Alpine salamander

from Wikipedia, the free encyclopedia
Alpine salamander
Alpine salamander (Salamandra atra)

Alpine salamander ( Salamandra atra )

Systematics
Order : Tail amphibian (caudata)
Superfamily : Salamander relatives (Salamandroidea)
Family : Real salamanders (Salamandridae)
Subfamily : Salamandrinae
Genre : True salamander ( Salamandra )
Type : Alpine salamander
Scientific name
Salamandra atra
Laurenti , 1768
Salamandra atra

The Alpine salamander ( Salamandra atra ) is a black-colored, terrestrial type of salamanders . It lives in a contiguous area in the Alps as well as in individual isolated populations in the Dinarides in medium to high mountain areas and prefers damp mixed deciduous forests , log heaps and alpine meadows in lime-rich habitats with numerous shelters. As a special adaptation to its habitat, the alpine salamander is one of the few viviparous amphibians and has probably the longest gestation period of all vertebrates.

description

size

Black Alpine Salamander ( Salamandra atra ) in the Berchtesgaden Alps

The males of the alpine salamander remain on average slightly smaller than the females, which can grow to around 15 cm long. On average, the animals are 11-12 cm long. 1444 measured animals showed lengths between 4.9 to 14.4 cm for males and up to 15.1 cm for females. Females of the species Lanza's Alpine salamander ( Salamandra lanzai ), which was previously confused with the Alpine salamander, can grow up to 17 cm long. Differences in size between the subspecies can mostly be traced back to differences in altitude, age structure and ecological factors such as the food available.

External features

Alpine salamander in Carinthia

The animals are - with the exception of the subspecies Salamandra atra aurorae , whose back has an irregularly defined dirty yellow color - uniformly black in color and appear a little less stocky than fire salamanders ( Salamandra salamandra ). In a healthy state, the animals are shiny. The color is lead-gray underneath and a longitudinal groove runs down to the tail. The male can be distinguished from the female externally by a slightly more bulging cloaca .

In addition to the black eyes, the protruding ear glands ( parotid glands ) are clearly visible on the head, which is clearly separated from the trunk and which is wider than it is long in shape and has a rounded snout . Along the body flanks the animals have a series of warty and rounded elevations that contain glandular exits. The trunk is laterally segmented by eleven to thirteen rib furrows. The tail is not as long as the trunk and is almost square in cross-section; it ends relatively pointed.

paleontology

In the Gamssulzenhöhle in the Dead Mountains , the alpine salamander was detected together with other amphibians on the basis of radiometric data for the late glacial period (14,000–10,000%).

Karyotype

As with all European salamandrids, the chromosome number is 2n = 24.

distribution

Distribution of the alpine salamander

The alpine salamander is an endemic of the Central European fauna with a distribution center in the Alps. Except in the Alps and their upstream flysch zone - for example from the Rhone Valley above Lake Geneva through the Central and Eastern Alps in Switzerland, Liechtenstein, Austria and northern Italy - the species can be found in the Dinaric Mountains as far as the Prokletije massif. The main focus of the German deposits is in the northern limestone Alps of Bavaria and represents the northernmost limit of distribution . The southeasternmost location in the Prokletije is in the Valbona valley in Dragobia in Albania and in the Raški do valley on the border between Montenegro and Serbia. In general, the distribution of the Alpine salamander is insufficiently known, even in the Alps, despite its sometimes frequent occurrence.

The lack of the alpine salamander in the dry parts of the Central Alps is striking . The species only reaches the Valais at two points on the northern edge and avoids the dry valley of the Rhone. Older details for Ticino are considered unassigned. Valais and Engadin represent gaps in distribution. In Switzerland, only the northern Alpine arc is continuously populated. It no longer occurs south of the Rhone and Bernese Alps. Individual, widely scattered observations in the Valais, however, require confirmation because of their isolated location. In western Switzerland, the occurrence extends to the national border. The Inn valley is nowhere populated. The species reappears on the calcareous slope of the Southern Alps, albeit rarely and in scattered isolates, from the Aosta Valley to Piedmont to Lombardy (south of the Valtellina). No reliable evidence is known from the Valtellina itself. Finds from the Adamello and the southern part of the Ortler massif are known further east. Evidence is missing from the Vinschgau, Stilfser Joch and the area between Münstertal and Puschlav.

