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Late Cretaceous marine reptiles from Malyy Prolom in Ryazan Oblast, Central
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DOI: 10.1016/j.cretres.2021.104946

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 Cretaceous Research 127 (2021) 104946

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Cretaceous Research
journal homepage: www.elsevier.com/locate/CretRes

Late Cretaceous marine reptiles from Malyy Prolom in Ryazan Oblast,

Central Russia
Sergey V. Solonin a, **, Alexey V. Vodorezov a, Benjamin P. Kear b, *
a
Department of Geography, Ecology and Nature Management, Ryazan State University named for S. Yesenin, Ulitsa Street 46, Ryazan 390000, Russian
Federation
b €gen 16, Uppsala, SE-752 36, Sweden
Museum of Evolution, Uppsala University, Norbyva

a r t i c l e i n f o a b s t r a c t

Article history: During the CenomanianeTuronian transition (~94 Ma), what is today Central Russia formed part of the
Received 17 February 2021 northern epicontinental margin of the Tethys Ocean. Diverse marine vertebrate faunas inhabited these
Received in revised form palaeoenvironments, but their fossils are incompletely documented. Here, we report the discovery of
20 May 2021
marine reptile remains, recovered together with pterosaur, chondrichthyan, and actinopterygian fish
Accepted in revised form 27 June 2021
Available online 3 July 2021
material from a basal-most glauconitic sand and gravel layer of the Dmitrov Formation. These strata are
exposed in an active quarry near the village of Malyy Prolom in the Shatsky District of Ryazan Oblast,
Central Russia. The Dmitrov Formation deposits are middleeupper Santonian, but unconformably con-
Keywords:
Brachaucheninae
tact the underlying loweremiddle Cenomanian Yakhroma Formation via a condensed boundary horizon
Elasmosauridae that contains the vertebrate fossils with bivalve shell fragments and siliceous and phosphatic clasts. Such
Polycotylidae sedimentary characteristics indicate a high-energy shoreface setting where the vertebrate teeth and
Pervushovisaurus bones were likely reworked during cyclical regressions commencing in the latest Cenomanianeearly
Mosasauroidea Turonian. Time-averaging is also evidenced by the mixed occurrences of brachauchenine pliosaurids,
Russian platform elasmosaurid and polycotylid plesiosauroids, ophthalmosaurid ichthyosaurians similar to Pervush-
ovisaurus, and a possible yaguarasaurine mosasauroid. These taxa are typical of CenomanianeTuronian
assemblages from across the northern peri-Tethys, and represent components of what were probably
palaeobiogeographically widespread marine reptile faunas.
© 2021 The Authors. Published by Elsevier Ltd. This is an open access article under the CC BY license
(http://creativecommons.org/licenses/by/4.0/).

1. Introduction Arkhangelsky et al., 2017; Zverkov et al., 2018a; Zverkov and

Late Cretaceous marine reptile fossils have been documented
from European Russia (Fig. 1A) for almost 180 years (e.g.,
Eichwald, 1842). Most of these specimens derive from localities
along the Volga River in Penza, Saratov, Ulyanovsk, and Volgograd
oblasts (¼ ‘provinces’), and the Chuvash Republic in the Volga
Federal District (e.g., Sintsov, 1872; Yakovlev, 1901; Bogolyubov,
1911; Tsaregradsky, 1926; Persson, 1963; Ochev, 1976; Nesov
et al., 1988; Yarkov, 1989, 1993; Arkhangelsky, 1998; Pervushov
et al., 1999; Storrs et al., 2000; Arkhangelsky et al., 2007a;
Arkhangelsky et al., 2007b; Grigoriev et al., 2009; Berezin, 2013,
2016; Fischer et al., 2014; Grigoriev, 2014; Grigoriev et al., 2015;
* Corresponding author.
** Corresponding author.
E-mail addresses: soloninserg.inbox@mail.ru (S.V. Solonin), a.vodorezov@365.
rsu.edu.ru (A.V. Vodorezov), benjamin.kear@em.uu.se (B.P. Kear).
Pervushov, 2020). By contrast, specimens from the neighbouring
Central Federal District (¼ Central Russia: Fig. 1A) have received
less research attention.
The earliest reported discoveries were made in 1848 by the
eminent palaeontologist Valerien A. Kiprianoff (Rozhdestvensky,
1973), who recovered ichthyosaurian and plesiosaurian bones and
teeth from Cenomanian phosphorite deposits exposed along the
Tuskar River in what was then the Kursk Governorate (now Kursk
Oblast). These specimens were published by Eichwald (1853, 1868),
although Kiprijanoff (1881, 1882, 1883) later identified a range of
AlbianeCenomanian marine reptile taxa from the “Kursk Osteolite”
of the Seversk Sandstone (see Welles, 1962; Bardet 1992; Storrs
et al., 2000, p. 201). These included the ophthalmosaurid ichthyo-
saurian Pervushovisaurus campylodon (Carter, 1846) (sensu Fischer,
2016; elsewhere referred to Myopterygius kiprijanoffi Romer, 1968
or Platypterygius campylodon sensu McGowan and Motani, 2003),
and various plesiosaurians, such as ‘Plesiosaurus’ helmerseni
Kiprijanoff, 1882 (associated with material of ‘Elasmosaurus’

