A preliminary comparison of cranial and external morphological variability of Southern Crested Newt (Triturus karelinii) from the Crimea and the Caucasus
The Triturus karelinii group of crested newts comprises the eastern, central and western lineages that differ by mitochondrial DNA. The eastern lineage largely retains the ancestral cranium shape and is distributed throughout the Crimean Peninsula, the Caucasus Isthmus, and the Caspian Sea southern...
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irk-123456789-1612402019-12-04T01:26:05Z A preliminary comparison of cranial and external morphological variability of Southern Crested Newt (Triturus karelinii) from the Crimea and the Caucasus Pysanets, Y. Kukushkin, O. Manuilova, O. The Triturus karelinii group of crested newts comprises the eastern, central and western lineages that differ by mitochondrial DNA. The eastern lineage largely retains the ancestral cranium shape and is distributed throughout the Crimean Peninsula, the Caucasus Isthmus, and the Caspian Sea southern shore in Iran. In this report the comparison of the Crimean and Trascaucasian T. karelinii cranium shape, differences in the dimensional characteristics variability and body colouring patterns is presented. The samples of T. dobrogicus from the Danube River Delta have been employed for outgroup comparison. Data of the differences between several populations within the eastern lineage suggest that it is not uniform and T. karelinii from the Crimean Peninsula is considered to be somewhat different. Група Triturus karelinii з числа гребінчастих тритонів включає східну, центральну і західну гілку, які відрізняються за мітохондріальною ДНК. Східна гілка значною мірою зберігає анцестральну форму черепної коробки і поширюється всім Кримським півостровом, Кавказьким перешийком і південним узбережжям Каспійського моря в Ірані. Проведено порівняння кримської і закавказької черепної форми Т. karelinii, вказано на відмінності мінливості розмірних характеристик і забарвлення тіла. Екземпляри Т. dobrogicus з дельти Дунаю були використані для порівняння. Дані про відмінності між декількома популяціями в межах східної гілки припускають, що вони не є однорідним, і Т. karelinii з Кримського півострова дещо відрізняється. Группа Triturus karelinii гребенчатых тритонов включает восточную, центральную и западную ветвь, которые отличаются митохондриальной ДНК. Восточная ветвь в значительной мере сохраняет анцестральную форму черепной коробки и распространена по всему Крымскому полуострову, Кавказскому перешейку и южному побережью Каспийского моря в Иране. Проведено сравнение крымской и закавказской черепной формы Т. karelinii, указано на отличия изменчивости размерных характеристик и окраски тела. Экземпляры Т. dobrogicus из дельты Дуная были использованы для сравнения. Данные об отличиях между несколькими популяциями в пределах восточной ветви допускают, что они неоднородны, и Т. karelinii с Крымского полуострова несколько отличаются. 2014 Article A preliminary comparison of cranial and external morphological variability of Southern Crested Newt (Triturus karelinii) from the Crimea and the Caucasus / Y. Pysanets, O. Kukushkin, O. Manuilova // Збірник праць Зоологічного музею. — 2014. — Вип. 45. — С. 46-56. — Бібліогр.: 23 назв. — англ. 0132-1102 http://dspace.nbuv.gov.ua/handle/123456789/161240 597.94(477.75:479.2) en Збірник праць Зоологічного музею Національний Науково-природничий музей НАН України |
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The Triturus karelinii group of crested newts comprises the eastern, central and western lineages that differ by mitochondrial DNA. The eastern lineage largely retains the ancestral cranium shape and is distributed throughout the Crimean Peninsula, the Caucasus Isthmus, and the Caspian Sea southern shore in Iran. In this report the comparison of the Crimean and Trascaucasian T. karelinii cranium shape, differences in the dimensional characteristics variability and body colouring patterns is presented. The samples of T. dobrogicus from the Danube River Delta have been employed for outgroup comparison. Data of the differences between several populations within the eastern lineage suggest that it is not uniform and T. karelinii from the Crimean Peninsula is considered to be somewhat different. |
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| author |
Pysanets, Y. Kukushkin, O. Manuilova, O. |
| spellingShingle |
