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The mounted skeleton of Anteophthalmosuchus hooleyi (IRSNB R47) from the Barremian-Lower Aptian coal mine of Bernissart, Belgium, in A, right lateral view and dorsal view B, with and C, without shield of osteoderms.
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Two specimens originally referred by Louis Dollo to Goniopholis simus from the Lower Cretaceous of Bernissart, Belgium, are described. They consist of fully articulated skeletons, one missing the skull and mandible. Comparison of these specimens with recently revised specimens from the Wealden of England allows confirmation that the Belgian specime...
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... beautiful specimen IRSNB R47 ( Fig. 1) was prepared, mounted in three dimensions, and exhibited in the Nassau Palace in Brussels in 1884, next to the first complete skeleton and holo- type specimen of Iguanodon bernissartensis (IRSNB R51) and the small Mantellisaurus atherfieldensis (IRSNB R57). The sec- ond specimen of Goniopholis IRSNB R290 (Fig. 2) was pre- pared on one ...
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... the Bernissart specimens are particularly dark and shiny, Figure 1 and Figures 3-7 have been executed by coating IRSNB R47 with ammonium chloride in order to give a neutral gray color and enhance the contrast and relief. The skull of IRSNB R47 is nearly complete, the tip of the rostrum being damaged (Fig. 3). ...
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... Abbreviations: azo, prezygapophysis; cp, capitulum; dp, dorsal process; hyp, hypapophysis; ns, neural spine; pzo, postzygapophysis; tb, tuberculum; trp, transverse process. elements of IRSNB R290 are consistent with the elements of IRSNB R47. Because postcranial elements have been completely freed from the matrix in IRSNB R47 (see Fig. 1 for a general view), the description below focuses on this ...
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... Vertebrae-Seven cervical vertebrae are present (e.g., Fig. 10A, B). All are amphicoelous, and with the exception of the heavily damaged second, all possess a concave ventral margin with a hypapophysis located near the anterior edge. The hypapophysis is a small knob that projects below the surface of the centrum and does not show shape variation through the cer- vical vertebrae. The first, and possibly ...
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... Ribs-The best example of a cervical rib is a right element from cervical 3 (Fig. 10A, B). It has a long posterior process and an anterior process that is only slightly shorter. The dorsal surface is concave, with a ventral surface that is flat and laminar anteriorly and assumes a rod-like shape ...
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... Vertebrae-There are 15 thoracic vertebrae (e.g., Fig. 10C-H). The only complete neural arches are those of the third and 10th thoracic vertebrae. Hypapophyses, consisting of short robust knobs, are visible on the first three thoracic verte- brae. The fourth bears a remnant of a hypapophysis, but all other vertebrae are devoid of it, even though these areas are well pre- served. All vertebrae are ...
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... pophyses. Thus, the capitulum and tuberculum are separated from each other on two proximal extensive branches at least on the first four vertebrae. Moving posteriorly, they become aligned in the same plane, with the tuberculum tapering further medially than the capitulum, a situation that tends to be less and less accentuated on posterior ribs (Fig. 10L, M). The anterior margin of the proximal shaft hosts a pronounced blade that progressively shifts distally. This blade is almost nonexistent in the 10th ...
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... sacral vertebrae are present and still fused to the ilia (Fig. 10I, J). Their neural arch is eroded. The ventral margin of the centrum is markedly concave. The first sacral ver- tebra shows the best-preserved transverse process contacting the ilia in anterior view. The sacral rib is arched, fused to the ilium, and the least deformed side is the right; its distal end is at the same level as the dorsal ...
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... Vertebrae-There are 20 preserved caudal vertebrae, but a few more may have been present (Fig. 10I-K). Until the fifth caudal, the posterior and anterior centrum surfaces are still slightly amphicoelous and circular in outline. Posterior to the fifth caudal position, all centra are flat, with a rectangular out- line, being mediolaterally compressed. All centra have a demar- cated concave ventral margin. In ventral view, the surface ...
