Turanosuchus aralensis, Kuzmin & Skutschas & Boitsova & Sues, 2019

Kuzmin, Ivan T, Skutschas, Pavel P, Boitsova, Elizaveta A & Sues, Hans-Dieter, 2019, Revision of the large crocodyliform Kansajsuchus (Neosuchia) from the Late Cretaceous of Central Asia, Zoological Journal of the Linnean Society 185 (2), pp. 335-387 : 361-368

KANSAJSUCHUS SP. CF. K. EXTENSUS

1988a Turanosuchus aralensis gen. et sp. nov. – Efimov

1988b Turanosuchus aralensis – Efimov

2000 Turanosuchus aralensis (Efimov) – Storrs & Efimov

2015 Neosuchia (? Goniopholididae ) indet. – Halliday et al.

2015 Kansajsuchus extensus ( Efimov, 1975) – Skutschas et al.

Referred material: Isolated cranial and postcranial fragments, teeth, osteoderms ( PIN collection number 2229, ZIN PH collection number 180; see Supporting Information, File S1 for the detailed list of specimens). Locality: Shakh Shakh locality, north-eastern Aral Sea Region, Kyzylorda Province, Kazakhstan; Bostobe Formation, Santonian–early Campanian.

Description

Cranium: Premaxilla ( Fig. 16A, D, E ): The premaxilla is represented by two partial specimens ( Fig. 16A, D, E ). Its posterodorsal process forms a sharp wedge posteriorly and contacts the nasal medially and the maxilla laterally. The suture with the nasal is oblique and oriented anteromedially.

Maxilla ( Fig. 16A–C ): The structure of the maxilla is partially documented by a number of specimens; PIN 2229/502 (a fragmentary anterior rostrum, right side) is the most anatomically informative specimen ( Fig. 16A–C ). Proportionally, the maxilla is relatively narrow lateromedially, indicating an elongated and narrow rostrum (also noted by Halliday et al., 2015). It contacts the premaxilla anteriorly and the nasal medially along a straight suture. The anterior maxillary alveoli gradually increase in size posteriorly, with, presumably, the fourth and the fifth being the largest ( Fig. 16B, C ). The maxillary alveoli are closely spaced. In dorsal view, the maxilla is clearly festooned in its anterior part. The lateral projection of the maxillary margin corresponds to the position of enlarged alveoli. In lateral view, the maxilla has a slightly sinuous shape; its anteriormost tip is curved upwards. The dorsal surface of the maxilla has pitted ornamentation; its alveolar margin is smooth.

Nasal ( Fig. 16A–G ): The nasal is mediolaterally narrow, anteroposteriorly elongated and has straight, sub-parallel lateral and medial margins ( Fig. 16A, D–G ). It is dorsoventrally flattened. The nasal tapers considerably at its contact with the premaxilla ( Fig. 16A, D ). The dorsal surface is ornamented with pits and grooves.

Jugal ( Fig. 16H–N ): The jugal is represented by five fragmentary specimens, which correspond to the anterior process ( Fig. 16H–J ) or the mid-section of the jugal near the base of the postorbital bar ( Fig. 16K–N ). The anterior process is dorsoventrally deep, mediolaterally flattened and plate-like. The medial surface of the process bears a facet for contact with the lacrimal anterodorsally ( Fig. 16I ). Posterior to this facet, the jugal forms the ventral orbital margin. The latter is ridge-like, with a shallow depression on the lateral surface of the jugal ventral to this ridge ( Fig. 16J ). The postorbital bar is inset from the lateral margin of the jugal. The base of the ascending process is rounded in cross-section ( Fig. 16N ). The jugal has a hollow recess inside at the base of the ascending process. This recess is connected to several openings on the external surface of the jugal (multiple foramina on the medial surface of the anterior process and a single opening posterior to the base of the ascending process; Fig. 16I, L, N ). The jugal bears a facet for contact with the ectopterygoid on its medial surface ( Fig. 16I, L ). The lateral surface of the jugal is covered with a pitted ornamentation and bears a longitudinal ridge ( Fig. 16H, M ). On the ventral edge of the anterior process, there is a series of four to six neurovascular foramina.

