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ISSN 00310301, Paleontological Journal, 2011, Vol. 45, No. 1, pp. 90–104. © Pleiades Publishing, Ltd., 2011
Original Russian Text © A.G. Sennikov, 2011, published in Paleontologicheskii Zhurnal, 2011, No. 1, pp. 82–96.
90
INTRODUCTION
The order Prolacertilia belongs to archosauro
morph diapsid reptiles and is subdivided into the fam
ilies Protorosauridae, Prolacertidae, and Tanystrop
heidae. Prolacertilians are almost globally distributed
and occur from the Late Permian to the end of the Tri
assic.
In Eastern Europe, prolacertilians are represented
by a relatively small number of taxa (Sennikov, 2008).
The Permian beds have only yielded individual frag
mentary specimens of a very specialized form from the
family Protorosauridae,
Eorasaurus olsoni
Sennikov,
1997. The Triassic of Eastern Europe has yielded sev
eral representatives of the family Prolacertidae,
namely,
Microcnemus efremovi
Huene, 1940,
Boreopri
cea funerea
Tatarinov, 1978, and
Vritramimosaurus
dzerzhinskii
Sennikov, 2005 from the Lower Triassic
beds and
Malutinisuchus gratus
Otschev, 1986 from the
Middle Triassic.
Among localities of Early Triassic vertebrates of the
Russian Platform and ForeUrals, the Donskaya Luka
locality occupies a special place (Rykov and Ochev,
1966; Sennikov, 1999, 2001; Novikov et al., 2001,
2002; Shishkin et al., 2006). This locality is situated
within a small field of Lower Triassic deposits on the
southeastern slope of the Voronezh Anteclise, on the
right slope of the Don River valley. The beds contain
ing vertebrate remains belong to the Lipovskaya For
mation of the Gamian Horizon of the Yarengian
Superhorizon of the Upper Olenekian Substage of the
Lower Triassic, that is, to the terminal Lower Triassic.
The Donskaya Luka locality has yielded a specific
local vertebrate fauna. Although vertebrate remains
are rather fragmentary, the faunal assemblage from this
locality is most diverse (taking into account both ter
restrial and marine taxa) and comprises the greatest
number of taxa among Early Triassic assemblages of
Eastern Europe. The Donskaya Luka Tetrapod
Assemblage includes the labyrinthodonts
Parotosu
chus panteleevi
Otschev,
Trematosaurus
sp.,
Inflecto
saurus amplus
Shishkin, and
Batrachosuchoides lacer
Shishkin; nondescribed Bystrowianidae (Novikov,
2006); the procolophonid
Orenburgia enigmatica
(Tchud. et Vjusch.); the presumable trilophosaurians
Coelodontognathus ricovi
Otschev,
C. donensis
Otschev,
Vitalia grata
Ivachnenko, and
Doniceps lipovensis
Otschev et Rikov; the eosauropterygian
Tanaisosaurus
kalandadzei
Sennikov; the rauisuchid thecodonts
Tsylmosuchus donensis
Sennikov and
Scythosuchus
basileus
Sennikov; probably, the erythrosuchid
Garjai
nia
sp. and nondescribed ctenosauriscids; and the
kannemeyeroid dicynodont
Putillosaurus sennikovi
Surkov. In addition, the Donskaya Luka locality has
yielded fragmentary vertebrate remains of uncertain
taxonomic position. In particular, Ochev (1976) men
tioned an unusual tooth tentatively determined as an
ichthyosaurian tooth.
The materials collected by expeditions of Saratov
State University and the Paleontological Institute of
the Russian Academy of Sciences (PIN) organized by
V.G. Ochev and M.A. Shishkin in Donskaya Luka
contain fragmentary remains of prolacertilians identi
fied as
Microcnemus
sp. (Rykov and Ochev, 1966;
Garyainov and Rykov, 1973) or as Prolacertidae gen.
New Tanystropheids (Reptilia: Archosauromorpha)
from the Triassic of Europe
A. G. Sennikov
Borissiak Paleontological Institute, Russian Academy of Sciences, Profsoyuznaya ul. 123, Moscow, 117997 Russia
email: sennikov@paleo.ru
Received December 22, 2009
Abstract
—A new prolacertilian species and genus,
Augustaburiania vatagini
gen. et sp. nov. (Reptilia: Archo
sauromorpha), from the Lower Triassic of the Don River Basin is described. It is the first representative of the
Tanystropheidae in the Eastern Europe and the world oldest member of this family. Another new genus
(
Protanystropheus
gen. nov.) from Central and Western Europe is also established. The diversity, systematics,
phylogeny, evolution, and stratigraphic and geographical distribution of prolacertilians are discussed. The
ecological role of prolacertilians in Early Triassic communities and adaptation to marine environments are
analyzed.
Keywords
: Archosauromorpha, Prolacertilia, Tanystropheidae, diversity, systematics, phylogeny, Triassic,
Eastern Europe.
DOI: 10.1134/S0031030111010151
PALEONTOLOGICAL JOURNAL Vol. 45 No. 1 2011
NEW TANYSTROPHEIDS (REPTILIA: ARCHOSAUROMORPHA) 91
nov. (Sennikov, 1999, 2001, 2005). As a result of recent
field studies, I.V. Novikov, A.G. Sennikov, and
M.G. Minikh obtained many new specimens, which
allow a new prolacertilian genus and species to be
described. Some specimens used in the present study
were kindly transferred to PIN by A.A. Yarkov (Volga
Humanitarian Institute, Volzhsk).
The new taxon is probably most closely related to
the primitive tanystropheids
“Tanystropheus” antiquus
and recently described
Amotosaurus rotfeldensis
(Fraser and Rieppel, 2006) from Central and Western
Europe. Thus,
Augustaburiania vatagini
gen. et sp. nov.
is the first tanystropheid from European Russia (Sen
nikov, 2008). It is also interesting that
Augustaburiania
is the world earliest (Early Triassic) member of this
family, which was recorded on other continents only
beginning from the Middle Triassic.
“Tanystropheus” antiquus
from Central Europe
probably deserves to be assigned to a new independent
genus, other than
Tanystropheus
.
