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Description

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Total length to 167 mm for males, up to 181 mm for females. Usually between 110 and 140 mm. Tail usually equal to or slightly shorter than snout-vent-length. Head is flat and somewhat longer than wide. 11-13 weakly defined costal grooves present at either side of the long, cylindrical body. There is a distinctive skin fold on the throat. Well-defined parotoid glands are present at the back of the head. Coloration in this species is highly variable and is subspecies specific. Currently, there are nine subspecies recognized, mainly on morphological grounds (see Comments section for a discussion on taxonomy). S. luschani helverseni: Only a few yellow spots on back, mainly concentrated middorsally. Back otherwise dark brown. Flanks are yellow. Parotoids are dark or with some yellow. Legs and tail are brown or black. Venter translucent, showing the internal organs. The underside of the tail is orange-yellow, the throat is yellowish-pink.S. luschani flavimembris: Back dark brown with very few, small silvery-white spots. The translucent venter is separated from the back by a lateral white stripe, consisting of a row of iridophore-accumulations. The dorsal side of the tail is brighter than that of the back. Extremities are pale orange or yellow. Gland openings on the parotoids and tail are visible as black spots. S. luschani fazilae: The dorsal side of the head, back and tail are reddish in this species. The general coloration moves more toward red with age. The pale venter is separated from the back by a white lateral stripe. The back bears brown to black blotches, that can flow together near the white lateral line.S. luschani luschani: The base color of the back is bright yellow or silver-white. The dark brown or black spots on the back however, can extend to fully cover the bright base color. The extremities are black, reddish or pale brown. The venter is sparsely pigmented and somewhat translucent.S. luschani basoglui: The base color of the back varies from brown to pale red, and is brighter on the parotoid glands. The brown or black spots that are distributed across the back are more extensive in females than in males. The head, tail and sides of the extremities are bright red-pink and sparsely covered with brown spots. The large yellow ovaria in females can be seen through the skin of the back.S. luschani finikensis: The base color of the back is very dark, and is covered with silver-white spots that have the tendency to form larger bright areas. The dorsal coloration completely lacks red or yellow. The white lateral band is sometimes dissolved into white blotches. The ventral side is pale, sometimes with white spots.S. luschani billae: The base color of the dorsal side can vary from salmon to black. The white spots that are distributed across the back are ordered regularly and can form two dorsolateral bands. The white lateral band that sharply separates the dorsal and the ventral side extends anterior to under the eyes. The openings of the dorsal, caudal and parotoid glands are visible as black spots.S. luschani antalyana: The dorsal side is yellow with dark spots, or vice versa. The lateral band is yellowish and not clearly defined. The ventral side is yellowish. The members of this subspecies are easily recognized by the conspicuous yellow parotoids and eye patch. S. luschani atifi: The dorsal side is brown to dark brown with very small white spots. Some individuals are wholly unspotted. The continuous bright lateral band that is present in most subspecies consists of white spots in this one. The ventral side is yellow, orange to red. This is the largest subspecies, reaching a total length of over 170 mm.There is distinct sexual dimorphism in this genus. In contrast to females, the males possess a spike-shaped protuberance on the dorsal surface of the tail base and nuptial pads on the forelegs which are best developed during the breeding season(Boehme et al 1999). The protuberance is thought to have a function in predisposing the females' cloaca for uptake of the spermatophore (Sever et al. 1997) (also see "Life history").This species was previously placed in the genus Mertensialla as sister species to M. caucasica. Weisrock et al (2001), subsequently placed the species within Salamandra based on mitochondrial DNA. They also identified six lineages, diagnosable via color pattern, which may represent distinct species. These are S. flavimembris, S. fazilae, S. luschani (including S. l. finikensis and S. l. basoglui), S. billae, S. anatalyana, and S. atifi.

