ArticlePDF Available

A new subspecies of Lyciasalamandra antalyana (Amphibia: Salamandridae) from the Lycian Coast, Turkey

A new subspecies of Lyciasalamandra antalyana from Turkey
All articles available online at
© 2014 Deutsche Gesellscha für Herpetologie und Terrarienkunde e.V. (DGHT), Mannheim, Germany
SALAMANDRA 50(3) 125–132 30 October 2014 ISSN 0036–3375
A new subspecies of Lyciasalamandra antalyana
(Amphibia: Salamandridae) from the Lycian Coast, Turkey
B A O G
1) Zoology Section, Department of Biology, Faculty of Science, Ege University, 35100 Bornova, Izmir, Turkey
2) Hauptstr. 33, 65527 Niedernhausen, Germany
Corresponding author: B A, email:
Manuscript received: 20 February 2014
Accepted 23 May 2014 by S L
Abstract. A new subspecies of the Lycian salamander Lyciasalamandra antalyana is described from Yağcavillage (Antalya
province) and Burdur province on the Lycian Coast, Turkey. It is distinguished from the nominotypical form by its dorsal
colouration, multivariate morphometrics, and mitochondrial molecular markers.
Key words. Urodela, Lyciasalamandra antalyana gocmeni ssp. n., SrDNA gene, Turkey.
Ten species and three subspecies have so far been de-
scribed (G et al. ) in the genus Lyciasalamandra
(V  S ). eir occurrence is limited to
a narrow area along the Lycian Coast in southern Turkey
and some Aegean islands. All species and subspecies have
relatively small and mostly allopatric distributions. In spite
of their geographic proximity, species and subspecies dier
signicantly in colouration, and populations may exhibit
substantial dierences colour and pattern even within taxa
(V  S ).
During the last two decades, several new species of
Lycia salamandra and a large number of new populations
have been recorded (e.g., V et al. , G et
al. , G  A ). Especially in Lyciasala-
mandra antalyana (B  B, ), the known
distribution was substantially extended towards the north.
Just a small distance to the north from the type locality at
Hurma village, recorded specimens are characterized by
pronounced dierences in terms of colouration (V et
al. ). Specimens from these newly discovered popu-
lations (e.g., Yağcavillage and ermessos) exhibit a ful-
ly striped yellow pattern, while topotypical animals from
Hurma are mainly yellow in the region of the eyes, nostrils
and parotoids (Fig. ).
A et al. () published a compilation of the
known L. antalyana populations and discussed if an as-
signment to subspecic status of the newly discovered yel-
low-coloured populations might be warranted. Based on
molecular data, multivariate analyses of morphometrics,
and the striking colouration, we here suggest the popula-
tions from Yağca village and farther north in the Burdur
province as a new subspecies.
Material and methods
Field trips took place between February of  and No-
vember of . Geographical coordinates for sample loca-
tions as given in Table  were computed with a Magellan XL
GPS. Digital colour photos were taken of all live specimens.
Specimens were euthanised by a  ethanol injection into
their body cavity and subsequently xed in  ethanol.
Additional specimens were collected from the type locality
of the species (Hurma/Antalya), as well as from Gökdere
and Hacısekililer for comparison with the new populations
from Yağca (Antalya) and Burdur. Specimens are deposited
at e Zoology Museum of Harran University, Şanlıurfa,
Turkey (ZMHRU; Tab. ). All currently known localities of
L. antalyana are illustrated in Figure .
We used the morphometric data published by A et
al. () to perform multiple discriminate analyses (DC)
with PAST version . (H et al. ). We prefer
the DC to a principle component analysis (PCA) since DC
is specically designed for testing the power of characters
to distinguish between pre-dened groups (here: poten-
tial subspecies; S R ). We studied males
and females separately to make provision for potential
dierences among sexes as was previously described for
other Lyciasalamandra species (e.g., G et al. ,
G  A ). Obvious juveniles could eas-
ily be distinguished from adults by their smaller size and
the lack of the hedonic gland; they were excluded from
B A  O G
analyses. We rst used the raw measurements of A
et al. (): rostrum–anus length (RA), tail length (TL),
nostril–eye distance (NED), distance between nostrils
(DBN), eye diameter (ED), head length (HL), head width
(HW), parotoid length (PL), parotoid width (PW), fore
limb length (FLL), hind limb length (HLL), and distance
between fore- and hindlimbs (DFHL). In a second analy-
sis, we employed ratios of these measurements calculated
against RA (PERCRA values) to correct for the overall size
of the specimens. We analysed a total of  and  males of
L. a. antalyana and L. a. gocmeni ssp. n., respectively, and
 females of each subspecies.
A segment of  base pairs (bp) in length of the Sr-
RNA gene was sequenced (for DNA extraction, primers,
PCR conditions and sequencing details, see V et al.
) for  specimens collected at new localities of the
new subspecies and provided by A et al. ():
Kavacık ( specimen), Kızılseki (), Kocaaliler  (), Ko-
caaliler () and Kırkgözhan, Yağca () (GenBank acces-
sion number of the single new haplotype: KJ; www. We aligned them with  homologous
sequences from V et al. () of additional Lycia-
salamandra populations as well as Salamandra infraim-
maculata, Mertensiella caucasica, Neurergus crocatus and
Pleuro deles poireti as outgroup taxa (GenBank accession
numbers: EU, EU, EU, EU,
EU, EU, EU, EU, EU,
EU, EU, EU, EU, EU,
EU, and EU) using the clustal W option in
Mega . (T et al. ). We used jModeltest, ver-
sion .. (D et al. ), to select the best t out of
 nested substitution models. e HKY+G model (gam-
ma shape parameter α = .) was selected based on the
Bayesian Information Criterion (BIC). However, HKY is
Figure 1. Map showing localities of Lyciasalamandra antalyana antalyana (solid circles) and L. a. gocmeni ssp. n. (asterisks); numbers
are according to Table 1.