SYSTEMATIC PALEONTOLOGY
Order Prolacertilia
Suborder Protorosauria
Family Tanystropheidae Gervais, 1858
Tanystropheidae: Gervais, 1858, p. 234; Romer, 1956, p. 658;
Tatarinov, 1964, p. 460; Kuhn, 1969, p. 71; Wild, 1973, p. 147;
Gow, 1975, p. 118; Chatterjee, 1980, p. 198; 1986, p. 311; Carroll,
1988, Vol. 3, p. 199.
Tribelesodontidae: von Nopcsa, 1923, p. 179.
Tanystrophaeidae: Camp, 1945, p. 97.
Askeptosauridae (pars): von Huene, 1956, p. 645.
Type genus.
Tanystropheus
H. von Meyer,
1855.
D i a g n o s i s. Squamosal without descending
process. Quadratojugal absent. Interpterygoid depres
sion short. Symphysis of lower jaw with expansion.
Marginal teeth of adults anisodont, subthecodont
(pleurothecodont), conical, pointed, curved slightly
posteriorly; in juveniles, teeth heterodont, with ante
rior teeth conical singleapical and posterior teeth
threeapical. Palatal teeth only developed on vomers.
Intercenters only present in atlas and axis. Number of
cervical vertebrae increased from eight to twenty
seven. Cervical vertebrae excessively elongated. Neu
ral spines of cervical vertebrae strongly elongated, very
low. Scapula very low and wide. Humerus with ectepi
condilar notch. Manus with two or three proximal and
two or three distal carpal elements. Phalangeal formula
of manus 2–3–4–4(5)–3. Pelvis with large thyroid
notch. Foot with two or three proximal and from one to
three distal tarsal elements. Fifth metatarsal short,
straight or slightly curved. First phalanx of fifth digit
elongated. Phalangeal formula of foot 2–3–4–5–4.
Generic composition.
Augustaburiania
gen. nov., Early Triassic of Eastern Europe;
Amotosau
rus
Fraser and Rieppel, 2006, Middle Triassic of Ger
many;
Protanystropheus
gen. nov., Middle Triassic of
Central and Western Europe;
Langobardisaurus
Renesto, 1994, Late Triassic of Italy;
Tanystropheus
H. von Meyer, 1855, Middle and Late Triassic of Cen
tral and Western Europe, North Africa, Near East,
North America, and China;
Tanytrachelos
Olsen,
1979, Late Triassic of North America; and
Dinocepha
losaurus
Li, 2003, Middle Triassic of China.
O c c u r r e n c e. Triassic; Eastern, Central, and
Western Europe, North America, North Africa, Near
East, and China.
Genus
Augustaburiania
Sennikov, gen. nov.
E t y m o l o g y. The genus is named in honor of the
Czech paleontologist J. Augusta and the animal
painter Z. Burian, who contributed greatly to popular
ization of paleontology.
Type species.
Augustaburiania vatagini
Senni
kov, sp. nov.
D i a g n o s i s. Mediumsized tanystropheid. Cer
vical vertebrae elongated, their centra 12–48 mm
long. Length to anterior height ratio of centrum of cer
vical vertebrae 2.8–6.8. Axis of cervical vertebral cen
tra positioned at
4°–6
°
to horizontal. Articular sur
faces of cervical vertebral centra low, wider than high.
Neural spines of cervical vertebrae long, low, in shape
of low inverted trapezium, or, less often, low crest low
ered at midlength and having sharp projecting anterior
and posterior margins. Postzygapophyses of cervical
vertebrae terminating in pointed horizontal projec
tions located posterior to their articular surfaces.
Femur strongly sigmoidal.
Species composition. Type species, Lower
Triassic of Eastern Europe.
Comparison.
Augustaburiania
differs from
other tanystropheids in the larger deviation from the
horizontal of the centrum axis of the cervical verte
brae, the higher neural spines, and inverted trapezoid
shape of the cervical vertebrae. It differs from
Amoto
saurus, Langobardisaurus
, and
Tanytrachelos
in the
larger absolute dimensions; and from
Tanystropheus
and
Dinocephalosaurus
in the smaller absolute dimen
sions. It differs from
Amotosaurus, Tanytrachelos
, and
Dinocephalosaurus
in the more elongated cervical ver
tebrae; from
Tanystropheus, Tanytrachelos
, and
Langobardisaurus
in the greater sigmoidal curvature of
femur; from
Tanystropheus
in the less elongated cervi
cal vertebrae and the postzygapophyses with more
pointed caudal projections posterior to their articular
surfaces; and from
Dinocephalosaurus
in the longer
and more gracile femur.
Remarks. The vertebrae of
Augustaburiania
are
already considerably elongated, transformed and typi
cal of early tanystropheids, but its femur is still most sim
ilar to that of prolacertids. In the evolution of tanystrop
heids, a key role was probably played by the transforma
tion and lengthening of the neck, and only then other
skeletal segments changed. Judging from the structure
of cervical vertebrae, the neck of
Augustaburiania
was
92
PALEONTOLOGICAL JOURNAL Vol. 45 No. 1 2011
SENNIKOV
somewhat raised and sigmoidal, i.e., it was intermedi
ate in structure between typical prolacertids and typi
cal tanystropheids.
Augustaburiania vatagini
Sennikov, sp. nov.
E t y m o l o g y. The species is named in honor of
the outstanding Russian animal painter V.A. Vatagin.
H o l o t y p e. PIN, no. 1043/587, cervical verte
bra; Volgograd Region, Ilovlyanskii District, right
slope of the Don River valley, village of Sirotinskaya,
Lipovskaya gully, Donskaya Luka locality; Lower Tri
assic, terminal Olenekian Stage, Yarengian Superhori
zon, Gamian Horizon, Lipovskaya Formation.
D e s c r i p t i o n. (Figs. 1–5). The epistropheus is
elongated, its centrum is 20 mm long. The length to
anterior height ratio of the vertebral centrum is 6.2–6.5.
