during irregular water flow, larvae may drift and move to new habitats. Dispersal occurs mostly by juvenile migration. Occasionally, however, adults may travel further distances and seek out new habitats. In particular, salamanders adapted to temporary puddles can occupy larger habitats and disperse long distances than individuals adapted to streams. In one study in Germany, stream-adapted fire salamanders were captured up to 500 m away, whereas fire salamanders adapted to temporary puddles were captured up to 1.9 km away. Whereas, the majority of animals in both populations were lazy to migrate and stayed within 200-300 m (Hendrix et al. 2017 ).
The adult diet includes snails, worms, arachnids, millipedes, isopods, earwigs, ground beetles, and other arthropods. The larvae also feed predatorily on small animals in the water such as stream fleas, water fleas, mayfly or stonefly larvae, mosquito larvae, woodlice, whirlpool worms and other arthropods that accidentally fall into the water.
In more isolated waters such as in caves with low food supply and at the same time high larval density, aggressive behavior among larvae and eventually cannibalism may occur. Whereby the larger larvae eat the smaller or later deposited ones (Degani et al. 1979 , Degani et al. 1980 ). However, in one study it was shown that only a temporary food shortage does not directly and immediately lead to cannibalism (Thiesmeier et al. 1990 ). The larvae also feed on other amphibian larvae including grass frog tadpoles or, more rarely, small mountain newt larvae.
Reproduction and development
At 5-6 years, the fire salamanders reach sexual maturity, the ♂♂ have then reached a length of over 15 cm and the ♀♀ of over 16 cm (Seifert 1991 ). Depending on the temperature or metabolic rate and food supply, sexual maturity can also be reached after 3-4 years, although certain descriptions also indicate 2 years. The mating of the fire salamander does not take place in the water but on land. In Central Europe mating takes place every year. (Thiesmeier et al. 2004 ), where the sex ratio between ♂♂ and ♀♀ is 1.3 to 1.0 (Thiesmeier 1992 ). The mating season for the banded subspecies is mostly between July and September, for the spotted subspecies the mating season starts a little later. Mating always takes place during the night. The ♂ pursues the ♀ and nudges it with its head. The ♂ tries to crawl under the ♀ from behind or sideways. If the ♂ is finally under the ♀, the ♂ reaches for the female’s front legs with its forelegs to hold on to them. Once the ♂ has the ♀ in a mating grip, it moves its head sideways back and forth and rubs the ♀’s chin, sometimes accompanied by undulating tail movements. At the end of the mating act, the ♂ rubs the female cloaca with the base of its tail and deposits a sperm packet (spermatophore) on the ground. The ♀ ultimately picks up dei spermatophore with her cloaca. Mating lasts half an hour or longer (Laufer et al. 2007 ). As a rule, the eggs are fertilized within the next few weeks (Thiesmeier 1992 ). Fertilization thus takes place independently of mating. The sperm taken up by the ♀ can remain capable of fertilization in a special seed pocket for up to two years. After a gestation period of 2-7 months, the ♀♀ give birth to their 2-3 cm long larvae between February and May. Sometimes the larvae are released in June and July or in autumn. Thus, the larvae can be set at practically any time of the year. Between 10 and 70 larvae have been counted per ♀, with most often between 20 and 40 larvae being set. Larvae weaned at the same time may descend from different fathers. Larval deposition is often completed in one night, but may be spread over several nights. Usually fire salamanders are viviparous, sometimes the larvae are still in the egg case, but then hatch immediately. In addition to viable larvae, unfertilized eggs, malformed or dead larvae are also repeatedly released (Thiesmeier 1992 ). Especially spinal crippling is not uncommon in freshly born larvae. There are cases where all laid eggs were unfertilized. The larvae, which are brown to grayish-black in color and have characteristic light yellowish spots only on the leg attachments, are deposited in shallow water places in forest streams, spring waters, wells, shady puddles, wagon tracks, and sometimes in mud collectors or even caves. Larvae are characterized by distinctive yellow spots on the upper legs, otherwise they resemble other caudate larvae. Larvae develop within 3-5 months depending on temperature and leave the water body as 4-7 cm long juveniles. In cave waters, development can take up to a year. The external gills are completely retracted when they come ashore. If the lung-breathing juveniles cannot reach land, they drown.