Pysanets, Y. Kukushkin, O. Manuilova, O. A preliminary comparison of cranial and external morphological variability of Southern Crested Newt (Triturus karelinii) from the Crimea and the Caucasus Збірник праць Зоологічного музею |
| author_facet |
Pysanets, Y. Kukushkin, O. Manuilova, O. |
| author_sort |
Pysanets, Y. |
| title |
A preliminary comparison of cranial and external morphological variability of Southern Crested Newt (Triturus karelinii) from the Crimea and the Caucasus |
| title_short |
A preliminary comparison of cranial and external morphological variability of Southern Crested Newt (Triturus karelinii) from the Crimea and the Caucasus |
| title_full |
A preliminary comparison of cranial and external morphological variability of Southern Crested Newt (Triturus karelinii) from the Crimea and the Caucasus |
| title_fullStr |
A preliminary comparison of cranial and external morphological variability of Southern Crested Newt (Triturus karelinii) from the Crimea and the Caucasus |
| title_full_unstemmed |
A preliminary comparison of cranial and external morphological variability of Southern Crested Newt (Triturus karelinii) from the Crimea and the Caucasus |
| title_sort |
preliminary comparison of cranial and external morphological variability of southern crested newt (triturus karelinii) from the crimea and the caucasus |
| publisher |
Національний Науково-природничий музей НАН України |
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2014 |
| url |
http://dspace.nbuv.gov.ua/handle/123456789/161240 |
| citation_txt |
A preliminary comparison of cranial and external morphological variability of Southern Crested Newt (Triturus karelinii) from the Crimea and the Caucasus / Y. Pysanets, O. Kukushkin, O. Manuilova // Збірник праць Зоологічного музею. — 2014. — Вип. 45. — С. 46-56. — Бібліогр.: 23 назв. — англ. |
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46 Збірник праць Зоологічного музею, 2014, № 45
Y. Pysanets, O. Kukushkin, O. Manuilova
Introduction
Crimean newt has been noticed by P.S. Pallas as a “water lizard” La-
certa aquatica: “Rossis a voice Ukola. In temperatioribus et australiori-
bus rarius in stagnis observatur; frequentior in Chersoneso Tauri-
ca...” (Pallas, 1811, p. 34). In turn, K. Kessler in travel notes of his Cri-
mean visit in September 1878 reported about the newt as “Triton cris-
tata” (Kessler, 1878, p. 212). In the monograph about vertebrates of the
Crimean peninsula (Никольский, 1895) newt species Molge cristata
has been referenced to as a common amphibian species of the peninsula.
Moreover, G. Boulenger also supported the validity of the “variation”
karelinii within the genus Molge, i.e. Molge cristata var. karelinii (Bou-
lenger, 1882). Five years later, B. Dürigen mentioned this “variation”
within the genus Triton as Triton cristatus var. karelinii (Dürigen,
1897). However, A. Brauner (Браунер) indicated newt from the Crimea
as “Molge cristata... var. Karelinii” (Strauch, 1870) (Браунер, 1904).
Wolterstorff described a new form taurica within the subspecies ka-
The Triturus karelinii group of crested newts comprises the eastern, central and western lineages that differ
by mitochondrial DNA. The eastern lineage largely retains the ancestral cranium shape and is distributed
throughout the Crimean Peninsula, the Caucasus Isthmus, and the Caspian Sea southern shore in Iran. In
this report the comparison of the Crimean and Trascaucasian T. karelinii cranium shape, differences in the
dimensional characteristics variability and body colouring patterns is presented. The samples of T. do-
brogicus from the Danube River Delta have been employed for outgroup comparison. Data of the differ-
ences between several populations within the eastern lineage suggest that it is not uniform and T. karelinii
from the Crimean Peninsula is considered to be somewhat different.
K e y w o r d s : Southern crested newt, Triturus karelinii, the Crimea, cranium, external morphology,
taxonomic status.
УДК 597.94(477.75:479.2)
Y. Pysanets1, O. Kukushkin2, O. Manuilova1
1 National Museum of Natural History NAS Ukraine,
B. Khmelnitsky Str., 15, Kyiv, 01601 Ukraine
2 Zoological Institute, Russian Academy of Sciences,
Universitetskaya nab. 1, St. Petersburg, 199034 Russian Federation
A PRELIMINARY COMPARISON OF CRANIAL
AND EXTERNAL MORPHOLOGICAL
VARIABILITY OF SOUTHERN CRESTED NEWT
(TRITURUS KARELINII) FROM THE CRIMEA
AND THE CAUCASUS
Збірник праць Зоологічного музею, 2014, № 45: 46–56.