A new subspecies of Lyciasalamandra antalyana from Turkey
not supported by Mega . and we therefore used the Ta-
mura-Nei substitution model (T  N ), since
the HKY model is a special feature of the Tamura-Nei
model that does not distinguish between substitution rates
for the two types of transformations.
According to V et al. (), a neighbour-joining
(NJ) tree of Lyciasalamandra mitochondrial genes pro-
duced equivalent results of maximum likelihood, maximum
parsimony, and Bayesian inference trees. We therefore only
calculated an NJ tree from , bootstrap replicates using
Mega . and applying the selected substitution model.
In terms of colour and pattern, specimens from Yağca
and Burdur populations dier from Hurma populations
(Figs, ). Yağca and Burdur populations have a more dis-
continuous pattern than Hurma populations. Additionally,
comparing the Yağca and Burdur populations, the discon-
tinuous dorsal pattern of Burdur specimens is dominated
by the dark background, while yellow is dominant in the
Yağca population. e dorsal colouration patterns of either
dier clearly from the nominotypical form. Yağca and Bur-
dur specimens have yellow dorsal markings that also occur
on the upper side of the tail. In contrast, specimens from
Hurma are yellow only in the region of the eyes, nostrils
and parotoids (Figs , ). All other taxa of Lyciasalamandra
do not exhibit any such yellow coloration.
Based on our raw data, the multiple discriminate func-
tion allowed to correctly classify both males and females
from Yağca and Burdur versus other populations only at 
and , respectively (Fig. ). e probability that males
and females of both taxa belong to the same group was
. and ., respectively. However, when using ra-
tios (Fig. ), the percentages of correctly classied speci-
mens of both sexes were  in males and  in females
(psame group = . + ., respectively).
Table 1. Geographic and some climatic information on the localities of Lyciasalamandra antalyana antalyana and L. a. gocmeni ssp. n.,
as well as museum numbers of the specimens used. e numbers in brackets correspond to the localities shown in Figure 1. Asterisk:
at the time of collecting.
Museum numbers
(ZMHRU) Locality Altitude
(m a.s.l.)
Collection date
number of specimens
2012/6 North of Kocaaliler [1] 750 37°20’N 30°42’E 29.03.2012
1 juv.
2012/5 Hamartaşı/Kocaaliler [2] 720 37°20’N 30°42’E 01.04.2012
5 (2♂♂, 3♀♀)
2012/42 Hamartaşı/Kocaaliler [2] 720 37°20’N 30°42’E 10.04.2012
2 (1, 1 juv.)
2012/9 Kavacık [3] 653 37°17’N 30°44’E 01.04.2012
2 (1, 1)
2012/44 Kavacık [3] 653 37°17’N 30°44’E 10.04.2012
1 juv.
2012/8 North of Kızılseki [4] 400 37°16’N 30°45’E 01.04.2012
4 (2♂♂, 1, 1 juv.)
2012/7 West of Kızılseki [5] 438 37°15’N 30°44’E 01.04.2012
2012/43 Ortacamevki/Kızılseki [6] 541 37°15’N 30°44’E 10.04.2012
1 juv.
2012/2 Kırkgözhan, Yağca [7] 348 37°06’N 30°34’E 07.03.2012
10 (2♂♂, 6♀♀, 2 juv.)
2012/3 Kırkgözhan, Yağca [8] 350 37°06’N 30°34’E 12.03.2012
2 (1, 1 juv.)
2012/4 Çığlık [9] 313 37°03’N 30°33’E 12.03.2012
3 (1, 1, 1 juv.)
2011/82 Hurma 99 36°51’N 30°35’E 25.02.2011
12 (1, 3♀♀, 8 juv.)
2012/1 Hurma 99 36°51’N 30°35’E 06.03.2012
17 (8♂♂, 8♀♀, 1 juv.)
2013/173 Gökdere 50 36.82925°N 30.55599°E 24.11.2013
1 (1 juv.)
2013/175 Hacısekililer 581 36.80321°N 30.48953°E 26.11.2013
3 (1, 1, 1 juv.)
B A  O G
Figure 2. (a) Male of Lyciasalamandra antalyana gocmeni ssp. n. from the type locality, Kırkgözhan, Yağca; (b) male, (c) female, and
(d)juvenile of L. a. gocmeni ssp. n. from Kızılseki.
Figure 3. Male (a), female (b) and juvenile (c) of Lyciasalamandra antalyana antalyana from the type locality, Hurma village; some
variation in colour pattern is shown in (d).
A new subspecies of Lyciasalamandra antalyana from Turkey
Of the  base pair positions, ve were diagnostic be-
tween the two L .antalyana lineages (=  sequence di-
vergence in uncorrected p-distances). Most of the new-
ly sequenced specimens shared the haplotype found by
V et al. () in specimens from Yağca village. Only
one additional haplotype with one mutation was found at
Kırkgözhan, Yağca village. In the NJ tree, the haplotypes
of the new subspecies form a clade of their own with 
bootstrap support (Fig. ).
Five diagnostic base positions within the studied -bp
fragment of the SrRNA mitochondrial gene, dierenc-
es surfacing from a multivariate analysis of morphomet-
ric ratios, and a distinctive colouration clearly separate the
suggested new subspecies from the nominotypical form.
Based on these results, we describe the Yağca and Burdur
populations as:
Lyciasalamandra antalyana gocmeni ssp. n.
Holotype and type locality (Figs a, ): Adult male, ZMHRU
/- from Kırkgözhan/Yağca, Antalya Province, Tur-
key,  m above sea level (°’.”N, °’.”E).
Leg.  March  by B. G, B. A, N. İ,
O.G and M. V.
Paratypes:  specimens collected from Antalya (Yağca and
Çığlık populations) and Burdur (Kavacık, Kızılseki, and
Kocaaliler populations) were deposited as paratypes. For
locality details and collection numbers see Table .