The centrum axis is positioned at an angle of
3°–4
°
to
the horizontal. The articular surfaces of the centrum
are low, wider than high. The centrum surface is
strongly ornamented, with many ridges. The synapo
physis is located at the anterior margin of the centrum,
weakly projects. The postsynapophyseal crest strongly
projects, extends throughout the extent of the vertebral
centrum up to its posterior edge, bifurcates in the mid
dle of the centrum length. Another crest extends from
the prezygapophysis to postzygapophysis along the
entire centrum. The spinal canal is low, wider than
high. The neural spine is low, long, only slightly
shorter than the vertebral centrum, in the shape of a
low arched crest, reaching the highest position in the
anterior part, with a thin upper margin. The prezyga
pophyses are short, with a small facet for articulation
with the neural arch of the atlas on the lateral side. The
postzygapophyses are positioned gently sloping, their
articular surfaces are wide and short. Between the
postzygapophyses, there is a narrow, wide, horizon
tally positioned bony plate. The keel is a narrow, thin,
strongly projecting crest, which extends over the entire
extent of the ventral surface of the centrum; it is singu
lar in the anterior part and bifurcates posteriorly. It
probably provided attachment area for the subverte
bralis muscle and pars capitis and cervicis muscles
(Wild, 1973).
The holotype is a cervical (probably the third) ver
tebra. It is elongated, the centrum is 24 mm long. The
length to anterior height ratio of the vertebral centrum
is 6.0. The centrum axis is positioned at an angle of
4°–5
°
to the horizontal. The articular surfaces of the
centrum are low, wider than high. The centrum surface
is strongly ornamented, with many ridges. The diapo
physes and parapophyses slightly project, are located
at the anterior margin of the centrum within its lower
half. The postdiapophyseal and postparapophyseal
0
(b)
(d)
(e)
(c)
1 cm
(а)
Fig. 1.
Augustaburiania vatagini
sp. nov.; (a–e) holotype PIN, no. 1043/587, cervical vertebra: (a) dorsal, (b) lateral (left), (c) ven
tral, (d) cranial, and (e) caudal views.
PALEONTOLOGICAL JOURNAL Vol. 45 No. 1 2011
NEW TANYSTROPHEIDS (REPTILIA: ARCHOSAUROMORPHA) 93
crests strongly project, extend over the entire extent of
the vertebral centrum to its posterior margin. A crest
deviates from the prezygapophysis, passes slightly
downwards, and comes in contact with the postdiapo
physeal crest in the posterior part of the centrum. One
more crest passes from the prezygapophysis to
postzygapophysis throughout the extent of the cen
trum. The spinal canal is low, wider than high; how
ever, posteriorly it is somewhat higher than anteriorly.
The neural spine is low, long, only slightly shorter than
the vertebral centrum, in the shape of a low inverted
trapezium, with a thin straight upper margin. The
prezygapophyses and postzygapophyses are posi
tioned gently sloping, their articular surfaces are wide
and short. Between the postzygapophyses, there is a
narrow, wide, horizontal bony plate. The postzygapo
physes terminate into pointed, horizontal projections
positioned posterior to the articular surfaces. The keel
is a single, narrow, thin, strongly projecting crestlike
bony plate, which extends over the entire extent of the
ventral surface of the centrum. On the sides of the keel,
the middle part of the vertebral centrum has two
slightly projecting crests.
Other middle cervical vertebrae differ in centrum
length, length to anterior height ratio of the centrum,
angle between the centrum axis and horizontal (Table 1),
(а) (b) (c) (d)
(e)
(f)
(g) (h) (i) (j)
(p)
(k)
(l) (m) (n) (o) (q)
01 cm
Fig. 2.
Augustaburiania vatagini
sp. nov.; (a, b) specimen PIN, no. 1043/585, posterior cervical vertebra: (a) cranial and (b) lateral
(left) views; (c, d) specimen PIN, no. 1043/589, middle pectoral vertebra: (c) lateral (right) and (d) cranial views; (e) specimen
PIN, no. 1043/598, neural arch of anterior pectoral vertebra, lateral (right) view; (f) specimen PIN, no. 1043/593, posterior pec
toral vertebra, lateral (left) view; (g, h) specimen PIN, no. 1043/595, sacral vertebra: (g) lateral (left) and (h) caudal views;
(i) specimen PIN, no. 1043/597, anterior caudal vertebra, lateral (left) view; (j, k) specimen PIN, no. 1043/596, posterior caudal
vertebra: (j) cranial and (k) lateral (left) views; (l, m) specimen PIN, no. 1043/649, left femur: (l) dorsal and (m) ventral views;
(n, o) specimen PIN, no. 1043/514, articulated distal femoral fragment and proximal tibial fragment (left): (n) lateral and
(o) dorsal views; (p, q) specimen PIN, no. 1043/671, proximal fragment of left femur: (p) proximal and (q) ventral views.
94
PALEONTOLOGICAL JOURNAL Vol. 45 No. 1 2011
SENNIKOV
and in the development of surface ornamentation,
including the keel shape, which is a thin, moderately
projecting crest usually divided in the middle and pos
terior part of the centrum into three crests.
Specimen PIN, no. 1043/840 is a middle (from
third to sixth) cervical vertebra. The postdiapophyseal
and postparapophyseal crests project moderately,
extend throughout the extent of the vertebral centrum
to its posterior margin. The neural spine is low, long,
only slightly shorter than the vertebral centrum, in the
shape of a low arched crest, with sharp, projecting
anterior and posterior margins and a slightly thickened
upper margin.
Specimen PIN, no. 1043/1141 is a middle (from
third to sixth) cervical vertebra. The postdiapophyseal
and postparapophyseal crests project weakly; only the
postparapophyseal crest extends over the entire length
of the vertebral centrum to its posterior edge. The neu
ral spine is low, long, in the shape of a low crest low
ered at the midlength.
0 1 cm
(а)
(b)
(c)
(d)
(e)
(f)
(g)
(h)
(i)
(j)
Fig. 3.
Augustaburiania vatagini
sp. nov., cervical vertebrae, right lateral view: (a) specimen PIN, no. 1043/842, epistropheus;
(b) specimen PIN, no. 1043/1142, posterior cervical vertebra; (c) specimen PIN, no. 1043/ 586, posterior cervical vertebra;
(d) specimen PIN, no. 1043/843, posterior cervical vertebra, (e) specimen PIN, no. 1043/585, posterior cervical vertebra,
(f) specimen PIN, no. 1043/1141, cervical vertebra; (g) specimen PIN, no. 1043/840, cervical vertebra; (h) specimen PIN,
no. 1043/1392, cervical vertebra; (i) specimen PIN, no. 1043/841, cervical vertebra; (j) holotype PIN, no. 1043/587, cervical ver
tebra.