https://doi.org/10.1016/j.cretres.2021.104946
0195-6671/© 2021 The Authors. Published by Elsevier Ltd. This is an open access article under the CC BY license (http://creativecommons.org/licenses/by/4.0/).
S.V. Solonin, A.V. Vodorezov and B.P. Kear
helmerseni and ‘Elasmosaurus’ kurskensis Bogolyubov, 1911:
Schroeder, 1884; Welles, 1962; Persson, 1963), which represents a
composite of indeterminate elasmosaurid remains (Welles, 1962;
Persson, 1963; Bardet and Godefroit, 1995; Storrs et al., 2000), and
‘Lütkesaurus’ (see Welles, 1962; Persson, 1963; Storrs et al., 2000),
which has been synonymised with the brachauchenine pliosaurid
Polyptychodon Owen, 1841 (designated a nomen dubium by Madzia,
2016).
Bogolyubov (1911) additionally described indeterminate poly-
cotylid plesiosaurian material as ‘Polycotylus’ epigurgitis
Bogolyubov, 1911 and ‘Polycotylus’ ichthyospondylus tanais
Bogolyubov, 1911 (¼ ‘Trinacromerum’ ichthyospondylum tanais:
Pravoslavlev, 1915) from an unspecified Cenomanian phosphorite
mine near Devitsy in Voronezh Governorate (¼ Voronezh Oblast).
More recently, the ichthyosaurian Platypterygius ochevi
Arkhangelsky et al., 2008 was recorded from an
AlbianeCenomanian glauconitic sand quarried at Semiluki on the
Veduga River in Voronezh Oblast (Arkhangelsky et al., 2008).
Finally, Starodubtseva et al. (2008) briefly mentioned Cen-
omanian plesiosaurian fossils from Moscow Oblast, and Eremin
(1998) noted the occurrence of an isolated mosasauroid vertebra
from undefined Cenomanian strata near Nikolskoye in the Ras-
skazovsky District of Tambov Oblast.
In this paper, we document an assemblage of plesiosaurian,
ichthyosaurian and mosasauroid dental fossils found in an active
quarry near the village of Malyy Prolom in the Shatsky District of
Ryazan Oblast, Central Russia. These deposits form part of the
Fig. 1. Diagrammatic locality maps. (A) The Russian Federation showing the approximate area of European Russia (mid-tone grey fill), the Central Federal District (light grey fill), and
Ryazan Oblast (black fill). (B) Ryazan Oblast with national highways including the E30 to Shatsk. (C) Enlargement indicating position of the Malyy Prolom quarry and mapped
distribution of Mesozoic sedimentary strata (modified from Fadeeva and Iosifova, 1998).
2
Cretaceous Research 127 (2021) 104946
Dmitrov Formation, which is a middleeupper Santonian unit
incorporating a condensed basal-most horizon containing
reworked vertebrate material (Drutskoi and Fadeeva, 2001). Our
survey evaluates the biostratigraphical and palaeobiogeographical
implications of the identifiable marine reptile remains. A pre-
liminary assessment of the associated elasmobranch assemblage
(which also includes a mention of some actinopterygian fish fossils)
has been presented elsewhere (Solonin et al., 2020).
1.1. Institutional abbreviations
CAMSM: Sedgwick Museum of Earth Sciences, University of
Cambridge, Cambridge, UK. PMU: Palaeontological Collection,
Museum of Evolution, Uppsala University, Uppsala, Sweden. RSU:
Ryazan State University, Department of Geography, Ecology and
Nature Management (DGENM), Ryazan, Russian Federation. SS:
Regional Museum of Earth Sciences, Saratov State University, Sar-
atov, Russian Federation.
2. Geological setting
The Malyy Prolom quarry is located on the on the northeastern
outskirts of Malyy Prolom village, ~170 km southeast of Ryazan
(Fig. 1B), and ~6 km northwest of the town of Shatsk (Fig. 1C). The
excavations, which currently extend to a depth of 11.2 m, cut into an
interfluve bordering the Shacha River and have been mined for
road base since the 1960s (Krivtsov et al., 2018). Geological surveys
S.V. Solonin, A.V. Vodorezov and B.P. Kear
carried out in 1998e1999, and updated in 2005, reported a ~4.5 m
thick succession of Upper Cretaceous deposits exposed across the
western, northern and eastern walls of the quarry. These are
overlain by up to 1.6 m of Quaternary glacial till capped by loam and
soil (Fig. 2). The Cretaceous strata are laterally continuous across
the southeastern Moscow depression, and form part of the Dmitrov
Formation (Kuzmin et al., 2015). This unit comprises fine-grained
quartz-glauconite ferruginous sandstones intercalated with clay-
silt layers towards the lower part of the section (Kuzmin et al.,
2015). At Malyy Prolom, the sandstone beds are further underlain
by a condensed ferruginous quartz sand horizon that is between
100 and 500 mm thick. This contains gravel, siliceous and phos-
phatic sandstone pebbles, bivalve shell fragments, and elasmo-
branch (selachimorphs and batoids), chimaeroid, actinopterygian,
pterosaur, and marine reptile fossils (see Krivtsov et al., 2018 for a
preliminary report). Subsurface lithostratigraphic samples from the
Karnauhovo borehole, ~3 km from Malyy Prolom, indicate that the
Dmitrov Formation is separated from the underlying Yakhroma
Formation by an unconformable erosion surface. Benthic in-
vertebrates including inoceramid bivalves and radiolarians suggest
that the Dmitrov Formation represents the middleeupper Santo-
nian (Sidorenko, 1971; Olferev and Alekseev, 2005). Conversely, the
Yakhroma Formation correlates with the loweremiddle Cen-
omanian (Sidorenko, 1971; Nikitin et al., 1984; Drutskoi and
Fadeeva, 2001; Kuzmin et al., 2015). The clastic basal-most hori-
zon of the Dmitrov Formation is thus likely enriched with winn-
owed material from of the Yakhroma Formation (Sidorenko, 1971),
Fig. 2. Diagrammatic stratigraphical column depicting the Yakhroma and Dmitrov
formation sedimentary succession and fossil bearing horizon within the Malyy Prolom
quarry exposure (compiled from field data by S.V.S. with information from Sidorenko,
1971; Kuzmin et al., 2015).
3
Cretaceous Research 127 (2021) 104946
implying that the accompanying fossils are probably reworked and
primarily of Cenomanian, but also Turonian and possibly Coniacian
in age (S.V.S. unpublished data; see also online poster presentation
by Solonin et al., 2020 DOI:10.13140/RG.2.2.25182.46404).
3. Materials and methods
The Malyy Prolom marine reptile fossils were recovered directly
from surface outcrop or via screen washing of bulk sediment.
Fieldwork was carried out by S.V.S., together with RSU staff and
other collaborators during multiple quarry visits in 2018e2020.
Some additional specimens were excavated by the avid fossil col-
lector Alexander Evsyutkin (Tambov), and have been registered in
the DGENM Geological Collection at RSU with digital images
archived at both RSU and PMU. The entire assemblage comprises 17
taxonomically diagnostic tooth crowns (see Supplementary
Information) and a number of unidentifiable reptilian bone frag-
ments. All of these skeletal remnants exhibit extensive surface
corrosion, pitting and edge-rounding consistent with transport and
abrasion prior to burial.
The fossils were photographed using a Nikon D7000 SLR or a
Canon EOS 650 D mounting 18e55 mm lenses. Image manipula-
tions and measurements were undertaken using ImageJ v. 1.47
(Schneider et al., 2012).
4. Survey of the marine reptile remains
4.1. Plesiosaurians
The plesiosaurian dental remains include shed crowns and
crown fragments. RSU DGE 2018 RO MP-33, RSU DGE 2018 RO MP-
49, RSU DGE 2018 RO MP-51, RSU DGE 2018 RO MP-54, RSU DGE
2019 RO MP-108, RSU DGE 2020 RO MP-02, RSU DGE 2020 RO MP-
05, RSU DGE 2020 RO MP-30, and RSU DGE 2020 RO MP-31 are all
robust curved and conical teeth with prominent apicobasal
enamel ridges. They are generally large, ranging up to ~35 mm in
maximum preserved height, although the most complete
example, RSU DGE 2018 RO MP-49, is 94 mm in maximum height,
79 mm in preserved enamel crown height, and 34 mm in
maximum basal diameter above the resorbed remnant of the root
(Fig. 3A, B). Enamel ridges encircle the crowns and typically
extend to the tooth apices (e.g., RSU DGE 2018 RO MP-51: Fig. 3C).
The ridges become densely-spaced across the lingual and distal
surfaces, which bear numerous interspersed “ridgelets” towards
the crown bases (sensu Zverkov et al., 2018b, p. 5). RSU DGE 2020
RO MP-30 and RSU DGE 2020 RO MP-31 exhibit enamel ridge
bifurcation and distinct medial and distal ridges (Fig. 3D). RSU
DGE 2020 RO MP-49 is unusual in possessing a smooth crown
apex, as well as an apparently pathological swelling and vermic-
ulate deformation of the midline enamel ridge on its distal surface
(Fig. 3A). There are no obvious enamel bands or undulations, but
the broken apex of RSU DGE 2019 RO MP-108 does have a flat-
tened labial surface delimited by prominent carinae (Fig. 3E, F).
RSU DGE 2019 RO MP-108 additionally possesses curving labial
ridges (Fig. 3E) similar to those described on a “pathological”
tooth crown (CAM SM B57333) referred to the European early
Aptian to possibly middle Santonian brachauchenine pliosaurid
‘Polyptychodon’ (see Madzia, 2016, p. 28, fig. 9).
Madzia (2016) characterised the different tooth morphologies of
‘Polyptychodon’ as sharing sub-circular to sub-oval crown cross-
sections and “coarse” (sensu Druckenmiller and Russell, 2008, p. 49,
character 91) apicobasal enamel ridges that are closely-spaced and
usually non-branching. Importantly, however, the teeth of ‘Poly-
ptychodon’ typically range up to ~50 mm in maximum crown height
(see Madzia, 2016), although much larger specimens have been
S.V. Solonin, A.V. Vodorezov and B.P. Kear Cretaceous Research 127 (2021) 104946

Fig. 3. Marine reptile dental fossils recovered from the Malyy Prolom quarry. Brachauchenine pliosaurid teeth: RSU DGE 2018 RO MP-49 in (A) mesiolabial view with deformed
midline enamel ridge (white arrowhead), and (B) lingual view; (C) RSU DGE 2018 RO MP-51 in mesolabial view; (D) RSU DGE 2020 RO MP-31 in distolingual view showing distinct
midline ridge (white arrowhead), and branching enamel ridges (black arrowhead); RSU DGE 2019 RO MP-108 in (E) lingual view with curving labial ridges (white arrowhead) and
carina (black arrowhead), and (F) labial view with carina (black arrowhead). Elasmosaurid tooth RSU DGE 2019 RO MP-107 in (G) mesolabial, and (H) lingual views. Polycotylid teeth
(I) RSU DGE 2018 RO MP-50 in mesolabial view showing branching apical ridge (black arrowhead), and (J) RSU DGE 2018 RO MP-52 in distolingual view with branching basal ridge
(white arrowhead). Ophthalmosaurid ichthyosaurian tooth fragments (K) RSU DGE 2019 RO MP-109, and (L) RSU DGE 2018 RO MP-53. Possible yaguarasaurine mosasauroid tooth
RSU DGE 2020 RO MP-32 in (M) mesolabial view, and (N) distolingual view with carina (white arrowhead). Scale bar equals 30 mm in A; 10 mm in C, EeM; 5 mm in D.

attributed (see Sachs, 2000; Kear et al., 2014). The dimensions of RSU
DGE 2018 RO MP-49 are thus comparable to the anisodont dentitions
of some larger-bodied brachauchenines, including Kronosaurus
queenslandicus Longman, 1924 from the upper Aptianeupper Albian
ramo-Fonseca et al., 2018 from
of Australia, and Sachicasaurus vitae Pa
the upper Barremian of Colombia d these taxa possess tooth crowns
that exceed ~80 mm in height (Kear, 2005; McHenry, 2009; Holland,
2018; P aramo-Fonseca et al., 2018). Interestingly, Zverkov and