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... are posteriorly directed, with a triangular outline. No evidence of a transverse process could be found behind the 13th caudal due to heavy damage. Nevertheless, caudal vertebra 20, which is complete, does not have a transverse process. The posteroventral margin of the centrum has two facets receiving chevrons. A few chevrons are preserved (Fig. 10N, O, P), with the most complete being just anterior to vertebrae 14 and 15. These chevrons possess a distal anteroposteriorly spatulate end as well as separated proxi- mal ...
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... scapulae are preserved. The right one is more complete, although still missing part of the blade on the anterior edge ( Fig. 11A-D). This bone consists of a dorsally narrow and proximodistally elongate blade. The blade shows a slight antero- posterior constriction where it merges with the proximal portion of the bone. In lateral view, the scapular blade is not perpendicu- lar but is oblique to the synchondrosis for the coracoid. A small crest for M. triceps brachii ...
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... shape of the coracoid is very similar to that of eusuchians in presenting a pendulous contribution for the gle- noid facet on its posterior side and a long ventral shaft expanding at the midpoint and becoming a fan-shaped blade with a convex edge ( Fig. 11E-H). In lateral or medial view, the coracoid blade is not perfectly symmetrical: the anterior edge has a pointed pro- cess. The anterior edge of the blade is markedly concave, whereas the posterior edge is straight to convex. An inward cur- vature characterizes the blade in anterior view. The circular cor- acoid foramen is close to the ...
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... interclavicle is preserved (Fig. 11I, J). The dorsal surface of the bone is slightly inflated and mediolaterally expanded at mid-length. The interclavicle bears a set of striations on the posterior half (same side as the ...
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... humerus (Fig. 12A-D) is 20.8 cm long and is less than a third longer than the radius and ulna (Fig. 13A-I). The ulna is 15.6 cm long, and the radius is 13.2 cm long. Distally, the ulna is not in line with the radius and ends more distally than the radius. This is reflected in the shape of the radiale. Humerus-Only the right humerus is complete; the left ...
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... humerus (Fig. 12A-D) is 20.8 cm long and is less than a third longer than the radius and ulna (Fig. 13A-I). The ulna is 15.6 cm long, and the radius is 13.2 cm long. Distally, the ulna is not in line with the radius and ends more distally than the radius. This is reflected in the shape of the radiale. Humerus-Only the right humerus is complete; the left one is preserved only proximally. It consists of a long straight shaft. The ...
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... radiale (Fig. 14E-H) is an elongate bone (unlike in eusuchians; e.g., Mook, 1921) with a nearly straight but slightly con- cave medial margin. The lateral edge of the shaft is shorter and more concave. The lateral facet for the ulna is remarkably long (1.6 cm) and hosts a spiny posterior process at its distal edge, serving as a con- tact with the ulnare. ...
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... proximal end of the ulnare (Fig. 14A-D) is as wide as the shaft and the distal end of the bone is fan-shaped; thus, the medial and lateral margins are widely concave. Laterally, just before mid-shaft, there is a small anterolateral scar. The distomedial facet contacting the radiale consists of a short peg. Proximally, the facet is rounded and convex. The distal facet is ...
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... right manus shows five digits, and although all metacarpals are preserved, only digits I, II, and V preserve pha- langes (Fig. 14). In digit I, the first phalanx is short and relatively robust, whereas in digit II, the first phalanx is gracile and about twice as long as the same phalanx in digit I. Metacarpal III is the longest of all but is only slightly longer than metacarpals II and IV. Metacarpal III possesses a globular callosity on its proximal head. The ...
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... ilia are incompletely preserved and variably affected by plastic deformation. The right one is heavily frac- tured, but the left one is not (Fig. 15). The dorsal margin of the blade is convex, has a low profile, and is moderately wasp-waisted posteriorly, as seen on the right element. This feature is barely detectable on the left ilium. Although a set of fractures have been filled with plaster on the lateral side, the dorsal margin and the medial side are not damaged and are fully ...
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... supraacetabular crest is restricted to the anterodorsal margin of the acetabulum (as in C. niloticus). Its shape is best preserved on the left ilium. Two anteroventral peduncles connect to the ischium and form the anterior and posterior margins of the hollow acetabulum, respectively (Fig. 15A). In ventral view, the anterior peduncle has a triangular outline. The posterior pedun- cle is mediolaterally enlarged and has the shape of a half moon. The medial surface of the ilium is only visible dorsally on the left element. It is smooth and slightly concave just below the rugose dorsal ...
