New Records of Proteid Salamanders (Amphibia, Caudata) from the Pliocene of Ukraine and Lower Pleistocene of Moldavia
Two isolated proteid vertebrae are described: the first is from the Pliocene Kotlovina locality, Ukraine, and is referred to Mioproteus sp., whereas the second is from the lower Pleistocene Chishmikioi locality in Moldavia (former USSR) and is referred to “Mioproteus” wezei...
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| Cite this: | New Records of Proteid Salamanders (Amphibia, Caudata) from the Pliocene of Ukraine and Lower Pleistocene of Moldavia / A.O. Averianov // Вестн. зоологии. — 2001. — Т. 35, № 1. — С. 43-46. — Бібліогр.: 13 назв. — англ. |
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irk-123456789-95112010-07-02T12:02:09Z New Records of Proteid Salamanders (Amphibia, Caudata) from the Pliocene of Ukraine and Lower Pleistocene of Moldavia Averianov, A.O. Фауна и систематика Two isolated proteid vertebrae are described: the first is from the Pliocene Kotlovina locality, Ukraine, and is referred to Mioproteus sp., whereas the second is from the lower Pleistocene Chishmikioi locality in Moldavia (former USSR) and is referred to “Mioproteus” wezei. This problematic species is distinct from the Miocene M. caucasicus and may pertain to a distinct genus that includes aquatic ancestors for the modern troglobitic Proteus. Описаны два позвонка, отнесенные к Mioproteus sp. и «Mioproteus» wezei и происходящие из соответственно плиоценового местонахождения Котловина в Украине и раннеплейстоценового местонахождения Чишмикиой в Молдавии. Последний вид отличен от миоценового M. caucasicus и, возможно, его следует выделить в особый род, объединяющий речных предков современных пещерных Proteus. 2001 Article New Records of Proteid Salamanders (Amphibia, Caudata) from the Pliocene of Ukraine and Lower Pleistocene of Moldavia / A.O. Averianov // Вестн. зоологии. — 2001. — Т. 35, № 1. — С. 43-46. — Бібліогр.: 13 назв. — англ. 0084-5604 http://dspace.nbuv.gov.ua/handle/123456789/9511 567.9 : 551.782.2(477, 478) en Інститут зоології ім. І.І. Шмальгаузена НАН України |
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Фауна и систематика Фауна и систематика Averianov, A.O. New Records of Proteid Salamanders (Amphibia, Caudata) from the Pliocene of Ukraine and Lower Pleistocene of Moldavia |
| description |
Two isolated proteid vertebrae are described: the first is from the Pliocene
Kotlovina locality, Ukraine, and is referred to Mioproteus sp., whereas the second is from the lower Pleistocene
Chishmikioi locality in Moldavia (former USSR) and is referred to “Mioproteus” wezei. This problematic
species is distinct from the Miocene M. caucasicus and may pertain to a distinct genus that includes aquatic
ancestors for the modern troglobitic Proteus. |
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Article |
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Averianov, A.O. |
| author_facet |
Averianov, A.O. |
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Averianov, A.O. |
| title |
New Records of Proteid Salamanders (Amphibia, Caudata) from the Pliocene of Ukraine and Lower Pleistocene of Moldavia |
| title_short |
New Records of Proteid Salamanders (Amphibia, Caudata) from the Pliocene of Ukraine and Lower Pleistocene of Moldavia |
| title_full |
New Records of Proteid Salamanders (Amphibia, Caudata) from the Pliocene of Ukraine and Lower Pleistocene of Moldavia |
| title_fullStr |
New Records of Proteid Salamanders (Amphibia, Caudata) from the Pliocene of Ukraine and Lower Pleistocene of Moldavia |
| title_full_unstemmed |
New Records of Proteid Salamanders (Amphibia, Caudata) from the Pliocene of Ukraine and Lower Pleistocene of Moldavia |
| title_sort |
new records of proteid salamanders (amphibia, caudata) from the pliocene of ukraine and lower pleistocene of moldavia |
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Інститут зоології ім. І.І. Шмальгаузена НАН України |
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2001 |
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| citation_txt |
New Records of Proteid Salamanders (Amphibia, Caudata) from the Pliocene of Ukraine and Lower Pleistocene of Moldavia / A.O. Averianov // Вестн. зоологии. — 2001. — Т. 35, № 1. — С. 43-46. — Бібліогр.: 13 назв. — англ. |
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AT averianovao newrecordsofproteidsalamandersamphibiacaudatafromtheplioceneofukraineandlowerpleistoceneofmoldavia |
| first_indexed |
2025-07-02T12:48:11Z |
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Vestnik zoologii, 35(1): 43—46, 2001
© 2001 A. O. Averianov
UDC 567.9 : 551.782.2(477, 478)
NEW RECORDS OF PROTEID SALAMANDERS
(AMPHIBIA, CAUDATA) FROM THE PLIOCENE OF UKRAINE AND
LOWER PLEISTOCENE OF MOLDAVIA
A. O. Averianov
Zoological Institute Russian Academy of Sciences, Universitetskaya nab., 1, St.-Petersburg, 199034 Russia
E-mail: sasha@AA1923.spb.edu
Accepted 24 April 2000
New Records of Proteid Salamanders (Amphibia, Caudata) from the Pliocene of Ukraine and Lower Pleistocene
of Moldavia. Averianov A. O. – Two isolated proteid vertebrae are described: the first is from the Pliocene
Kotlovina locality, Ukraine, and is referred to Mioproteus sp., whereas the second is from the lower Pleistocene
Chishmikioi locality in Moldavia (former USSR) and is referred to “Mioproteus” wezei. This problematic
species is distinct from the Miocene M. caucasicus and may pertain to a distinct genus that includes aquatic
ancestors for the modern troglobitic Proteus.