Diagnosis: Lyciasalamandra antalyana gocmeni ssp. n. dif-
fers from all other Lyciasalamandra species and subspecies
by having yellow dorsal and supracaudal markings in life.
Figure 4. Multiple discriminate analysis of males and females of L. a. antalyana (black) and L. a. gocmeni ssp. n. (grey) based on raw
data; males: Hotelling’s t² = 54.834, F = 1.924; females: Hotelling’s t² = 10.81, F = 0.4504.
Figure 5. Multiple discriminate analysis of males and females of L. a. antalyana (black) and L. a. gocmeni ssp. n. (grey) based on ratios;
males: Hotelling’s =58.741, F = 3.0916; females: Hotelling’s t² = 43.601, F = 2.5764.
B A  O G
In contrast, specimens of L . a. antalyana are yellow only
in the region of the eyes, nostrils and parotoids. No other
taxon of Lyciasalamandra shows any such yellow coloura-
Description of the holotype: e body shape is equivalent
to other species of Lyciasalamandra. Head at, longer than
broad (HW/HL .). Snout rounded. Parotoids long and
narrow (PW/PL .), the posterior part broader than the
anterior part. Gular fold distinct. e cloacal region shows
a very slight swelling, and the pads on the upper arm are
quite developed. In prole, the tiny thorns on the dorsal
side are visible. e nger-like projection above the base
of the tail measures about . mm tall; it is pointed and
curved forward at its free end.
In life, the colouration of the dorsum including the head
and upper jaws was brownish-red, especially on the paro-
toids that were dotted with  black dermal pores on each
side. On each upper eyelid, there was a thin black cross bar.
e interparotoid, interorbital and internasal spaces were
yellow with brown ecks (Fig. a).
e ground colour of the dorsal side of the trunk had
an interrupted light yellow pattern. is pattern extended
backwards into the median region of the trunk. Two broad
brown stripes that were intersected by transverse brown
bars extended along the dorsolateral sides of the trunk.
Yellow was dominant on the dorsum. e legs and the tail
were esh-coloured. e proximal half of each leg had a
Figure 6. NJ tree of the Tamura-Nei distances of Lyciasalamandra antalyana, showing the monophyletic position of L. a. gocmeni;
numbers at branches indicate bootstrap support ≥ 70% for 2,000 replicates.
Figure 7. Type locality of Lyciasalamandra antalyana gocmeni
ssp.n. at Kırkgözhan, Yağca.
A new subspecies of Lyciasalamandra antalyana from Turkey
light yellow maculation, and a light brown reticulation was
found on elbows and knees. Tiny ecks were present on
the dorsal side of some nger joints. On the dorsal side of
the tail, which had rows of black dermal pores, there were a
light brown and yellow maculation. e lower parts of the
trunk, tail and legs were esh-coloured. e throat region
had a yellow tinge.
Measurements of the holotype (in mm): Total length
(ToL) .; RA and length of trunk (LT) . and .,
respectively; TL .; NED .; DBN .; ED .; HL
.; HW .; PL .; PW .; FLL .; HLL .;
DFHL .
Paratype variation: Variation observed in some morpho-
metric characters and ratios vis-à-vis the holotype are
summarized separately for adults and juveniles in Table .
Sexual dimorphism was observed within the population
(p ≤ .) regarding HL, PL, PW and FLL in raw data or
PERCRA values. In addition, the projection at the base of
the tail in nine male specimens ranged between . and
. mm with an average of . mm. As far as the colour
pattern is concerned, the description given for the holo-
type largely applies to the other males and females as well
(Fig.). e discontinuous light yellow parts in the pattern
of adult females are broader and more pronounced than
that of adult males. Juveniles sport a darker ground colour
with a more intensely yellow and brown pattern on their
dorsum. From this series and the specimens observed in
the eld, it would appear that the pattern as well as the dor-
sal background colour change with age and depending on
sex. Both females and juveniles lack any protuberance at
their tail bases and have smooth or less swollen cloacae.
Habitat, geographic distribution, and ecology: We found
Lyciasalamandra antalyana gocmeni ssp. n. under rocks in
a karst-dominated landscape. In the dry months of sum-
mer, the animals use hollow spaces in the karst or heaps of
rock for sheltering. e hillside is usually vegetated with
plane (Platanus orientalis) and pine trees (Pinus brutia).
Activity outside of the shelters is highly dependent on tem-
peratures and humidity. All localities recorded lie with-
in the potential distribution predicted by R et al.
(). e habitat of the new subspecies extends into the
Taurus mountain range at the western border of the An-
talya lowlands. In the east, the area is bordered by the Aksu
valley. In the west and north, the forest-free tablelands of
the Taurus do not allow further expansion.
Derivatio nominis: e name of the new subspecies has
been chosen in honour of the Turkish herpetologist, Pro-
fessor Dr. B G. Besides his contributions to
herpetology, he is a role model and a good friend.
anks to M V for providing unpublished DNA data.
A, B., B. G, N. İ,  D. Y (): Range
extension of Lyciasalamandra antalyana. –Biharean Biologist,
: –.
B, M.  I. B (): e subspecic status of the
population of Mertensiella luschani (Steindachner) in the An-
talya region of Southwestern Anatolia. – Ege Üniversitesi Fen
Fakültesi İlmi Raporlar Serisi, : –.
D, D., G. L. T, R. D D. P ():
jModel Test . More models, new heuristic and parallel com-
puting. – Nature Methods, : .
Table 2. Some morphometric characters (in mm) and ratios of
Lyciasalamandra antalyana gocmeni ssp. n. 1: Values in raw data;
2: Values in PERCRA; N: number of specimens; SD: Standard
deviation; the other abbreviations of characters are given in Ma-
terial and methods.