The Ötztal Alps are also a distribution gap. It was not until 2007 that a find was recorded in the northern Austrian part. The rarity of the alpine salamander in the Ötztal Alps is due on the one hand to the structure of the mountains made of crystalline slate, on the other hand to the rather dry regional climate. The northern Ötztal Alps are more rainy with 110 rainy days, which is used as a possible explanation for the new finds.

The southernmost point of the contiguous settlement area in the Alps is the Ternowaner Forest in Slovenia. To the south are isolated occurrences of the middle and high Dinarides. The species is relatively common in the Krainer Schneeberg and Kapela Mountains in Croatia. After a (real?) Distribution gap of 200 km, the species colonizes the massifs on both sides of the Neretva , the Prenj -planina and the Čvrsnica in Bosnia and Herzegovina. Another (real?) Distribution gap of 200 km is enough to Prokletije . Here the alpine salamander was first described only from the eastern part in the silicate mountain massifs in Montenegro, Kosovo and Albania. As recently as 2017, four more sites were reported from the western part of the Albanian Bjeshket e Namuna / Prokletije Mountains, which is made up of alkaline rocks. A surprising discovery comes from the Mediterranean Orjen Mountains in Montenegro. Here the alpine salamander was discovered in a large paleodoline at an altitude of 1570 m only 15 km from the Mediterranean. In the depression of the Opuvani do not only the alpine salamander but also glacial relics of the flora have been preserved. This microrefugium is decoupled from the general climate of the atmosphere by the formation of a cold air lake. Periglacial process dynamics are pronounced due to longer snow cover, meltwater and frequent changes in frost. The alpine salamander finds suitable habitats for it in the periglacial debris layers formed in this way and in the arctic-alpine and alpine dwarf shrub heather with silver arum and blunt-leaved willow .

Despite intensive research, the species could no longer be detected in the Bosnian Treskavica Mountains, where it is documented from conservative finds from the 19th century. The presumed distribution in the cool and humid Volujak, Bioč and Maglić is not supported by missing finds during several intensive collection campaigns. According to this, the distribution area in Bosnia is currently limited to the limestone mountains Prenj and Čvrsnica, which are built up from sedimentary stones, as well as in the border area of ​​Montenegro, Albania and Kosovo in Prokletije to massifs with metamorphic bedrock. To what extent the geology and the underground rock as well as the resulting geomorphology and habitat configuration are significant for the distribution in the Dinarides cannot be clearly clarified at the moment. It is noticeable that the frequent occurrence in the Prenj between 1700 and 2000 m seems to be encouraged by countless hiding places in alpine karst forms.

habitat

The Opuvani do in Montenegro is home to the southernmost and Mediterranean populations of alpine salamanders and silverwort in Europe

The habitats of the alpine salamander are located in karst areas and high mountain gorges. It occurs in the Alps mostly from heights of 1000  m , regionally even from 800  m above sea ​​level ; specimens are rarely found in lower altitudes (details from the literature: 420  m near Walensee / Switzerland, 430  m in Austria). Sometimes such finds can also be traced back to the drifting of animals with rapid streams. The highest individual locations are at 2800  m in Carinthia / Austria and 2400  m in Switzerland; in the German Alps the species seldom rises above 1600  m , exceptionally up to 1900  m . Moist deciduous and mixed mountain forests are predominantly settled near mountain streams , and above the tree line biotopes such as moist alpine pastures, dwarf shrub heaths and rubble heaps . Here the alpine salamander can be found under stones or dead wood.

Favorable habitats for dense populations are stabilized block and rubble heaps, fat meadows, forest edges and mountain forests. Within the forests, pure coniferous forest areas are rather avoided, deciduous-coniferous mixed forests and especially deciduous forest areas are preferred. The greatest densities are found along small forest streams or in spray zones at the foot of waterfalls. Gorges also offer good living conditions.