Key wo rd s : Amphibia, Caudata, Proteidae, Mioproteus, Pliocene, Pleistocene, Ukraine, Moldavia.
Íîâûå íàõîäêè ïðîòåèäíûõ õâîñòàòûõ àìôèáèé (Amphibia, Caudata) èç ïëèîöåíà Óêðàèíû è íèæíåãî
ïëåéñòîöåíà Ìîëäàâèè. Àâåðüÿíîâ À. Î. – Îïèñàíû äâà ïîçâîíêà, îòíåñåííûå ê Mioproteus sp. è
«Mioproteus» wezei è ïðîèñõîäÿùèå èç ñîîòâåòñòâåííî ïëèîöåíîâîãî ìåñòîíàõîæäåíèÿ Êîòëîâèíà â
Óêðàèíå è ðàííåïëåéñòîöåíîâîãî ìåñòîíàõîæäåíèÿ ×èøìèêèîé â Ìîëäàâèè. Ïîñëåäíèé âèä îòëè÷åí
îò ìèîöåíîâîãî M. caucasicus è, âîçìîæíî, åãî ñëåäóåò âûäåëèòü â îñîáûé ðîä, îáúåäèíÿþùèé ðå÷íûõ
ïðåäêîâ ñîâðåìåííûõ ïåùåðíûõ Proteus.
Êëþ÷åâûå ñëîâà : Amphibia, Caudata, Proteidae, Mioproteus, ïëèîöåí, ïëåéñòîöåí, Óêðàèíà, Ìîëäà-
âèÿ.
Introduction
Proteids (Proteidae) are a family of specialized, paedomorphic, perennibranchiate salamanders with an
elongate body. Extant proteids are confined to Europe and North America, but in the past the family’s range
extended as far eastwards as Kazakhstan or, possible, Kirghisia. The fossil record for the Proteidae is limited (Estes,
1981), particularly from the territory of the former Soviet Union. Badly preserved vertebrae of Proteidae? were
reported from the lower Eocene Andarak II locality in Kirghisia (×õèêâàäçå, 1984). The middle Miocene (middle
Sarmatian, Vallesian) Mioproteus caucasicus Estes, Darevsky, 1977 was described on the basis of isolated vertebrae
and some cranial elements from the Maikop locality in the North Caucasus, Russia (Estes, Darevsky, 1977).
Mioproteus sp. was reported from the Miocene Kentyubek locality in western Kazakhstan (Áåíäóêèäçå, ×õèêâàäçå,
1976; ×õèêâàäçå, 1984; Estes, 1981). Chkhikvadze (×õèêâàäçå, 1981: 152) stated that “probably all the salamander
amphicoelous vertebrae from the [Mio-Pliocene Moldavian and Ukrainian] localities Buzhory, Male,
Novoelisavetovka, Novaya Emetovka, Cherevichnoe, and Kuchurgan should be referred to the genus Mioproteus.”