References

  • Sever, D. M., Sparreboom, M., and Schultschik, G. (1997). "The dorsal tail tubercle of Mertensiella caucasica and M. luschani (Amphibia: Salamandridae)." Journal of Morphology, 232, 93-105.
  • Weisrock, D. W., Macey, J. R., Ugurtas, I. H., Larson, A. and Papenfuss, T. J. (2001). ''Molecular phylogenetics and historical biogeography among salamandrids of the “true” salamander clade: Rapid branching of numerous highly divergent lineages in Mertensiella luschani associated with the rise of Anatolia.'' Molecular Phylogenetics and Evolution, 18(3), 434-448.

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Distribution and Habitat

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The distribution of S. luschani is centered around southwestern Turkey. The Turkish distribution contains eight subspecies. Their distribution is scattered and sharply defined. In addition to mainland Turkey, the species is also found on four Turkish islands and one (politically) Greek island off the Turkish coast. The Greek island of Kastellorizon is inhabited by S. luschani basoglui. This subspecies also occurs on Kekova. Tersane and Domuz Adese are both inhabited by S. luschani fazilae. S. luschani flavimembris lives on the island of Bogaz Adasi. The European distribution, from a zoogeographic perspective, consists of the occurrence of S. luschani helverseni on the islands Karpathos, Kassos and Saria. The main limitations to the distribution of S. luschani seem to be abiotic factors like temperature and precipitation. The greatest part of its currently known distribution lies in an area that receives more than 1000 mm of precipitation annually and where the average temperature in January lies above freezing. The species is highly specialized and seems to be closely associated with the Karst-area. The cave systems in the limestone offer the salamanders a cool and humid retreat during the dry and hot summer. Not only natural caves are inhabited. Various authors report the occurrence of this species near human settlements, inhabiting loose rock walls and ruins. Their natural habitat however ranges from pinewoods at sea-level to dry vegetation with Quercus coccifera. The specialized reproductive biology of S. luschani allows it to inhabit areas devoid of surface water. (see "Life History")
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Life History, Abundance, Activity, and Special Behaviors

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S. luschani is mostly active during the cooler winter months, and can then be quite abundant. Mating also takes place during this period. These salamanders are nocturnal and seem to be more active at the surface during and after rainfall and at dropping atmospheric pressure. This species reproduces independent of water. The protuberance on the tail-base is rubbed against the female cloaca during the ventral amplexus, but its precise function is unknown (Sever et al. 1997). One egg develops in each horn of the uterus. The larvae feed through intrauterine oophagy. There is no evidence of a zona trophica and therefore no epithelofagy. The two young emerge fully developed after 5 to 8 months. They can measure as long as 7 cm upon birth, and weigh up to 2 grams. Sexual maturity is reached at an age of three years in captivity. Longevity is estimated at over ten years (Boehme et al 1999).
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Life History, Abundance, Activity, and Special Behaviors

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Small island populations of S. luschani encounter the same problems as all other rare species. The population on Kasos island is the most endangered. Main threats to the mainland populations are habitat destruction and compulsive collectors (Gasc 1997). Species with such narrow ecological demands are easily threatened by small changes in their environment. Increasing popularity of the Turkish Mediterranean coast as a tourist destination can pose a serious threat for this species (Boehme et al 1999).
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Luschan's salamander

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Luschan's salamander or Lycian salamander (Lyciasalamandra luschani) is a species of salamander in the family Salamandridae. It is found in the southwestern Anatolia in Turkey and adjacent Greece, in the island of Kastellorizo and its satellites.[1][2]

Description

Luschan's salamander is brown with small yellow dots dorsally, yellow or whitish laterally, and flesh-colored ventrally. The eyes and the narrow paratoid glands are prominent. Its tail is thin. Males have a protruding "spike" at the base of their tails on the dorsal surface. Adults may grow to 13 cm (5.1 in) in length.[3] The Luschan’s salamander is a nocturnal, land-dwelling animal, most active during the rainy or wet season, which lasts from November to April in the Mediterranean region.[4] Males and females have similar weights.[4]