The alpine salamander tends to occur in locations with a soil-based reaction and high pH values. This also explains why sites with acidic soil reactions are problematic for him. This also explains why it occurs more frequently in deciduous than in coniferous forests with low pH values. Fertilization has a positive effect on its occurrence, as fat meadows are more humid than lean meadows. The alpine salamander is true to location; its living space is only a few square meters. Rapid habitat change is therefore unfavorable for long-term establishment. Since salamander populations have a very slow generation change, adaptations to strong habitat changes can only be successful after several decades.

The habitat of the fire salamander is roughly allopatric; their areas complement each other and can only overlap in a strip a few kilometers wide in the northern Alps. The two salamanders inhabit the same forests along the foothills of the Alps at an altitude of 700–900. For example, both species were picked up under the same stone slab.

ecology

The essential boundary conditions are given by temperature, humidity (which should ideally be around 85 percent) and soil structure. Gorges, Bachtobel, spring corridors and spray zones are only populated as far as the soil structure prevents waterlogging. Up to now it is not known whether a local seasonal migration of the animals takes place in places with periodic inundations like floods after snowmelt. A protective blanket of snow in winter seems necessary, as southern slopes are hardly or only sparsely populated. The alpine salamander does not tolerate dehydration and avoids locations exposed to wind and locations with high temperatures. In sparse deciduous forest zones, up to 120 individuals / ha is estimated, in dense vegetation 64 individuals / ha. The largest occurrences, calculated per area, were found in alpine pastures, dwarf shrub heaths and overgrown debris cones with deep, fissured and void-rich soil structures and, at 2,000 to 3,000 individuals / ha, are close to the local capacity limit. After landslides, alpine salamanders find themselves relatively quickly with good populations. In the Elm landslide area (September 9, 1888) in the canton of Glarus, a slate / flysch zone, good populations were found 100 years later. The subspecies aurorae lives almost exclusively in loose white fir stands and mixed forests with grassy subsoil, where z. Sometimes quite high individual densities of 770 individuals / ha can be achieved. Most of the animals there live underground and hardly come to the surface from the karst fissures.

Lifestyle and diet

Alpine salamanders are predominantly nocturnal and feed on animal prey such as insects, spiders, various larvae , woodlice, snails and earthworms. The animals also come out of their hiding places during the day, especially after rainfall. Depending on the altitude, they can usually be found between April and October, while in the winter months they spend a long time slumbering in underground hiding places.

As predators mainly come magpies and chough in question, as well as occasionally the viper . As a defense, Alpine salamanders, like fire salamanders, excrete a poisonous skin secretion . They also go into a threatening position in which they lift their heads and bend backwards.

Reproduction and development

Alpine salamander in Slovenia

Alpine salamanders mate outside of any body of water. The development of one larva at a time takes place in both uteri of the female until metamorphosis is complete and takes about two years (up to three years at higher altitudes). Depending on the weather and temperature conditions, the animals are courtship-active in late spring to early summer . Mating on land occurs during the alpine spring. The males do not have an anatomically intrusive organ for mating. After ingesting one to three spermatophores from the male, the females can store the sperm for up to two years in the "Siebold's tubes", special seed bags ( spermatheks ), capable of being fertilized . After ingesting the sperm, the females ready for it ovulate. Usually only one egg per side is provided with a stately gelatinous shell as it passes through the oviduct. On average, only these eggs, located caudally in the filling uterus, develop. The remaining eggs, often misshapen in shape, are, if at all, only sparsely surrounded by a gelatinous shell and can soon lose their shape completely and disintegrate. The embryo hatched from the egg shell feeds itself actively on the remaining eggs as soon as it is anatomically able to do so. This oophage (egg-eating) stage lasts about 1–2 years, depending on climatic conditions.