Giving birth to fully developed fire salamanders
Most fire salamanders are ovoviparous (give birth to larvae), although there are exceptions with viviparity (give birth to fully developed salamanders). Most populations of S. s. bernardezi and most populations of S. s. fastuosa are viviparous. (Buckley et al. 2009 ). In S. s. gallaica, only the populations of the two small islands of San Martiño and Ons of Galicia are viviparous (Velo-Antón et al. 2007 ). Viviparous fire salamanders, like alpine salamanders, do not require bodies of water for reproduction. Decades earlier, viviparity among Fire Salamanders was reported from Spain but also from France. An observation from Mont de Bedat (Département Hautes-Pyrénées) is quite interesting. In one litter 24 normal larvae and 4 already fully developed salamanders were born (Wolterstorff 1928 ). Researchers reported that S. s. bernadezi gives birth to up to 25 fully developed young ranging in length from 27 to 51 mm (Thiesmeier et al. 1990 ).
Adults have few natural enemies such as hedgehogs, badgers, wild boars, or rats because of their warning pattern or toxicity. Some other animals eat salamanders occasionally or by accident. For example, there are observations of grass snakes surviving the consumption of a fire salamander while other snakes choke out the salamander or even die. Whereas the grass snake encounters the fire salamander rather rarely. The effect of the fire salamander’s warning pattern is currently being studied in more detail (Dr. Benedikt R. Schmidt, personal communication 02 May 2022). During the spawning season of common toads and grass frogs, clasping of male frogs can occur, which can lead to drowning of adult fire salamanders. Since typical spawning waters often differ, this rarely occurs, but is documented relatively frequently due to its absurdity. The non-poisonous larvae are consumed by fish, especially brown and rainbow trout, but also by bullheads. With a favorable bank structure, with niches inaccessible to trout, the fire salamanders can also coexist with the brown trout (Blau 2002 ). Other enemies of the larvae may include dragonfly larvae, stone crayfish, and less commonly, yellow firefly larvae. Water shrews (Neomys fodiens) are known to dive prey on fire salamander larvae. Observations indicate that when extremities are missing or damaged, the water shrews are responsible, not conspecifics or dragonfly larvae (Thiesmeier 1990 ) or brown and rainbow trout (Dr. Benedikt R. Schmidt, personal communication 02 May 2022). Freshly converted shorebirds can also become easy prey for blackbirds, song thrushes and dippers (Kneis 1989 ). Cannibalism among larvae is present at high larval densities in combination with food shortage, a constellation that occurs mainly in cave waters.
The fire salamander secretes a skin toxin from its powerful ear glands and many dorsal glands. The venom also protects the fire salamander from bacterial infections. The venom consists mainly of the steroid alkaloids samandarin (C19H31NO2) and samandaron (C19H29NO2) and other steroid alkaloids such as samandaridin (C21H31NO3), samandenon (C22H31NO2), samandinin (C24H39NO3), etc., but these cannot always be detected. In the alkaloid mixture of salamander toxins, moreover, cholesterol is present in larger quantities as a concomitant substance. Cholesterol serves as a starting substance for the biosynthesis of steroid alkaloids (Becker 1986 , Lüddecke 2019 ). In the liver, testicles and ovaries, these alkaloids can also be found. The sharp yellow stain pattern actually already warns the attacker about the toxicity and usually repels him. The yellow content does not correlate with the content of alkaloids (Preißler 2019 ). The larvae are completely free of alkaloids and thus non-toxic. The toxic effect in young salamanders is still weak and develops only with increasing age.
Effect of poisons on other animals
All animals, including salamanders, react extremely violently to salamander alkaloids if they enter the bloodstream. Salamander alkaloids cause diastolic cardiac arrest or respiratory paralysis. Other poisonous effects include: Restlessness, epileptic convulsions, change in pupil diameter, weakened or absent reflexes, weakened breathing, irregular heartbeat, paralysis of hind legs, convulsions, vomiting (Becker 1986 ). Vomiting is actually desirable and can be induced intentionally, for example by drinking salt water. It also helps to rinse the mouth, especially if the salamander has not been swallowed. The mere touching of the fire salamander does not usually pose a danger to humans, in the case of skin wounds it may sting a little. Often the fire salamanders are not even attacked because of the warning colors, or if they are, they are immediately spat out because of the taste. Far more dangerous for dogs and cats are the dangers of wood tick and tick bites, which often ambush potential hosts in the meadows.