Juveniles (N = 10)
Mean±SD (Range)
Adults (N = 22)
Mean±SD (Range)
ToL 1 86.50±9.76 (77.00–108.00) 130.68±15.65 (101.00–153.00)
2 1.79±0.08 (1.72–1.93) 1.79±0.06 (1.66–1.89)
RA 1 48.40±5.52 (42.00–60.00) 72.95±7.67 (60.00–86.00)
LT 1 33.07±3.36 (29.13–40.12) 48.25±5.39 (39.50–58.17)
2 0.69±0.05 (0.61–0.76) 0.66±0.02 (0.61–0.69)
TL 1 38.10±5.02 (33.00–48.00) 57.73±8.49 (40.00–68.00)
2 0.79±0.08 (0.72–0.93) 0.79±0.06 (0.66–0.89)
NED 1 2.27±0.32 (1.67–2.92) 2.92±0.41 (2.22–3.68)
2 0.05±0.01 (0.04–0.06) 0.04±0.00 (0.03–0.05)
DBN 1 3.72±0.32 (3.24–4.26) 5.09±0.54 (4.13–5.92)
2 0.08±0.00 (0.07–0.08) 0.07±0.01 (0.06–0.08)
ED 1 3.28±0.42 (2.84–4.14) 4.27±1.27 (3.26–9.78)
2 0.07±0.01 (0.05–0.09) 0.06±0.02 (0.05–0.16)
HL 1 13.58±0.87 (12.41–15.37) 17.61±1.49 (15.12–20.94)
2 0.28±0.02 (0.26–0.32) 0.24±0.01 (0.22–0.26)
HW 1 8.75±0.74 (7.61–9.89) 11.42±0.88 (9.88–12.84)
2 0.18±0.02 (0.16–0.21) 0.16±0.01 (0.14–0.18)
PL 1 6.37±0.77 (5.28–7.25) 8.59±0.78 (7.31–10.29)
2 0.13±0.02 (0.11–0.16) 0.12±0.01 (0.10–0.14)
PW 1 2.24±0.39 (1.68–2.77) 3.22±0.46 (2.61–4.11)
2 0.05±0.01 (0.04–0.06) 0.04±0.00 (0.04–0.05)
FLL 1 15.94±1.87 (13.82–19.49) 21.72±1.65 (18.20–24.75)
2 0.33±0.02 (0.31–0.36) 0.30±0.02 (0.26–0.34)
HLL 1 18.10±2.75 (15.00–23.47) 26.19±2.16 (21.79–28.92)
2 0.37±0.03 (0.34–0.41) 0.36±0.02 (0.34–0.40)
DFHL 1 26.83±2.86 (22.23–31.52) 39.91±4.84 (31.54–47.10)
2 0.56±0.05 (0.50–0.65) 0.55±0.03 (0.49–0.58)
HW/HL 1 0.64±0.04 (0.56–0.71) 0.65±0.04 (0.59–0.74)
TL/TBL 1 0.44±0.02 (0.42–0.48) 0.44±0.02 (0.40–0.47)
PW/PL 1 0.35±0.03 (0.28–0.38) 0.37±0.04 (0.30–0.44)
NED/HL 1 0.17±0.02 (0.13–0.19) 0.17±0.02 (0.13–0.20)
B A  O G
G B., H. A  D. Y(): A new Lycian
salamander, threatened with extinction, from the Göynük
Canyon (Antalya, Anatolia), Lyciasalamandra irfani n. sp.
(Urodela: Salamandridae). North-Western Journal of Zoo-
lo gy, : –.
G, B.  B. A (): Lyciasalamandra arikani n. sp.
& L. yehudahi n. sp. (Amphibia: Salamandridae), two new Ly-
cian salamanders from Southwestern Anatolia. – North-West-
ern Journal of Zoology, : –.
G, B., M. V, B. A, O. G, N. İ,  M.
A. O (): New records of the Turkish Lycian salaman-
ders (Lyciasalamandra, Salamandridae). North-Western
Journal of Zoology, : –.
H, Ø., D. A. T. H,  P. D. R (): PAST: Pale-
ontological statistics soware package for education and data
analysis. – Palaeontologia Electronica, : –.
R, D., S. L, M. Ö, S. B, K. E
 M. V (): A novel method to calculate climatic niche
similarity among species with restricted ranges – e case of
the terrestrial Lycian salamanders. – Organisms, Diversity and
Evolution, : –.
S, R. R.  F. J. R (): Biometry: the principles and
practice of statistics in biological research. ird edition. –
Freeman, New York.
T, K. M. N (): Estimation of the number of nu-
cleotide substitutions in the control region of mitochondrial-
DNA in humans and chimpanzees. – Molecular Biology and
Evolution, : –.
T, K., G. S, D. P, A. F  S. K
(): MEGA: Molecular evolutionary genetics analysis ver-
sion .. – Molecular Biology and Evolution, : –.
V M., S. S, R. Z, A. S A. M
(): A molecular phylogeny of ‘true’ salamanders (family
Salamandridae) and the evolution of terrestriality of repro-
ductive modes. – Journal of Zoological Systematics and Evo-
lutionary Research, : –.
V, M., I. B, O. G, A. K, M. Ö M. R.
T (): A revision of population designation and geo-
graphic distribution of the Lycian salamander Mertensiella
luschani (Steindachner, ). Zoology in the Middle East,
: –.
V, M.  S. S (): When non-monophyly re-
sults in taxonomic consequences – the case of Mertensiella
within the Salamandridae (Amphibia: Urodela). Salaman-
dra, : –.
V, M., E. L, M. Ö, A. K, I. B, R. M. P-
 S. S (): Cracking the nut: geographi-
cal adjacency of sister taxa supports vicariance in a polytomic
sala mander clade in the absence of node support. – Molecular
Phylogenetics and Evolution, : –.