Ectopterygoid ( Fig. 16K ): The structure of the ectopterygoid is incompletely known from a couple of specimens( ZINPH 5/180,fragmentofaleftectopterygoid; PIN 2229/508, partial left ectopterygoid preserved in an anatomical association with a fragment of the jugal). Additional information on the ectopterygoid can be inferred from the articular facet on the jugal ( Fig. 16I, L ). The ectopterygoid is posteroventrally deflected as in other neosuchians. Its anterior process is elongated and contacts the maxilla anterolaterally and the jugal posterolaterally ( Fig. 16I ). The posterior process of the ectopterygoid contacts the jugal at the base of the postorbital bar ( Fig. 16K ). The ascending process does not contribute significantly to the postorbital bar. The contact with the descending process of the postorbital is present on the medial surface of the postorbital bar but is not extensive ( Fig. 16K ).

Frontal ( Fig. 17A–E ): The unpaired frontal is represented by three partial specimens of different sizes, which preserve the posterior interorbital and skull table sections ( Fig. 17A–E ). Its anterior process is relatively wide mediolaterally (even in the smallest specimen, ZIN PH 27/180; Fig. 17D ) and only slightly narrower between orbits than the skull table portion of the frontal. The frontal forms the anteromedial corners of the supratemporal fenestrae. The lateral margins of the anterior process preserve the facet for the prefrontals. The degree of the frontal participation in the formation of the medial orbital margin differs among the available specimens. The smallest specimen ( ZIN PH 27/180; Fig. 17D, E ) shows considerable frontal participation in the medial orbital margin, comparable to that in Paralligator spp. The orbital margins are ridge-like and distinctly upturned ( Fig. 17E ). In the largest specimen, ZIN PH 26/180 ( Fig. 17A, B ), the frontal forms only the small posteromedial corners of the orbits (bordered by facets for the postorbital and prefrontal). A medium-sized specimen ( PIN 2229/510; Fig. 17C ) shows an intermediate degree of frontal participation. As ontogenetic and intraspecific variability of this feature has been observed (see comparison below), we consider the difference among specimens to be a size- and, possibly, age-related feature. The frontal bears a pronounced sagittal ridge on the dorsal surface of its anterior process. The dorsal surface of the frontal is densely pitted.

Parietal ( Fig. 17I–K ): The unpaired parietal forms the interfenestral bar between the supratemporal fenestrae, which were likely moderately enlarged, as suggested by the sizes of corresponding notches on the lateral margins of the parietal ( Fig. 17I ). The parietal contacts the frontal anteriorly along a transversely straight suture and the supraoccipital ventrally. Facets for contact with the laterosphenoid and prootic are present on the ventral surface of the parietal ( Fig. 17J ). The dorsal surface has pitted ornamentation. A slight sagittal ridge is present on the dorsal surface but it is damaged. The parietal pneumatic recess is apparently present and connected to the intertympanic recess in the supraoccipital by two pairs of foramina – larger anterolateral, situated near the parietal-prootic-supraoccipital contact, and smaller slit-like posteromedial foramina ( Fig. 17J ).

Squamosal ( Fig. 17F–H ): The squamosal is represented by a partial anterior process of the right element, articulated with a fragment of the postorbital ( ZIN PH 36/180). The dorsoventrally deep lateral lamina of the anterior process projects anteriorly and overlaps the postorbital laterally ( Fig. 17H ). Ventrally, there is a facet for contact with the quadrate ( Fig. 17F ). The dorsal surface of the anterior process is densely ornamented, whereas the lateral surface is smooth, with a groove for the muscles associated with an external ear flap ( Fig. 17G, H ).