These records were repeated in Chkhikvadze (×õèêâàäçå, 1984) and Chkhikvadze & Lungu (×õèêâàäçå, Ëóíãó,
1984), who referred a fragmentary vertebra from the middle Sarmatian (Vallesian, MN 9) Moldavian Buzhory
locality to Mioproteus sp. However, amphicoelous vertebrae are also characteristic for many salamander families,
including the Cryptobranchidae and Hynobiidae, both of which are also known from the Neogene of Eurasia
(Roček, 1994; Averianov, Tjutkova, 1995; Venczel 1999). For example, the cryptobranchid Andrias [sp.] was
reported from the Ruscinian (MN 14) Kuchurgan locality in Moldavia (×õèêâàäçå, 1981, 1984) and some Miocene
localities in Moldavia (×õèêâàäçå, 1984). The ZIN PH collection contains an undescribed dentary fragment of a
cryptobranchid from the lower Ruscinian (MN 14) Antipovka locality, Voronezh Region of Russia. None of the
vertebrae from the localities mentioned by Chkhikvadze has been described or illustrated and their attribution to
proteid salamanders remains to be demonstrated; these records are not considered future here. The only reliable,
published records of fossil European Proteidae are the following: M. caucasicus from the middle Miocene of Maikop,
western Russia and Appertshofen, Germany; Orthophyia longa Meyer, 1845 from the upper Miocene of Oeningen,
Germany; “Mioproteus” wezei (Estes in Młynarski et al. 1984) from the lower Pliocene of Węże II, Poland and up-
per Pliocene of Balaruc II, France; and Proteus bavaricus Brunner, 1956 from the Pleistocene of Kleinen
A. O. Averianov 44
Teufelshöle, Germany (Dehm 1961; Estes, Darevsky, 1977; Estes, 1981; Młynarski et al., 1984 and references
therein). The proteid vertebrae from the Miocene of Kentyubek, western Kazakhstan (Áåíäóêèäçå, ×õèêâàäçå,
1976) are probably referable to M. causcasicus (Estes, 1981: 27), but they still not described.
In this note I describe two isolated proteid vertebrae: one is from the Pliocene Kotlovina locality, Ukraine, and
is referred to Mioproteus sp.; the second vertebra is from the lower Pleistocene Chishmikioi locality, Moldavia
(former USSR Republic), and is referred to “Mioproteus” wezei.
The materials described herein are housed in the Paleoherpetological collection of the Zoological Institute,
Russian Academy of Sciences, St.-Petersburg (ZIN PH).
Order CAUDATA Oppel, 1811
Family Proteidae Hogg, 1838
Mioproteus Estes & Darevsky, 1977
Mioproteus sp. (fig. 1)
Ma t e r i a l . ZIN PH 1/14, trunk vertebra. Kotlovina, Odessa Province, Ukraine. Pliocene.
Descr ip t ion . The vertebra is relatively broad transversely and short anteroposteriorly.
The centrum is amphicoelous, without peripheral calcification in the cotyles. The
ventromedian keel is shallow. Weak posterior basapophyses are present. The centrum is
deeply excavated to either side and below the transverse processes. There are two separate
subcentral foramina on the right side and large single foramen on the left side. The
transverse process is unicipital, without a trace of the rib articulation surface. There is a
prominent, flange-like and horizontal ventral lamina on the transverse process. The posterior
edge of the lamina extends posterolaterally. A prominent vertical lamina originates from the
underside of the interzygapophyseal ridge and extends posteroventrally to join with the
ventral lamina at the posterior edge of the latter. Just anterior and posterior to the vertical
lamina there are two foramina that are
confluent with the subcentral foramen.
The neural arch is flattened. The neural
spine is a poorly developed keel. The
posterior surface of the neural arch is
deeply incised, but lacks the posteriorly
directed “forks” that are characteristic for
trunk vertebrae in M. caucasicus. The
centrum length is 6.20 mm.
Compar i son. The vertebra de-
scribed is referred to Proteidae by the
following combination of characters:
1) transverse process unicipital; 2) there
is no trace of rib articulation surface;
3) transverse process with prominent ven-
tral and vertical laminae. It is referred to
Mioproteus by posterior basapophyses pre-
sent, neural arch relatively wide, and
ventral surface of centrum flattened. The
specimen described is within the size
range of the type species, M. caucasicus
(centrum length 2.50—6.65 mm, Estes,
Darevsky, 1977: 165). It differs from the
latter in lacking a distinct “fork” at the
posterior margin of neural arch and in
lacking an anteriorly projecting process
arising from the ventral lamina of the
transverse process.
Comments . There are three fos-
siliferous levels at the Kotlovina locality,
one Ruscinian and two early Villafran-
Fig. 1. Mioproteus, ZIN PH 1/14, trunk vertebra. Kotlovina,
Odessa Province, Ukraine. Pliocene: A – dorsal; B – ventral;
C – anterior; D – posterior; E – right lateral views. Scale –
1 mm.
Ðèñ. 1. Mioproteus ÇÈÍ PH 1/14, òóëîâèùíûé ïîçâîíîê.
Êîòëîâèíà, Îäåññêàÿ îáë, Óêðàèíà. Ïëèîöåí: A – äîð-
ñàëüíî; B – âåíòðàëüíî; C – ñïåðåäè; D – ñçàäè; E –
ñïðàâà. Ìàñøòàá – 1 ìì.