© Y. PYSANETS, O. KUKUSHKIN, O. MANUILOVA, 2014
Збірник праць Зоологічного музею, 2014, № 45 47
A preliminary comparison of cranial and external morphological variability of Southern Crested Newt...
relinii ─ Triton cristatus karelinii forma taurica (Wolterstorff, 1923). Nevertheless, its
validity has been never confirmed, since the International Code of Zoological Nomen-
clature does not recognize infra-subspecies taxa (art. 1. 3. 4, 45.5 & 45.6). Furthermore,
Mertens and Müller (1928) obviously included the subspecies T. cristatus karelinii in
the European amphibians and reptiles checklist. Finally, N. Shcherbak (1966) used this
name for newt found at the Crimean Peninsula and summarized all available data re-
garding its biology, distribution and variation. Based on the comparative analysis of ge-
nomic characters as well as on the results of cytogenetic studies of hybrids, Bucci-
Innocenti et al. (1983) proposed to consider its species level as specific (Triturus ka-
relinii). Later it was supposed (Litvinchuk et al., 1999), the species divide on two sub-
species, where T. karelinii karelinii range includes the Crimea, the Caucasus, Northern
Iran and partially Anatolia, while T. karelinii arntzeni Litvinchuk, Borkin, Džukić and
Kalezić, 1999 is attributed to the Balkan Peninsula. The taxonomic status of the last
taxon later has been changed to the species level. However, the name T. arntzeni Lit-
vinchuk, Borkin, Džukić, Kalezić, 1999 appiared a junior synonym of T. macedonicus
(Karaman, 1922) that inhabits the western and north-western parts of the Balkan Penin-
sula (Wielstra et al., 2014).
The recent study of T. karelinii sensu lato based on nuclear and mitochondrial
DNA revealed the existence of three deeply diverged lineages: earstern, central and
western (Wielstra et al., 2010; Wielstra and Arntzen, 2011). These lineages comprised at
least two species: T. karelinii from Northern Iran, the Caucasus, and the Crimean Penin-
sula, and T. ivanbureschi Arntzen & Wielstra, 2013 from the south-eastern Balkans and
western and northern parts of Anatolia (Aegean Sea, Marmara Sea, and apparently the
Black Sea regions) (Wielstra et al., 2013 a, 2014).
At the same time, some authors indicated that the skull shape does not clearly
support the presence of three discrete geographical groups as suggested by mitochon-
drial DNA data (Ivanović et al., 2013, p. 269). Besides that, they have registrated no
significant sexual dimorphism in cranium size and shape within eastern lineage (T. kare-
linii sensu stricto), while the newts of central and western lineages (T. ivanbureschi)
shows the statistically significant sexual dimorphism in cranium size/shape (Ivanović et
al., 2013). The external morphology study of southern crested newt in the Crimean and
Asia Minor Peninsulas, the Caucasus and the Balkans “revealed the relative uniformity
of populations of the species by dimensional characteristics” (Litvinchuk and Borkin,
2009, p. 81). In addition, the stability of the spotted colour throat patterns and their geo-
graphical variation were also found in this newt group (Wielstra et al., 2013 a). Hence,
previous authors compared mitochondrial DNA data of the newts from three discrete
geographical groups representing two different species, T. ivanbureschi and T. karelinii.
Some differences between these species in mitochondrial DNA variability, sexual di-
morphism іn cranium (skull) characteristics and features of body coloration have been
also indicated.
Therefore, the aim of our study was to compare the variability of cranium archi-
tecture (skull structure), dimensional characreristics of the external morphology and
some features of the body coloration within the representatives of eastern lineage from
the Crimea and the Caucasus.
Materials and Methods
Species localities, samples sizes, number of males and females are presented in
figure 1, a, and table 1. As outgroup we employed T. dobrogicus (Kiritzescu, 1903)
sample (cranium and external morphology). Skulls of adult newts were cleared with
Na2CO3 (solution ─ 10−12%, 12−14 hours) and put into hydrogen peroxide (3%) up to
20−30 min. Images of dorsal, ventral, lateral and posterior sides were obtained with
digital camera with the cranium roof positioned parallel to the photographic plane. The
48 Збірник праць Зоологічного музею, 2014, № 45
Y. Pysanets, O. Kukushkin, O. Manuilova
Table 1. Summary of Triturus karelinii and T. dobrogicus (referred to as outgroup) characteristics:
locality; decimal degrees (latitude (N) specifies the north-south position, longitude (E) ─ east-west
position); No.F ─ females number; No.M ─ males number; skulls; morphometry; throat color
pattern.