The fire salamander (Salamandra salamandra sp.) is distributed in large parts of Central, Eastern and Southern Europe. In the former Soviet Union, it is known only from the mountains and foothills of the Ukrainian Carpathians (compare Figure 4). Populations in the Iberian Peninsula are very fragmented. Fire salamanders were also discovered in Turkey not long ago, but they belong to a different fire salamander species (Salamandra infraimmaculata) and are therefore not shown on the map below. Fire Salamander (Salamandra salamandra sp.) is found in the following countries: Albania, Andorra, Austria, Belgium, Bosnia and Herzegovina, Bulgaria, Croatia, Czech Republic, France, Germany, Greece, Hungary, Italy, Liechtenstein, Luxembourg, Montenegro, Netherlands, Northern Macedonia, Poland, Portugal, Romania, San Marino, Serbia, Slovakia, Slovenia, Spain, Switzerland, Ukraine.
Distribution of fire salamanders in Germany
In Germany, the fire salamander occurs in the western, southwestern and central low mountain ranges, in the Harz and Ore Mountains, as well as in northern and eastern Bavaria. The fire salamander is mostly absent in the North German lowlands and in the Allgäu.
The two subspecies occurring in Germany, the banded (S. s. terrestris) as well as the spotted fire salamander (S. s. salamandra), have a 140 km hybrid zone extending from the Hunsrücken mountains in the west to the more eastern Spessart low mountain range. Within this hybrid zone there is intermixing of both subspecies (Compare Figure 5 / Veith 1992 ).
Distribution of fire salamanders in Austria
The largest distribution area extends over the northeastern Limestone Alps from the north of Salzburg to the Vienna Woods. Heaped occurrences can also be found in the East Styrian Hills in the southeast of Styria. In the province of Carinthia, the fire salamanders can be found in all districts. Most occurrences are attributed to the spotted fire salamander (S. s. salamandra). The distribution of the banded fire salamander or their records are limited to the western Tyrol and Vorarlberg (Cabela et al. 2001 ).
Distribution of fire salamanders in Switzerland
North of the Alps lives the banded fire salamander (S. s. terrestris) and south of the Alps in Ticino and the southern valleys of Grisons (Misox, Bergell, Puschlav) the spotted fire salamander (S. s. salamandra).
Distribution of fire salamanders in France
In France, most of the records are of the banded fire salamander (S. s. terrestris). The spotted fire salamander (S. s. salamandra) is found less in France, occurring exclusively in the extreme southeast of France (Hautes-Alpes, Alpes-de-Haute-Provence, Alpes-maritimes, Var). Bordering Spain in the departments of Pyrénées-Atlantiques and Hautes-Pyrénée, the subspecies S. s. fastuosa can be found from 500 m altitude upwards
Distribution of the subspecies on the Iberian Peninsula
Most subspecies of the fire salamander S. s. are found on the Iberian Peninsula (compare Figure 9 / Pereira et al. 2016 ). The natural boundary of S. s. terrestris with respect to the other subspecies is the Pyrenees, the mountain range that separates the Iberian Peninsula from the rest of Europe. The two small islands of Ons and San Martiño of Galicia are missing from the distribution map.
Skin fungus Bsal (Batrachochytrium salamandrivorans) introduced from Asia -> see the article on this subject Salamanderpest / Batrachochytrium salamandrivorans (Bsal), Rezoning of forestry land to agricultural land, general destruction of forest, conversion of deciduous forest to coniferous forest (spruce forests, pine forests), water pollution by agricultural overfertilization, new “unnecessary”, roads and paths in forest areas, hydraulic engineering measures such as canalized or underground “streams” in pipes, Destruction of spring creeks, reservoir creeks with hardened water bottoms without scour, artificial fish stocking of larval waters, obstructions, death traps such as ventilation shafts, light wells, mud collectors, spring catchments and old ruins are sometimes also death traps.
Artificial fish stocking
Juvenile trout are farmed and released selectively in Europe. The release often takes place in side streams where the young trout can grow protected from the adults. This is consistent with natural behavior, where trout seek out waters or side streams for spawning that provide protection for the fry. However, not all waters where the small trout are released are accessible to the large trout on their own. Without pronounced scour, the large trout cannot get enough “run” to overcome higher obstacles, so such places can be quickly identified where a release may not actually be carried out from a natural point of view. Since the releases take place every year at the same places, an amphibian-specific examination is actually necessary only once. Optimally, solutions are worked out with the communities concerned and the relevant fishing and hunting associations, from which both species or, if possible, all stream inhabitants can benefit.
The German name allegedly has nothing to do with the yellow-black spot pattern, but is based on the superstition that the fire salamander can survive fire and its poisonous skin secretion can even extinguish the fire. Supposedly, the animals used to be thrown into the fire during fires.
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