Male and female L. luschani are sexually dimorphic.[4] Males can be distinguished by a protuberance near the tail absent in both females and juvenile salamanders. The females, however, have a cloaca that can be used to determine their sex. Juvenile salamanders possess neither. One study has also shown that females tended to have a larger snout to vent length on average than males, but males had a significantly larger maximum snout-vent-length in comparison.[4]

One population of the Lyciasalamandra luschani was found at the Dodurga village in southwestern Anatolia. The population was split into two types. Type 1 had a light brown dorsum, while type 2 had a light brown dorsum that is orangish. The other main differences can be seen in the upper eyelids such that they have a greenish light yellow color in type 2 salamanders. These eyelids also had a darker mid part. The type 2 specimens had dark brown spots all over them.[5] Both type 1 and type 2 specimens have visible viscera, a wider posterior parotid gland, and an unpigmented ventral trunk. Males may sometimes exhibit a defensive posture with an arched back. It is worth noting that both type 1 and type 2 secretory adenoid glands can be found randomly distributed throughout the partoids.[6]

Subspecies

The three subspecies are:[2][7]

Genetics & phylogeny

At least eight subspecies of Mertensiella luschani have been described: M. l.billae, M. l. fazilae, M. l. luschani, M. l. atif, M. l. finikensis, M. l. flavimembris, M. l. antalyana, and M. l. basoglui.[10] M. luschani overall was supported to be a well evidenced sister taxon to the Salamandra clade with a boot-strap of 100% and decay index of 41.[10] Scientists also described that M. l. finikensis, M. l. luschani, and M. l. basoglui seem to form a monophyletic group with a bootstrap of 100% and a decay index of 15.[10]

Scientists have also worked to categorize M. luschani into its appropriate phylogenetic positions based on different DNA sequences it possesses.[11] A study on the 16,650 base pair nucleotide sequence of the salamander’s mitochondrial DNA revealed that it has similar features to other vertebrate genomes.[11] For example, the line of genetic code from the mitochondria produces 22 transfer RNAs, 2 ribosomal RNAs, and 13 peptides.[11] The nucleic acid breakdown is the following: 15% guanine, 32% adenine, 29% thymine, and 24% cytosine. It is worth mentioning that there are two non coding repeats of 124 base pairs that interrupt the typical vertebrate mitochondrial consensus genome sequence.[11]

It has also been shown that the control region of this mitochondrial genome is 922 base pairs long. The origin of light strand replication has also been noted to form a secondary structure known as a stem-loop.[11] The 16S and 12S ribosomal RNA genes are 1567 and 921 nucleotides in length. As with other genetic sequences, the majority of genes that produce proteins began with the ATG start codon. Exceptions to this include COI, ND3, and ND6. Based on genomic data and subsequent phylogenetic results, some scientists have argued that hagfish and lamprey should not be used as outgroup taxa.[11]

Cytogenetic analyses

One analysis on genetic composition of the L. l. luschani subspecies showed hetermorphic nucleolus organizing regions. Surprisingly, these were all found on a single homolog of chromosome pairs.[12] It was shown to have similar karytompes of 2n = 24 biarmed chromosomes. And this was consistent with other species in the Lyciasalamandra genus. In addition, the scientists had observed that there were solid C-bands near the centromere of the chromosomes. They had also seen heterochromatin on the telomeres as well.[12]

Habitat and distribution

L. luschani's natural habitats are temperate forests and arid Mediterranean environments.[13] During active seasons, they live under shelters (i.e., rocks, wood) during the day, and at night, they forage.[13] L. lushani populations are threatened by habitat loss. One of its subspecies, M. 1.helverseni, is known to be found within a pine forest on the island of Carpathos.[14] M. luschani has also been reported to inhabit about 350 kilometers of southwestern Turkish coast.[10]

Conservation

One source, the Red Data Book of Threatened Animals of Greece, classifies the salamander, particularly L. luschani basoglui, as vulnerable.[4] Another source, the Red Data Book of the International Union for Conservation of Nature (IUCN) considers the salamander as an endangered species.[4]