... mm and 19.0°C, respectively (Turkish Meteorological Stations, 2022). We gathered 254 occurrence records for the six species from the published literature (Başoğlu and Atatür, 1974;Mutz and Steinfartz, 1995;Veith, 2001;Öz et al., 2004;Johannesen et al., 2006;Beukema et al., 2009;Akman et al., 2011Akman et al., , 2013Göçmen and Akman, 2012;Göçmen et al., 2011Göçmen et al., , 2013Göçmen et al., , 2018Akman and Godmann, 2014;Oğuz et al., 2014;Şenol, 2015;Üzüm et al., 2015;Yıldız and Akman, 2015;Dereağzı, 2016;Godmann et al., 2016;Tok et al., 2016b;Göçmen and Karış, 2017;Arslan et al., 2018Arslan et al., , 2020Veith et al., 2016Veith et al., , 2020Bülbül and Özkan, 2021;Dilbe, 2021). We confirmed about 80% of these records during our previous fieldwork between 2004 and 2021. ...
Full-text available
Lycian salamanders, genus Lyciasalamandra, are a group of seven allopatric, endangered species restricted to the Mediterranean coast of Anatolia, Turkey and certain adjacent Aegean islands. In this study, we created maps to prediction the potential ranges for six of these species using a maximum-entropy algorithm to identify the most significant environmental factors that shape their distribution. These projections were created under current climate conditions and for three future periods (2011–2040, 2041–2070, and 2071–2100), using two shared socioeconomic pathways (SSPs), middle-of-the-road (SSP370) and pessimistic (SSP585). We found that temperature and precipitation are the most important factors influencing the distribution of these salamanders in southwestern Turkey. Under both SSP scenarios, we found that five Lyciasalamandra species are likely to shift or reduce their ranges in the near future, while one species (L. antalyana) was predicted to expand its range. For the five negatively affected species, whose ranges are already highly ranges, this predicts that climate change combined with high anthropogenic pressure poses a serious risk of extinction. Significant population declines are predicted, particularly for L. billae and L. luschani. We hope that our findings can serve as a resource for the long-term conservation of these species and aid decision-makers in the development of effective management strategies.
... Field studies were carried out at intervals in November-February between 2010-2020 and all records of the species were confirmed from previously published literature (Başoğlu & Atatür, 1974;Mutz & Steinfartz, 1995;Veith et al., 2001;Öz et al., 2004;Johannesen et al., 2006;Beukema et al., 2009;Akman et al., 2011;Göçmen & Akman 2012;Göçmen et al., 2013;Akman & Godmann, 2014;Üzüm et al., 2015;Göçmen & Karış, 2017;Arslan et al., 2018;Oğuz et al., 2020;Veith et al., 2020). A total of 249 presence records were gathered for the six species. ...
Full-text available
The Lycian salamanders consist of seven allopatric endangered and endemic species restricted to Mediterranean Turkey and some adjacent Aegean islands. Mega forest fires occurred consecutively over a prolonged period of time in the distribution areas of six of the species in 2021, leading to habitat fragmentation and habitat loss. In this study, we find that a total of 751.9 hectares of Lycian salamander habitats were lost due to the mega-fires that occurred in summer 2021. Our results suggest that L. atifi is the most vulnerable with a loss of 285.84 hectares of habitat area, followed by L. flavimembris with 242.54 hectares, and L. antalyana with 124.16 hectares. L. fazilae is the species which suffered the least habitat loss, at 25.83 hectares. L. billae and L. luschani suffered habitat losses of 30.83 and 43.40 hectares respectively. When the transformation of morphological classes was examined, a significant decrease of all species was observed in the core areas which ensure spatial connectedness, and the edge magnitude, which was taken as an indicator of fragmentation, increased. Bridges providing connectedness were observed to have increased for some species. This indicates that while existing connections in habitats were fragmented due to the forest fires, potential connections may be formed after the forest fires. When the fragmentation values were examined according to the results of pattern analysis, the most notable marginal increase in fragmentation before and after the fires was found to have occurred in the habitat of L. atifi. In addition, we discuss recommendations for the sustainability of species populations in habitat restoration and forest management.
... Field studies were carried out at intervals in November-February between 2010-2020 and all records of the species were confirmed from previously published literature (Başoğlu & Atatür, 1974;Mutz & Steinfartz, 1995;Veith et al., 2001;Öz et al., 2004;Johannesen et al., 2006;Beukema et al., 2009;Akman et al., 2011;Göçmen & Akman 2012;Göçmen et al., 2013;Akman & Godmann, 2014;Üzüm et al., 2015;Göçmen & Karış, 2017;Arslan et al., 2018;Oğuz et al., 2020;Veith et al., 2020). A total of 249 presence records were gathered for the six species. ...
... The systematic evaluations reporting new species or subspecies of the genus Lyciasalamandra were performed in previous studies (Veith et al., 2001;Göçmen et al., 2011;Göçmen & Akman, 2012;Göçmen et al., 2013;Akman & Godmann, 2014;Yıldız & Akman, 2015;Godmann et al., 2016). Populations of the same taxon, which occur only a few kilometers apart, often show distinct color and pattern differences (Veith & Steinfartz, 2004). ...
Full-text available
The Bille’s Lycian Salamander, Lyciasalamandra billae is an endemic salamander species of Turkey and it has a very narrow distribution area in the Antalya province of the country. A limited number of the reported populations of this critically endangered species are known. The present study aims to show that the distribution of the species extends towards the northeast of Antalya province. Two adult individuals (1 ♂ and 1 ♀) were caught from the Sarısu (Antalya, Turkey) population. We recorded a new locality of the species located about 11 km northeast of Gedeller village.
... (Nematocystis bayramiMallik & Bandyopadhyay, 2009), an endemic species of spiders living in Turkey (Typhlonesticus gocmeniRibera, Elverici, Kunt & Özkütük, 2014), a species of scorpions endemic to Turkey (Euscorpius gocmeni Tropea, Yağmur & Yeşilyurt, 2014), a terrestrial salamander race (Lyciasalamandra antalyana gocmeniAkman & Godmann 2014), another subspecies of a terrestrial salamander (Lyciasalamandra atifi bayrami, and two species of grasshoppers (Bradyporus gocmeniÜnal, 2017; Eupholidoptera gocmeni Ünal, 2019). ...
