AmphibiaWeb - Lyciasalamandra billae


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Lyciasalamandra billae (Franzen & Klewen, 1987)
Bay Lycian Salamander
family: Salamandridae
subfamily: Salamandrinae
genus: Lyciasalamandra
Species Description: Franzen, M., Klewen, R. (1987). "Mertensiella luschani billae ssp. n. —eine neue Unterart des Lykischen Salamanders aus SW- Anatolien." Salamandra 23: 132–141.
Taxonomic Notes: Elevated from subspecies to full species by Veith M, Steinfartz S (2004). "When non-monopyly results in taxonomic consequences - the case of Mertensiella within the Salamandridae (Amphibia:Urodela)." Salamandra 40:67-80.
Lyciasalamandra billae
© 2015 Dr. Joachim Nerz (1 of 48)
Conservation Status (definitions)
IUCN Red List Status Account Critically Endangered (CR)
National Status None
Regional Status None


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Lyciasalamandra billae is a Turkish salamander composed of at least five subspecies: L. b. billae, L. b. arikani, L. b. eikeae, L. b. irfani, and L. b. yehudahi. As a whole, their snout-vent length ranges from 55.0 – 82.0 mm, total length from 94.0 – 146.0 mm, and tail lengths range from 36.0 – 65.00 mm (Franzen and Klewen 1987, Göçmen et al. 2011, Göçmen and Akman 2012, Godmann et al. 2016). Their tails usually equal 80 to 90 percent of their snout-vent length (Veith and Steinfartz 2004). Lyciasalamandra billae is generally similar in external morphology to other members of the Lyciasalamandra genus. More specifically, their heads are slightly to significantly longer than wide (Franzen and Klewen 1987, Göçmen et al. 2011, Göçmen and Akman 2012, Godmann et al. 2016) and appear flattened. Their snout is rounded and their nostrils are small. The species has long, curved, narrow paratoid glands that are wider on the posterior end. There is also a distinct gular fold. The tips of the digits are thickened. Males of the species have a protuberance at the base of their tail on the dorsal surface that ranges from 1.59 – 2.49 mm in height and have swollen vents that are characteristic of the genus (Göçmen et al. 2011, Göçmen and Akman 2012, Godmann et al. 2016).

The genus Lyciasalamandra is closely related to the genera Chioglossa, Mertensiella, and Salamandra but can be distinguished by several morphological features. From Salamandra and Chioglossa, Lyciasalamandra can be differentiated by having an additional phalange on the first digit of the hand and foot with the phalange formula for Lyciasalamandra being 2, 2, 3, 2 on the hand and 2, 2, 3, 3, 2 on the foot. Additionally, Salamandra and Chioglossa lack the keratinized epidermal projections covering the entirety of the dorsal part of the body present in Lyciasalamandra. Specific differences from Chioglossa, include Lyciasalamandra having fully developed lungs unlike the reduced lungs of Chioglossa and Lyciasalamandra lacking the tongue-projection mechanism common to Chioglossa. Lyciasalamandra can be further distinguished from Salamandra based on osteology. Mertensiella, from which Lyciasalamandra was split, has a larger tail to body ratio and darker dorsal background coloration in juveniles than Lyciasalamandra (Veith and Steinfartz 2004).

Most subspecies of L. billae can be differentiated from other members of its genus by coloration, patterning, and locality (see Distribution section below). Although some individuals of L. b. billae can be dorsally black, the other subspecies have a lighter background coloration than L. atifi, L. flavimembris, L. helverseni, and L. luschani . Furthermore, L. atifi has a yellow upper eyelid with a thin black strip and L. luschani has a white upper eyelid that differentiates it from the typically black upper eyelid in most L. billae subspecies. However, L. b. arikani has a yellowish to white-green upper eyelid and L. b. eikeae has a yellow upper eyelid, but neither have a black strip. Most subspecies of L. billae also lacks yellow spots that L. helverseni and L. antayana have, black dorsal spot that L. fazilae has, and marbling like some populations of L. luschani, however, L. b. yehudahi does have some yellowish flecks and dots that are irregularly scattered on the dorsum (Franzen and Klewen 1987, Göçmen et al. 2011, Göçmen and Akman 2012, Goodman et al. 2016). Geographically, L. billae is parapatric to L. antalaya and near the range of L. luschani (Göçmen and Akman 2012, Goodman et al. 2016) making it distinguishable by geography from L. atifi, L. fazilae, L. flavimebris, and L. helverseni.