4
S.V. Solonin, A.V. Vodorezov and B.P. Kear
Pervushov (2020) reported a brachauchenine cervical centrum (SSU
02/212) from probable middle Cenomanian deposits in Saratov Oblast
and extrapolated a maximum estimated body-length of ~10e11 m.
Zverkov (2015) and Zverkov et al. (2018b) also described brachau-
chenine teeth ~75 mm in crown height from BerriasianeValanginian
strata in the Republic of Crimea and Kirov Oblast, respectively. These
specimens had flattened labial surfaces and trihedral crown cross-
sections similar that of RSU DGE 2019 RO MP-108. Notably, the bra-
chauchenines Makhaira rossica Fischer et al., 2015 and Luskhan itilensis
Fischer et al., 2017, both from the upper Hauterivian of Ulyanovsk
Oblast, have diagnostically trihedral to “subtrihedral” (sensu Zverkov
et al., 2018b, p. 832) tooth cross-sections (Fischer et al., 2015, 2017), as
does Stenorhynchosaurus munozi Pa ramo-Fonseca, Go  mez-Pe
rez,
Noe , and Etayo-Serna, 2016 from the upper Barremian of Colombia
(P
aramo-Fonseca et al., 2019). Finally, the branching enamel ridges of
RSU DGE 2020 RO MP-31 are compatible with teeth of the middle
Turonian brachauchenines Brachauchenius lucasi Williston, 1903 and
Megacephalosaurus eulerti Schumacher, Carpenter, Everhart, 2013
from Kansas and Utah in the USA (Carpenter, 1996; Albright et al.,
2007a; Madzia et al., 2019), and Morocco (Angst and Bardet, 2016).
Madzia (2016) and Madzia and Machalski (2017) also described some
examples of ‘Polyptychodon’ teeth with branching enamel ridges
distributed towards the crown-root junction.
RSU DGE 2019 RO MP-107 is a broken crown (28 mm in
maximum height) that is labiolingually compressed like the “oval to
elliptical” cross-sections of elasmosaurid teeth (Druckenmiller and
Russell, 2008, p. 49e50, character 92). The distolabial enamel sur-
faces of RSU DGE 2019 RO MP-107 are smooth, but the mesiolabial
surfaces bear fine, anastamosing apicobasal ridges that extend to the
apex (Fig. 3G, H). At 11.5 mm in maximum basal diameter, RSU DGE
2019 RO MP-107 is stouter than the thin ‘needle-like’ teeth of aris-
tonectine elasmosaurids (see Otero et al., 2014; O'Keefe et al., 2017),
and otherwise resembles the more robust anisodont dentitions of
non-aristonectines (Kear et al., 2017), such as the middle Cen-
omanian Thalassomedon haningtoni Welles, 1943 (Sachs et al., 2021),
and uppermost CenomanianeTuronian Libonectes morgani (Welles,
1949) from Colorado, Nebraska, and Texas, USA (Welles, 1943,
1949; Sachs and Kear, 2015), and Morocco (Buchy, 2005; Sachs and
Kear, 2017; Allemand et al., 2017, 2018, 2019).
RSU DGE 2018 RO MP-50, RSU DGE 2018 RO MP-52, and RSU
DGE 2020 RO MP-04 (29.5 mm, 36 mm, and 32 mm in maximum
height, respectively) are slender, recurved teeth with sub-circular
basal cross-sections. They have fine apicobasal enamel ridges that
encircle the crown base, but are widely-spaced over the distolabial
surfaces and towards the apex (Fig. 3I, J). The enamel ridges also
bifurcate near the crown base on RSU DGE 2018 RO MP-50, and at
the apex on RSU DGE 2018 RO MP-52.
Schmeisser McKean (2012) described teeth of the early Turonian
polycotylid Dolichorhynchops tropicensis Schmeisser McKean, 2012
from Utah in the USA, which are similar to RSU DGE 2018 RO MP-
50, RSU DGE 2018 RO MP-52, and RSU DGE 2020 RO MP-04.
Bardet et al. (2003a) and Fischer et al. (2018) also reported elon-
gate, recurved, and carinate tooth crowns in the Moroccan middle
Turonian polycotylids Thililua longicollis Bardet et al., 2003a and
Manemergus anguirostris Buchy et al., 2005. Carinae are absent in
RSU DGE 2018 RO MP-50, RSU DGE 2018 RO MP-52, and RSU DGE
2020 RO MP-04 but reduction of the enamel ridging is character-
istic of many polycotylids as noted by Carpenter (1996), Bardet et al.
(2003a), Sato (2005), Albright et al. (2007b), Schmeisser McKean
(2012), O'Gorman and Gasparini (2013), and Fischer et al. (2018).
4.2. Ichthyosaurians
RSU DGE 2018 RO MP-53, RSU DGE 2019 RO MP-109, and RSU
DGE 2019 RO MP-110 (36 mm, 16.5 mm, and 18 mm in maximum
5
Cretaceous Research 127 (2021) 104946
preserved height, respectively) are reminiscent of ichthyosaurian
teeth. RSU DGE 2019 RO MP-109 is a squat, conical crown whose
thin enamel layer has worn away but still preserves faint traces of
regularly-spaced apicobasal ridges (Fig. 3K). RSU DGE 2018 RO MP-
53 is a tall straight crown (Fig. 3L). RSU DGE 2019 RO MP-110
alternatively comprises only a fragment, but both specimens bear
coarsely ridged enamel, which borders a dense root surface
consistent with the acellular cementum layer on the teeth of
Cretaceous ophthalmosaurids, such as Platypterygius von Huene,
1922 (Maxwell et al., 2011). Fischer et al. (2014) referred all of the
Cenomanian Platypterygius specimens from western Russia to Per-
vushovisaurus Arkhangelsky, 1998 (see also McGowan and Motani,
2003), which is distinguished by “straight, non-recurved tooth
crowns” (Fischer, 2016, p. 8) and “rugose” enamel (Fischer, 2016, p.
12) identical to the conditions in RSU DGE 2018 RO MP-53, RSU DGE
2019 RO MP-109, and RSU DGE 2019 RO MP-110.
4.3. Aquatic squamates
RSU DGE 2020 RO MP-32 is a shed tooth with flared base and
deeply inset pulp cavity comparable to the replaced functional
teeth of mosasauroids (e.g., Caldwell, 2007). The crown is 19.5 mm
in maximum height and distally recurved with a sub-circular basal
cross-section. The thin enamel is cracked and mostly worn away,
but was clearly smooth with at least a non-serrated anterior carina;
there are no obvious enamel ridges or facets (Fig. 3M, N).
VonLoh and Bell (1998) and Shimada et al. (2006) documented
(although substantially smaller at less than ~2 mm in maximum
height) recurved crowns with sub-circular basal cross-sections
from middleeupper Cenomanian deposits in South Dakota and
Colorado, USA. These were attributed to “basal mosasauroids” (see
Shimada et al., 2006: 29), but lacked carinae, which differs from
RSU DGE 2020 RO MP-32. Pa ramo-Fonseca (2000) alternately
described dimensionally larger carinate teeth in the yaguarasaurine
(sensu Palci et al., 2013) Yaguarasaurus columbianus Pa ramo, 1994
from the Turonian of Colombia. Palci et al. (2013), Street and
Caldwell (2014), and Sachs et al. (2018) likewise reported conical
carinate teeth in yaguarasaurines from the loweremiddle Turonian
strata of Italy (see also Romano et al., 2019), England, and Poland,
respectively. Notably, most of these specimens exhibited enamel
ridging (an exception being Romeosaurus fumanensis Palci et al.,
2013 from the middle Turonian of Italy), which is not visible in
RSU DGE 2020 RO MP-32, but might have originally been present
across its now abraded basal enamel surface.
5. Biostratigraphical and palaeobiogeographical implications
The Russian Platform was inundated by a shallow epicontinental
seaway throughout most of the Late Cretaceous (Vinogradov, 1975).
Connectivity along this peri-Tethyan oceanic margin extended into
the marine basins of Western Europe, northwards towards the West
Siberia Boreal Basin, and southwards into the Caucuses and Kopet
Dagh Basin of what is today Central Asia (Vinogradov, 1975;
Baraboshkin et al., 2003). Widespread marine transgression during
the earlyemiddle Cenomanian was followed by a marked regression
in the latest Cenomanianeearly Turonian; this exposed and eroded
later Cenomanian deposits, and was succeeded by multiple
transgressiveeregressive cycles during the late TuronianeConiacian
and late ConiacianeSantonian (Baraboshkin et al., 2003).
At Malyy Prolom, the unconformity between the Yakhroma
Formation and Dmitrov Formation corresponds with the onset of
these regional depositional hiatuses (Baraboshkin et al., 2002;
Baraboshkin et al., 2003; Baraboshkin, 2008). The recovered
vertebrate fossils further accord with this interpretation by incor-
porating a mixed CenomanianeSantonian assemblage:
S.V. Solonin, A.V. Vodorezov and B.P. Kear Cretaceous Research 127 (2021) 104946

elasmobranchs (see Solonin et al., 2020), such as johnlongine
odontaspidids (Johnlongia aff. J. allocotodon Siverson, 1996), carda-
biodontids (Cardabiodon sp. Siverson, 1999 and Dwardius sp.
Siverson, 1999), otodontids (Cretalamna appendiculata [Agassiz,
1843] type: sensu Siversson et al., 2015), cretoxyrhinids (Cretox-
yrhina mantelli [Agassiz, 1835]), anacoracids (Squalicorax aff. kaupi
[Agassiz, 1843]), mitsukurhinids (Anomotodon aff. A. plicatus
Arambourg, 1952), and ptychodontids (various species of Ptychodus
Agassiz, 1835); the pachycormiform fish Protosphyraena Leidy,
1857; ornithocheirid pterosaurs (Solonin et al., 2021); and marine
reptiles including brachauchenine pliosaurids, ophthalmosaurid
ichthyosaurians (Pervushovisaurus), and a possible yaguarasaurine
mosasauroid. Collectively, these taxa imply a cosmopolitan fauna
(Fig. 4) with proximal palaeobiogeographical links across European
Russia, Central and Western Europe, and as far as the northern
Gondwanan shelves of Morocco and Colombia, and the Western
Interior Basin of North America (e.g., Everhart, 2005; Siverson and
Lindgren, 2005; Ifrim et al., 2007; Cavin et al., 2010; Cumbaa et al.,
2010; Underwood et al., 2011; Cappetta, 2012; Shimada, 2012; Palci
et al., 2013; Shimada et al., 2015; Fischer, 2016; Madzia, 2016; Case
et al., 2019; Frey et al., 2020).
Among the marine reptiles, brachauchenine pliosaurids are
ubiquitous in CenomanianeTuronian assemblages from Russia
(e.g., Storrs et al., 2000; Zverkov and Pervushov, 2020), Poland
(Madzia and Machalski, 2017; Sachs et al., 2018), the Czech Republic
(Kear et al., 2014), Germany (Sachs, 2000; Sachs et al., 2016; Sachs
et al., 2017), England (Madzia 2016), and potentially Belgium,
Portugal and elsewhere (Bardet and Godefroit, 1995; Madzia 2016).
Trans-Atlantic species occurrences have been evidenced from the
Lower Chalk of Dorking, England, Goulima region of southern
Morocco, and Western Interior Basin of Utah, Kansas and Texas in
the USA (Benson et al., 2013; Angst and Bardet, 2016). Distribution
further eastwards along the Asian peri-Tethys and palaeo-Pacific
margin is also inferred by enigmatic records of ‘Polytpychodon-
like’ teeth from Japan (see Sato et al., 2012). However, the putative
account of ‘Polpytychodon patagonicus’ Ameghino, 1893 from the
southern Atlantic margin of Argentina (see Madzia, 2016) has been
reinterpreted as an indeterminate plesiosaurian (O'Gorman and
Varela, 2010).
Like brachauchenine pliosaurids, elasmosaurids and poly-
cotylids were geographically ubiquitous Late Cretaceous plesio-
saurian clades, but with numerous CenomanianeTuronian records
(including trans-Atlantic species ranges, e.g., Libonectes morgani:
Sachs and Kear, 2017) from European Russia to Western Europe
(e.g., Persson, 1963; Milner, 1987; Bardet and Godefroit, 1995; Storrs
et al., 2000; Kear et al., 2014; Sachs et al., 2016, 2017, 2018), North
Africa (e.g., Bardet et al., 2003a; Allemand et al., 2017; Sachs and
Kear, 2017; Allemand et al., 2018; Fischer et al., 2018; Allemand
et al., 2019), and North America (e.g., Carpenter, 1996, 1997, 1999;
Storrs, 1999; Sachs and Kear, 2015; Sachs et al., 2021). Similarly, Late
Cretaceous ophthalmosaurid ichthyosaurians were globally
distributed, and apparently diversified within the epicontinental
seas covering what is now the Russian Platform prior to their final
extinction at the end of the Cenomanian (Fischer et al., 2016).
Lastly, the occurrence of a possible yaguarasaurine (sensu
Madzia and Cau, 2017) is important because these and other basally