Concerns about Luschan's salamander conservation are rooted in 1) habitat loss due to development, and 2) subspecies isolation.[4]

Diet

When comparing adult and juvenile L. luschani basoglui salamanders, scientists found that there were statistically significant differences in the measurements of body length, body weight, snout to vent length, head width, mean prey length, and mean prey volume.[4] This study showed that juvenile salamanders tended to have a diet heavily centered on collembolans and gastropods, whereas adult males tended to mostly eat beetles, and adult females mostly ate beetles and Hymenoptera. The researchers also discovered that L. l basoglui's snout-vent-length and the head with were significant predictors of both mean prey length and mean prey volume.[4]

Reproduction

Luschan's salamanders are viviparous. Females give birth to two offspring that are produced from each of the female's two oviducts. Gestation time ranges from 5–8 months.[15]

Using 9 salamanders from the M.l. antalyna subspecies, scientists discovered the basic reproductive biology of M. luschani.[16] The period for L. luschani birthing spans around two seasons, from the end of September to the end of March.[16]

This partially due to the fact that a preferred humidity and temperature are desired before the salamander gives birth. Amplexus was observed frequently in April, suggesting that springtime may be its mating season.[16]

Development and life cycle

It has been discovered that soon after the metaphosphoric deposits of “periosteal parallel-fibred” bone can be found near the embryonic bone. L. l luschani may also have a rested growth period each year that tends to occur during the winter.[4]

M. luschani achieve maturation at about 3 years old and are considered to be juveniles beforehand. Their life expectancy was shown to be about 5.4 years with some males living up to 8 years and some females living up to 10 years.[4]

Anatomy and morphology

M. luschani is said to have a flat and wide head. Its general trunk and body is orange with brown spots.[14]

In one of the earliest papers describing the morphology of M. luschani, its anatomy was compared between two seemingly similar salamanders: Mertensiella caucasica and Salamandra salamandra.[14] M. luschani’s skull is larger than that of M. caucasica but similar in size to that of S. salamandra. M. luschani’s skull also has two short premaxillae. Its jaw has been described to be somewhere in between the skull of Mertensiella caucasica and Salamandra salamandra. It has big nasal bones as well. The frontal bones of the skull have a pointed end on the side closest to the tail.[14]

Lyciasalamander luschani basoglui has been described to have fairly rough skin, about 68 nanometers thick, that allows it to conserve water for extended time periods.[17] The skin is mentioned to also have deep pits of 141 nanometers. It was found to have rougher skin than Salamndra salamandra. This suggests that L. l. basoglui has a slower rate of water vaporization through their skin than other salamanders.[17]

Additionally, the salamander possesses one cervical vertebra, about sixteen trunk vertebrae, one sacral vertebra, three caudosacral vertebrae, and at least two dozen caudal vertebrae.[14] The number of caudal vertebrae was as high as thirty in some species. The hips of M. luschani are said to be in between the size of M. caucasica and S. salamandra.[14] Part of this pelvic girdle is made up of cartilage, but ossification starts centrally and then spreads laterally.

M. luschani has a stomach made up of the pylorus and fundus, like other salamanders.[18] The stomach is composed of four layers: tunica serosa, tunica muscularis, submucosa, and tunica mucosa. The tunica mucosa can be further subdivided into the lamina muscularis, lamina propria, and lamina epitheliasis. As for the specific cell types present in each layer, the mucous layer has simple columnar epithelium. Vasculature as well as collagen can be found in the submucosa layer. Glands found in the pylorus appear to be branched and tubular, while glands in the fundus are of a simple tubular form.[18]

The small intestine is made up of similar layers and lacks the muscularis mucosa layer.[18] Goblet cells are present within the pseudostratified ciliated epithelium in the mucous membrane of the intestine. The small intestine is also made up of folds called villi. The study that characterized this also found that M. luschani had significantly greater lamina epitheliasis thickness and lumen area compared to the intestines of the Southern crested newt, Triturus karelinii.[18]