Supraoccipital ( Fig. 17I–K ): The supraoccipital is represented by a single specimen ZIN PH 28/180, which lacks the ventral portion of the element. It contacts the parietal dorsally; the contacts with the prootic anteroventrally and the otoccipitals posteroventrally are not preserved. The supraoccipital apparently extends on to the skull roof ( Fig. 17I, K ); the suture is clearly seen in posterior view, but is barely traceable on the dorsal surface of the specimen. The occipital (posterior) surface of the supraoccipital is strongly compressed, but was apparently vertical and lacked a prominent sagittal ridge. The supraoccipital contains the large intertympanic recess ( Fig. 17J ).

Mandible: Dentary ( Fig. 18A–G ): The structure of the dentary is known from a number of specimens: PIN 2229 /507 (the mandibular symphysis, originally designated as the holotype of ‘ Turanosuchus aralensis ’; Fig.18A–C ), PIN 2229 /506 and PIN 2229 /515 (two partial specimens, corresponding to the mid-posterior portion of the dentary; Fig. 18D–G ). The mandibular symphysis is anteroposteriorly elongated, lateromedially narrow and relatively flat. The dentary symphysis extends posteriorly to the level of the seventh or eighth alveolus ( Fig. 18A ). The anterior alveoli gradually increase in size up to the fourth, which is the largest. This alveolus is distinct in both dorsal and lateral views ( Fig. 18A, B ). The lateral margin of the dentary is festooned in dorsal view, corresponding to changes in alveolar size. The alveolar arrangement is linear, as there are no two neighbouring alveoli positioned in the same transverse plane. The alveoli are closely spaced, with small interalveolar distances posterior to the fourth alveolus. Other fragmentary specimens ( PIN 2229 /506 and PIN 2229 /515) also show this feature. They are dorsoventrally deeper than PIN 2229 /507. This difference is considered to reflect changes in the depth of the dentary along the mandibular ramus, rather than a characteristic feature of a different neosuchian taxon (contra Halliday et al., 2015) .

Splenial ( Fig. 18H, I ): The splenial is a dorsoventrally deep and lateromedially thin bone. It participates in the mandibular symphysis, as is evidenced by a small splenial fragment in PIN 2229/507 ( Fig. 18C ; thin lamina of bone attached to the medial surface of a left dentary) and by a corresponding facet on the medial surface of ZIN PH 32/180 (anterior fragment of a right splenial; Fig. 18H ). The dorsal margin of the splenial is slightly convex and smooth; its ventral margin is sharp and ridge-like. There is an anteroposteriorly extending facet for contact with the dentary on the lateral surface of the splenial, divided by the centrally located Meckelian groove ( Fig. 18I ).

Angular ( Fig. 18M–O ): The angular smoothly arches posterodorsally. Its anteroventral process preserves facets for contact with the dentary both medially and laterally ( Fig. 18M, O ). Most of the laterodorsal margin, which corresponds to the contacts with the dentary and surangular, respectively, is damaged. The angular forms the floor of the adductor cavity; posterior to it, the facet for the contact with the articular is present ( Fig. 18N ). The lateral surface of the angular is covered by pits. A prominent, obliquely longitudinal ridge extends posteroventrally ( Fig. 18O ).

Surangular ( Fig. 18J–L ): A fragment of the surangular is present in anatomical articulation with the articular ( PIN 2229/504). The surangular forms the lateral margin of the glenoid fossa ( Fig. 18J ). A facet for contact with the angular is present on the ventral margin of the surangular ( Fig. 18L ). The surangular extends posteriorly to the base of the retroarticular process. Its lateral surface has pitted ornamentation.

Articular ( Fig. 18J–L ): The structure of the articular is partially documented by a single specimen ( PIN 2229/504). The glenoid facet of the craniomandibular joint is mostly formed by the articular, with the surangular participating only in its lateral corner. A prominent transverse ridge bounds the glenoid posteriorly ( Fig. 18J, L ). Only the base of the retroarticular process is preserved. Considering the shape of the preserved portion, the retroarticular process was relatively horizontal and not as curved posterodorsally as in extant crocodylians.