Fig. 4. Late CenomanianeTuronian palaeogeographical reconstruction showing locations of cited marine reptile assemblage occurrences. Russia (red stars): Malyy Prolom. Dmitrov
Formation and unspecified Cenomanian strata, Ryazan, Moscow and Tambov oblasts; 1. Seversk Sandstone, Kursk Oblast; 2. Unspecified Cenomanian strata, Voronezh Oblast; 3.
Melovatskaya Formation and other Cenomanian units, Saratov Oblast. Western Europe (white circles): 4. White Chalk Subgroup, southern England, UK; 5. Do € lzschen and Strehlen
formations, Dresden, Germany; 6. Bíla Hora, Jizera, Teplice formations, Bohemian Cretaceous Basin, Czech Republic; 7. Marl-limestone units, Opole, Poland; 8. Scaglia Rossa Veneta
Formation and other units, Italy; 9. Vale de Figueira, Portugal. North America (yellow squares): 10. Britton Formation and other units, Texas, USA; 11. Plattenkalk deposits, Vallecillo,
Mexico; 12. Graneros Shale, Carlile Shale and other equivalent units, Kansas, USA; 13. Graneros Shale and Greenhorn Formation, Colorado, USA; 14. Greenhorn Formation, South
Dakota, USA; 15. Carlile Shale and Belle Fourche Formation, Wyoming, USA; 16. Tropic Shale, Utah, USA. Africa (orange hexagons): 17. Akrabou Formation, Goulmima, Morocco; 18.
Itombe Formation, Iembe, Angola. South America (green diamond): 19. Villeta Formation, Neiva, Colombia. Australia (grey pentagon): 20. Molecap Greensand, Western Australia,
Australia. Japan (pink triangle): 21. Middle Yezo Group, Hokkaido, Japan. Palaeogeographical map (90 Ma Mollweide projection) reproduced under license from Global Paleoge-
ography and Tectonics in Deep Time ©2016 Colorado Plateau Geosystems Inc. (For interpretation of the references to color in this figure legend, the reader is referred to the Web
version of this article.)