Physiology

Scientists have determined that the liver esterases present in M. l. luschani are all carboxylesterases.[19] Since the esterases were all inhibited by diisopropylfluoro-phosphate (DFP), it was clear that such esterases were present. They also appeared to be stable in higher temperatures up to 45 degrees Celsius. There were around ten molecular forms of esterases present based on isoelectric focusing and gel electrophoresis. The main two molecular weights seen were typically 70,000 and 230,000 [19]

Parasites

Lyciasasalamandra luschani, also known as Mertensiella luschani, has been shown to be a parasitic host for certain nematodes or helminths.[20] Angiostoma aspersae have been found in certain salamander carcasses. Of the salamanders observed, about 92 percent of them had this nematode infecting their bodies.[20]

References

  1. ^ a b Lymberakis, P.; Kaska, Y.; Kumlutaş, Y.; Avci, A.; Üzüm, N.; Yeniyurt, C.; Akarsu, F.; Tok, V.; Ugurtas, I.H.; Sevinç, M.; Crochet, P.-A.; Papenfuss, T.; Sparreboom, M.; Kuzmin, S.; Anderson, S. & Denoel, M. (2016) [errata version of 2009 assessment]. "Lyciasalamandra luschani". IUCN Red List of Threatened Species. 2009: e.T41241A86525768. doi:10.2305/IUCN.UK.2009.RLTS.T41241A10422896.en.
  2. ^ a b c Frost, Darrel R. (2018). "Lyciasalamandra luschani (Steindachner, 1891)". Amphibian Species of the World: an Online Reference. Version 6.0. American Museum of Natural History. Retrieved 7 June 2018.
  3. ^ Arnold, E.N and J.A. Burton. 1978. A Field Guide to the Reptiles and Amphibians of Britain and Europe. Collins. London ISBN 0002199645.
  4. ^ a b c d e f g h i j k Polymeni, Rosa-Maria; Radea, Canella; Papanayotou, Costis (2011). "Diet Composition of the salamander Lyciasalamandra luschani basoglui on the Greek Island of Kastellorizo in the Southeast Aegean Sea" (PDF). Asian Herpetological Research. 2 (3): 155–160. doi:10.3724/SP.J.1245.2011.00155.
  5. ^ Karış, Mert; Göçmen, Bayram; Mermer, Ahmet. "axonomical and Biological Comparison of Two Luschan's Lycian Salamander, Lyciasalamandra luschani (Steindachner, 1891) (Urodela: Salamandridae) Populations from Southwestern Anatolia". 6: 107–136. {{cite journal}}: Cite journal requires |journal= (help)
  6. ^ Karış, Mert; Göçmen, Bayram; Mermer, Ahmet. "axonomical and Biological Comparison of Two Luschan's Lycian Salamander, Lyciasalamandra luschani (Steindachner, 1891) (Urodela: Salamandridae) Populations from Southwestern Anatolia". 6: 107–136. {{cite journal}}: Cite journal requires |journal= (help)
  7. ^ Veith, Michael & Steinfartz, Sebastian (2004). "When non-monophyly results in taxonomic consequences – the case of Mertensiella within the Salamandridae (Amphibia: Urodela)". Salamandra. 40 (1): 67–80. Archived 2013-09-21 at the Wayback Machine
  8. ^ a b c Budak, A. & Göçmen, B. (2005). Herpetology. Ege Üniversitesi Fen Fakültesi Kitaplar Serisi, No. 194, Ege Üniversitesi Basimevi, Bornova-Izmir
  9. ^ a b c (in Turkish) Bayram Göçmen Archived 2012-12-21 at archive.today. Fen.ege.edu.tr. Retrieved on 2013-01-03.
  10. ^ a b c d Weisrock, David W; Macey, J. Robert; Ugurtas, Ismail H; Larson, Allan; Papenfuss, Theodore J (1 March 2001). "Molecular Phylogenetics and Historical Biogeography among Salamandrids of the "True" Salamander Clade: Rapid Branching of Numerous Highly Divergent Lineages in Mertensiella luschani Associated with the Rise of Anatolia". Molecular Phylogenetics and Evolution. 18 (3): 434–448. doi:10.1006/mpev.2000.0905.