N
Samples (from museum funds
lables)
L
atitude (N
)
L
ongitude (E
)
No.F. No.M.
skulls
m
orphom
etry
throat color
pattern
skulls
m
orphom
etry
throat color
pattern
1
T. dobrogicus, Ukraine, Odesa
Province, Vilkovo town, 1974
45°23′57″ 29°35′37″ 8 10 10 4 18 18
2
T. karelinii, Crimea, Sevastopol
City Council, Bakhchisaraiskiy
District, Ternovka village,
Southern Mangup, June 8−9, 2003
44°34′55″ 33°48′30″ 3 11 11 3 10 10
3
T. karelinii, Crimea, Krymskyi
Nature Reserve, Mt. Chuchel, 1959
44°39'12" 34°14'8" 6 6 8 8
4
T. karelinii, Crimea, Alushta City
Council, 2 km S-SW from Angar
Pass, Lake Kutuzovskoe near the
foot of SE slope of Mt. Chatyrdag,
870 m a. s. l., 2015
44°44'21" 34°20'9" 13 15
5
T.karelinii, Crimea, near Alushta,
Luchistoe village, April 30 2004,
2005, 2005
44°44′40″ 34°22′15″ 15 15 7 7
6
T. karelinii, Georgia, Borzhomskiy
District, Lake Kakhisy, Chobis-
Khevi village, July 6−9, 1979
41°51′15″ 43° 24′ 45″ 1 23 23 1 16 16
7
T. karelinii, Azerbaijan, Lerikskiy
District, Mts. Talysh, Almu
settlement, 1974
38° 42′00″ 48°18′00″ 1 16 16 1 18 18
8
T. karelinii, Russia, Krasnodarskyi
krai, Adigeya, Afinsip settlement,
07.04.1975
44°54′00″ 38°50′00″ 1 2
9
T. karelinii, Russia, Krasnodarskyi
krai, Novorossiysk City, Abrau-
Dyurso, 27.05.1961
44°42′10″ 37°36′06″ 3 1
10
T. karelinii, Russia, Krasnodarskyi
krai, Goryachiy Klyuch,
10.04.1975
44°38′ 39°08′ 2
In total 19 81 94 12 77 92
Збірник праць Зоологічного музею, 2014, № 45 49
A preliminary comparison of cranial and external morphological variability of Southern Crested Newt...
positions and names of cranium bones were identified following special protocols
(Dunaev, 1996; Ivanović et al., 2008; 2013). The external 18 morphometric
characteristics were measured with dial calliper (0,1 mm scale) (Писанець, 2007;
Litvinchuk and Borkin, 2009; Naumov and Tzankov, 2009; Çiçek et al., 2010; Писа-
нець, 2012 with some modifications): SVL (snout-vent. post length) ─ measured from
the snout to the posterior edge of the base of the cloaca; L (snout-vent. ant length) ─
measured from the snout to the anterior edge of the base of the cloaca; Lcd (tail length )
─ measured from the anterior edge of the base of the cloaca to the tail tip; TL ─ total
length (L.+L.cd.); Lcd1 (tail length1) ─ measured from the posterior edge of the base of
the cloaca to the tail tip; LiE (distance between fore and hindlimbs) ─ measured from
the anterior base of posterior forelimb to the posterior base of hindlimb; L.pc. (distance
between snout and hind limbs) ─ measured from snout tip to the anterior base of hind
limbs; L. m. (distance between snout and jaw articulation) ─ measured from snout tip to
the jaw articulation; Lt.c. (head width) ─ measured between the posterior of the mouth
jaw articulations; Alt.c. (head height) ─ measured between the upper and below head
sufaces after the eyes; P.a. (length of the forelimb) ─ measured between the base to the
tip of forelimbs; P.p (length of the hindlimb) ─ measured between the base to the tip of
hindlimbs; C.l. (length of the cloaca) ─ measured from anterior and posterior edges of
cloaca; D.t. (lenght of the the third toe of forelimb) ─ measuring from the base of toe to
its tip; Sp.in. ─ distance between the nostrils; D.n.o.(distance between the eye and the
nostril) ─ measuring between the anterior corner of the eyes aperture and the nostril;
L.o. ─ horizontal length of the eyes aperture; Sp.ino.1 ─ distance between the anterior
margins of the eyes; Sp.ino.2 ─ distance between the posterior margins of the eyes.
Means of replicates were subjected to statistical analysis using Statistica 6.0
software package. Pattern of geographical variability was analyzed by principial compo-
nent analysis (PCA) on standardized morphometric characters (see above).