In life, the dorsal background color and patterning of L. billae varies with subspecies enough that several subspecies were thought to be full species before molecular analysis determined their genetic distance.

Lyciasalamandra. b. arikani have honey yellow to brown dorsal surfaces on the head (excepting the upper eyelids), legs, and tail with irregularly scattered red to brown flecks. The area around the snout and up to the eyes is lighter and more orange. The upper eyelids are yellowish to whitish green and have indistinct brownish or blackish dots. The paratoids are light orange, similarly to the snout, and the gland openings of the paratoids, back, and tail are black. There are yellowish-white flecks that start below the eyes, but do not form a distinct line, along the dorsolateral region of their flanks that separates the dorsum from the ventrum. Some individuals may have blackish blotches in the area near the flanks. In the skin of the ventrum on the lateral sides, legs, cloaca, and tail, is pale yellow. The throat of the ventrum lacks spots and is slightly transparent, showing the internal organs, with a reddish glow of the blood vessels (Göçmen and Akman 2012).

The dorsal base color of the head, body, and tail of L. b. billae ranges from salmon to black, with silvery-white iridophore spots up to 1 mm in diameter dispersed regularly across the back and often in two obvious dorsolateral bands that start below the nostrils and extends to the insertion of the hind limbs, with an interruption at the insertion of the forelimbs, that separates the dorsal and ventral sides. The dark brown to black coloration on the dorsal surface is the result of dense melanophores. Small dark spots are present on the nostrils. The upper eyelids are black but have a high density of the silvery white iridophores. The paratoids have the same background color as the dorsum but the gland openings on the parotoid, dorsum, and tail are black. There are dark brown melanophores spots above the elbows and knees. The abdominal skin is translucent, with a whitish iridophore marbling effect along the dorsolateral line that is denser toward the anterior region and between the forelimbs. As a result of the translucent ventrum, the abdominal organs such as the liver are dimly visible. The ventral surface of the limbs is salmon colored with the slivery-white iridophores. The surfaces of the hands and feet are white but the toes are tinted dark red due to strong vascularization. The underside of the tail is a bright salmon color with no melanophores or iridophores except round the cloaca, which has many iridophores (Franzen and Klewen 1987).

The dorsal surface of L. b. eikeae is light salmon-orange with adult males having irregular white flecks while females are patternless. The upper eyelids are yellow with a darker middle region. The paratoids are also yellowish with the gland openings of the paratoids, back, and tail being black. The dorsolateral lines are not continuous. The throat of the ventrum is slightly to moderately transparent with a reddish glow of the blood vessels. The ventrum of L. b. eikeae is flesh colored (Godmann et al. 2016).

The dorsum of L. b. irfani is aubergine reddish brown with white flecks or spots that have tiny dark brown spots. The area around the eyes, in particular the upper eyelids, are blackish with some small, indistinct white dots. The posterior quarter of the paratoids are reddish yellow and the gland openings of the paratoids, back, and tail are black. There are continuous dorsolateral lines formed by white flecks on their flanks that start at under the eyes and separates the dorsum from the ventrum. There may also be a lighter red vertebral stripe on some individuals. The throat is unmarked and somewhat transparent, showing the internal organs and having a reddish glow of blood vessels. The rest of the ventrum is reddish, especially on the ventral and lateral sides of the legs, clocal region, and tail (Göçmen et al. 2011).

The dorsal background color of L. b. yehudahi is brown. The dorsum has irregularly scattered yellowish-white flecks or dots that themselves have minuscule brown dots. The upper eyelids are almost black, but also have small, indistinct whitish dots that form a narrow band at the margin of the eyelids and the area around the anterior region of the eyes has a blackish tinge. The paratoids are orange and the gland openings of the paratoids, back, and tail are black. The limbs are also a lighter orange and sparsely covered with yellowish-white spots like the dorsum. There are concentrations of yellowish-white flecks that form an incomplete dorsolateral line that start under the eye and extending to separate the dorsum from the ventrum. The ventrum is flesh colored but transitions to orange on the lateral edges of the limbs, cloaca, and tail. The throat is slightly translucent, without any markings but with a reddish glow of the blood vessels (Göçmen and Akman 2012).

There is variation in coloration and patterning between (see above) and within subspecies. The species also displays sexual dimorphism with adult males having a curved spiny protuberance on the dorsal side at the base of their tails and a swollen cloaca. There is further sexual dimorphism within some subspecies (Göçmen et al. 2011, Göçmen and Akman 2012, Godmann et al. 2016).