PALEONTOLOGICAL JOURNAL Vol. 45 No. 1 2011
NEW TANYSTROPHEIDS (REPTILIA: ARCHOSAUROMORPHA) 95
Specimen PIN, no. 1043/841 is a cervical (from
fifth to eighth) vertebra. The diapophyses and parapo
physes are located in the upper part of the anterior
margin of the centrum, project relatively strongly. The
postdiapophyseal and postparapophyseal crests
project moderately, extend throughout the length of
the vertebral centrum to its posterior edge. The neural
spine is low, long, in the shape of an inverted trape
zium; its upper margin is not thickened.
The posterior cervical vertebrae are relatively
shorter than the middle cervical vertebrae (Table 1).
Specimen PIN, no. 1043/1142 is a posterior (probably
eighth or ninth) cervical vertebra. The articular sur
faces of the centrum are low, slightly wider than high.
The diapophyses and parapophyses are located in the
upper part of the anterior margin of the centrum,
project moderately. The postdiapophyseal crest
projects moderately, extend throughout the length of
the vertebral centrum to its posterior edge. The post
parapophyseal crest and keel are undeveloped. The
spinal canal is low, wider than high; however, it is
somewhat higher posteriorly than anteriorly. The neu
ral spine is low, long, only slightly shorter than the ver
tebral centrum, looks like a low inverted trapezium,
with a thin straight upper margin. The prezygapophyses
and postzygapophyses are positioned gently sloping,
their articular surfaces are wide and short. Between the
postzygapophyses, there is a small, thin, horizontal
Fig. 4.
Augustaburiania vatagini
sp. nov., reconstruction of the cervical and anterior pectoral regions of the vertebral column based
on holotype PIN, no. 1043/587 and specimens PIN, nos. 1043/842, 1141, 840, 841, 1392, 1142, 843, 589, 598, and 1310; skull
outline of
Tanystropheus
is after Nosotti (2007).
Table 1. Proportions of vertebral centra in Augustaburiania vatagini
Augustaburiania vatagini, specimen no.
Centrum length
of cervical vertebrae, mm
Elongation of centrum
of cervical vertebrae
Angle between centrum
axis of cervical vertebrae
and horizontal
Epistropheus, PIN, no. 1043/842
20 6.2–6.5 3
°
–4
°
Middle cervical vertebrae
.
Holotype PIN, no. 1043/587
24 6.0 4
°
–5
°
PIN, no. 1043/840
40 5.4–5.6 4
°
–5
°
PIN, no. 1043/841
32 4.6–4.8 4
°
–5
°
PIN, no. 1043/1141
47–48 6.3–6.5 4
°
–5
°
PIN, no. 1043/1392
41 6.8 5
°
–6
°
Posterior cervical vertebrae
.
PIN, no. 1043/585
12 3.0 5–6
°
PIN, no. 1043/843
12 3.8–4.0 4
°
–5
°
PIN, no. 1043/586
17 2.8–3.0 5
°
PIN, no. 1043/1142
23 3.8 ?
Pectoral vertebrae
.
PIN, no. 1043/1310
14 1.6 0
°
–2
°
PIN, no. 1043/589
15 1.9 2
°
–3
°
PIN, no. 1043/593
11 1.8 2
°
–3
°
PIN, no. 1043/845
81.62
°
–3
°
96
PALEONTOLOGICAL JOURNAL Vol. 45 No. 1 2011
SENNIKOV
0 1 cm
(а) (b) (c) (d)
(e) (f) (g) (h)
Fig. 5.
Augustaburiania vatagini
sp. nov., left femora: (a–d) dorsal and (e–h) ventral views: (a, e) specimen PIN, no. 1043/650;
(b, f) specimen PIN, no. 1043/649; (c, g) specimen PIN, no. 1043/851; (d, h) specimen PIN, no. 1043/671, proximal fragment
and specimen PIN, no. 1043/512, distal fragment.
PALEONTOLOGICAL JOURNAL Vol. 45 No. 1 2011
NEW TANYSTROPHEIDS (REPTILIA: ARCHOSAUROMORPHA) 97
bony plate. In specimen PIN, no. 1043/843, the upper
margin of the neural spine is arched rather than
straight, projects somewhat upwards at the midlength.
The postdiapophyseal and postparapophyseal crests are
moderately developed. Specimens PIN, nos. 1043/585
and 843 have a keel in the shape of a single narrow
strongly projecting crest.
The pectoral vertebrae are elongated to a lesser
extent than cervical vertebrae (Table 1). The articular
surfaces of the centrum are high, higher than wide.
The spinal canal is moderately high, its height is equal
to, or slightly greater than, width, somewhat higher
posteriorly than anteriorly. The neural spines are mod
erately high. The prezygapophyses and postzygapo
physes are positioned somewhat obliquely, their artic
ular surfaces are wide and short. The keel is undevel
oped.
In the anterior pectoral vertebrae (specimen PIN,
no. 1310), the strongly projecting diapophysis and
parapophysis are located at the base of the neural arch;
the parapophysis is at a small distance from the ante
rior edge of the centrum, the diapophysis is at the
midlength of the centrum. The diapophysis is con
nected by narrow crests to the perezygapophysis,
postzygapophysis, parapophysis, and posterior edge of
the centrum.
The neural spine of the anterior pectoral vertebra
(specimen PIN, no. 598) expands dorsally in the cran
iocaudal direction, with a slightly thickened arched
upper margin.
The middle and posterior pectoral vertebrae (spec
imens PIN, nos. 589, 593, etc.), with a strongly pro
jecting synapophysis, are located at the base of the
neural arch in the anterior part of the centrum length
at a greater or lesser distance from the anterior edge.
The neural spine of the posterior pectoral vertebrae is
inclined posteriorly, does not expand dorsally.
Thus, the diapophysis and parapophysis gradually
change their positions and shape in the caudal direc
tion in the vertebral column. From the anterior to pos
terior cervical vertebrae and, then, to the pectoral ver
tebrae, they move dorsally (from the lower part of the
centrum to the upper part and, then, onto the neural
arch) and posteriorly (from the anterior edge of the
centrum to the midlength). In the middle of the pec
toral region and further posteriorly, the diapophysis
and parapophysis are fused to form the synapophysis.