In contrast to most other amphibians, which lay their eggs ( spawn ) in bodies of water and whose offspring go through a larval stage with gill breathing, the viviparous alpine salamanders give birth to one or two fully developed, approximately four centimeters large and lung-breathing, independent young animals that immediately are viable on land. The young are usually born in summer. Nutrition takes place first through the yolks contained in the own egg , then through the yolks of other eggs that do not develop. When the entire yolk is used up, the mother animal produces a cellular substance in a tissue in the transition area from the fallopian tube to the uterus for further feeding of the larvae. The larvae have large, red gills - comparable to other tail salmonella larvae that grow freely in the water - but they regress before the birth process and replace them with lungs. This prenatal gill regression distinguishes the species from the ovoviviparous fire salamander, whose Central European subspecies give birth to gill-bearing larvae that depend on a body of water for their further development. The alpine salamander is the only Central European amphibian that can exist independently of surface water - an adaptation to the often arid living conditions in the high mountains.

The reproductive cycles include gestation periods of at least two or more years, depending on climatic conditions. Four-year reproductive cycles were found at an altitude of 1700 m, but since the species rises to 2500 m, a reproductive cycle of 5 years or longer can be assumed at extreme alpine altitudes. The alpine salamander is therefore likely to have the longest gestation period of all vertebrates.

behavior

The main activity phases fall in the early morning hours (3 to 7 a.m.), as observed in June / July in the Üblital in the canton of Glarus at an altitude of 1500 m. The activity is determined less by the temperature than by the relative humidity. From temperatures of 3–5 ° C animals could be observed in the open air, larger activities were recorded at 7–15 ° C and over 90% humidity. When the humidity is below 70%, alpine salamanders can rarely be seen. Dew or new rain, but not sedentary fog, were the best moisturizers for the animals to appear. The animals don't like wind at all, but daylight (no direct sun) seems to disturb them only moderately if the humidity is sufficient. If the rain persists for days, the activity decreases sharply. A particularly large number of animals can be recorded after dry periods followed by a mild thunderstorm. Day hiding places are predominantly hollow stones and stone structures, followed by small mammals, tree wood, moss cushions and under accumulated leaves.

Activity radii are 10 m (between 4 and 22 m). The animals seem to know their environment and now and then change their daytime hiding place. Observations during or shortly after the snowmelt are extremely rare. Even at low altitudes around 600 m, they hardly appear on the surface before the end of April. They probably only become active on the surface weeks after the snow melts. In June the number of active Alpine salamanders observed increases significantly and reaches its peak in midsummer, in July / August. Snowfall in summer interrupts activity for a short time, but can completely and abruptly end surface activity in late summer. With September the frequency of observation decreases again; in the first half of October only a few animals are still active.

Systematics

Salamandra atra aurorae
A new discovery of the subspecies ssp. prenjensis in the Mediterranean Orjen Mountains in Montenegro took place in 2018.

The alpine salamander occurs in the greater part of its range in its nominate form Salamandra atra atra . The aurora alpine salamander ( Salamandra atra aurorae ) is considered a subspecies and occurs only in the Italian province of Vicenza at altitudes between 1300  m and 1500  m . This is characterized by the fact that it has a yellow mark in variable proportions. All animals between the Pokljuka in the southeastern Alps via Snežnik, Velika Kapela, Ćrvsnica , Prenj and Prokletije are genetically different from the nominate form and form a clade. If their subspecies status is retained, they will become Salamandra atra ssp. prenjensis posed; however, this status is controversial, as the differences to the nominate form lie within the normal range of variation of the animals. The questionable subdivision has not yet been fully clarified by the analysis of phylogenetic relationships of peripheral populations between the southern Alps and the central Dinarides, published in 2018. A satisfactory taxonomic differentiation therefore requires further research.

In 2018, a relic population of the Dinaric “subspecies” was discovered within the sub-Mediterranean zone on the edge of the Mediterranean Sea in the Orjen Mountains in southern Montenegro. It covers a previous 200 km, "actual" (?), Distribution gap between the mountains of Prenj and Prokletije. The presumed survival of this presumably only small residual population within a microclimatic / mesotopographical niche of a cold air lake since the end of the last Ice Age is discussed, as it occurs here with other glacial relics in the flora. These relics are no longer widespread outside the relevant karst sinkhole in the immediate vicinity or in the area of ​​discovery .