In L. b. arikani there is differences between the sexes in paratoid size (length and width) and head length. Adult males are also darker than females and females have bulkier bodies (Göçmen and Akman 2012).

In L. b. irfani, adult females and juveniles have bulkier bodies and darker colorations with larger white flecks. Juveniles also have more yellowish paratoids than adults (Göçmen et al. 2011).

In L. b. yehudahi the ovaries of the female can be seen through the transparent skin, giving the posterior part of the ventrum a yellowish color. Like L. b. arikani and L. b. irfani, the females and juveniles of this subspecies also have bulkier bodies. Some juveniles have blacker colorations although others are colored similarly to adults but have scattered yellowish-orange blotches along the dorsal midline (Göçmen and Akman 2012).

Distribution and Habitat

Country distribution from AmphibiaWeb's database: Turkey

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As a whole, L. billae can be found on the western Anatolian coast of Turkey, starting in the Taurus Mountains northwest of Antayla (Godmann et al. 2016) and extending along the costal mountains south to the mountains between Beycik and Kumlucayazi (Göçmen and Akman 2012). The species is associated with rocky karstic limestone outcrops and is often found in pine or marqui woodlands that range from sea level to 200 meters above sea level (Veith et al. 2001). The arid Mediterranean habitat in which they live usually receives less than 1000 mm of rainfall per year (Çiçek et al. 2007), with typical rainfall between 400 - 600 mm per year (Göçmen et al. 2013). The species does not live in modified habitats (Veith et al. 2001).

The subspecies have non-overlapping geographic ranges that are isolated from each other. Lyciasalamandra b. arikani has the southern most range and appears to be restricted to 25 - 30 km2 area around Erentepe Mt. However its range may include the southern slopes of Tahtali Mountain and on the slopes and plains of “Ulupinar” between Beycik and Kumlucayazi at altitudes between 425 – 782 m a.s.l. They are found under stones on the humid and rocky slopes with Pinus brutia (Brutian pine), Quercus ilex (Holly oak), and Arbuthus andrachne (Greek strawberry trees) and other Mediterranean plants (Göçmen and Akman 2012).

The nominotypical subspecies, L. b. billae can be found 25 km southwest of Antalya on the eastern mountain slopes of Beydaglari Mountain Range (Franzen and Klewen 1987) and extending to north of Tahtali Mountain (Göçmen and Akman 2012). The soil they are found on is mostly covered with dense pine needles and a rich, base soil (pH = 8) (Franzen and Klewen 1987). The most northern subspecies, L. b. eikeae, is found northwest of Antalya in the Taurus mountains around Geyikbayırı at altitudes above 880 m a.s.l. This subspecies is separated from L. b. billae by and nested in populations of L. antalyana (Godmann et al. 2016).

Lyciasalamandra b. irfani can be found exclusively in Göynük Canyon in southwestern Anatolia at altitudes ranging from 90 - 385 m a. s. l. Individuals can be found under stones along rocky slopes with vegetation such as Platanus orientalis (Oriental plane), Pinus brutia (Brutian pine), Arbuthus andrachne (Greek strawberry trees), and Mediterranean shrubs (Göçmen et al. 2011).

Lyciasalamandra b. yehudahi is found on the rainy spring days on the eastern slopes of Tahtali Mountain (including Gedelme on the northwest slope) between Kemer stream and Tekirova, at altitudes of 87 – 646 m a.s.l. Their habitat is in calcareous rocky areas with woodlands composed of Platanus orientalis (Oriental plane), Pinus brutia (Brutian pine), Arbuthus andrachne (Greek strawberry trees), and Mediterranean shrubs (Göçmen and Akman 2012).

Life History, Abundance, Activity, and Special Behaviors
Lyciasalamandra billae is a terrestrial and nocturnal species, like many others in its genus. Their activity is dependent on temperature and humidity. During the day they often hide under stones, whereas in the evening they are active. They also appear during mild rain and cooler temperatures but are known to take refuge from dehydration deep inside the system of cracks in the limestone during the summer (Veith et al. 2001).

The ratio of males to females is 2.4:1 (Franzen and Klewen 1987).

This species is viviparous and produce one to two metamorphosed young. The gestation period lasts about one year (Stuart et al. 2008). Parturition time of Lycian salamanders is believed to be between the end of autumn and spring (September to March), corresponding with a higher humidity and average rainfall. The mating season is in the spring, predominantly in April (Polymeni 1994).