Dentition: All available teeth show similar crown structure: they are coarsely striated and have prominent distal and mesial keels (bicarinate condition sensu Storrs & Efimov, 2000; double-ridged pattern sensu Halliday et al., 2015). The teeth vary in shape from tall and conical to blunt and short ( Fig. 17L, M ). The larger teeth have robust crowns comparable to those of K. extensus (see above). Most of the teeth have rounded bases; however, lateromedially flattened teeth (corresponding to the posterior cheek region) are also present ( Fig. 17M ). The latter morphotype has a finer pattern of striae on the crown surface.

Postcranial skeleton: Vertebrae ( Fig. 19A–H ): The structure of the dorsal vertebrae is documented by a number of specimens ( Fig. 19A–H ; see also Supporting Information, File S1). All recovered vertebrae have amphicoelous centra with slightly concave to almost flat posterior articular surfaces. The centrum is anteroposteriorly longer than dorsoventrally deep. The pedicles of the neural arches are dorsoventrally low and anteroposteriorly elongated ( Fig. 19C, F ). The neural spine base is anteroposteriorly elongated and extends for most of the dorsal surface of the neural arch ( Fig. 19D ). ZIN PH 11/180 ( Fig. 19G, H ) is dorsoventrally slightly compressed compared to other recovered dorsals. It has paired articular surfaces posterolaterally. The shape of its centrum and the presence of these additional articular surfaces suggest that ZIN PH 11/180 is probably a posteriormost dorsal vertebra, immediately anterior to the first sacral.

Osteoderms ( Fig. 19I – O ): A large number of osteoderms and osteoderm fragments of relatively large size was recovered. The available material is similar in its structure to that described in detail for K. extensus . Two morphotypes are represented: keeled paravertebral dorsal osteoderms ( Fig. 19I– N ) and unkeeled polygonal ventral osteoderms ( Fig. 19O ). The paravertebral dorsal elements are mostly rectangular, with keels restricted to the posterior margins of the osteoderms. Paravertebral osteoderms lack the anterolateral processes but have lateromedially extensive, unornamented anterior processes (for imbrication with adjacent elements in a longitudinal row). The dorsal osteoderms were arranged in at least four longitudinal rows, as indicated by the presence of paired articular facets on lateral and medial margins of several elements (e.g. ZIN PH 15/180; Fig. 19K, L ). ZIN PH 16/180 is strongly curved dorsally ( Fig. 19K ). Most likely this specimen reflects the structure of dorsal osteoderms from a particular shield area, which is not otherwise sampled in the material from the Kansai locality. The ventral osteoderms were sutured together and formed the ventral shield, as indicated by their straight margins with sutural facets ( Fig. 19O ). All recovered osteoderms are similarly densely pitted on their external surfaces.

Figures

Figure 1. Premaxillae of Kansajsuchus extensus from the Yalovach Formation (Santonian), Tadzhikistan. PIN 2399/301 (holotype), isolated right premaxilla in dorsal (A), ventral (B), lateral (C), and anterodorsal (D), views. PIN 2399/303, referred left partial premaxilla in dorsal (E), and ventral (F), views. Abbreviations: adp, anterodorsal process; dp, dorsal process; if, incisive foramen; mf, maxillary facet; nf, nasal facet; no, narial opening; nvf, neurovascular foramen; pdp, posterodorsal process; pmf, premaxillary facet; pvc, postvestibular canal; vsf, ventral subnarial foramen.Arabic numerals correspond to numbers of premaxillary alveoli, Roman numerals correspond to numbers of occlusal pits for dentary teeth. Scale bar = 1 cm.

Figure 2. Reconstruction of the skull of Kansajsuchus extensus from the Yalovach Formation (Santonian), Tadzhikistan, in dorsal view. Shorter dashes and schematic bone ornamentation correspond to the extent of actual preserved specimens. Solid lines and longer dashes correspond to the preserved and unpreserved anatomical borders. Grey areas represent overall sampled parts of the skull. The bone surface texture comprises subcircular or elongated pits (empty circles), slight ridges or wrinkles (curved lines), unornamented areas (dots) and major ridges on the frontal, prefrontal, jugal, quadrate and squamosal (solid black lines). Note neurovascular foramina (solid black dots) and unornamented alveolar margins of the premaxilla and the maxilla. Scale bar = 10 cm.