A relative of the alpine salamander is Lanza's alpine salamander ( Salamandra lanzai ), which is native to the Cottian Alps in western Piedmont and looks very similar to the alpine salamander, but is about two to three centimeters larger and has a more robust body and a rounded tail end. Occasionally, Salamandra lanzai is still described as a subspecies of the Alpine salamander, since from a genetic point of view the variability for a "real" species seems to be very low.

In some regions the alpine salamander is alternatively referred to as a mountain salamander , in the eastern alpine region (Bavaria and Austria) also as a mountain mandl or a Wegmandl .

Hazard and protection

Alpine salamander in Switzerland

The low rate of reproduction of the alpine salamander is sufficient for the species to survive, as adult animals are protected by their poisonous skin secretions and therefore hardly have any natural predators.

Due to their regional rarity, alpine salamanders are under strict protection in Germany, Austria and Switzerland. Sometimes they are even quite common in their range, so that the species is not endangered. Rather, they are regionally threatened by the destruction of their habitat, which only exists within the Alps and few other European high mountains and is accordingly unique. Mainly due to the very limited distribution areas, the subspecies Salamandra atra aurorae and the French populations of the Alpine salamander are particularly threatened and must be protected accordingly.

Legal protection status (selection)

National Red List classifications (selection)

literature

  • Andreas Nöllert, Christel Nöllert: The amphibians of Europe . Franck'sche Verlagsbuchhandlung, Stuttgart 1992, ISBN 3-440-06340-2 .
  • Rainer Günther (Ed.): The amphibians and reptiles of Germany . Gustav Fischer Verlag, Jena 1996, ISBN 3-437-35016-1 .
  • Axel Kwet: Reptiles and Amphibians of Europe . Franck'sche Verlagsbuchhandlung, Stuttgart 2005, ISBN 3-440-10237-8 .
  • Alexandre Riberon, Effimia Sotiriou, Claude Miaud, Franco Androne & Pierre Taberlet: Lack of Genetic Diversity in Salamandra lanzai Revealed by Cytochrome b Gene Sequences. Copeia , (1), 2002, pp. 229-232.
  • Magdalena Meikl: On the trail of Bergnarr and Regenmandl: Alpine and fire salamanders as indicators of intact nature. ANLiegen Natur (36/2), Laufen 2014, pp. 75–81 ( PDF; 0.8 MB ).