The adult Lycian salamander diet is composed of terrestrial invertebrates, especially larvae and adult beetles and centipedes (Çiçek et al. 2007).

Using phalangeal skeletochronology, males were found to reach an age of eight years, whereas females reached ten years of age. Juveniles ranged from 1 to 3 years of age, with sexual maturity estimated at 3 years of age. Overall, adult life expectancy is 5.4 years and annual survivorship is at 0.79 (Olgun et al. 2001).

Mertensiella luschani, the genus and species under which L. billae was initially listed, has been found to use chemical scents deposited on substrates to not only relocate previously used burrows but also to identify the occupants of a shelter as either male or female. These chemical tracks provide social rather than sexual information and do not play a role in sexual attraction (Gautier et al. 2006). It is unclear if L. billae also exhibits this character.

This species reproduces via live birth (Stuart et al. 2008).

Trends and Threats
As of January 2019, the IUCN Redlist has not assessed L. billae as a whole species. However, as four of the subspecies were considered full species until 2016, assessments or assessment recommendations of “Critically Endangered” have been made for those subspecies (Göçmen et al. 2011, Göçmen and Akman 2012, Kaska et al. 2017).

Despite begin found in the Çaliticak Protected Area, L. b. billae has an IUCN Redlist threat status of “Critically Endangered” (Kaska et al. 2017). The species is declining, a state attributed to its immensely restricted habitat that is decreasing in the quality and extent. There is a low human population in this subspecies’ range, but increasing development including tourism in the region, scientific collection of the species, and forest fires all threaten L. b. billae (Stuart et al. 2008, Kaska et al. 2017).

The type locality for L. b. arikani is not a Special Protected Area and has experienced forestry and urbanization activity, especially around the Ovacık plateau and Dağdibi mevkii. Given the small extent of this area, 25 - 30 km2, the subspecies fits the IUCN species’ criteria for a listing of “Critically Endangered (Göçmen and Akman 2012).

The Göynük Canyon, where L. b. irfani is found is a Special Protected Area, which is open to tourism and contains historic Lycian roads, traits which attract a decent number of tourists to the region. As L. b. irfani is only found in a 4 - 5 km2 area, it could be classified as “Critically Endangered”, according to the criteria of IUCN Red list Annex-2 (area of occupancy is less than 10 square km) (Göçmen et al. 2011).

Similarly, although L. b. yehudahi is found in a Special Protected Area, it is open to tourism traffic and has experienced unplanned urbanization around Servel Çukuru and Kuzdere, that is degrading the habitat. Because this area is only 35 – 40 km2, this subspecies would also fit the IUCN criteria for “Critically Endangered” (Göçmen and Akman 2012).

Relation to Humans
The species is potentially over collected for scientific research purposes (Stuart et al. 2008). This species is not specifically important to the local community or to tourism.

Possible reasons for amphibian decline

General habitat alteration and loss
Habitat modification from deforestation, or logging related activities
Subtle changes to necessary specialized habitat
Loss of genetic diversity from small population phenomena
Intentional mortality (over-harvesting, pet trade or collecting)

The species authority for L. billae is: Franzen, M. and Klewen, R. (1987). “Mertensiella luschani billae ssp. n. – eine neue Unterart des Lykischen salamanders aus SW-Anatolien (Caudata: Salamandridae).” Salamandra 23 (2/3), 132-141.

The subspecies authority for L. b. arikani and yehudahi is: Göçmen, B., Akman, B. (2012). “Lyciasalamandra arikani n. sp. and L. yehudahi n. sp. (Amphibia: Salamandridae), two new Lycian salamanders from southwestern Anatolia.” North-Western Journal of Zoology. Oradea, Romania 8: 181–194.

The subspecies authority for L. b. eikeae is: Godmann, O., Kariş, M., Göçmen, B. (2016). “Geographic nestedness of Lyciasalamandra billae (Amphibia: Salamandridae) populations within L. antalyana and description of a new subspecies.” Zoology in the Middle East. Heidelberg 62: 255–260.

The subspecies authority for L. b. irfani is: Göçmen, B, Arikan, H, Yalçinkaya, D. (2011). ''A new Lycian salamander, threatened with extinction, from the Göynük Canyon (Antalya, Anatolia), Lyciasalamandra irfani n. sp. (Urodela: Salamandridae)”. North-Western Journal of Zoology 7 (1): 151-160

Lyciasalamandra billae was initially described by Michael Frazen and Reiner Klewen in 1987 as a subspecies of L. luschani called Mertensiella luschani billae. It was elevated to species level by Michael Veith and Sebastian Steinfartz in 2004 through molecular analysis, though the authors maintain that the species is morphologically distinct from L. luschani as well (Veith and Steinfartz 2004).