Figure 3. Referred maxillary remains of Kansajsuchus extensus from the Yalovach Formation (Santonian), Tadzhikistan. PIN 2399/307, anterior fragment of the right maxilla in ventral (A), and dorsal (B), views. PIN 2399/443, anteriormost fragment of the right maxilla in dorsal (C), and ventral (D), views. PIN 2399/451, posteriormost fragment of the right maxilla in laterodorsal (E), and lateral (F), views. PIN 2399/445, mid-section maxillary fragment in cross-sectional (G), and dorsal (H), views. Abbreviations: jf, jugal facet; nf, nasal facet; nvc, neurovascular canal; nvf, neurovascular foramen; pmf, premaxillary facet. Arabic numerals correspond to numbers of maxillary alveoli. Scale bar = 1 cm.

Figure 4. Referred cranial specimens of Kansajsuchus extensus from the Yalovach Formation (Santonian), Tadzhikistan. PIN 2399/446, posterior fragment of the left nasal with a lacrimal piece in association, in dorsal (A), and ventral (B), views. PIN 2399/462, mid-section nasal fragment in medial view (C). PIN 2399/306, posterior fragment of the right nasal in dorsal (D), and ventral (E), views. Abbreviations: frf, frontal facet; lac, lacrimal; lacf, lacrimal facet; mf, maxillary facet; nf, nasal facet; pfrf, prefrontal facet. Scale bar = 1 cm.

Figure 5. Referred circumorbital elements of Kansajsuchus extensus from the Yalovach Formation (Santonian), Tadzhikistan. PIN 2399/349, left prefrontal and anterior process of frontal in dorsal (A), ventral (B), and left lateral (C), views. PIN 2399/355, partial right prefrontal in ventral (D), dorsal (E), medial (F), and lateral (G), views. PIN 2399/310, an almost complete frontal in dorsal (H), ventral (I), oblique antero-dorso-lateral (J), and right lateral (K), views. Abbreviations: ccf, cristae cranii frontales; d, depression; fr, frontal; frf, frontal facet; g, groove; lacf, lacrimal facet; n, nasal; nf, nasal facet; olfd, depression, corresponding to the olfactory region of the nasal cavity; orb, orbit; pfr, prefrontal; pfrc, prefrontal crest; pfrf, prefrontal facet; pfrp, prefrontal pillar; pof, postorbital facet; sgc, sagittal crest; spef, sphenethmoid facet; tic, transverse interorbital crest; I, groove corresponding to the olfactory tract. Scale bar = 1 cm.

Figure 6. Referred cranial specimens of Kansajsuchus extensus from the Yalovach Formation (Santonian), Tadzhikistan. PIN 2399/308, articulated posterior cranial portion (dorsal piece) in dorsal (A, D), and right lateral (E), views, with details of the right supratemporal fossa (B, C). PIN 2399/449, isolated posterolateral process of the right squamosal in lateral view (F). PIN 2399/311, partial left squamosal in posterior (G), and dorsal (H), views. Abbreviations: alp, anterolateral process; antp, anterior process; CrA’, crest A’; dsp, descending process; elg, earlid groove; fll, flared lobule; fr, frontal; ls, laterosphenoid; lsq, lateral squamosal lamina; medp, medial process; oi, otic incisure; p, parietal; pf, parietal facet; plp, posterolateral process; po, postorbital; pro, prootic; q, quadrate; qjf, quadratojugal facet; sf, subtympanic foramen; so, supraoccipital; sq, squamosal; stfe, supratemporal fenestra; stfo, supratemporal fossa. Scale bar = 1 cm; B and C not to scale.