Full-text available
If there was something written to my destiny: "Biology / Zoology is not a profession for me, it is my lifestyle. My feelings, approaches, and reactions are coming out of this lifestyle. If you want to learn something, experience it! If you take nature as an example, there is no chance of making a mistake! That is the right way!"-Prof. Dr. Bayram GÖÇMEN. We (and many people around the world who know him) are in deep sorrow to have lost Prof. Dr. Bayram GÖÇMEN, who passed away on 22 March 2019. He will always be remembered with his passion to biology, his invaluable works, and the students he left behind. After the untimely loss of Prof. Dr. Bayram GÖÇMEN, who was one of the leading zoologists of Turkey and the Turkish Republic of Northern Cyprus, the first sentence said by many people from Turkish and world scientific community as well as many nature lovers was "Reptiles and amphibians are orphaned". This sentence summarizes his passion for his profession (biology/zoology) and science. With the large number of herpetological research he conducted in his last years, the taxa he described, and graduate students he supervised , Prof. Dr. Bayram GÖÇMEN was among the leading herpetologists in Turkey. He did not do merely scientific research. As an active social media user, he also founded an amphibian-reptile photo sharing web site, AdaMerOs Herptil Türkiye (The Amphibians and Reptiles Monitoring and Photography Society in Turkey) and gave opportunity to hundreds of people to know amphibians and reptiles more closely. In this sense, he made an exemplary effort to transfer scientific knowledge to the public and he led this internet community which is as valuable as the scientific articles. In addition to the herpetological studies which he concentrated on during the last years of his life, he was a world-renowned scientist in the field of protozoology-parasitology as well with his scientific research, many taxa he described, and the students he trained. He was a scientist who read immensely and had knowledge about not only in the fields of herpetology and protozoology but also in all other branches of biology and even in other fields of natural sciences such as geology and paleontology. His death was timeless for those who love him as he still had so much to give to the world. However, as we realize again while writing this article with sadness, he embraced his profession with passion throughout his life and left behind an important scientific heritage of many valuable works and trained scientists. Prof. Dr. Bayram GÖÇMEN was born on 23 December, 1965 in the village of Çanakkale (Kantou) in Limassol (Ley-mosun), Cyprus. He completed his primary and secondary Figure 1. at his office in Ege University, 09.04.2013, Bornova, İzmir, Turkey. education in Cyprus. In 1983, he started undergraduate education at Gazi University, Faculty of Engineering and Architecture, Department of Architecture and continued for two semesters. Then, he dropped out and returned to Cy-prus for 24 months of military service. In 1985, he came to Izmir and began to study Biology as an undergraduate student at Ege University, Faculty of Science, Department of Biology. He graduated in 1989 as the top student of the department and faculty with a high grade of 91 out of 100. He
... And if eventually small-scale dispersal may have occurred, such founder events again should have reduced local genetic variability. All this may have led to today's small-scale differences of pattern and colour among populations, forming the basis for the description of numerous new taxa within the last years (e.g., [9][10][11][12][13][14][15][16]). Unfortunately, up to know data supporting such a scenario solely stem from organelle data, so male-biased dispersal, which may erode local population differentiation, can only be detected by also analysing nuclear DNA. ...
Full-text available
Lycian salamanders (genus Lyciasalamandra) constitute an exceptional case of micro-endemism of an amphibian species on the Asian Minor mainland. These viviparous salamanders are confined to karstic limestone formations along the southern Anatolian coast and some islands. We here study the genetic differentiation within and among 118 populations of all seven Lyciasalamandra species across the entire genus’ distribution. Based on circa 900 base pairs of fragments of the mitochondrial 16SrDNA and ATPase genes, we analysed the spatial haplotype distribution as well as the genetic structure and demographic history of populations. We used 253 geo-referenced populations and CHELSA climate data to infer species distribution models which we projected on climatic conditions of the Last Glacial Maximum (LGM). Within all but one species, distinct phyloclades were identified, which only in parts matched current taxonomy. Most haplotypes (78%) were private to single populations. Sometimes population genetic parameters showed contradicting results, although in several cases they indicated recent population expansion of phyloclades. Climatic suitability of localities currently inhabited by salamanders was significantly lower during the LGM compared to recent climate. All data indicated a strong degree of isolation among Lyciasalamandra populations, even within phyloclades. Given the sometimes high degree of haplotype differentiation between adjacent populations, they must have survived periods of deteriorated climates during the Quaternary on the spot. However, the alternative explanation of male biased dispersal combined with a pronounced female philopatry can only be excluded if independent nuclear data confirm this result.
... (Nematocystis bayramiMallik & Bandyopadhyay, 2009), an endemic species of spiders living in Turkey (Typhlonesticus gocmeniRibera, Elverici, Kunt & Özkütük, 2014), a species of scorpions endemic to Turkey (Euscorpius gocmeni Tropea, Yağmur & Yeşilyurt, 2014), a terrestrial salamander race (Lyciasalamandra antalyana gocmeniAkman & Godmann 2014), another subspecies of a terrestrial salamander (Lyciasalamandra atifi bayrami, and two species of grasshoppers (Bradyporus gocmeniÜnal, 2017; Eupholidoptera gocmeni Ünal, 2019). ...
Full-text available
If there was something written to my destiny: "Biology / Zoology is not a profession for me, it is my lifestyle. My feelings, approaches, and reactions are coming out of this lifestyle. If you want to learn something, experience it! If you take nature as an example, there is no chance of making a mistake! That is the right way!" ‒ Prof. Dr. Bayram GÖÇMEN. We (and many people around the world who know him) are in deep sorrow to have lost Prof. Dr. Bayram GÖÇMEN, who passed away on 22 March 2019. He will always be remembered with his passion to biology, his invaluable works, and the students he left behind.