Parsimony analysis of 20 mitochondrial DNA sequences revealed that Lyciasalamandra was significantly different from the genus Mertensiella, however the authors chose to place the species with in the genus Salamandra (Weisrock et al. 2001). Additional allozyme analysis of 18 presumptive nuclear gene loci revealed enough genetic differences that the genus Lyciasalamandra was created (Veith and Steinfartz 2004). And further analysis of 4500 base pairs of mitochondrial DNA revealed that previously described species, L. arikani, L. irfani, and L. yehudahi were subspecies of L. billae. It also showed that L. billae is sister to L. atifi. The clade composed of the two species is sister to the clade composed of L. antalyana, L. fazilae and L. helverseni and the genus Lyciasalamandra is sister to the genus Salamandra (Veith et al. 2016).

Lyciasalamandra billae is known to hybridize with L. antalyana (Johannesen et al. 2006).

The meaning for the genus Lyciasalamandra can be broken up into two parts: “Lycia” is the name of a historical Roman province in the south of Turkey and refers to the salamanders’ geographical distribution. The second half of the genus implies a close phylogenetic relationship with Lyciasalamandra’s sister genus, Salamandra (Veith and Steinfartz 2004).

The species epithet, billae, was given in honor of Bille Nicolai, the girlfriend of Michael Franzen, a co-author of the description, for her energetic support of during amphibian surveys (Franzen and Klewen 1987).

The subspecies name, arikani, was given to honor Hüseyin Arikan, a Turkish herpetologist who made valuable contributions to Turkish herpetological knowledge (Göçmen and Akman 2012).

The subspecies name, eikeae, was given in honor of one of the author’s mother, Mrs. Eike Bettina Godmann (Godmann et al. 2016).

The subspecies name, irfani, was given in honor of the senior author's late father, Irfan Göçmen (Göçmen et al. 2011).

The subspecies name, yehudahi, was given to honor Yehudah Leopold Werner, the senior author’s second academic father and close friend who is a Professor Emeritus in the Department of Ecology, Evolution, and Behavior at the Hebrew University of Jerusalem, Israel and who made valuable contributions to Middle Eastern herpetological knowledge (Göçmen and Akman 2012).

Molecular clock analysis of the genus Lyciasalamandra places the species split into seven major lineages at 12.3 – 10.2 million years ago, which coincides with the emergence of the final Aegean trench. However, the Messinian Salinity Crisis is likely to be responsible for first divergence events of subspecies within L. luschani, L. fazilae and L. flavimembris. The diversification of other subspecies can be attributed to the Pliocene and Pleistocene glaciations (Vieth et al. 2016).

The subspecies, L. b. eikeae likely arose after L. b. billae experienced a range expansion into the Geyikbayiri Montains but was later isolated by climatic conditions. Some time after L. b. eikeae's isolation, L. antalyana colonized the area between the two subspecies, further isolating L. b. eikeae. At the time of the subspecies description it was unknown if L. b. eikeae has contact with L. antalyana (Godmann et al. 2016)

Three helmith species of the phylum Nematoda were found on Lyciasalamandra billae: Aplectana acuminata, Cosmocerca longicauda, and Angiostoma aspersae. This represents the first time helmith species have been found to parasitize any species within Lyciasalamandra (Yildirimhan and Öz 2008).


Franzen, M., Klewen, R. (1987). ''“Mertensiella luschani billae ssp. n. – eine neue Unterart des Lykischen salamanders aus SW-Anatolien (Caudata: Salamandridae).'' Salamandra , 23(2/3), 132-141.

Frost, D. R. (2014). “Lyciasalamandra Veith and Steinfartz, 2004.”Amphibian Species of the World: an Online Reference. Version 6.0 (Feb. 12, 2014). Electronic Database accessible at American Museum of Natural History, New York, USA.

Gautier, P., Olgun, K., Uzum, N., Miaud, C. (2006). ''Gregarious behaviour in a salamander: Attraction to conspecific chemical cues in burrow choice.'' Behavioral Ecology and Sociobiology , 59(6), 836-841.