6
S.V. Solonin, A.V. Vodorezov and B.P. Kear
branching mosasauroids (e.g., tethysaurines) are known from
Turonian strata throughout Europe (Palci et al., 2013; Kear et al.,
2014; Street and Caldwell, 2014; Sachs et al., 2018; Romano et al.,
2019), the Western Interior Basin of North America (Bell and
Polcyn, 2005; Polcyn and Bell, 2005; Schumacher, 2011), and the
palaeo-Atlantic margins of Africa (Lingham-Soliar, 1994; Bardet
et al., 2003b; Jacobs et al., 2006; Mateus et al., 2012; Jime nez-
Huidobro et al., 2017), and South America (Pa ramo-Fonseca,
2000; Jime nez-Huidobro et al., 2017). Jime nez-Huidobro et al.
(2017) cited evidence of CenomanianeTuronian mosasauroids
from the southern Tethys in Australia but these remains are
contentious (Kear et al., 2005; Kear, 2016; Kear et al., 2018). Putative
late Cenomanian mosasauroid fossils from Russia have been
posited as “the earliest mosasaur” (sensu Grigoriev et al., 2009, p.
315), but as in the case of the Malyy Prolom assemblage more
feasibly infer time-averaging, with both late Cenomanian and
stratigraphically younger marine reptile material being reworked
during regional marine regressions and deep sediment winnowing
(see Sidorenko, 1971).
6. Conclusions
 We describe an assemblage of Late Cretaceous marine reptile
fossils recovered from an active quarry near the village of Malyy
Prolom in the Shatsky District of Ryazan Oblast, Central Federal
District, Russian Federation. The source unit is a condensed sand
and gravel deposit representing the basal-most horizon of the
middleeupper Santonian Dmitrov Formation (Sidorenko, 1971;
Olferev and Alekseev, 2005). However, these sediments are
extensively winnowed and reworked from the unconformably
underlying Yakhroma Formation, which is loweremiddle Cen-
omanian (Sidorenko, 1971; Nikitin et al., 1984; Drutskoi and
Fadeeva, 2001; Kuzmin et al., 2015).
 Our survey of the Malyy Prolom marine reptile remains identi-
fied: (1) brachauchenine pliosaurids similar to the classic Eu-
ropean genus ‘Polyptychodon’ (Madzia, 2016), and possibly
another larger-bodied taxon (see Zverkov and Pervushov, 2020);
(2) indeterminate elasmosaurid and polycotylid plesiosaurians;
(3) ophthalmosaurid ichthyosaurians comparable with Per-
vushovisaurus d reported elsewhere from the loweremiddle
Cenomanian of Russia and Western Europe (Fischer et al.,
2014; Fischer, 2016); (4) a possible yaguarasaurine mosasauroid.
 The Malyy Prolom marine reptile assemblage thus appears to be
time-averaged, which is consistent with regional marine re-
gressions and widespread erosion that commenced during the
latest Cenomanianeearly Turonian (Sidorenko, 1971). Palae-
obiogeographical affinities are concordant with Cenomanian
and Turonian taxa from along the peri-Tethyan margins of
Russia and Central and Western Europe (e.g., Storrs et al., 2000;
Fischer et al., 2014; Kear et al., 2014; Madzia, 2016; Sachs et al.,
} et al., 2019), the palaeo-Atlantic rims of
2016, 2017, 2018; Osi
northern Africa (Bardet et al., 2008) and South America (e.g.,
Paramo-Fonseca, 2000; Jime nez-Huidobro et al., 2017), and the
Western Interior Seaway of North America (e.g., VonLoh and
Bell, 1998; Carpenter, 1999; Storrs, 1999; Polcyn et al., 2008;
Maxwell and Kear, 2010; Albright et al., 2013; Van Vranken,
2018). This suggests both dispersal and compositional homo-
geneity among low to mid-palaeolatitude (up to ~50 N) faunas
from this timeframe.
Acknowledgements
Alexander Evsyutkin (Tambov) provided generous access to his
collection and personal photographs. Vyacheslav Krivtsov (RSU),
Johan Lindgren (Lund University), David Martill (University of
7
Cretaceous Research 127 (2021) 104946
Portsmouth), and Robert Nied zwiedzki (University of Wroclaw)
contributed interpretive discussions. Eduardo Koutsoukos, the
Editor-in-Chief of Cretaceous Research, Daniel Madzia (Polish
Academy of Sciences) and Sven Sachs (Naturkunde-Museum Bie-
lefeld) provided constructive comments. BPK acknowledges
funding from a Swedish Research Council Project Grant (2020-
3423).
References
Agassiz, J.L.R., 1833e1843. Reserches sur les poissons fossils. Imprimerie de Patit-
pierre, Neuchatel, p. 1420.
Albright, B.L., Gillette, D.D., Titus, A.L., 2007a. Plesiosaurs from the Upper Cretaceous
(Cenomanian-Turonian) Tropic Shale of southern Utah. Part 1: new records of
the pliosaur Brachauchenius lucasi. Journal of Vertebrate Paleontology 27,
41e58.
Albright, B.L., Gillette, D.D., Titus, A.L., 2007b. Plesiosaurs from the Upper Creta-
ceous (CenomanianeTuronian) Tropic Shale of southern Utah, Part 2: Poly-
cotylidae. Journal of Vertebrate Paleontology 27, 41e58.
Albright, L.B., Gillette, D.D., Titus, A.L., 2013. Fossil vertebrates from the Tropic Shale
(Upper Cretaceous), Southern Utah. In: Titus, A.L., Loewen, M.A. (Eds.), At the
Top of the Grand Staircase: the Late Cretaceous of Southern Utah. Indiana
University Press, Bloomington and Indianapolis, pp. 536e562.
Allemand, R., Bardet, N., Houssaye, A., Vincent, P., 2017. Virtual reexamination of a
plesiosaurian specimen (Reptilia, Plesiosauria) from the Late Cretaceous
(Turonian) of Goulmima, Morocco, using computed tomography. Journal of
Vertebrate Paleontology 37, e1325894.
Allemand, R., Bardet, N., Houssaye, A., Vincent, P., 2018. New plesiosaurian speci-
mens (Reptilia, Plesiosauria) from the Upper Cretaceous (Turonian) of Goul-
mima (Southern Morocco). Cretaceous Research 82, 83e98.
Allemand, R., Houssaye, A., Bardet, N., Vincent, P., 2019. Endocranial anatomy of
plesiosaurians (Reptilia, Plesiosauria) from the Late Cretaceous (Turonian) of
Goulmima (southern Morocco). Journal of Vertebrate Paleontology 39,
e1595636.
Ameghino, F., 1893. Sobre la presencia de vertebrados de aspecto mesozoico en la
formacio  n Santacrucen ~ a de la Patagonia aus- tral. Revista del Jardín Zoolo  gico
de Buenos Aires 1, 76e84.
Angst, D., Bardet, N., 2016. A new record of the pliosaur Brachauchenius lucasi
Williston, 1903 (Reptilia: Sauropterygia) of Turonian (Late Cretaceous) age,
Morocco. Geological Magazine 153, 449e459.
bre
s fossils des gisements de phosphates (Maroc-
Arambourg, C., 1952. Les verte
rie-Tunisie). Notes et Me
Alge moires du Service Ge ologique du Maroc 92, 1e372.
Arkhangelsky, M.S., 1998. On the genus of ichthyosaurs Platypterygius. Paleon-
tologicheskii Zhurnal 32, 611e615.
Arkhangelsky, M.S., Pervushov, E.M., Zozyrev, N.Y., 2007a. About the first find of the
remains of a Mosasaurus from the Santonian deposits of Russia. In: Izvestiia
Vysshykh Uchebnykh Zavedenii. Geologiia i razvedka, vol. 2, pp. 3e5 (in
Russian).
Arkhangelsky, M.S., Averianov, A.O., Pervushov, E.M., 2007b. Short-necked plesio-
saurs of the Family Polycotylidae from the Campanian of the Saratov Region.
Paleontologicheskii Zhurnal 6, 62e66.
Arkhangelsky, M.S., Averianov, A.O., Pervushov, E.M., Ratnikov, V.Y., Zozyrev, N.Y.,
2008. Ichthyosaur remains from the Cretaceous of the Voronezh region.
Palaeontologicheskii Zhurnal 42, 287e291.
Arkhangelsky, M.S., Zverkov, N.G., Ivanov, A.V., 2017. On the first find of polycolid
plesiosaur from the Upper Santonian of Saratov region (Russia). In: Collection of
the Works of the All-Russian Scientific Conference Dedicated to the Memory of
Professor Vitaly Georgievich Ochev. Problems of Paleoecology and Historical
Geoecology. Kuznitsa Reklamy, Saratov, pp. 83e89 (in Russian).
Baraboshkin, E.I., 2008. Condensed sections: terminology, types, conditions of for-
mation. In: Essays on regional geology. Collection of scientific work, vol. 3.
Izdatelskii Tsentr Nauka, pp. 20e33 (in Russian).
Baraboshkin, E.I., Veimarn, A.B., Kopaevich, L.F., Naidin, D.P., 2002. Study of breaks
in stratigraphic sequence in the production of geological surveys. Guidelines.
Moskovskii Gosudarstvennyi Universitet, Moskva, p. 163 (in Russian).
Baraboshkin, E.I., Alekseev, A.S., Kopaevich, L.F., 2003. Cretaceous palaeogeography
of the North-Eastern Peri-Tethys. Palaeogeography, Palaeoclimatology, Palae-
oecology 196, 177e208.
Bardet, N., 1992. Stratigraphic evidence for the extinction of the ichthyosaurs. Terra
Nova 4, 649e656.
Bardet, N., Godefroit, P., 1995. Plesiosaurus houzeaui Dollo, 1909 from the Upper
Campanian of Ciply (Belgium) and a review of the Upper Cretaceous plesiosaurs
from Europe. Bulletin de l’Institut royal des Sciences naturelles de Belgique,
Sciences de la Terre 65, 179e186.
Bardet, N., Pereda Suberbiola, X., Jalil, N.-E., 2003a. A new polycotylid plesiosaur
from the Late Cretaceous (Turonian) of Morocco. Comptes Rendus Palevol 2,
307e315.
Bardet, N., Pereda Suberbiola, X., Jalil, N.-E., 2003b. A new mosasauroid (Squamata)
from the Late Cretaceous (Turonian) of Morocco. Comptes Rendus Palevol 2,
607e616.
Bardet, N., Houssaye, A., Rage, J.-C., Pereda Suberbiola, X., 2008. The
CenomanianeTuronian (Late Cretaceous) radiation of marine squamates