The sacral vertebra (probably the second) has small
sacral ribs deviating laterally at the midlength of the
centrum.
The anterior caudal vertebrae (specimens PIN,
nos. 597, 847) are moderately elongated. The articular
surfaces of the centrum are high, higher than wide.
The spinal canal is moderately high; its height is equal
to, or slightly greater than, the width, somewhat
higher posteriorly than anteriorly. The prezygapophy
ses and postzygapophyses are positioned obliquely,
their articular surfaces are wide and short. The trans
verse processes strongly project, are located at the
midlength of the centrum. The neural spines are mod
erately high, expand dorsally in the caudal direction,
with a slightly thickened arched upper margin. The
keel is undeveloped.
From the anterior to middle caudal vertebrae and,
then, to the posterior caudal vertebrae, the following
changes are observed: (1) elongation of the centrum;
(2) posterior displacement and shortening of the neu
ral spine and, then, reduction of its height down to
complete disappearance; and (3) reduction of trans
verse processes to complete disappearance.
In humeral fragments, the proximal epiphysis is
strongly widened and the diaphysis is narrow.
The femora are gracile, strongly sigmoidal. The
proximal articular surface is flat or slightly concave,
that is, the proximal head of the bone poorly ossified
and was formed of cartilage. The internal trochanter
and intertrochanteric fossa are well developed. The
lateral condyle is somewhat larger than the medial
condyle and projects ventrolaterally to a greater
extent. In the proximal region of the left femur (spec
imen PIN, no. 1043/851), the internal trochanter dis
plays traces of antemortem damage (probably a bite),
with partial healing manifested in thickened margins
(Fig. 5).
The tibiae are long, with a strongly expanded prox
imal epiphysis.
Va r i a b i l i t y. Particular bones of
Augustaburi
ania
show strikingly wide variability in size. The largest
cervical and pectoral vertebrae are approximately
twice longer than the smallest (Fig. 3). The femora dif
fer in size by a factor of 2–6 (Fig. 5). At the same time,
the massiveness, shape, and proportions of the bones
vary only slightly, i.e., even adult
Augustaburiania
con
tinued growing and increased several times in size
without changes in proportions and shape of the body.
This is typical for aquatic vertebrates, the constructive
requirements to the body of which during the growth
and increase in size and weight are less rigid compared
to terrestrial tetrapods, since they dwell in liquid envi
ronments in a suspended state. In general, marine rep
tiles are characterized by great age variations in size
(Sennikov, 2001). The largest adult
Augustaburiania
probably reached two meters of length. The wide size
variation of this East European tanystropheid con
firms its aquatic mode of life, dwelling in the zone of
coastal banks.
The great variability of
Augustaburiania
in both size
and fine morphological features confirms the assump
tion of a transitional position of this taxon between
prolacertids and tanystropheids. For example, the
shape of the neural spines of cervical vertebrae varies
widely from a very low inverted trapezium to slightly
projecting flat crest. This covers the variation range of
two prolacertilian families, partly prolacertids and
partly tanystropheids. Another changeable character
98
PALEONTOLOGICAL JOURNAL Vol. 45 No. 1 2011
SENNIKOV
is ornamentation, the extent to which ridges are devel
oped on the centrum surface of cervical vertebrae.
M a t e r i a l. In addition to the holotype, the type
locality has yielded specimen PIN no. 1043/842, epis
tropheus; PIN, nos. 1043/840, 841, 1141, 1327, 1328,
1392, cervical vertebrae; PIN, nos. 1043/588, 651,
fragments of cervical vertebrae; PIN, nos. 1043/585,
586, 843, 1142, posterior cervical vertebrae; PIN,
nos. 1043/589–594, 844, 845, 1143, 1144, 1310, pec
toral vertebrae; PIN, nos. 1043/397, 398, 598, neural
arches of pectoral vertebrae; PIN, no. 1043/595, sac
ral vertebra; PIN, nos. 1043/596, 597, 847–850,
1145–1147, caudal vertebrae; PIN, no. 1043/648,
fragment of right humerus; PIN, no. 1043/771, frag
ment of left humerus; PIN, nos. 1043/649, 650, 851,
left femora; PIN, nos. 1043/603, 671, proximal frag
ments of left femora; PIN, no. 1043/606, proximal
fragment of right femur; PIN, nos. 1043/601, 604,
641, 642, distal fragments of femora; PIN,
no. 1043/514, articulated distal fragment of femur and
proximal fragment of tibia (left); PIN, nos. 1043/512,
513, 644–646, 770, proximal fragments of tibiae;
PIN, no. 1043/643, distal fragment of tibia; PIN,
no. 1043/647, proximal fragment of fibula; and PIN,
nos. 1043/602, 1393, metatarsals.
R e m a r k s. The Fedorovka locality (Kirov
Region, Nagorskii District; Lower Triassic, terminal
Olenekian Stage, Yarengian Superhorizon, Fedorovka
Horizon, Fedorovka Formation) has yielded a frag
ment (posterior part) of an elongated cervical vertebra
of a prolacertilian (specimen PIN, no. 953/390),
which is similar in structure to that of
Augustaburiania
vatagini.
However, the incomplete preservation of this
isolated specimen prevents confident assignment of
the prolacertilian from Fedorovka to the new species.
The KheiYaga 4 locality (Arkhangelsk Region,
Nenets National District; Lower Triassic, terminal
Olenekian Stage, Yarengian Superhorizon, Lestan
shorskaya Formation, Lower Subformation) has
yielded a strongly elongated cervical vertebra of a pro
lacertilian (specimen PIN, no. 4370/10) which is also
similar in structure to that of
Augustaburiania vatagini.
The vertebral centrum is about 30 mm long. The
length to anterior height ratio of the vertebral centrum
is about 6.0. The axis of the vertebral centrum is at an
angle of approximately
5
°
to the horizontal. The verte
bra is enclosed in very firm sandstone, which compli
cates preparation. This vertebra possibly belongs to
Augustaburiania vatagini
; however, it is impossible to
prove this statement based on available material.