Figure 7. Referred jugal remains of Kansajsuchus extensus from the Yalovach Formation (Santonian), Tadzhikistan. PIN 2399/322, almost complete left jugal of subadult individual in lateral (A), and medial (B), views. PIN 2399/316, mid-section of the right jugal in lateral (C), and medial (D), views. PIN 2399/452, partial anterior process of the right jugal in lateral view (E). PIN 2399/325, posterior process of the left jugal in lateral (F), and medial (G), views.Abbreviations: asp, ascending process; ectf, ectopterygoid facet; lacf, lacrimal facet; latr, lateral ridge; orbr, orbital ridge; qjf, quadratojugal facet. Scale bar = 1 cm.

Figure 8. Details of the braincase region of Kansajsuchus extensus from the Yalovach Formation (Santonian), Tadzhikistan. PIN 2399/308, articulated posterior cranial portion (dorsal piece) in ventral view (A, B). PIN 2399/466, posterior portion of the right quadrate of subadult individual in posterior (C), ventral (D), and dorsal (E), views. Abbreviations: adp, anterodorsal process; CrA, crest A; CrA’, crest A’; dsp, descending process; ecc, endocranial cavity; fae, foramen aereum; fr, frontal; latc, lateral condyle; ls, laterosphenoid; lsb, laterosphenoid bridge; lsbt, laterosphenoid buttress; lscp, laterosphenoid capitate process; medc, medial condyle; p, parietal; po, postorbital; pro, prootic; q, quadrate; qjf, quadratojugal facet; r, ridge; so, supraoccipital; tgf, trigeminal fossa; II, notch corresponding to the optic cranial nerve passage; IV, foramen for the trochlear nerve; VI–V tymp, passages of various trigeminal nerve branches. Scale bar = 1 cm.

Figure 9. Details of the braincase region of Kansajsuchus extensus from the Yalovach Formation (Santonian), Tadzhikistan. PIN 2399/308, articulated posterior cranial portion (dorsal piece) in posterior view (A, E), with selected details of the right prootic region in postero-ventro-medial (B), posterior (C), and dorsolateral (D), views. Black rod is inserted into the anterior semicircular canal and the ampullary recess of the right inner ear capsule. Blue rod passes through the canal corresponding to the passage of the facial (VII) nerve. Red rod is inserted into the canal corresponding to the passage of the tympanic branch of the trigeminal nerve (Vtymp). White arrows reflect communications of the prootic pneumatic recess (housed inside the prootic) with surrounding pneumatic recesses. Abbreviations: aar, anterior ampullary recess; asc, anterior semicircular canal; cf, foramen corresponding to the communication between two pneumatic recesses; ecc, endocranial cavity; itr, intertympanic pneumatic recess; ls, laterosphenoid; lsb, laterosphenoid bridge; oi, otic incisure; p, parietal; phr, pharyngotympanic pneumatic recess; po, postorbital; pro, prootic; ProR, prootic pneumatic recess; q, quadrate; qr, quadrate pneumatic recess; so, supraoccipital; sq, squamosal; stfo, supratemporal fossa; tgf, trigeminal fossa; VI, passage of the ophthalmic branch of the trigeminal nerve; Vtymp, passage of the tympanic branch of the trigeminal nerve; VII, facial nerve passage. Scale bar = 1 cm; B–D not to scale.

Figure 16. Referred cranial specimens of Kansajsuchus sp. cf. K. extensus from the Bostobe Formation (Santonian- Campanian), Kazakhstan. PIN 2229/502, articulated fragmentary rostrum in dorsal (A), right lateral (B), and ventral (C), views. ZIN PH 3/180, left premaxillary dorsal process and anterior nasal fragment in dorsal (D), and ventral (E), views. ZIN PH 2/180, posterior nasal fragment in ventral (F), and dorsal (G), views. ZIN PH 4/180, anterior process of the right jugal in lateral (H), medial (I), and oblique postero-dorso-lateral (J), views. PIN 2229/508, mid-section of the left jugal with ectopterygoid fragment in association, in medial view (K). ZIN PH 30/180, mid-section of the left jugal in medial (L), lateral (M), and dorsal (N), views. Abbreviations: asp, ascending process; ect, ectopterygoid; ectf, ectopterygoid facet; fo, foramen; j, jugal; lacf, lacrimal facet; latr, lateral ridge; m, maxilla; mf, maxillary facet; n, nasal; nf, nasal facet; orb, orbit; orbr, orbital ridge; pm, premaxilla; pof, postorbital facet. Arabic numerals correspond to numbers of maxillary alveoli. Scale bar = 1 cm.