Results and Discussion
Cranial characteristics and their variation. The general view of T. karelinii female
skull structure are presented on figure 1, b. External views of female and male skulls of
T. karelinii from the Crimea (Sevastopol City Council, Ternovka village vicinity) are
shown on figure 1, c. Even though male and female skulls look much alike (fig. 1, c), vi-
sual comparison of cranium dorsal sides indicates that males have: (1) a much wider an-
terior fontanel; (2) a shorter distance between the rear end of the premaxilla and the
front end of the pterygoideum. At the same time, others characteristics for the compa-
rison between males and females skulls (e.g., ventral and lateral sides) have not revealed
any significant differences.
Generally, the comparison of T. karelinii skulls from the Crimea and the Cauca-
sus shown similarities in the shape and positioning of the majority of elements, how-
ever, with some differences. Pronounced sexual dimorphism in the linear dimensions of
Transcaucasian newts (especially in the sample from South-Eastern Azerbaijan,
Mts.Talysh) characterized by larger skulls has been revealed (fig. 2, a). The structure of
the Crimean and Caucasian T. karelinii skulls differs clearly from T. dobrogicus ones
taken for outgroup comparison (fig. 2, b), first of all, in smaller size of the latter and the
lack of postfrontal processes that has been indicated earlier in other studies (Litvinchuk
and Borkin, 2009). Some unexpected differences in several characteristics of skulls have
been revealed in southern crested newt from the Crimea compared with the Transcauca-
sian T. karelinii. For instance, the quadratum of the Crimean T. karelinii were smaller
than those of the Caucasian species, what affects relative cranium height that was much
lower at the Crimean T. karelinii in most cases (fig. 2, c).
At the same time, any clear diagnostic characteristics of the cranium shape able to
distinguish evidently T. karelinii from the Crimean Peninsula and continental newts
50 Збірник праць Зоологічного музею, 2014, № 45
Y. Pysanets, O. Kukushkin, O. Manuilova
Fig. 1. a ─ T. karelinii localities in terms of skull differentiation, morphometrics measurements,
coloration patterns. Solid circles (1, 2, 3, ,5, 6, 7), colorless circles (4) and squares (8, 9, 10) indicate
analyzed samples number (see Table 1). b ─ T. karelinii female skull, right ─ dorsal view, left ─ ventral
view. Location: Crimea, Sevastopol City Council, Bakhchisaraiskiy District, environs of Ternovka village.
Names of skull elements and abbreviations (by Dunaev, 1996; Ivanović et al., 2008; Ivanović et al., 2013
with some modifications): PA ─ parietal, OC ─ occipital, SQ ─ squamosum, MX ─ maxilla, F ─ frontal,
FP ─ frontalis processus, N ─ nasal, PRF ─ prefrontal, PMX ─ premaxilla; Q ─ quadratum, OCC ─
occipital condyle, PT ─ pterygoideum, OF ─ oval fenestra, V (VPAL) ─ vomer (vomero-palatine), FI ─
fenestra intermaxillaris, PS ─ parasphenoid, CHO ─ choane. c ─ Dorsal (up) and ventral (down) view of
T. karelinii female and male skulls (left and right, respectively). Location the same as of b. Arrows
indicate the differences in anterior fontanelle size and form (i), and distance between pterygoideum and
maxilla (ii).
from the Caucasus have not been found at the current level of studies, and the differen-
ces described above have been proven only statistically. Another osteological examina-
tion revealed species-specific cranium shape of T. karelinii from the Caucasus (Dunaev,
1996). It was emphasised one year later that Karelin’s newt differs from other three taxa
of T.cristatus complex in cranium shape by 6 characteristics: “Cavum internasale or in-
termaxillare (dorsal view)... Processus postfrontalis (dorsal and ventral views)... Proces-
sus occipetalis (dorsal and ventral views)... Foramen magnum (caudal view)... Posterior
margin of parietale (dorsal view)... Basal part of pterigoid...” (Litvinchuk, Borkin, 2009,
p. 532). It has been also noticed that all species of the complex T. cristatus are well dif-
ferentiated from each other on the cranium structure (Litvinchuk, Borkin, 2009).
Differences in male and female craniums shape of southern crested newt have
been also mentioned in another special study of Ivanović et al. (2013). Osteological
characteristics analysis of 197 specimens of T. karelinii from different parts of species
range (including 16 individuals from the Crimea) showed that among three phylogenetic
branches of this taxon (central, eastern and western, fig. 1, a) the statistically significant
differences in the size and configuration of the cranium between males and females
were inherent only for the representatives of the first two groups. Ivanović et al. (2013)
also emphasized that the variability of cranium size was statistically significant only for
cranium dorsal side in males.