Godmann, O., Kariş, M., Göçmen, B. (2016). ''Geographic nestedness of Lyciasalamandra billae (Amphibia: Salamandridae) populations within L. antalyana and description of a new subspecies.'' Zoology in the Middle East, 62, 255-260.

Göçmen B, Arikan H, Yalçinkaya D. (2011). ''A new Lycian salamander, threatened with extinction, from the Göynük Canyon (Antalya, Anatolia), Lyciasalamandra irfani n. sp. (Urodela: Salamandridae).'' North-western Journal of Zoology Romania, 7, 151-160.

Göçmen, B., Akman, B. (2012). ''Lyciasalamandra arikani n. sp. and L. yehudahi n. sp. (Amphibia: Salamandridae), two new Lycian salamanders from southwestern Anatolia.'' North-Western Journal of Zoology. Oradea, Romania, 8(1), 181–194. [link]

Göçmen, B., Veith, M., Akman, B., Godmann, O., İğci, N., Oğuz, A. (2013). ''New records of the Turkish Lycian salamanders (Lyciasalamandra, Salamandridae).'' North-Western Journal of Zoology, 9(2), 319-328.

Johannesen, J., Johannesen, B., Griebeler, E. M., Baran, I., Tunç, M. R., Kiefer, A., Vieth, M. (2006). ''Distortion of symmetrical introgression in a hybrid zone: evidence for locus-specific selection and uni-directional range expansion.'' Journal of Evolutionary Biology, 19(3), 705-716.

Kaska, Y., Kumluta, 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., Denoël, M. (2017). ''Lyciasalamandra billae ssp. billae (amended version of 2009 assessment).'' The IUCN Red List of Threatened Species 2017: e.T61915A112693602. Downloaded on 17 January 2019.

Olgun, K., Miaud, C., Gautier, P. (2001). ''Age, growth, and survivorship in the viviparous salamander Mertensiella luschani from southwestern Turkey.'' Canadian Journal of Zoology , 79, 1559-1567.

Polymeni, R.M. (1994). ''On the biology of Mertensiella luschani (Steindachner, 1891): a review.'' Mertensiella, 4, 301-314.

Stuart, S., Hoffmann, M., Chanson, J., Cox, N., Berridge, R., Ramani, P., Young, B. (eds) (2008). Threatened Amphibians of the World. Lynx Edicions, IUCN, and Conservation International, Barcelona, Spain; Gland, Switzerland; and Arlington, Virginia, USA.

Veith, M., Baran, I., Godmann, O., Oz, M., Tunc, M.R. (2001). ''A revision of population designation and geographic distribution of the Lycian Salamander Mertensiella luschani (Steindachner, 1891).'' Zoology in the Middle East , 22(1), 67-82.

Veith, M., Göçmen, B., Sotiropoulos, K., Kieren, S., Godmann, O., Steinfartz, S. (2016). ''Seven at one blow: the origin of major lineages of the viviparous Lycian salamanders (Lyciasalamandra Veith and Steinfartz, 2004) was triggered by a single paleo-historic event.'' Amphibia-Reptilia , 37, 373-387.

Veith, M., Steinfartz, S. (2004). ''When non-monophyly results in taxonomic consequences – the case of Mertensiella within the Salamandridae (Amphibia: Urodela).'' Salamandra, 40(1), 67-80.

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.

Yildirimhan, H.S., Oz, M. (2008). ''Helminth fauna of Lyciasalamandra billae (Franzen & Klewen) (Luschan salamander) collected from Antalya.'' Turkish Society for Parasitology, 32(4), 390-392.

Çiçek, K., Tok, C.V., Mermer, A., Tosungolu, M., Ayaz, D. (2007). ''Food Habits of the Lycian Salamander, Lyciasalamandra fazilae (Başoğlu and Atatür, 1974): Preliminary Data on Dalyan Population.'' North-Western Journal of Zoology, 3(1), 1-8.

Originally submitted by: Bryanna Mariel Andrews, Ayala Noga Berger, Emily Lauren Wong (first posted 2019-01-17)
Larva by: Ann T. Chang (updated 2023-02-23)

Edited by: Ann T. Chang and John Cavagnaro (2023-02-23)

Species Account Citation: AmphibiaWeb 2023 Lyciasalamandra billae: Bay Lycian Salamander <> University of California, Berkeley, CA, USA. Accessed Jun 15, 2024.

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Citation: AmphibiaWeb. 2024. <> University of California, Berkeley, CA, USA. Accessed 15 Jun 2024.

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