Until new data are obtained, the distribution of
Augustaburiania vatagini is restricted to the Donskaya
Luka locality. Nevertheless, prolacertilians from
Fedorovka and KheiYaga 4 suggest that early
tanystropheids (possibly Augustaburiania) had wider
geographical and stratigraphic ranges in Eastern
Europe.
Genus
Protanystropheus
Sennikov, gen. nov.
Tanystropheus
: von Huene, 1905, p. 349; 1908, p. 223; 1931,
p. 72; Peyer, 1931, p. 93; Wild, 1973, p. 151; Wild and Oosterink,
1984, p. 143; Fraser and Rieppel, 2006, p. 869.
Etymology. From the Greek
pro
(pre) and the
generic name
Tanystropheus.
Type species.
Tanystropheus antiquus
von
Huene, 1905.
D i a g n o s i s. Large massive tanystropheid. Neck
consisting of nine vertebrae. Cervical vertebrae elon
gated, their centra 24–93 mm long. Length to anterior
height ratio of cervical vertebral centra 3.7–6.5. Axis
of cervical vertebral centra positioned at
2°–5
°
to hor
izontal. Articular surfaces of cervical vertebral centra
relatively high, slightly higher than wide. Neural
spines of cervical vertebrae low, elongated, in shape of
flat crest, most projecting at midlength. Postzygapo
physes elongated, terminating in pointed projections
projecting posterior to their articular surfaces. Femur
sigmoidal.
Species composition. Type species, Mid
dle Triassic of Central and Western Europe.
Comparison.
Protanystropheus
differs from
other tanystropheids in the more massive cervical ver
tebrae; from
Amotosaurus
,
Langobardisaurus, Tany
trachelos
, and
Dinocephalosaurus
in the more elon
gated cervical vertebrae; from
Tanystropheus, Tanytra
chelos
, and
Dinocephalosaurus
in the fewer cervical
vertebrae; from
Langobardisaurus
in the greater num
ber of cervical vertebrae; from
Augustaburiania, Amo
tosaurus
, and
Langobardisaurus
in the larger absolute
body size and the lower and more weakly projecting
neural spines of cervical vertebrae in the shape of a low
arched crest; from
Tanystropheus, Tanytrachelos
, and
Langobardisaurus
in the greater sigmoidal curvature of
the femur; from
Tanystropheus
in the smaller absolute
body size, the shorter cervical vertebrae with relatively
higher neural spines more strongly projecting in the
middle part, the greater angle between the axis of the
cervical vertebral centrum and horizontal, and the
more elongated postzygapophyses with more pointed
caudal projections positioned posterior to their articu
lar surfaces; from
Augustaburiania
in the smaller angle
between the axis of the cervical vertebral centrum and
horizontal and the higher articular surfaces of these
vertebrae; and from
Dinocephalosaurus
in the longer
and more gracile femur.
Protanystropheus antiquus
(von Huene, 1905)
Tanystropheus antiquus
: von Huene, 1905, p. 349; 1908, p. 223,
textfigs. 246–249, pl. 93, figs. 1–6, pl. 94, figs. 2–5; 1931, p. 72,
textfigs. 6–17; Peyer, 1931, p. 93, textfig. 28; Wild, 1973, p. 151;
Wild and Oosterink, 1984, p. 143, textfigs. 1 and 3; Fraser and
Rieppel, 2006, p. 869.
Thecodontosaurus primus
: von Huene, 1908, p. 217, pl. 92,
figs. 8 and 9.
L e c t o t y p e. Staatliches Museum für
Naturkunde Stuttgart, no. 16687, cervical vertebra;
Poland, Upper Silesia, Krappitz (Krapkowi e) local
c
‹
PALEONTOLOGICAL JOURNAL Vol. 45 No. 1 2011
NEW TANYSTROPHEIDS (REPTILIA: ARCHOSAUROMORPHA) 99
ity; Middle Triassic, Anisian Stage, lower shell lime
stone.
D e s c r i p t i o n (Figs. 6, 7). See the generic diag
nosis.
M a t e r i a l. In addition to the lectotype, a series
of cervical and pectoral vertebrae from the Gogolin
and Sosnowi e localities (Poland, Upper Silesia;
Middle Triassic, Anisian Stage, lower shell limestone)
and from the lower Muschelkalk (shell limestone) of
Germany, Austria, and Holland (see Wild and Ooster
ink, 1984).
THE ORIGIN OF TANYSTROPHEIDS
AND ADAPTATION OF PROLACERTILIANS
TO MARINE BIOTOPES
As mentioned above,
Augustaburiania
is intermedi
ate in morphology between prolacertids and tanystro
pheids. It is not surprising that this form was originally
identified as
Microcnemus
sp. or Prolacertidae gen.
nov. Only with accumulation of new data, it has
become clear that it is an early primitive tanystrop
heid. The discovery of
Augustaburiania
provides a new
c
‹
insight into the system and phylogeny of prolacertilians.
The elongated cervical vertebrae and elongated neck are
diagnostic characters of all members of this group. At
the same time, distinctions in the morphology of cervi
cal vertebrae of prolacertilians are evidence of different
lifestyles of different genera (Table 2).
The family Prolacertidae comprises typical prolac
ertilians. Their marginal teeth are pointed, isodont,
curved slightly posteriorly; seven elongated cervical
vertebrae are present; the neural spines of cervical ver
tebrae are elongated, with expansion in the upper part,
in the shape of an inverted trapezium; the manus
includes three proximal and four distal carpals; the
foot includes three proximal and four distal tarsals, the
fifth metatarsal is elongated hooked. Prolacertids were
probably terrestrial reptiles capable of efficient swim
ming.
The most primitive and smallest representative of
the family Prolacertidae is
Boreopricea
.
Boreopricea
is
at the beginning of the morphological series continued
by more advanced
Microcnemus
or
Prolacerta
and,
then, large specialized
Malutinisuchus
,
Vritramimo
saurus, Pamelaria
, and
Malerisaurus
. Their cervical
1 cm
(b)
(c)
(а)
Fig. 6.
Protanystropheus antiquus
(von Huene), 1905; lectotype SMNS, no. 16687, cervical vertebra: (a) cranial, (b) lateral (left),
and (c) ventral views (after Peyer, 1931).
0 20 cm
Fig. 7.
Reconstruction of skeleton of
Protanystropheus antiquus
(von Huene), 1905 (after Wild and Oosterink, 1984).