Web links

Commons : Alpine Salamander  - Collection of images, videos and audio files

Individual evidence

  1. ↑ a b c d Gaston D. Guex, Kurt Grossenbacher: Salamandra atra Laurenti, 1768 - Alpensalamander . In: B. Thiesmeier, K. Grossenbacher (Eds.): Handbook of the reptiles and amphibians of Europe . Volume 4 / IIB. Tail amphibian (Urodela) IIB, Salamandridae III: Triturus 2, Salamandra (pp. 975-1028). Aurla-Verlag, Wiebelsheim 2004.
  2. ↑ Gaston D. Guex, Kurt Grossenbacher: Salamandra atra Laurenti, 1768 - Alpensalamander . 2004, here p. 990.
  3. ↑ a b Kurt Grossenbacher 1988: Distribution atlas of the amphibians in Switzerland. Documenta Faunitsticae Helveticae, 7 (PDF)
  4. ↑ a b Gaston D. Guex, Kurt Grossenbacher: Salamandra atra Laurenti, 1768 - Alpine salamander . 2004, here p. 984.
  5. ↑ Cabela A., Hill J. (2007): Salamandra atra Laurenti, 1768 in the Ötztaler Alpen Massif (Austria, Italy). Herpetozoa, 20, 88-91. (PDF)
  6. ↑ Márton Szabolcs, Edvárd Mizsei, Daniel Jablonski, Balázs Vági, Béla Mester, Zsolt Végvári, Szabolcs Lengyel 2017: Distribution and diversity of amphibians in Albania: new data and foundations of a comprehensive database. Amphibia-Reptilia 38 (2017): 435-448 (PDF)
  7. ↑ Pavle Cikovac & Katarina Ljubisavljevic 2020: Another isolated relic population of the Alpine Salamander (Salamandra atraLaurenti, 1768) (Amphibia: Caudata: Salamandridae) in the Balkans. Russian Journal of Herpetology, Vol. 27/2: 109-112 (PDF)
  8. ↑ Pavle Cikovac & Ingo Hölzle 2018: On glacial microrefugia Opuvani do - Mt. Orjen. 7th Balkan Botanical Congress, University of Novi Sad 10-14 September 2018. (PDF)
  9. ↑ https://www.rufford.org/files/17459-2%20Mid%20Term%20Report_0.pdf Prenjensis conservation project - mid term report
  10. ↑ https://www.researchgate.net/publication/301232231_Ecological_characteristics_and_population_structure_of_the_alpine_salamander_from_Mt_Prenj Ecological characteristics and population structure of the alpine salamander from Mt. Prenj
  11. ↑ Klemens Fritz, Peter Sowig: Alpensalamander - Salamandra atra Laurenti, 1768. In: Laufer, Fritz, Sowig: The amphibians and reptiles of Baden-Württemberg. Ulmer, Stuttgart 2007, ISBN 978-3-8001-4385-6 , pp. 159-170.
  12. ↑ Geiger C. (2006): Ecological requirements of the Alpine Salamander Salamandra atra: assessing the effects of current habitat structure and landscape dynamics on local distribution. B. Sc. Thesis, Faculty of Science, University of Bern, Switzerland. (PDF)
  13. ↑ Kurt Grossenbacher 1988.
  14. ↑ Kurt Grossenbacher 1988
  15. ↑ Gaston D. Guex, Kurt Grossenbacher: Salamandra atra Laurenti, 1768 - Alpensalamander. In: B. Thiesmeier, K. Grossenbacher (Eds.): Handbook of the reptiles and amphibians of Europe. Volume 4 / IIB. Tail amphibian (Urodela) IIB, Salamandridae III: Triturus 2, Salamandra (pp. 975-1028). Aula-Verlag, Wiebelsheim 2004.
  16. ↑ Marvalee H. Wake: Evolution of oviductal gestation in amphibians . In: Journal of Experimental Zoology . Volume 266, No. 5, pp. 394-413, August 1, 1993, doi : 10.1002 / jez.1402660507 .
  17. ↑ David M. Sever: Comparative anatomy and phylogeny of the cloacae of salamanders (Amphibia: Caudata) IV. Salamandridae . In: The Anatomical Record . Volume 233, No. 2, 1992, pp. 229-244, doi : 10.1002 / ar.1092330206 .
  18. ↑ A. Razpet, E. Šunje, B. Kalamujic, U. Tuli, N. Pojskic, D. Stankovic, S. Maric: Genetic differentiation and population dynamics of Alpine salamanders (Salam atra Laurenti, 1768) in South Eastern Alps and Dinarides . In: Herpetological Journal 26, 2016, pp. 111–119.
  19. ↑ Lucio Bonato, Andrea Corbetta, Giovanni Biovine, Enrico Romanazzi, Emina Šunje, Cristiano Vernesi, Barbara Crestanello: Diversity among peripheral populations: genetic and evolutionary differentiation of Salamandra atra at the southern edge of the Alps . In: Journal of Zoological Systematic and Evolutionary Research . April 2018, pp. 1–16 ( limited preview ).
  20. ↑ Pavle Cikovac & Katarina Ljubisavljević 2020: Another isolated relic population of the Alpine Salamander ( Salamandra atra Laurenti, 1768) (Amphibia: Caudata: Salamandridae) in the Balkans. Russian Journal of Herpetology, Vol. 27/2: 109-112, April 25, 2020 (PDF) .
  21. ↑ Fred Langer: Herbal Medicine: The Power of Green Herbs. In: Geo . November 2008, accessed February 5, 2018 .
  22. ↑ Federal Agency for Nature Conservation (ed.): Red list of endangered animals, plants and fungi in Germany 1: Vertebrates. Landwirtschaftsverlag, Münster 2009, ISBN 978-3-7843-5033-2 .
This version was added to the list of articles worth reading on February 25, 2006 .