Variation of the external morphology. It has been established that Wolterstorff
index (P.a./LiE) and relative head width (Lt.c./L.) had the greatest value among the
T. cristatus complex (Litvinchuk, Borkin, 2009). Comparative characteristics of T. ka-
relinii (the Crimea and the Caucasus) and in T. dobrogicus (Ukraine, Odesa Province)
are shown in tables 2, 3 and 4.
The data mentioned above indicate that both indexes clearly distinguish T. dobro-
gicus from T. karelinii from the Crimea and the Caucasus; in turn, T. karelinii from
these two regions show reliable difference on the basis of Lt.c./L ratio. Our results cor-
roborate to some extent with the geographical variation data for T. karelinii that indicate
variation in the range of 0.68–0.88 (males) and 0.59–0.79 (females) for P.a./LiE, and
0.18–0.22 (males) and 0.17–0.20 (females) for Lt.c./L. (Litvinchuk and Borkin, 2009).
These data are presented on page 83 (Litvinchuk and Borkin, 2009); on page 484 other
indeces are indicated: 0.56–0.97 (0.73±0.07) (males) and 0.50–0.79 (0.62±0.06) (fema-
les) for Pa/LiE; 0.15–0.22 (0.19±0.01) (males) and 0.15–0.23 (0.18±0.01) (females) for
Ltc / L.
The Crimean T. karelinii and representatives of this group from six different regi-
ons including the Caucasus, the Balkans, and Asia Minor were analysed by multivariate
statistical approach using “indices of high taxonomic interest” (Litvinchuk and Borkin,
2009, p. 79) and the logarithmic conversion of characteristics, however, unfortunately,
without the outgroup comparison. Comparative uniformity of populations by morhpo-
metric characteristics has been emphasised (Litvinchuk, Borkin, 2009).
Збірник праць Зоологічного музею, 2014, № 45 51
A preliminary comparison of cranial and external morphological variability of Southern Crested Newt...
a
b
c
52 Збірник праць Зоологічного музею, 2014, № 45
Y. Pysanets, O. Kukushkin, O. Manuilova
Fig. 2. a ─ Female and male T. karelinii skulls (left and right, respectively) dorsal (right) and ventral (left)
view. Location: Azerbaijan, Lerikskiy District , Mts.Talysh, Almu settlement. b ─ Female and male T. ka-
relinii skulls (left and right, respectively) dorsal (right) and ventral (left) view. Location: Ukraine, Odesa
Province , Vilkovo town. c ─ T. karelinii, male skulls, posterior side. Left three skulls ─ Sevastopol City
Council, Balaklava Distrcit, environs of Ternovka village; fourth (right) ─ Azerbaijan, Lerikskiy District,
Mts.Talysh, Almu settlement. Arrows indicate quadratum bones. d ─ Throat coloration patterns of alive
T. karelinii from different populations in the Crimea: A, B ─ Lake Kutuzovskoe, Alushta City Council;
C, D ─ near Luchistoe village, Alushta City Council.
a
b
d
c
Збірник праць Зоологічного музею, 2014, № 45 53
A preliminary comparison of cranial and external morphological variability of Southern Crested Newt...
Table 2. Comparison of Wolterstorff index (P.a./LiE) and relative head width (Lt.c./L.) in
representatives of different taxa and populations in the newts from genera Triturus.
Table 3. Coefficients of differences reliability (t) for Wolterstorff index (P.a./LiE) and relative head
width (Lt.c./L.) in males of different taxa and populations of the newts from genera Triturus
(significant difference p < 0.05 marked bold).
Table 4. Coefficients of differences reliability (t) for Wolterstorff index (P.a./LiE) and relative head
width (Lt.c./L.) in females of different taxa and populations of the newts from genera Triturus
(significant difference p < 0,05 marked bold).
Species, samples, number
T. karelinii,
The Crimea,
n= 25
T. karelinii,
Transcaucasia: Georgia and
Azerbaijan (Talysh Mts.)
together, n = 34
T. dobrogicus,
Odesa Region,
n= 18
Pa/LiE
T. karelinii, The Crimea, n = 25 0 0.4 8.8
T. karelinii, Transcaucasia:
Georgia and Azerbaijan
(Talysh Mts.) together, n = 34
0.4 0 7.9
T. dobrogicus, Odesa Province,
n = 18 8.8 7.9 0
Ltc/L
T. karelinii, The Crimea, n = 25 0 4.1 10.3
T. karelinii, Transcaucasia :
Georgia and Azerbaijan
(Talysh Mts.) together, n = 34
4.1 0 18.4
T. dobrogicus, Odesa Province,
n = 18 10.3 18.4 0
T. karelinii
The Crimea
T. karelinii
Transcaucasia: Georgia and
Azebaijan (Talysh Mts.)