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... ischia are preserved, although the left more completely preserves the articular facets (Fig. 15B-E). The proximal area is anterolaterally expanded and is divided into two facets that articulate with the ilium. In proximal view, the anterior and posterior iliac processes are not aligned with the anteroposterior axis of the ischial blade. These two processes are separated by a marked notch corresponding to the acetabulum, which is ...
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... pubes are complete (Fig. 15F), but the antero- medial margin of the right pubis is damaged. The bone is flat- tened throughout. The proximal facet for the ischium is ovoid, being wider than tall. In dorsal view, its proximal area is depressed. The distal blade is spatulate, with a wing-shaped end that expands rapidly after the constriction of the proximal por- tion ...
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... femur is about one-third longer than the tibia and fibula (Figs. 16, 17). The femur has a proximodistal length of 20.1 cm, the tibia is 14.4 cm long, and the fibula 13.8 cm ...
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... femora are preserved and complete. The femur has a sigmoid shaft in lateral view and resembles the femur of Bernissartia fagesii (IRSNB R46) and other derived neosuchians. The proximal head of the femur is mediolaterally flattened and bears on its lateral surface a shallow concave area, and on its posteromedial side a knob-like process (Fig. 16B, C). As opposed to the great trochanter, the fourth trochanter is well FIGURE 18. Astragalus and calcaneum of Anteophthalmosuchus hoo- leyi (IRSNB R47) from the Barremian-Lower Aptian coal mine of Ber- nissart, Belgium, in A, B, posterior, C, D, ventral, E, F, dorsal, and G, H, lateral views. A, C, E, G, photographs; B, D, F, H, ...
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... Distally, the condyles for tibia and fibula are individualized, of comparable dimension, and are nearly aligned with each other. On the anterior surface, the intercondylar groove is visible but remains shallow and restricted to the distal- most area of the femur. On the posterior surface, the two con- dyles are separated by a deep popliteal fossa (Fig. ...
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... tibia is a long straight bone with moderately expanded distal and proximal ends. The anterior margin of the shaft is rounded, and the posterior margin is flat. Near both the proximal and distal ends, facets are visible on the posterolateral margins of the tibia to accommodate the fibula (Fig. 17B, ...
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... and Calcaneum-These two bones (Fig. 18) are similar to those of eusuchians in shape and proportions (e.g., C. niloticus IRSNB 20.150). The right calcaneum is preserved and is pressed against the astragalus and two rounded ossifications that correspond to the distal tarsals. The calcaneum is well visible lat- erally, showing the outline of the anterior ball and the posterior ...
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... right pes (Fig. 19) is the better preserved and is less deformed than the left one. No phalanges are preserved. Two distal tarsals consist of small rounded bones that connect between the astragalus-calcaneum complex and the proximal articulation of metatarsals I-III. From I to V, all metatarsals overlap each other by projecting a small proximolateral peg ...
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... consequence of such work allows the recognition that European Goniopholididae are composed of four to five genera: Anteophthalmosuchus, Goniopholis, Hul- kepholis, Nannosuchus, and possibly Vectisuchus. Although Nan- nosuchus has sometimes been viewed as a juvenile form of Goniopholis (Salisbury, 2002), recent interpretations seem to recognize this genus as valid (Andrade et al., 2011). Concerning Vectisuchus, it was originally viewed as a longirostrine goniopho- lidid (Buffetaut and Hutt, 1980) and recently recovered it as part of the Pholidosauridae. ...
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... compared with Cretaceous derived neosuchians, this shape resembles that of Isisfordia duncani (Salisbury et al., 2006: fig. 2), Susisuchus anatoceps (Salisbury et al., 2003: fig. 3), or Pachycheilosuchus trinquei (Rogers, 2003: fig. 6). The scapula does not resemble that of Thalattosuchia, which is rod-like (e.g., Steneosaurus; Andrews, 1913a: fig. 41). However, the scapula of IRSNB R47 drastically differs from that of basal mesoeucrocodylians, which do pos- sess a relatively short and dorsally wide or fan-shaped scapular blade, as seen in some notosuchians, including Simosuchus clarki The coracoid morphology is similar to that of eusuchians and is not that different from that of the ...