Figure 17. Referred cranial specimens and teeth of Kansajsuchus sp. cf.K.extensus from the Bostobe Formation (Santonian –Campanian), Kazakhstan. ZIN PH 26/180, partial frontal of an adult individual in dorsal (A), and oblique antero-dorsolateral (B), views. PIN 2229/510, partial frontal in dorsal view (C). ZIN PH 27/180, partial frontal of a subadult individual in dorsal (D), and oblique antero-dorso-lateral (E), views. ZIN PH 36/180, skull roof fragment preserving pieces of the right postorbital and squamosal in articulation, in ventral (F), dorsal (G), and lateral (H), views. ZIN PH 28/180, skull fragment preserving the parietal and supraoccipital in articulation, in dorsal (I), ventral (J), and posterior (K), views. ZIN PH 8/180, conical tooth in mesial or distal view (L). ZIN PH 7/180, blunt-crowned tooth in lingual view (M).Abbreviations: acf, anterior communicative foramen of the parietal pneumatic recess; elg, earlid groove; frf, frontal facet; itr, intertympanic pneumatic recess; k, keel; lsf, laterosphenoid facet; lsq, lateral squamosal lamina; orb, orbit; p, parietal; pcf, posterior communicative foramen of the parietal pneumatic recess; pfrf, prefrontal facet; po, postorbital; pof, postorbital facet; prof, prootic facet; qf, quadrate facet; sgc, sagittal crest; so, supraoccipital; sq, squamosal; stfe, supratemporal fenestra; stfo, supratemporal fossa. Scale bar = 1 cm.

Figure 18. Referred mandibular remains of Kansajsuchus sp. cf. K. extensus from the Bostobe Formation (Santonian– Campanian), Kazakhstan. PIN 2229/507, mandibular symphysis (holotype of ‘Turanosuchus aralensis’) in dorsal (A), left lateral (B), and ventral (C), views. Mid-section dentary fragments: PIN 2229/515, in dorsal (D), and medial (E), views; PIN 2229/506, in dorsal (F), and lateral (G), views. ZIN PH 32/180, anterior fragment of the right splenial in medial (H), and lateral (I), views. PIN 2229/504, fragmentary left surangular and articular in articulation, in dorsal (J), medial (K), and lateral (L), views. PIN 2229/501, partial left angular in medial (M), dorsal (N), and lateral (O), views. Abbreviations: anf, angular facet; ar, articular; arf, articular facet; df, dentary facet; gf, glenoid fossa; Mcg, Meckelian groove; nvf, neurovascular foramen; r, ridge; rap, retroarticular process; san, surangular; spf,splenial facet. Arabic numerals correspond to numbers of dentary alveoli. Scale bar = 1 cm.

Figure 19. Referred postcranial remains of Kansajsuchus sp. cf. K. extensus from the Bostobe Formation (Santonian – Campanian), Kazakhstan. PIN 2229/520, dorsal vertebra in oblique anterior (A), posterior (B), left lateral (C), and dorsal (D), views. ZIN PH 12/180, fragmentary dorsal vertebra in anterior (E), and right lateral (F), views. ZIN PH 11/180, fragmentary dorsal vertebra (likely, the posteriormost from the section) in posterior (G), and left lateral (H), views. Presacral paravertebral osteoderms: ZIN PH 16/180, in dorsal (I), and anterior (J), views; ZIN PH 15/180, in dorsal (K), and ventral (L), views; PIN 2229/503, in dorsal (M), and ventral (N), views. ZIN PH 34/180, ventral hexagonal osteoderm in ventral view, O. Abbreviations: aas, anterior articular surface; anr, annular rugosity; antp, anterior process; k, keel; ns, neural spine; pas, posterior articular surface. Scale bar = 1 cm.