T. dobrogicus
Ukraine, Odesa Province
Index males n=25 females n=21 males n=34 females n=39 males n=18 females n=10
Wolterstorff
index
P.a./LiE
0.70 ±0.011
0.60−0.82
0.58 ±0.008
0.53−0.67
0.70 ±0.011
0.59−0.93
0.62± 0.008
0.52−0.75
0.57±0.08
0.51−0.64
0.40 ±0.027
0.21−0.49
Relative head
width Lt.c./L.
0. 21 ±0.003
0.16−0.22
0.18 ±0,002
0.16−0.20
0.21 ±0.002
0.19−0.24
0.21 ±0.002
0.19−0.23
0.16 ±0.002
0.14−0.17
0.14 ±0.004
0.12−0,17
Species, samples, number
T. karelinii,
The Crimea,
n = 21
T. karelinii,
Transcaucasia: Georgia and
Azerbaijan (Talysh Mts.)
together, n = 39
T. dobrogicus,
Odesa Region,
n = 10
Pa/LiE
T. karelinii, The Crimea, n = 21 0 0,3 8.2
T. karelinii, Transcaucasia:
Georgia and Azerbaijan (Mt.
Talysh) together, n = 39
0.3 0 10.9
T. dobrogicus, Odesa Region,
n = 10 8.2 10.9 0
Ltc/L
T. karelinii, The Crimea, n = 21 0 6.0 8.1
T. karelinii, Transcaucasia:
Georgia and Azerbaijan (Mt.
Talysh) together, n = 39
6.0 0 14.4
T. dobrogicus, Odesa Region,
n = 10 8.1 14.4 0
54 Збірник праць Зоологічного музею, 2014, № 45
Y. Pysanets, O. Kukushkin, O. Manuilova
Moreover, we have compared the external morphology variability of the Crimean
southern crested newt and the Caucasian species by the methods of multivariate
statistics (fig. 3) while the Danube newt (T. dobrogicus) from South-Western Ukraine
(Odesa Province,Vilkovo) was employed as the taxon for outgroup comparison. It was
found that the variability of characteristics was determined by the first three principal
components, namely taxonomic status, gender, and geographical variation. The first
principal component characterizes the distant location of T. dobrogicus and clearly indi-
cates the importancy of taxonomic status on the graph. The second principal component
is different males and females location. It emphasizes the significance of gender. The
third principal component characterizes presence of geographical variation as samples
distribution from differents regions do not offen coincide.
It has to be noticed that the current data allow to assume the interpopulation level
of differences between T. karelinii from the Crimea and the Caucasus. Generally, males
were smaller comparing to females (e. g., sample from Ternovka village had the follow-
ing parameters for females (n=16) L. = 62.2±0.7 mm and L.cd. = 50.2±1.13; males
(n=15) L. = 58.2±1.19 and L.cd. = 47.2±1.09 mm). Furthermore, the special difference
was revealed between sizes of the newts collected in the Crimea in the whole by
N. Szczerbak in 1950’s and those collected near Ternovka village by the first author of
this paper in 2003. The first sample had the following parameters: L. = 63.33±7.04 mm,
L. cd. = 58.73±7.97, P. a. = 24.62±4.01 mm, P. p. = 25.98±5.27 mm, while the sample
collected later had other values: L. = 60.3±0.76 mm, L. cd. = 48.7±0.82, Lt. c. =
12.3±0.14, P. a. = 20.4±0.27, P. p. = 22.1±0.27, DiE = 32.4±0.58, IW = 63.2±0.98. It
might be suggested that the difference is influenced by males that represented about
72% of the first sample (Щербак, 1966).
The exclusive attention has to be paid to the specificity of the throat coloration
(spotted pattern) of T. karelinii from the Crimea. The particular features of this charac-
teristic in the representatives of different taxa of T. karelinii sensu lato and the pattern of
its geographical variation have been studied before. All researchers agreed that southern
crested newt has the numerous spots on the throat (Olgun et al., 2001; Litvinchuk and
Borkin, 2009; Çiçek et al., 2010; Wielstra et al., 2013 a). At the same time, some of
them pointed out “to the absence of the distinctive T. karelinii throat pattern...” on the s.
l. in holotype and paratypes of T. arntzeni (Wielstra et al., 2013 a, p. 444).