  11. ^ a b c d e f Zardoya, Rafael; Malaga-Trillo, Edward; Veith, Michael; Meyer, Axel (23 October 2003). "Complete nucleotide sequence of the mitochondrial genome of a salamander, Mertensiella luschani". Gene. 317 (1–2): 17–27. doi:10.1016/S0378-1119(03)00655-3. PMID 14604788.
  12. ^ a b Mezzasalma, Marcello; Odierna, Gaetano; Petraccioli, Agnese; Veith, Michael; Guarino, Fabio Maria (8 June 2021). "Karyological Diversification in the Genus Lyciasalamandra (Urodela: Salamandridae)". Animals. 11 (6): 1709. doi:10.3390/ani11061709.
  13. ^ a b Gautier, Patrick; Olgun, Kurtulus; Uzum, Nazan; Miaud, Claude (2006-04-01). "Gregarious behaviour in a salamander: attraction to conspecific chemical cues in burrow choice". Behavioral Ecology and Sociobiology. 59 (6): 836–841. doi:10.1007/s00265-005-0130-8. ISSN 1432-0762. S2CID 39664728.
  14. ^ a b c d e f Özeti, Neclâ (1967). "The Morphology of the Salamander Mertensiella luschani ( Steindachner ) and the Relationships of Mertensiella and Salamandra". American Society of Ichthyologists and Herpetologists ( ASIH ): 287–298.
  15. ^ Vitt, Laurie J.; Caldwell, Janalee P. (2014). Herpetology: An Introductory Biology of Amphibians and Reptiles (4th ed.). Academic Press. p. 169.
  16. ^ a b c Özeti̇, Neclâ (1979). "Reproductive Biology of the Salamander Mertensiella luschani antalyana". Herpetologica. 35 (3): 193–197. JSTOR 3891686.
  17. ^ a b Polymeni, R.; Spanakis, E.; Argiropoulos, A.; Rhizopoulou, S. (2010). "Aspects on the relief of living surfaces using atomic force microscopy allow "art" to imitate nature". Integrative Zoology. 5 (3): 218––25. doi:10.1111/j.1749-4877.2010.00207.x. PMID 21392340.
  18. ^ a b c d Başimoğlu Koca, Yücel; Karakahya, Feryal (2015). "The Structure of Stomach and Intestine of Triturus karelinii (Strauch, 1870) and Mertensiella luschani (Steindachner, 1891) (Amphibia: Urodela): Histological and Histometrical Study". Cumhuriyet University Faculty of Science Science Journal. 36 (1): 1–16.
  19. ^ a b Tzannetatou-Polymeni, R.; Haritos, A.A. (January 1989). "Physicochemical characterization and tissue distribution of esterases in two salamandridae species (Mertensiella luschani and Salamandra salamandra)". Comparative Biochemistry and Physiology Part B: Comparative Biochemistry. 92 (3): 469–475. doi:10.1016/0305-0491(89)90118-1. PMID 2706937.
  20. ^ a b Yildirimhan, Hikmet S.; Tunc, Mehmet R.; Sumer, Nurhan; Incedogan, Sezen; Bursey, Charles R. (2011). "Nematode parasites of Lyciasalamandra antalyana and Lyciasalamandra luschani (Caudata: Salamandridae) from Turkey". Comparative Parasitology. 78 (2): 375–377. doi:10.1654/4484.1. S2CID 86225907.
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Luschan's salamander: Brief Summary

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Luschan's salamander or Lycian salamander (Lyciasalamandra luschani) is a species of salamander in the family Salamandridae. It is found in the southwestern Anatolia in Turkey and adjacent Greece, in the island of Kastellorizo and its satellites.

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