S.V. Solonin, A.V. Vodorezov and B.P. Kear
(Reptilia): the role of the Mediterranean Tethys. Bulletin de la Socie te
 convergence in the evolution of short-necked plesiosaurs. Current Biology 27,
Geologique de France 179, 605e622.
Bell, G.L.J., Polcyn, M.J., 2005. Dallasaurus turneri, a new primitive mosasauroid from
the Middle Turonian of Texas and comments on the phylogeny of Mosasauridae
(Squamata). Netherlands Journal of Geosciences 84, 177e194.
Benson, R.B.J., Evans, M., Smith, A.S., Sassoon, J., Moore-Faye, S., Ketchum, H.F.,
Forest, R., 2013. A giant pliosaurid skull from the Late Jurassic of England. PLos
One 8, e65989.
Berezin, A.I., 2013. Plesiosaurs (Sauropterygia: Plesiosauria) from the Mesozoic
deposits of Chuvashia. In: Geoecological Research in Chuvashia and Adjacent
Regions: Materials of Reports of the Regional Scientific Practical Conference.
Novoe Vremia, St Petersburg, pp. 13e19 (in Russian).
Berezin, A.I., 2016. Cretaceous plesiosaurs of Russia. In: Cretaceous System of Russia
and Near Abroad: Problems of Stratigraphy and Paleogeography. Izdatelskii
Dom, pp. 64e66 (in Russian).
Bogolyubov, N.N., 1911. From the history of plesiosaurs in Russia. Scientific notes of
Moscow University. Tipografiia Imperatorskogo Moskovskogo Universiteta,
Moskva, p. 412 [in Russian]).
Buchy, M.C., 2005. An elasmosaur (Reptilia: Sauropterygia) from the Turonian
(Upper Cretaceous) of Morocco. Carolinea 63, 5e28.
Buchy, M.-C., Me tayer, F., Frey, E., 2005. Osteology of Manemergus anguirostris n.
gen. et sp., a new plesiosaur (Reptilia, Sauropterygia) from the Upper Creta-
ceous of Morocco. Palaeontographica 272, 97e120.
Caldwell, M.W., 2007. Ontogeny, anatomy and attachment of the dentition in mo-
sasaurs (Mosasauridae: Squamata). Zoological Journal of the Linnean Society
149, 687e700.
Cappetta, H., 2012. Part 3E. Chondrichthyes. Mesozoic and Cenozoic Elasmo-
branchii. Verlag Dr. Friedrich Pfeil, München, p. 512.
Carpenter, K., 1996. A review of the short-necked plesiosaurs from the Cretaceous of
the Western Interior, North America. Neues Jahrbuch für Geologie und
Pal€aontologie - Abhandlungen 201, 259e287.
Carpenter, K., 1997. Comparative cranial anatomy of two North American plesio-
saurs. In: Callaway, J.M., Nicholls, E.L. (Eds.), Ancient Marine Reptiles. Academic
Press, San Diego, pp. 191e216.
Carpenter, K., 1999. Revision of North American elasmosaurs from the Cretaceous of
the Western Interior. Paludicola 2, 148e173.
Carter, J., 1846. On the occurrence of a new species of Ichthyosaurus in the Chalk.
London Geological Journal 1, 7e9.
Case, G.R., Cook, T.D., Kightlinger, T., 2019. Middle Campanian euselachian diversity
of the southern region of the Atlantic coastal plain of North America. Vertebrate
Anatomy Morphology Palaeontology 7, 69e82.
Cavin, L., Tong, H., Boudad, L., Meister, C., Piuz, A., Tabouelle, J., Aarab, M., Amiot, R.,
Buffetaut, E., Dyke, G., Hua, S., Le Loeuff, J., 2010. Vertebrate assemblages from
the early Late Cretaceous of southeastern Morocco: An overview. Journal of
African Earth Sciences 57, 391e412.
Cumbaa, S., Shimada, K., Cook, T.D., 2010. Mid-Cretaceous vertebrate faunas of the
Western Interior Seaway of North America and their evolutionary, paleo-
biogeographical, and paleoecological implications. Paleogeography, Paleocli-
matology, Paleoecology 295, 199e214.
Druckenmiller, P.S., Russell, A.P., 2008. A phylogeny of Plesiosauria (Sauropterygia)
and its bearing on the systematic status of Leptocleidus Andrews, 1922. Zootaxa
1863, 1e120.
Drutskoi, S.V., Fadeeva, L.I., 2001. Explanatory memorandum to the geological maps
of Quaternary and pre-Quaternary deposits of the Ryazan region at a scale of 1:
500,000. Mezhregionalnyi Tsentr po geologicheskoi kartografii, Moskva, p. 57
(in Russian).
Eichwald, E.I., 1842. Über Ichthyosaurier und Ceratiten Russlands. Bulletin de
l'Acade mie impe riale des sciences de St.-Pe tersbourg 9, 1196e1285.
Eichwald, E.I., 1853. Einige palaeontologische Bemerkungen über den Eisensand von
Kursk. Bulletin de la Socie  te
 imperiale des naturalistes de Moscou 2, 209e231.
Eichwald, E.I., 1868. Classe neuvie me. Reptiles. In: Lethaea Rossica ou Pale
ontologie
de la Russie, de crite et figure e, vol. 2. E. Schweizerbart (E. Koch), Stuttgart,
pp. 1256e1285.
Eremin, A.V., 1998. Geology of Tambov Region. Two Volumes. Volume 1. Precam-
brian. Paleozoic. Mesozoic. Izdatelstvo TGU imeni G.R. Derzhavina, Tambov,
p. 112 (In Russian).
Everhart, M.J., 2005. Oceans of Kansas. Indiana University Press, Bloomington, p. 322.
Fadeeva, L.I., Iosifova, I.I., 1998. Geological map of pre-Quaternary deposits of the
Ryazan region. Scale 1:500,000. Mezhregionalnyi tsentr po geologicheskoi
kartografii, Moskva (in Russian).
Fischer, V., 2016. Taxonomy of Platypterygius campylodon and the diversity of the
last ichthyosaurs. PeerJ 4, e2604.
Fischer, V., Arkhangelsky, M.S., Naish, D., Stenshin, I.M., Uspensky, G.N., Godefroit, P.,
2014. Simbirskiasaurus and Pervushovisaurus reassessed: implications for the
taxonomy and cranial osteology of Cretaceous platypterygiine ichthyosaurs.
Zoological Journal of the Linnean Society 171, 822e841.
Fischer, V., Arkhangelsky, M.S., Stenshin, I.M., Uspensky, G.N., Zverkov, N.G.,
Benson, R.B.J., 2015. Peculiar macrophagous adaptations in a new Cretaceous
pliosaurid. Royal Society Open Science 2, 150552.
Fischer, V., Bardet, N., Benson, R.B.J., Arkhangelsky, M.S., Friedman, M., 2016.
Extinction of fish-shaped marine reptiles associated with reduced evolutionary
rates and global environmental volatility. Nature Communications 7, 1e11.
Fischer, V., Benson, R.B.J., Zverkov, N.G., Soul, L.C., Arkhangelsky, M.S., Lambert, O.,
Stenshin, I.M., Uspensky, G.N., Druckenmiller, P.S., 2017. Plasticity and
8
Cretaceous Research 127 (2021) 104946
1667e1676.
Fischer, V., Benson, R.B.J., Druckenmiller, P.S., Ketchum, H.F., Bardet, N., 2018. The
evolutionary history of polycotylid plesiosaurians. Royal Society Open Science
5, 172177.
Frey, E.D., Stinnesbeck, W., Martill, D.M., Rivera-Sylva, H.E., Múzquiz, H.P., 2020. The
geologically youngest remains of an ornithocheirid pterosaur from the late
Cenomanian (Late Cretaceous) of northeastern Mexico with implications on the
paleogeography and extinction of Late Cretaceous ornithocheirids. Palae-
overtebrata 43, 1e12.
Grigoriev, D.V., 2014. Giant Mosasaurus hoffmanni (Squamata, Mosasauridae) from
the Cretaceous (Maastrichtian) of Penza, Russia. Proceedings of the Zoological
Institute of the Royal Academy of Sciences 318, 148e167.
Grigoriev, D.V., Averianov, A.O., Arkhangelsky, M.S., Pervushov, E.M., Zozyrev, N.Yu,
2009. A mosasaur from the Cenomanian of Russia. Paleontologicheskii Zhurnal
3, 76e82.
Grigoriev, D.V., Arkhangelsky, M.S., Merkulov, S.M., 2015. A record of Clidastes
propython Cope (Squamata, Mosasauridae) in the Upper Cretaceous of the
Saratov Region, Russia. Paleontologicheskii Zhurnal 5, 60e68.
Holland, T., 2018. The mandible of Kronosaurus queenslandicus Longman, 1924
(Pliosauridae, Brachaucheniinae), from the Lower Cretaceous of Northwest
Queensland, Australia. Journal of Vertebrate Paleontology 38, e1511569.
Ifrim, C., Stinnesbeck, W., Frey, E., 2007. Upper Cretaceous (CenomanianeTuronian
and TuronianeConiacian) open marine plattenkalk deposits in NE Mexico.
Neues Jahrbuch fu€r Geologie und Pal€ aontologie-Abhandlungen 245, 71e81.
Jacobs, L.L., Mateus, O., Polcyn, M.J., Schulp, A.S., Antunes, M.T., Morais, M.L., da Silva
Tavares, T., 2006. The occurrence and geological setting of Cretaceous dinosaurs,
mosasaurs, plesiosaurs, and turtles from Angola. Paleontological Society of
Korea 22, 91e110.
Jimenez-Huidobro, P., Simo ~es, T.R., Caldwell, M.W., 2017. Mosasauroids from
Gondwanan continents. Journal of Herpetology 51, 355e364.
Kear, B.P., 2005. Marine reptiles from the Lower Cretaceous (Aptian) deposits of
White Cliffs, southeastern Australia: implications of a high-latitude cold water
assemblage. Cretaceous Research 26, 769e782.
Kear, B.P., 2016. Cretaceous marine amniotes of Australia: perspectives on a decade
of new research. Memoirs of Museum Victoria 74, 17e28.
Kear, B.P., Long, J.A., Martin, J.E., 2005. A review of Australian mosasaur occurrences.
Netherlands Journal of Geosciences 84, 307e313.
Kear, B.P., Ekrt, B., Prokop, J., Georgalis, G.L., 2014. Turonian marine amniotes from
the Bohemian Cretaceous Basin, Czech Republic. Geological Magazine 151,
183e198.
Kear, B.P., Larsson, D., Lindgren, J., Kundrat, M., 2017. Exceptionally prolonged tooth
formation in elasmosaurid plesiosaurians. PLos One 12, e0172759.
Kear, B.P., Fordyce, E., Hiller, N., Siversson, M., 2018. A palaeobiogeographical syn-
thesis of Australasian Mesozoic marine tetrapods. Alcheringa: An Australasian
Journal of Palaeontology 42, 461e486.
