Mertensiella Luschani
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Luschan's salamander or Lycian salamander (''Lyciasalamandra luschani'') is a species of
salamander Salamanders are a group of amphibians typically characterized by their lizard-like appearance, with slender bodies, blunt snouts, short limbs projecting at right angles to the body, and the presence of a tail in both larvae and adults. All ten ...
in the family
Salamandridae Salamandridae is a family of salamanders consisting of true salamanders and newts. Salamandrids are distinguished from other salamanders by the lack of rib or costal grooves along the sides of their bodies and by their rough skin. Their skin is v ...
. It is found in the southwestern
Anatolia Anatolia, tr, Anadolu Yarımadası), and the Anatolian plateau, also known as Asia Minor, is a large peninsula in Western Asia and the westernmost protrusion of the Asian continent. It constitutes the major part of modern-day Turkey. The re ...
in
Turkey Turkey ( tr, Türkiye ), officially the Republic of Türkiye ( tr, Türkiye Cumhuriyeti, links=no ), is a list of transcontinental countries, transcontinental country located mainly on the Anatolia, Anatolian Peninsula in Western Asia, with ...
and adjacent
Greece Greece,, or , romanized: ', officially the Hellenic Republic, is a country in Southeast Europe. It is situated on the southern tip of the Balkans, and is located at the crossroads of Europe, Asia, and Africa. Greece shares land borders with ...
, in the island of
Kastellorizo Kastellorizo or Castellorizo (; el, Καστελλόριζο, Kastellórizo), officially Megisti ( ''Megísti''), is a Greece, Greek island and Communities and Municipalities of Greece, municipality of the Dodecanese in the Eastern Mediterranean ...
and its satellites.


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 in length. 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. Males and females have similar weights. Male and female ''L. luschani'' are sexually dimorphic. 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. 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. 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.


Subspecies

The three
subspecies In biological classification, subspecies is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics (morphology), but that can successfully interbreed. Not all species ...
are: *''L. l. basoglui'' (Baran & Atatür, 1980)Budak, A. & Göçmen, B. (2005). Herpetology. Ege Üniversitesi Fen Fakültesi Kitaplar Serisi, No. 194, Ege Üniversitesi Basimevi, Bornova-IzmirBayram Göçmen
. Fen.ege.edu.tr. Retrieved on 2013-01-03.
*''L. l. finikensis'' (Basoglu & Atatür, 1975) *''L. l. luschani'' (
Steindachner Franz Steindachner (11 November 1834 in Vienna – 10 December 1919 in Vienna) was an Austrian zoologist, ichthyologist, and herpetologist. He published over 200 papers on fishes and over 50 papers on reptiles and amphibians. Steindachner describ ...
, 1891)


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''. ''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. 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. Scientists have also worked to categorize ''M. luschani'' into its appropriate phylogenetic positions based on different DNA sequences it possesses. 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. For example, the line of genetic code from the mitochondria produces 22 transfer RNAs, 2 ribosomal RNAs, and 13 peptides. 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. 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. 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.


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. 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.


Habitat and distribution

''L. luschani's'' natural
habitat In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...
s are temperate
forest A forest is an area of land dominated by trees. Hundreds of definitions of forest are used throughout the world, incorporating factors such as tree density, tree height, land use, legal standing, and ecological function. The United Nations' ...
s and arid Mediterranean environments. During active seasons, they live under shelters (i.e., rocks, wood) during the day, and at night, they forage. ''L. lushani'' populations are threatened by
habitat loss Habitat destruction (also termed habitat loss and habitat reduction) is the process by which a natural habitat becomes incapable of supporting its native species. The organisms that previously inhabited the site are displaced or dead, thereby ...
. One of its subspecies, M. 1.helverseni, is known to be found within a pine forest on the island of Carpathos. ''M. luschani'' has also been reported to inhabit about 350 kilometers of southwestern Turkish coast.


Conservation

One source, the Red Data Book of Threatened Animals of Greece, classifies the salamander, particularly ''L. luschani basoglui'', as vulnerable. Another source, the Red Data Book of the International Union for Conservation of Nature (IUCN) considers the salamander as an endangered species. Concerns about Luschan's salamander conservation are rooted in 1) habitat loss due to development, and 2) subspecies isolation.


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. 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.


Reproduction

Luschan's salamanders are
viviparous Among animals, viviparity is development of the embryo inside the body of the parent. This is opposed to oviparity which is a reproductive mode in which females lay developing eggs that complete their development and hatch externally from the m ...
. Females give birth to two offspring that are produced from each of the female's two
oviduct The oviduct in mammals, is the passageway from an ovary. In human females this is more usually known as the Fallopian tube or uterine tube. The eggs travel along the oviduct. These eggs will either be fertilized by spermatozoa to become a zygote, o ...
s.
Gestation Gestation is the period of development during the carrying of an embryo, and later fetus, inside viviparous animals (the embryo develops within the parent). It is typical for mammals, but also occurs for some non-mammals. Mammals during pregna ...
time ranges from 5–8 months. Using 9 salamanders from the M.l. antalyna subspecies, scientists discovered the basic reproductive biology of ''M. luschani''. The period for ''L. luschani'' birthing spans around two seasons, from the end of September to the end of March. 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.


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. ''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.


Anatomy and morphology

''M. luschani'' is said to have a flat and wide head. Its general trunk and body is orange with brown spots. 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''. ''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. ''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. 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. Additionally, the salamander possesses one cervical vertebra, about sixteen trunk vertebrae, one sacral vertebra, three caudosacral vertebrae, and at least two dozen caudal vertebrae. 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''. 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. 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. The small intestine is made up of similar layers and lacks the muscularis mucosa layer. 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''.


Physiology

Scientists have determined that the liver esterases present in ''M. l. luschani'' are all carboxylesterases. 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


Parasites

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


References

{{Taxonbar, from=Q2343974 Lyciasalamandra Amphibians of Europe Amphibians of Turkey Amphibians described in 1891 Taxa named by Franz Steindachner Taxonomy articles created by Polbot