Our analysis of seven T. karelinii samples from the Crimea, the North (Ciscauca-
sia) and the South Caucasus (Transcaucasia) regions as well as T. dobrogicus (Ukraine,
Odesa Region) revealed that only about 10% of the Crimean population has no
distinctive throat spotted pattern (fig. 2, d).
Fig. 3. Females (a) and males (b) distribution in coordinates of the first two principal components
according to the results of the discriminant analysis based on the external morphomeric characteristics
(females numbers are presented in Table 1).
Збірник праць Зоологічного музею, 2014, № 45 55
A preliminary comparison of cranial and external morphological variability of Southern Crested Newt...
Conclusions
Our data confirm that T. karelinii from the Crimea is related to Karelin’s newt from the
Caucasus that belongs to the same (eastern) mitochondrial DNA gene pool lineage
(Wielstra et al., 2013 b). On the other hand, the newts from these regions demonstrated
some differences in skull structure, external morphometric characteristics and throat
coloration patterns. Further detailed studies of T.karelinii from the Crimea are required.
We wish to express our gratitude to Dr. Yu.A. Krasylenko (Ukraine, Kyiv) for the im-
provement of the English text of manuscript prior to publication and M. Tokar (New
York) as well.
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Є. Писанець, О. Кукушкін, О. Мануілова
ПОПЕРЕДНЄ ПОРІВНЯННЯ КРАНІАЛЬНОЇ І ЗОВНІШНЬОЇ МОРФОЛОГІЧНОЇ МІНЛИВОСТІ
ТРИТОНА КАРЕЛІНА (TRITURUS KARELINII) З КРИМУ ТА КАВКАЗУ
Група Triturus karelinii з числа гребінчастих тритонів включає східну, центральну і західну гілку, які
відрізняються за мітохондріальною ДНК. Східна гілка значною мірою зберігає анцестральну форму
черепної коробки і поширюється всім Кримським півостровом, Кавказьким перешийком і південним
узбережжям Каспійського моря в Ірані. Проведено порівняння кримської і закавказької черепної
форми Т. karelinii, вказано на відмінності мінливості розмірних характеристик і забарвлення тіла.
Екземпляри Т. dobrogicus з дельти Дунаю були використані для порівняння. Дані про відмінності
між декількома популяціями в межах східної гілки припускають, що вони не є однорідним, і Т. ka-
relinii з Кримського півострова дещо відрізняється.
Ключ о в і с л о в а : тритон Кареліна, Triturus karelinii, Крим, Кавказ, череп, зовнішня морфологія,
таксономічний статус.
Е. Писанец, О. Кукушкин, О. Мануилова
ПРЕДВАРИТЕЛЬНОЕ СРАВНЕНИЕ КРАНИАЛЬНОЙ И ВНЕШНЕЙ МОРФОЛОГИЧЕСКОЙ ИЗ-
МЕНЧИВОСТИ ТРИТОНА КАРЕЛИНА (TRITURUS KARELINII) ИЗ КРИМА И КАВКАЗА
Группа Triturus karelinii гребенчатых тритонов включает восточную, центральную и западную ветвь,
которые отличаются митохондриальной ДНК. Восточная ветвь в значительной мере сохраняет анце-
стральную форму черепной коробки и распространена по всему Крымскому полуострову, Кавказ-
скому перешейку и южному побережью Каспийского моря в Иране. Проведено сравнение крымской
и закавказской черепной формы Т. karelinii, указано на отличия изменчивости размерных характери-
стик и окраски тела. Экземпляры Т. dobrogicus из дельты Дуная были использованы для сравнения.
Данные об отличиях между несколькими популяциями в пределах восточной ветви допускают, что
они неоднородны, и Т. karelinii с Крымского полуострова несколько отличаются.
Ключ е вы е с л о в а : тритон Карелина, Triturus karelinii, Крым, Кавказ, череп, внешняя
морфология, таксономический статус.
Contact:
Yevgen Pysanets,
National Museum of Natural History NAS Ukraine,
15 B. Khmelnitsky Str., Kyiv, 01601 Ukraine.
E-mail: yevgen_pysanets@yahoo.com
Контакт:
Євген Писанець,
Національний науково-природничий музей НАН України,
вул. Б. Хмельницького, 15, Київ, 01601 Україна.
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