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... Goniopholididae is a well-known extinct group of neosuchian crocodyliform taxa that bear resemblance, externally in general shape and lifestyle, to extant crocodylians Martin et al., 2016), ranking from the Early Jurassic Calsoyasuchus valliceps, Tykoski, Rowe, Ketcham, and Colbert, 2002 as the oldest representative, until the Late Cretaceous with Denazinosuchus kirtlandicus Wiman, 1932 as the youngest representative of the clade (Sullivan and Lucas, 2003). This clade was common in the Northern Hemisphere, where fossils are mostly found in North America (Allen, 2012) and Europe (Ristevski et al., 2018), with some remains found also in Africa (Goodwin et al., 1999) and Asia (Maisch et al., 2003;Cuny et al., 2010). ...
... This species was described based on a fragmentary skull and isolated postcranial remains. Another two goniopholidid genera, Anteophthalmosuchus and Hulkepholis (Buscalioni et al., 2013;Puértolas-Pascual et al., 2015;Arribas et al., 2019) were described from Teruel (Spain) as being closer looking species from the same genera found in Wessex region (United Kingdom) (Salisbury and Naish, 2011;Ristevski et al., 2018) and Bernissart (Belgium) (Martin et al., 2016). These Early Cretaceous Iberian species are: Anteophthalmosuchus escuchae Buscalioni, Alcalá, Espílez and Mampel, 2013, a partial skeleton and a disarticulated skull, plus several isolated teeth; and Hulkepholis plotos Buscalioni, Alcalá, Espílez, and Mampel, 2013 and Hulkepholis rori Arribas, Buscalioni, Royo Torres, Espílez, Mampel, and Alcalá, 2019, also as partial skeletons and skull remains. ...
... There are well-marked maxillary depressions at the posterolateral-most region of the maxilla (Figures 2-3), a feature shared with some pholidosaurids and most goniopholidids (e.g., Martin and Buffetaut, 2012;Martin et al., 2016;Yoshida et al., 2021). Nannosuchus, Vectisuchus, and the putative goniopholidid Denazinosuchus, do not show any maxillary depressions. ...
Plenty of goniopholidid species from the Mesozoic have been found in the Iberian Peninsula. A previous goniopholidid taxon, Goniopholis baryglyphaeus Schwarz, 2002, from the Late Jurassic of the Guimarota coal mine (Leiria, central Portugal) was described. This taxon corresponds to a partial skull and some postcranial material, and it marked the oldest and first ever record of goniopholidid and Goniopholis species described for the Iberian Peninsula. Here we present a well-preserved, almost complete, skull of a new species, Ophiussasuchus paimogonectes gen. et sp. nov. from Upper Jurassic deposits of Praia de Paimogo (near Lourinhã, central west Portugal). The specimen corresponds to a mesorostrine, platyrostral skull of a medium-sized goniopholidid coming from the upper Kimmeridgian within the Lourinhã Formation. Phylogenetically, the new species is recovered as the sister taxon of the Early Cretaceous European clade made by Hulkepholis and Anteophthalmosuchus. Although its position is well-resolved, this new taxon displays intermediate morphological traits, sharing characteristics with Jurassic Asian and American basal goniopholidids (e.g., presence but lesser development of the secondary choana with the nasopharyngeal duct partially open), as well as more derived characters shared with Cretaceous European taxa such as Hulkepholis (e.g., the shape of the supratemporal fenestra and the palatines). As a result, Ophiussasuchus paimogonectes gen. et sp. nov. exhibits characteristics suggesting a reversion to primitive goniopholidid conditions or intermediate states between the goniopholidid taxa of North America and Europe. These findings support the shared Late Jurassic fauna between the Morrison and Lourinhã Formations, while also having high endemism of taxa.