Kiprijanoff, W.A., 1881. Studien über die fossilen Reptilien Russlands. Theil 1, Gat-
tung Ichthyosaurus Ko €nig aus dem Severischen Sandstein oder Osteolith der
Kreidegruppe. Me moires de l'Acade mie impe riale des Sciences de St.-
Petersbourg, VIIe serie 28, 1e103.
Kiprijanoff, W.A., 1882. Studien über die fossilen Reptilien Russlands. 2. Theil.
Gattung Plesiosaurus Conybeare aus dem Severischen Sandstein oder Osteolith
der Kreidegruppe. Me moires de l'Acade mie impe riale des Sciences de St.-
Petersbourg, VIIe serie 30, 1e55.
Kiprijanoff, W.A., 1883. Studien uber die fossilen Reptilien Russlands. 3. Theil.
Gruppe Thaumatosauria n. aus der Kreide-Formation und dem Moskauer Jura.
Me moires de l'Acade mie impe riale des Sciences de St.-Pe
tersbourg, VIIe serie
31, 1e57.
Krivtsov, V.A., Vodorezov, A.V., Solonin, S.V., Tobratov, S.A., 2018. The potential of
creating and using protected areas of geological and geomorphological interest
in the Ryazan Region. Vestnik Ryazanskogo Gosudarstvennogo Universiteta
imeni S.A. Yesenina 3, 108e119 (in Russian).
Kuzmin, A.N., Kirikov, V.P., Lukianova, N.V., Maksimov, A.V., Kossovaia, O.L.,
Evdokimova, I.O., Gorbatsevich, N.R., Savanin, V.V., Kotliar, G.V., Samsonov, A.V.,
Ianovskii, A.S., Pugovkin, A.A., Engalychev, S.I., Latypov, R.K., Leontev, D.I.,
Petrov, B.V., Mazurkevich, K.N., 2015. State Geological Map of the Russian
Federation at a scale 1: 1,000,000. Sheet N-37 (Moscow). Kartograficheskaia
fabrika VSEGEI, SPb, p. 496 (in Russian).
Leidy, J., 1857. Remarks on Saurocephalus and its allies. Transactions of the American
Philosophical Society 11, 91e95.
Lingham-Soliar, T., 1994. The mosasaur “Angolasaurus” bocagei (Reptilia: Mosa-
sauridae) from the Turonian of Angola re-interpreted as the earliest member of
the genus Platecarpus. Pala €ontologische Zeitschrift 68, 267e282.
Longman, H.A., 1924. A new gigantic marine reptile from the Queensland Creta-
ceous. Memoirs of the Queensland Museum 8, 26e28.
Madzia, D., 2016. A reappraisal of Polyptychodon (Plesiosauria) from the Cretaceous
of England. PeerJ 4, e1998.
Madzia, D., Cau, A., 2017. Inferring ‘weak spots’ in phylogenetic trees: application to
mosasauroid nomenclature. PeerJ 5, e3782.
Madzia, D., Machalski, M., 2017. Isolated pliosaurid teeth from the
AlbianeCenomanian (Cretaceous) of Annopol, Poland. Acta Geologica Polonica
67, 393e403.
Madzia, D., Sachs, S., Lindgren, J., 2019. Morphological and phylogenetic aspects of
the dentition of Megacephalosaurus eulerti, a pliosaurid from the Turonian of
S.V. Solonin, A.V. Vodorezov and B.P. Kear
Kansas, USA, with remarks on the cranial anatomy of the taxon. Geological
Magazine 156, 1201e1216.
Mateus, O., Polcyn, M.J., Jacobs, L.L., Araújo, R., Schulp, A.S., Marinheiro, J., Pereira, B.,
Vineyard, D., 2012. Cretaceous amniotes from Angola: dinosaurs, pterosaurs,
mosasaurs, plesiosaurs, and turtles. V Jornadas Internacionales sobre Paleon-
tología de Dinosaurios y su Entorno 71e105.
Maxwell, E.E., Kear, B.P., 2010. Postcranial anatomy of Platypterygius americanus
(Reptilia: Ichthyosauria) from the Cretaceous of Wyoming. Journal of Vertebrate
Paleontology 30, 1059e1068.
Maxwell, E.E., Caldwell, M.W., Lamoureux, D.O., 2011. Tooth histology in the
Cretaceous ichthyosaur Platypterygius australis, and its significance for the
conservation and divergence of mineralized tooth tissues in amniotes. Journal
of Morphology 272, 129e135.
McGowan, C., Motani, R., 2003. Part 8. Ichthyopterygia. Verlag Dr. Friedrich Pfeil,
München, p. 173.
McHenry, C.R., 2009. ‘Devourer of Gods’. The Palaeoecology of the Cretaceous
Pliosaur Kronosaurus queenslandicus. Ph.D. thesis. University of Newcastle,
Newcastle, Australia, p. 616.
Milner, A.C., 1987. Reptiles. In: Owen, E., Smith, A.B. (Eds.), Fossils of the Chalk.
Palaeontological Association, London, pp. 266e280.
Nesov, L.A., Ivanov, A.O., Khozatsky, L.I., 1988. On the finds of the remains of ich-
thyosaurs in the USSR and the problem of fauna change in the middle Creta-
ceous. In: Vestnik Leningradskogo Universiteta, Seriia 7, Geologiia i Geografiia 1,
pp. 15e25 (in Russian).
Nikitin, S.N., Izrailev, V.M., Nazarova, T.I., Karaulova, E.A., 1984. Report on aerial
photo-geological mapping at a scale of 1: 200,000 of the Oka-Tsninsk swell
area. Moskovskaia Geologorazvedochnaia ekspeditsiia, Moskva, p. 334 (in
Russian).
Ochev, V.G., 1976. New plesiosaur from the Upper Cretaceous of the Penza Region.
Paleontologicheskii Zhurnal 2, 135e138 (in Russian).
O'Gorman, J.P., Gasparini, Z., 2013. Revision of Sulcusuchus erraini (Sauropterygia,
Polycotylidae) from the Upper Cretaceous of Patagonia, Argentina. Alcheringa:
An Australasian Journal of Palaeontology 37, 163e176.
O'Gorman, J.P., Varela, A.N., 2010. The oldest lower Upper Cretaceous plesiosaurs
(Reptilia, Sauropterygia) from southern Patagonia, Argentina. Ameghiniana 47,
447e459.
Olferev, A.G., Alekseev, A.S., 2005. Stratigraphic diagram of the Upper Cretaceous
deposits of the East European Platform. Explanatory memorandum. Paleon-
tologicheskii Institut RAN, Moskva, p. 204 (in Russian).
O'Keefe, F.R., Otero, R.A., Soto-Acun ~ a, S., O'Gorman, J.P., Godfrey, S.J., Chaterjee, S.,
2017. Cranial anatomy of Morturneria seymourensis from Antarctica, and the
evolution of filter feeding in plesiosaurs of the Austral Late Cretaceous. Journal
of Vertebrate Paleontology 37, e1347570.
} A., Szabo, M., Kollmann, H., Wagreich, M., Kalma
Osi, r, R., Maka di, L., Szentesi, Z.,
Summesberger, H., 2019. Vertebrate remains from the Turonian (Upper Creta-
ceous) Gosau Group of Gams, Austria. Cretaceous Research 99, 190e208.
Otero, R.A., Soto-Acun ~ a, S., O'Keefe, F.R., O'Gorman, J.P., Stinnesbeck, W.S.,
Suarez, M.E., Rubilar-Rogers, D., Salazar, C., Quinzio-Sinn, L.A., 2014. Aristonectes
quiriquinensis, sp. nov., a new highly derived elasmosaurid from the upper
Maastrichtian of central Chile. Journal of Vertebrate Paleontology 34, 100e125.
Palci, A., Caldwell, M.W., Papazzoni, C.A., 2013. A new genus and subfamily of
mosasaurs from the Upper Cretaceous of northern Italy. Journal of Vertebrate
Paleontology 33, 599e612.
ramo, M.E., 1994. Posicio
Pa  n Sistematica de un Reptil Marino con Base en los Restos
 siles Encontrados en Capas del Creta
Fo cico Superior en Yaguara  (Huila). Revista
de la Academia Colombiana de Ciencias Exactas, Físicas y Naturales 72, 63e80.
ramo-Fonseca, M.E., 2000. Yaguarasaurus columbianus (Reptilia, Mosasauridae), a
Pa
primitive mosasaur from the Turonian (Upper Cretaceous) of Colombia. His-
torical Biology 14, 121e131.
ramo-Fonseca, M.E., Benavides-Cabra, C.D., Gutie
Pa rrez, I.E., 2018. A new large
Pliosaurid from the Barremian (Lower Cretaceous) of Sa chica, Boyaca , Colombia.
Earth Sciences Research Journal 22, 223e238.
ramo-Fonseca, M.E., Benavides-Cabra, C.D., Gutie
Pa rrez, I.E., 2019. A new specimen
of Stenorhynchosaurus munozi P aramo-Fonseca et al., 2016 (Plesiosauria, Plio-
sauridae), from the Barremian of Colombia: new morphological features and
ontogenetic implications. Journal of Vertebrate Paleontology 39, e1663426.
Persson, P.O., 1963. A revision of the classification of the Plesiosauria with a syn-
opsis of the stratigraphical and geographical distribution of the group. Lunds
Universitets Årsskrift 59, 1e59.
Pervushov, E.M., Arkhangelsky, M.S., Ivanov, A.V., 1999. Catalog of the locations of
the remainders of sea reptiles in the Jurassic and Cretaceous of the Lower Volga
Region. Saratov University, Saratov.
Polcyn, M.J., Bell Jr., G.L., 2005. Russellosaurus coheni n. gen., n. sp., a 92 million-year-
old mosasaur from Texas (USA), and the definition of the parafamily Russello-
saurina. Netherlands Journal of Geosciences 84, 321e333.
Polcyn, M.J., Bell Jr., G.L., Shimada, K., Everhart, M.J., 2008. The oldest North
American mosasaurs (Squamata: Mosasauridae) from the Turonian (Upper
Cretaceous) of Kansas and Texas with comments on the radiations of major
mosasaur clades. In: Everhart, M.J. (Ed.), Proceedings of the Second Mosasaur
Meeting, Fort Hays Studies. Fort Hays State University, Hays, Kansas,
pp. 137e155.
Pravoslavlev, P.A., 1915. Restes d’un jeune Plesiosaurus trouve s dans le cre tace
 from the Upper Cretaceous Niobrara Formation of central North America, 11.
superieur du basin de la rivie re Liski, province du Don. Annuaire Ge ologique et
Mine ralogique de la Russie 17, 1e18.
9
Cretaceous Research 127 (2021) 104946
Romano, M., Manni, R., Venditti, E., Nicosia, U., Cipriani, A., 2019. First occurrence of
a tylosaurine mosasaur from the Turonian of the Central Apennines, Italy.
Cretaceous Research 96, 196e209.
Romer, A.S., 1968. An ichthyosaur skull from the Cretaceous of Wyoming. Contri-
butions to Geology 7, 27e41. Wyoming University.
Rozhdestvensky, A.K., 1973. Study of Cretaceous reptiles in Russia. Paleon-
tologicheskii Zhurnal 2, 90e99 (in Russian).