... During the last decade, several new populations of Lyciasalamandra have been found and some of them has been described as new subspecies (Göçmen, Arıkan, & Yalçınkaya, 2011;Göçmen & Akman, 2012;Göçmen et al., 2013;Akman & Godmann, 2014;Godmann, Karış, & Göçmen, 2016;Oğuz, Göçmen, & Yalçınkaya, 2016;Tok, Afsar, & Yakın, 2016;Göçmen & Karış, 2017). Up to now, only the Greek L. helverseni and the Turkish Fazila's Lycian Salamander, L. fazilae, remained monotypic, although for both substantial intraspecific genetic variation has been reported (e.g., Eleftherakos, Sotioropoulos, & Polymeni, 2007;Veith et al., 2008Veith et al., , 2016. ...
Full-text available
A new subspecies of the Fazila’s Lycian Salamander Lyciasalamandra fazilae is described based on material from ten localities in the Köyceğiz, Ortaca and Dalaman area in south-western Turkey. It is distinguished from the nominotypical subspecies by differences in the colouration pattern, morphometry and the mitochondrial molecular marker 16S rRNA. The distribution area of the new subspecies is located mainly in the western part of Dalaman River except for two seemingly intermediate populations (Şerefler and Sarsala-Kapıkargın). New localities for the species are reported.
... These were: [Lyciasalamandra luschani luschani (Steindachner, 1891; L. atifi atifi (Başoğlu 1967); L. fazilae (Başoğlu & Atatür 1974); L. luschani finikensis (Başoğlu & Atatür 1975); L. antalyana antalyana (Başoğlu & Baran 1976); L. luschani basoglui (Baran & Atatür 1980); L. billae billae (Franzen & Klewen 1987); L. flavimembris flavimembris (Mutz & Steinfartz 1995); L. irfani Göçmen et al. 2011;L. arikani Göçmen & Akman 2012; L. yehudahi ; L. antalyana gocmeni Akman & Godmann 2014; L. atifi bayrami Yıldız & Akman 2015; L. flavimembris ilgazi Üzüm et al. 2015; L. billae eikeae Godmann et al. 2016; L. atifi oezi Tok et al. 2016]. According to recently published paper by Veith et al. (2016), based on levels of molecular differentiation L. arikani, L. irfani and L yehudahi were considered as subspecies of L. billae. ...
Full-text available
We compared the isolated populations of Lyciasalamandra atifi, (Başoğlu, 1967) a salamander endemic to the historic Lycia region of Turkey, that is found across a range from Antalya/Selge (Altınkaya) to Antalya/Gazipaşa. Along this distance, we determined eight isolated populations (Selge, Fersin, Dikmen, Güzelbağ, Türbelinaz, Gündoğmuş, Cebireis, Gazipaşa) in 2013 and used morphology and serology to compare them. The collected specimens were registered under the ZMADYU (Zoology Museum of Adıyaman University), and a total of 237 (59 ♂♂, 96 ♀♀, 82 juv.) specimens were studied. As a result of our research, three new subspecies are described: Lyciasalamandra atifi godmanni n. ssp. from Selge, Lyciasalamandra atifi veithi n. ssp. from Dikmen and Lyciasalamandra atifi kunti n. ssp. from Güzelbağ. Except Cebireis (L. a. bayrami) and Gazipaşa (L. a. oezi) populations, the other isolated populations were incorporated in the nominat subspecies due to morphological and serological similarities.
... We analyzed specimens of all major Lyciasalamandra lineages as identified by Veith et al. (2008), as well as newly described species and subspecies (Göçmen et al., 2011(Göçmen et al., , 2012Akman and Godmann, 2014;Üzüm, 2015) (see table S1 in the online supplementary material). For hierarchical outgroup rooting, as well as for molecular clock calibration, we added homologous gene fragments from complete mitochondrial genomes of further Salamandridae species from Zhang et al. (2008): Salamandra salamandra (EU880331), Chioglossa lusitanica (EU880308), Mertensiella caucasica (EU880319), Pleurodeles poireti (EU880329), Pleurodeles waltl (EU880330), Euproctus platycephalus (EU880317) and Euproctus montanus (EU880316). ...
Full-text available
The number of tectonic and climatic events that are used to explain speciation processes in the eastern Mediterranean region is low compared to the western Mediterranean. Among them, the emergence of the mid-Aegean trench and the Messinian Salinity Crisis (MSC) often concurred with speciation time estimates that were inferred from molecular data. We here present a dated molecular phylogeny of Lyciasalamandra from Turkey and Greece based on ca. 4500 bp of the mitochondrial genome (3000 bp of three nuclear genes appeared to be completely inconclusive due to their extremely low degree of variation among taxa). Seven major lineages emerged simultaneously from a basal hard polytomy. A scenario that dates this polytomy to 12.3 and 10.2 million years ago, around the final emergence of the mid-Aegean trench, appears to be most plausible. The MSC can be made responsible for first intraspecific divergence events within L. luschani, L. fazilae and L. flavimembris. Further diversification can be explained by Pliocene and Pleistocene glaciations. Based on levels of molecular differentiation we suggest the recently described species L. arikani, L. irfani and L. yehudahi to be treated as subspecies of L. billae.
Full-text available
Nine subspecies of the viviparous Lycian Salamander Mertensiella luschani live along the coast of southwestern and southern Turkey and on some islands (e.g. Kastellorizon, Meis, Kekova, and Carpathos). The species is locally very common. We describe seven new localities and add precise co-ordinates for most known populations. Existing confusion in the literature regarding locality names is discussed. Two type localities are re-defined. The actual and potential distribution of the species is analysed from ecological data. The range of M. luschani is restricted to karstic limestone with precipitation exceeding 800 mm annual rainfall. Most localities are below 500 m a.s.l., with a maximum of up to 1340 m a.s.l. The typical habitat of the species is pine forest on northerly exposed slopes.