100
PALEONTOLOGICAL JOURNAL Vol. 45 No. 1 2011
SENNIKOV
vertebrae had relatively high neural spines, and the
centrum axis deviated considerably from the horizon
tal. These characters indicate that the neck of these
animals was long, relatively massive, deviating consid
erably from the horizontal, with a strong Sshaped
curvature and welldeveloped musculotendinous
apparatus, primarily epaxial muscles; the head is
raised, providing some advantage during searching for
prey and hunting on land. It is evident that this was a
lineage of specialization of relatively terrestrial prolac
ertids.
In the prolacertid
Macrocnemus
, the neural spines
of cervical vertebrae are long and low and the number
of distal tarsals is reduced to two. In addition, this ani
mal is usually found in coastal beds. This suggests that
Macrocnemus
is a prolacertid most adapted for dwell
ing in aquatic environments and, hence, resembles
tanystropheids. It is possible that, from
Microcnemus
through
Macrocnemus
, a different prolacertid lineage
leads to tanystropheids.
The family Tanystropheidae is characterized by
excessively elongated cervical vertebrae, with very long
and low, reduced neural spines, and the centrum axis
of vertebrae positioned close to the horizontal. The
number of cervical vertebrae increased. This suggests
that the neck of tanystropheids was very long, without
an Sshaped curvature and with a modified, partly
reduced epaxial part of the musculotendinous appara
tus. The neck and head are positioned close to the hor
izontal plane. The limb structure also changed; the
femur has become straighter, the manus and foot are
less consolidated and poorly ossified, and the number
of distal carpals and tarsals is reduced. These morpho
logical features of tanystropheids are usually associ
Table 2. Proportions of cervical vertebral centra of prolacertilians
Taxon
Centrum length
of cervical vertebrae,
mm
Elongation
of centrum
of cervical vertebrae
Angle between
centrum axis
of cervical vertebrae
and horizontal
Protorosauridae
Eorasaurus olsoni
20 2.0 17
°
–18
°
Protorosaurus speneri
(
after
Seeley, 1887; von Meyer, 1856) 24–34 2.6–3.4 7
°
–10
°
Prolacertidae
Prolacerta broomi
(
after
Gow, 1975) 19–22 2.7–2.8 5
°
–6
°
Boreopricea funerea
5–6 2.0–2.2 10
°
–12
°
Macrocnemus bassanii
(
after
Peyer, 1937) 12–23 2.9–3.2 5
°
–7
°
Microcnemus efremovi
10–15 2.6–3.4 6
°
–8
°
Rhombopholis scutulata
(
after
Benton, Walker, 1996) 12 2.6 5
°
Vritramimosaurus dzerzhinskii
60 3.0 10
°
Malutinisuchus gratus
42–46 2.6–3.8 7
°
–9
°
Pamelaria dolichotrachela
(
after
Sen, 2003) 74–78 2.7–2.9 10
°
–12
°
Malerisaurus robinsonae
(
after
Chatterjee, 1980) 27–32 2.7–3.5 9
°
–10
°
Malerisaurus langstoni
(
after
Chatterjee, 1986) 29–31 3.1–3.5 12
°
–13
°
Ta n y s t r op he i da e
La
ngobardisaurus pandolfii
(
after
Renesto, 1994) 17–22 3.9–6.0 3
°
–5
°
Amotosaurus rotfeldensis
(
after
Fraser, Rieppel, 2006) 16–21 4.0–5.5 3
°
–5
°
Augustaburiania vatagini
12–48 2.8–6.8 4
°
–6
°
Protanystropheus antiquus
(
after
von Huene, 1908, 1931; Pey
er, 1931; Wild, 1980a,b; Wild, Oosterink, 1984)
24–93 3.7–6.5 2
°
–5
°
Tanystropheus longobardicus
(
after
Peyer, 1931; Wild, 1973) 27–260 6.0–10.0 0
°
–3
°
Tanystropheus conspicuus
(
after
Wild, 1973) 160–260 7.0–14.0 0
°
–3
°
Tanystropheus meridensis
(
after
Wild, 1980a) 28–44 11.0–12.0 0
°
–3
°
Tanystrachelos ahynis
(
after
Olsen, 1979) 5–6 2.4–2.8 ?2
°
–4
°
Dinocephalosaurus orientalis
(
after
Li, 2003; Rieppel, Fraser, 2008)
38–48 3.0–3.4 ?0
°
–4
°
PALEONTOLOGICAL JOURNAL Vol. 45 No. 1 2011
NEW TANYSTROPHEIDS (REPTILIA: ARCHOSAUROMORPHA) 101
ated with adaptation to semiaquatic or even aquatic
mode of life (Wild, 1973; Tschanz, 1985; Renesto,
2005; Nosotti, 2007). This assumption is confirmed by
the fact that the majority of tanystropheids come from
coastal beds. Specialization to aquatic (marine) mode
of life is most pronounced in
Dinocephalosaurus
(Renesto, 2005; Rieppel and Fraser, 2008).
Wild (1980a, 1980b; Wild and Oosterink, 1984) and
Tschanz (1988) have already indicated that “
Tanystro
pheus
”
antiquus
is primitive, close to prolacertids and
should be assigned to a separate tanystropheid genus.
It is interesting that Renesto (1994; Renesto and Dalla
Vecchia, 2000) also marked that
Langobardisaurus
and
“
Tanystropheus
”
antiquus
are intermediate between
prolacertids and typical tanystropheids, but assigned
them to prolacertids.
Langobardisaurus
,
Protanystro
pheus, Amotosaurus
, and
Augustaburiania
are similar
in a number of morphological characters, such as the
number of cervical vertebrae, extent of elongation of
their centra, and reduction of neural spines, interme
diate between typical prolacertids and typical
tanystropheids. All of these taxa are probably early
tanystropheids, most closely related to prolacertids.
Thus, the following morphological series is con
structed: from
Microcnemus
through
Macrocnemus
to
Langobardisaurus
,
Augustaburiania, Amotosaurus
and,
then, to
Protanystropheus
, and from
Protanystropheus
to
Tanystropheus
, a typical and most widespread
tanystropheid. At the same time, true phylogenetic
relationships were more complex.