Sachs, S., 2000. Ein Pliosauride (Sauropterygia: Plesiosauria) aus der Oberkreide von
Anro €chte in Westfalen. Geologie und Pala €ontologie in Westfalen 56, 25e33.
Sachs, S., Kear, B.P., 2015. Postcranium of the paradigm elasmosaurid plesiosaurian
Libonectes morgani (Welles, 1949). Geological Magazine 152, 694e710.
Sachs, S., Kear, B.P., 2017. Redescription of the elasmosaurid plesiosaurian Libonectes
atlasense from the Upper Cretaceous of Morocco. Cretaceous Research 74,
205e222.
Sachs, S., Hornung, J.J., Knüppe, J., Wilmsen, M., Kear, B.P., 2016. 15. Reptilien.
Geologica Saxonica 62, 169e179.
Sachs, S., Wilmsen, M., Knüppe, J., Hornung, J.J., Kear, B.P., 2017. Cenomanian-
Turonian marine amniote remains from the Saxonian Cretaceous Basin of
Germany. Geological Magazine 154, 237e246.
Sachs, S., Jagt, J., Nied zwiedzki, R., Ke˛ dzierski, M., Yazykova, E., Kear, B.P., 2018.
Turonian marine amniotes from the Opole area in southwest Poland. Creta-
ceous Research 84, 578e587.
Sachs, S., Lindgren, J., Madzia, D., Kear, B.P., 2021. Cranial osteology of the mid-
Cretaceous elasmosaurid Thalassomedon haningtoni from the Western Interior
Seaway of North America. Cretaceous Research. https://doi.org/10.1016/
j.cretres.2021.104769.
Sato, T., 2005. A new polycotylid plesiosaur (Reptilia: Sauropterygia) from the
Upper Cretaceous Bearpaw Formation in Saskatchewan, Canada. Journal of
Paleontology 79, 969e980.
Sato, T., Konishi, T., Hirayama, R., Caldwell, M.W., 2012. A review of the Upper
Cretaceous marine reptiles from Japan. Cretaceous Research 37, 319e340.
Schmeisser McKean, R., 2012. A new species of polycotylid plesiosaur (Reptilia:
Sauropterygia) from the Lower Turonian of Utah: extending the stratigraphic
range of Dolichorhynchops. Cretaceous Research 34, 184e199.
Schneider, C.A., Rasband, W.S., Eliceiri, K.W., 2012. NIH Image to ImageJ: 25 years of
image analysis. Nature Methods 9, 671e675.
Schroeder, H., 1884. Saurierreste aus der baltischen oberen Kreide. Jahrbuch der
Preussischen Geologischen Landesanstalt 1884, 293e333.
Schumacher, B.A., 2011. A ‘woollgari-Zone mosasaur’ (Squamata; Mosasauridae)
from the Carlile Shale (lower middle Turonian) of central Kansas and the
stratigraphic overlap of early mosasaurs and pliosaurid plesiosaurs. Trans-
actions of the Kansas Academy of Science 114, 1e14.
Schumacher, B.A., Carpenter, K., Everhart, M.J., 2013. A new Cretaceous pliosaurid
(Reptilia, Plesiosauria) from the Carlile Shale (middle Turonian) of Russell
County, Kansas. Journal of Vertebrate Paleontology 33, 613e628.
Shimada, K., 2012. Dentition of Late Cretaceous shark, Ptychodus mortoni (Elasmo-
branchii, Ptychodontidae). Journal of Vertebrate Paleontology 32, 1271e1284.
Shimada, K., Schumacher, B.A., Parkin, J.A., Palermo, J.M., 2006. Fossil marine ver-
tebrates from the lowermost Greenhorn Limestone (Upper Cretaceous: middle
Cenomanian) in southeastern Colorado. Journal of Paleontology 80, 1e45.
Shimada, K., Popov, E.V., Siversson, M., Welton, B.J., Long, D.J., 2015. A new clade of
putative plankton-feeding sharks from the Upper Cretaceous of Russia and the
United States. Journal of Vertebrate Paleontology 35, e981335.
Sidorenko, A.V.,1971. Geology of the USSR. Volume IV. Center of the European part of the
USSR. Geological description. Izdatelstvo Nedra, Moskva, p. 742 (in Russian).
Sintsov, I.F., 1872. Jurassic and Cretaceous fossils of the Saratov Province. Volume 4.
Materials for the geology of Russia. Tipografiia Imperatorskoi Akademii Nauk,
St. Petersburg, p. 127 (in Russian).
Siverson, M., 1996. Lamniform sharks of the mid-Cretaceous Alinga Formation and
Beedagong Claystone, Western Australia. Palaeontology 39, 813e849.
Siverson, M., 1999. A new large lamniform shark from the uppermost Gearle Silt-
stone (Cenomanian, Late Cretaceous) of Western Australia. Transactions of the
Royal Society of Edinburgh Earth Sciences 90, 49e66.
Siverson, M., Lindgren, J., 2005. Late Cretaceous sharks Cretoxyrhina and Carda-
biodon from Montana, USA. Acta Palaeontologica Polonica 50, 301e314.
Siversson, M., Lindgren, J., Newbrey, M.G., Cederstro € m, P., Cook, T.D., 2015. Cen-
omanianeCampanian (Late Cretaceous) mid-palaeolatitude sharks of Creta-
lamna appendiculata type. Acta Palaeontologica Polonica 60, 339e384.
Solonin, S.V., Martill, D.M., Smith, R.E., Vodorezov, A.V., 2021. First occurrence of
ornithocheirid pterosaur teeth in the Dmitrov Formation (Santonian) of Ryazan
Oblast, Russia. Cretaceous Research. https://doi.org/10.1016/
j.cretres.2021.104943.
Solonin, S.V., Shell, R., Vodorezov, A.V., Nied zwiedzki, R., 2020. Preliminary report of
an Upper Cretaceous elasmobranch fauna from Ryazan Oblast, Russia, 52.
Geological Society of America Abstracts with Programs, p. 6.
Starodubtseva, I.A., Sennikov, A.G., Soroka, I.L., 2008. Geological History of the
Moscow Region in the Collections of the Natural Museums of the Russian
Academy of Sciences. In: Gosudarstvenny Geologicheskiy Muzey imeni V.I.
Vernandskogo RAN. Paleontologichesky Institut RAN, Izdatelskii Tsentr Nauka,
Moskva, p. 229 (In Russian).
Storrs, G.W., 1999. An examination of the Plesiosauria (Diapsida; Sauropterygia)
University of Kansas Paleontological Contributions, pp. 1e15 (New Series).
Storrs, G.W., Arkhangelsky, M.S., Efimov, F.V., 2000. Mesozoic marine reptiles of
Russia and other former Soviet republics. In: Benton, M.J., Shishkin, M.A.,
S.V. Solonin, A.V. Vodorezov and B.P. Kear
Unwin, D.M., Kurochkin, E.N. (Eds.), The Age of Dinosaurs in Russia and
Mongolia. Cambridge University Press, Cambridge, pp. 187e210.
Street, H.P., Caldwell, M.W., 2014. Reassessment of Turonian mosasaur material
from the ‘Middle Chalk’ (England, UK), and the status of Mosasaurus gracilis
Owen, 1849. Journal of Vertebrate Paleontology 34, 1072e1079.
Tsaregradsky, V.A., 1926. Remains of mosasaurs from Saratov Province. Izvestiia
Geologicheskogo Komiteta 45, 563e572 (in Russian).
Underwood, C.J., Goswami, A., Prasad, G.V.R., Verma, O., Flynn, J.J., 2011. Marine
vertebrates from the ‘middle’ Cretaceous (early Cenomanian) of South India.
Journal of Vertebrate Paleontology 31, 539e552.
Van Vranken, N.E., 2018. An overview of ichthyosaurian remains from the Creta-
ceous of Texas, USA. Alcheringa: An Australasian Journal of Palaeontology 42,
509e515.
Vinogradov, A.P., 1975. Paleogeography of the USSR. In: Explanatory memorandum
to the Atlas of lithological and paleogeographic maps of the USSR (in four
volumes), vol. 3. Nedra, Moskva, p. 200 (in Russian).
VonLoh, J.P., Bell Jr., G.L., 1998. Fossil reptiles from the Late Cretaceous Greenhorn
Formation (late Cenomanianemiddle Turonian) of the Black Hills region, South
Dakota. Dakoterra 5, 28e38.
Welles, S.P., 1943. Elasmosaurid plesiosaurs with a description of new material from
California and Colorado, vol. 13. University of California Publications in
Geological Sciences, pp. 125e215.
Welles, S.P., 1949. A new elasmosaur from the Eagle Ford Shale of Texas. Part I.
Systematic description. Fondren Science Series 1, 1e28.
Welles, S.P., 1962. A new species of elasmosaur from the Aptian of Colombia, and a
review of the Cretaceous plesiosaurs, vol. 44. University of California Publica-
tions in Geological Sciences, pp. 1e96.
10
View publication stats
Cretaceous Research 127 (2021) 104946
Williston, S., 1903. North American plesiosaurs, Part 1. Field Colombian Museum
Publication (Geology) 73, 1e77.
Yakovlev, N.N., 1901. Remains of mosasaurs from the Upper Cretaceous deposits of
Southern Russia. Izvestiia Geologicheskogo Komiteta 20, 507e518 (in Russian).
Yarkov, A.A., 1989. Poluninskie finds of marine reptiles. Local history notes, vol. 6.
Nizhne-Volzhskoe Knizhnoe Izdatelstvo, pp. 207e216 (in Russian).
Yarkov, A.A., 1993. History of the study of mosasaurs in Russia and some comments
on their taxonomy. Izdatelstvo Saratovskogo Universiteta 8, 26e40 (in Russian).
Zverkov, N.G., 2015. On a typically Late Jurassic pliosaur from the Lower Cretaceous
of Crimea. In: Baraboshkin, E.Y., Bykov, D.E. (Eds.), The International Scientific
Conference on the Jurassic/Cretaceous Boundary. September 7e13, 2015.
Togliatti, Kassandra, Sarama, Russia, pp. 89e94.
Zverkov, N.G., Pervushov, E.M., 2020. A gigantic pliosaurid from the Cenomanian
(Upper Cretaceous) of the Volga Region, Russia. Cretaceous Research 110,
104419.
Zverkov, N.G., Averianov, A.O., Popov, E.V., 2018a. Basicranium of an elasmosaurid
plesiosaur from the Campanian of European Russia. Alcheringa: An Australasian
Journal of Palaeontology 42, 528e542.
Zverkov, N.G., Fischer, V., Madzia, D., Benson, R.B.J., 2018b. Increased pliosaurid
dental disparity across the JurassiceCretaceous transition. Palaeontology 61,
825e846.
Appendix A. Supplementary data
Supplementary data to this article can be found online at https://doi.org/10.
1016/j.cretres.2021.104946.