New Records of Proteid Salamanders... 45
chian (Òîïà÷åâñêèé, Íåñèí, 1989). The stratigraphic provenance ZIN PH 1/14 is uncer-
tain because it is not known from which level the specimen was collected.
“Mioproteus” wezei (Estes in Młynarski et al. 1984) (fig. 2)
Ma t e r i a l . ZIN PH 2/14, anterior trunk vertebra. Chishmikioi, Moldavia. Lower Pleistocene.
Descr ip t ion . The vertebra amphicoelous, lightly ossified, and relatively elongate. The
prezygapophysis is large, projects anterolaterally, and bears a well defined articular facet. The
ventromedian keel is flattened anteriorly, but the form of the more posterior part of the keel
is unknown because the remainder of the centrum is broken. Along the middle part the crest
was probably sharply crested. The centrum is deeply excavated in the region of the transverse
processes. There are two separate subcentral foramina on the left side and large single
foramen on the right side. The transverse process is unicipital, without a trace of the rib
articulation surface. There is a prominent ventral lamina on the transverse process, with the
posterior edge of the lamina directed posterolaterally. In lateral view the interzygapophyseal
ridge is bent dorsally to form an arc. A prominent vertical lamina originates from the
interzygapophyseal ridge and extends posteroventrally to join with the ventral lamina at the
posterior edge of the latter. Just anterior and posterior to the vertical lamina there are two
foramina that are confluent with the subcentral foramen. The neural arch is flattened. The
neural crest is a low keel and the neural spine is a low, posterodorsally projecting process.
The posterior edge of the neural arch is not incised and has no “forks”. The centrum length
is 8.05 mm.
Compar i son. The specimen de-
scribed is referred to the Proteidae us-
ing the same characters listed above for
ZIN PH 1/14. ZIN PH 2/14 is most
similar to vertebrae of the Pliocene
“Mioproteus” wezei having delicate,
elongate, and lightly ossified vertebrae,
a deeply amphicoelous centrum, lateral
cavities delimited by ventral and
vertical laminae deep and voluminous,
and zygapophyses protuberant. The
most striking similarity between the
Polish and Moldavian specimens is a
great dorsal curvature of the interzy-
gapophyseal ridge, which may be a
diagnostic feature at the specific or
generic level. As in a supposed anterior
vertebra of “Mioproteus” wezei (Mły-
narski et al., 1984: 212) in ZIN PH
2/14 the neural arch is not forked pos-
teriorly; instead the neural spine is a
single median projection. ZIN PH 2/14
thus appears to be an anterior trunk
vertebra. The specimen described
exceeds the known size range for
“Mioproteus” wezei vertebrae from Węże
II (centrum length 4.3—7.5, M=5.6
mm, Młynarski et al., 1984: 212).
Comments . “Mioproteus” wezei
apparently was not a troglobitic but
normally aquatic animal, judging from
the associate fauna and from lack of
Fig. 2. “Mioproteus” wezei, ZIN PH 2/14, anterior trunk verte-
bra. Chishmikioi, Moldavia. Early Pleistocene: A – dorsal; B –
ventral; C – anterior; D – posterior; E – right lateral views.
Scale – 1 mm.
Ðèñ. 2. “Mioproteus” wezei, ÇÈÍ PH 2/14, ïåðåäíåòóëîâèù-
íûé ïîçâîíîê. ×èøìèêèîé, Ìîëäàâèÿ. Ðàííèé ïëèîöåí.
A – äîðñàëüíî; B – âåíòðàëüíî; C – ñïåðåäè; D – ñçàäè;
E – ñïðàâà ñáîêó. Ìàñøòàá – 1 ìì.
A. O. Averianov 46
mountains close to the locality areas. “Mioproteus” wezei differs from M. caucasicus in having
more delicate and elongate vertebrae and, for these reasons, the former species probably
should be referred to another genus. This undescribed genus possibly was the aquatic
ancestor for the modern troglobitic Proteus.
The late Miocene Orthophyia longa from Germany resembles M. caucasicus in having
long and narrow vertebrae and relatively short skull. These resemblances may indicate that
O. longa and M. caucasicus are congeners or even the same species (Estes, 1981: 29).
However, the relatively more elongate vertebrae in O. longa seem to preclude this possibility.
Nevertheless, Orthophyia may prove to be the appropriate name for the aquatic European
Plio-Pleistocene proteids that are ancestral to modern Proteus.
Acknowledgements
I am grateful to Igor Danilov (ZIN) and James Gardner (University of Alberta, Canada) for reading of the
manuscript and providing useful comments.
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