Microcnemus
existed simultaneously with
Tanystropheus
, i.e., too
late to be an ancestor of tanystropheids. Apparently,
Langobardisaurus
is also the latest known representa
tive of archaic tanystropheids. Probably, it is only pos
sible to reduce
Tanystrachelos
and
Dinocephalosaurus
to a common prolacertid ancestor of tanystropheids
(of the specialization level of
Microcnemus
) rather
than derive them from the early tanystropheids
Augustaburiania, Amotosaurus
, and
Protanystropheus
.
As a result of a mass extinction (ecological crisis) at
the Permian–Triassic boundary, continental commu
nities of the very beginning of the Triassic were
extremely impoverished and included a few tetrapod
taxa of small and weakly specialized forms. Archosau
romorphs were represented by small lizardlike prolac
ertilians and primitive thecodonts. During the Early
Triassic, the diversity of terrestrial biota was gradually
restored. At the end of this epoch, large specialized
vertebrates appeared. The community structure had
restored to the former, precrisis level of complexity.
The appearance of
Vritramimosaurus
, a large terres
trial prolacertid predator, and
Augustaburiania
, a rela
tively large semiaquatic coastal tanystropheid, is evi
dence of complexity and diversification of vertebrate
communities of that time in Eastern Europe.
The Triassic was the time of mass return of tetra
pods into aquatic environments and adaptation to
dwelling in the sea. Transition of prolacertilians to
semiaquatic and, then, aquatic and even marine mode
of life at the end of the Early Triassic was probably
connected with diversification of the terrestrial biota,
niche packing, increasing competitive pressure within
continental communities, and an increase in the press
of rapidly evolving progressive predators, early archo
saurs. Gracile prolacertilians with a long neck and
small head were undoubtedly disadvantageous relative
to more massive and strong predatory thecodonts of
the end of the Early Triassic, i.e., Erythrosuchidae and
Rauisuchidae, which occupied a top position in the
food chain. Therefore, prolacertilians could not real
ize the ecological type of large terrestrial predators and
were displaced into the sea (tanystropheids, which
were up to 6 m long) or the ecological niche of a
mediumsized lizardlike terrestrial or coastal–semi
aquatic predator (late prolacertids). The last large ter
restrial prolacertids were 3–4mlong
Pamelaria
from
the Anisian of India and 2–3mlong
Malutinisuchus
from the Ladinian of the southern ForeUrals.
Adaptive radiation of prolacertids and, then,
tanystropheids occurred in the Early Triassic.
Tanystropheids represent a Triassic generation of
marine reptiles, as well as thalattosaurs, aquatic thec
odonts, and crocodiles. In marine environments,
tanystropheids met strongly specialized ichthyosaurs
and early sauropterygians, which certainly persisted
from the Permian Time, although the first peak of
these groups occurred at the end of the Early Triassic
and in the Middle Triassic. Even earlier, in the Early
Permian, marine reptile habitats were mastered by
mesosaurs and, at the end of this period, by eosu
chians. In my opinion, transition of these reptile
groups into the sea was not caused by progressive
adaptation to new biotopes and food objects alone.
Such an abrupt shift in adaptive strategy was probably
caused by a considerable increase in diversity of terres
trial tetrapods, packing of terrestrial niche space, a
sharp increase in competitive pressure of more
advanced groups and regulatory effect of dominant
predators.
From the end of the Early Triassic, tanystropheids
probably occupied specific ecotopes and realized an
unusual adaptive zone, which allowed them to survive
up to the end of the Triassic. They, but for
Dinocepha
losaurus
, were adapted to marine environments to a
lesser extent than early sauropterygians. Tanystrop
heids probably specialized in boundary biotopes of
marine coasts, littoral, and banks (Fig. 8). This was
probably the reason why they reached a peak in the
Middle Triassic epicontinental marine basin of Cen
tral Europe, with shallow banks, indented coastline,
many islands, gulfs, and lagoons, and abundant
marine biota. In the continental basins (North Amer
ica) and on the coasts of the Tethys (from Mediterra
nean to China), they were relatively rare, i.e., oceanic
or continental water environments and the coasts were
undoubtedly less favorable for these reptiles.
By the end of the Early Triassic, the spatial differ
entiation, diversity of regional vertebrate faunas of
102
PALEONTOLOGICAL JOURNAL Vol. 45 No. 1 2011
SENNIKOV
Eastern Europe had reached maximum. The Late
Olenekian fauna of the southern region (Voronezh
Anteclise, Donskaya Luka), with the early tanystrop
heid
Augustaburiania
, differs sharply from the fauna of
the southeastern region (southern ForeUrals and the
southeastern Russian Platform), wherefrom the pro
lacertid
Vritramimosaurus
comes. The vertebrate fauna
from the Donskaya Luka locality is more similar to
central European and even Euramerican faunas in, for
example, the sharp prevalence of rauisuchids, the
absence of proterosuchids and erythrosuchids among
thecodonts, and the presence of cymatosaurids,
ctenosaurids, and tanystropheids. These distinctive
features are probably accounted for by the distribution
of this fauna at the northern coast of the Tethys within
Eastern Europe. Various tetrapod groups could
migrate along the oceanic coasts from Europe to
North America and up to China; this provided intense
faunal exchange. Semiaquatic and aquatic coastal
tanystropheids display circumtethyan distribution (on
northern and southern coasts) and occur in the Cen
tral European Basin, i.e., the United States, Holland,
Germany, Poland, Switzerland, Italy, Romania,
southern European Russia, Morocco, Israel, Saudi
Arabia, and China. Distinctions of the Donskaya Luka
Fauna are probably connected with specific features of
coastal marine biotopes compared with inland
biotopes; this provided certain advantage to some tet
rapod groups relative to others for intense dispersal
and caused faunal and ecological isolation of the
region under study.
ACKNOWLEDGMENTS
I am grateful to A.A. Yarkov for making the speci
mens available for this study.
The study was supported by the Russian Founda
tion for Basic Research (project nos. 080500526a,
070500069a, 100500611a), the programs of the
Presidium of the Russian Academy of Sciences no. 15
“Origin of the Biosphere and Evolution of Geobiologi
cal Systems,” Subroutine II; and American Paleonto
logical Society PalSIRP, Sepkoski grant, Project
RUG11648XX06.
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