Full-text available
During fieldwork conducted between end of February and mid-April 2012 14 new localities were ascertained for four different taxa of the amphibian genus Lyciasalamandra (L. l. basoglui, L. l. finikensis, L. arikani and L. atifi). This paper represents the first record of L. l. basoglui from Muğla province (Saklıkent, Fethiye). Unlike the common belief in previous researches based on literature we determined that there are almost no gaps between the distributional areas of the known subspecies of L. luschani, moreover the subspecies were found in contact. The recently discovered populations of L. arikani and L. atifi were found to have some distinctive morphological differences compared to other populations of the respective species.
Full-text available
Two new species of Lycian salamanders, Lyciasalamandra arikani n. sp. and L. yehudahi n.sp. are described and their relationships with similar and neighbouring taxa are discussed. Both taxa originate from areas new for the genus, with the former from around Erentepe Mt. (Kumluca, Antalya) and the latter from Tahtalı Mt. (Kemer, Antalya) in the southern parts of Beydağları Mountain range in southwestern Anatolia. Some information about their habitats and behaviour are reported.
Full-text available
A new species of the Lycian Salamander, Lyciasalamandra irfani n. sp. is described and its relation with similar previously known taxa is discussed. The new species characterized by having rather darkly col-oured head part and also an aubergine reddish brown ground colour on dorsum with irregularly scattered white flecks. It originates from Göynük Canyon (Antalya) in southwestern Anatolia. At present, the distribu-tion is limited to its type locality. Some information is added in regard to its habitat.
Full-text available
A comprehensive, but simple-to-use software package for executing a range of standard numerical analysis and operations used in quantitative paleontology has been developed. The program, called PAST (PAleontological STatistics), runs on standard Windows computers and is available free of charge. PAST integrates spreadsheettype data entry with univariate and multivariate statistics, curve fitting, time-series analysis, data plotting, and simple phylogenetic analysis. Many of the functions are specific to paleontology and ecology, and these functions are not found in standard, more extensive, statistical packages. PAST also includes fourteen case studies (data files and exercises) illustrating use of the program for paleontological problems, making it a complete educational package for courses in quantitative methods.
Full-text available
We announce the release of an advanced version of the Molecular Evolutionary Genetics Analysis (MEGA) software, which currently contains facilities for building sequence alignments, inferring phylogenetic histories, and conducting molecular evolutionary analysis. In version 6.0, MEGA now enables the inference of timetrees, as it implements our RelTime method for estimating divergence times for all branching points in a phylogeny. A new Timetree Wizard in MEGA6 facilitates this timetree inference by providing a graphical user interface (GUI) to specify the phylogeny and calibration constraints step-by-step. This version also contains enhanced algorithms to search for the optimal trees under evolutionary criteria and implements a more advanced memory management that can double the size of sequence data sets to which MEGA can be applied. Both GUI and command-line versions of MEGA6 can be downloaded from free of charge.
Full-text available
We report 17 specimens of the Lycian salamander Lyciasalamandra antalyana, endemic to Turkey from Burdur province, in the western Mediterranean region of Anatolia. Previously all known localities for L. antalyana were in Antalya province. These new records extend the known distribution area of the species to Burdur province in Turkey. These new specimens were compared with L. antalyana specimens from some other known localities (Hurma and Yağca populations) in terms of morphological and serological features. According to coloration and metric characters, our specimens can be included in L. antalyana. This paper represents a considerable range extension for this species, 32 km air distance to the north-east and reports L. antalyana from Burdur province for the first time.
Within the framework of the present study we test whether climatic niche similarity can be identified in a monophyletic group of species inhabiting remarkably restricted ranges by pooling presence data of all species into a single concatenated data set and subsequently jackknifing single species. We expect that, when the jackknifed species differs markedly in its climatic niche from all other species, this approach will result in increased niche homogeneity, allowing assessments of niche diver-gence patterns. To test our novel jackknife approach, we developed species distribution models for all members of Lycian salamanders (genus Lyciasalamandra), native to Turkey and the adjacent Aegean islands using Maxent. Degrees of niche similarity among species were assessed using Schoener's index. Significance of results was tested using null-models. The degree of niche similarity was generally high among all seven species, with only L. helverseni differing significantly from the others. Carstic lime stones providing specific microhabitat features may explain the high degree of niche similarity detected, since the variables with the highest explanative power in our models (i.e. mean temperature, and precipitation of the coldest quarter) corresponded well with salamander natural history observations, supporting the biologically plausibility of the results. We conclude that the jackknife approach presented here for the first time allows testing for niche similarity in species inhabiting restricted ranges and with few species records available. Our results strongly support the view that detailed natural history information and knowledge of microhabitats is crucial when assessing possible climate change impacts on species.
Muscle weakness is commonly cited as a cause of crouch gait in individuals with cerebral palsy; however, outcomes after strength training are variable and mechanisms by which muscle weakness may contribute to crouch gait are unclear. Understanding how much muscle strength is required to walk in a crouch gait compared to an unimpaired gait may provide insight into how muscle weakness contributes to crouch gait and assist in the design of strength training programs. The goal of this study was to examine how much muscle groups could be weakened before crouch gait becomes impossible. To investigate this question, we first created muscle-driven simulations of gait for three typically developing children and six children with cerebral palsy who walked with varying degrees of crouch severity. We then simulated muscle weakness by systematically reducing the maximum isometric force of each muscle group until the simulation could no longer reproduce each subject's gait. This analysis indicated that moderate crouch gait required significantly more knee extensor strength than unimpaired gait. In contrast, moderate crouch gait required significantly less hip abductor strength than unimpaired gait, and mild crouch gait required significantly less ankle plantarflexor strength than unimpaired gait. The reduced strength required from the hip abductors and ankle plantarflexors during crouch gait suggests that weakness of these muscle groups may contribute to crouch gait and that these muscle groups are potential targets for strength training.