... The hexagonal shape is commonly observed in ventral osteoderms of Mesoeucrocodylia (Benton and Clark, 1988;Pol and Gasparini, 2009;Pinheiro et al., 2011;de Araújo Carvalho et al., 2021). Additionally, the polygonal morphology is generally consistent with non-eusuchian mesoeucrocodylians (Stevens et al., 2013), particularly in the ventral osteoderms of Neosuchia, such as Goniopholididae, Pholidosauridae, and Dyrosauridae (Wu et al., 1996(Wu et al., , 2001Schwarz, 2003;Martin et al., 2016a;Jouve and Jalil, 2020). The osteoderm shapes observed in M8 closely resemble the most completely ventral osteoderms of the goniopholid Sunosuchus junggarensis from the Late Jurassic in the Junggar Basin of Xinjiang, China (Wu et al., 1996). ...
... The pentagonal and hexagonal osteoderms also resemble the mid-ventral osteoderm in the Thai goniopholidid Siamosuchus phuphokensis from the Early Cretaceous, Sao Khua Formation (Lauprasert et al., 2007), and other neosuchian taxa, such as Goniopholis baryglyphaeus (Schwarz, 2002;Puértolas-Pascual and Mateus, 2020), Sunosuchus sp. from Kirghisia (Averianov, 2001), Anteophthalmosuchus cf. escuchae (Puértolas-Pascual et al., 2015), and Anteophthalmosuchus hooleyi (Martin et al., 2016a). Moreover, the pentagonal and hexagonal osteoderms also discovered in ventral osteodermal shield of other Mesoeucrocodylia taxa, Hsisosuchus from the Late Jurassic of Yunnan, China (Wu et al., 2023) and Hsisosuchus chowi from the Late Jurassic of southern Sichuan, China (Peng and Shu, 2005). ...
... The pits on their external surface are uniformly small, round to ovoid in shape, and well separated in random arrangements. These morphological characteristics are reminiscent of ventral osteoderms found in Mesoeucrocodylia (Averianov, 2001;Wu et al., 1996Wu et al., , 2001Wu et al., , 2023Schwarz, 2002Schwarz, , 2003Peng and Shu, 2005;Lauprasert et al., 2007;Puértolas-Pascual et al., 2015;Martin et al., 2016a;Jouve and Jalil, 2020;Puértolas-Pascual and Mateus, 2020;Wu et al., 2023). At this stage, the M8 is likely belongs to a mesoeucrocodylid other than Indosinosuchus and their exact assignment is pending on further research. ...
... The teleosaurid Peipehsuchus teleorhinus and the goniopholidid Sunosuchus shunanensis do not have the bones preserved. However, the humerus is longer than the ulna or the radius in other genera of the Teleosauridae such as a teleosauroid Aeolodon priscus (Foffa et al. 2019: figure 10) and other taxa of the Goniopholididae such as Anteophthalmosuchus hooleyi (Martin et al. 2016: figure 1). In addition, a different species (Sunosuchus junggarensis from the Upper Jurassic of western China) of Sunosuchus has the left humerus preserved (Wu et al. 1996: figure 14). ...
Hsisosuchus, with three known species, was the most common genus of crocodyliforms in the Sichuan Basin during the Jurassic. However, the overall shape of the ventral trunk shield of osteoderms has not been clarified due to the lack of complete specimens. A new specimen of Hsisosuchus recently recovered from the Upper Jurassic of Yunnan has the nearly complete ventral trunk shield preserved. This specimen provides an example for the first time for us to understand the shape and the way of the arrangement of the osteoderms in the ventral trunk shield in the genus. Compared with the articulated part of the ventral trunk shield preserved in two of the three species of the genus, the osteoderms of the ventral trunk shield of the new specimen are different in both shape and way of arrangement, indicating that the pattern of the ventral trunk shield is not constant within Hsisosuchus. With no skull and much of the postcranial skeleton poorly preserved as well as no corresponding element that can be used to compare with all the three species, the new specimen was considered as an indeterminate species of Hsisosuchus, awaiting more materials to clarify the problem.
... The trapezoid shape of LPUFS 5854 is also a feature present in the mesoeucrocodilian dermal plates: according to Pinheiro et al. (2011) it refers to the ventral part of the body (gastral or ventral shield). Such shape of the osteoderm is also observed in some gastral dermal plates of semiaquatic goniopholidids neosuchian such as Anteophthalmosuchus (Martin et al., 2016;Ristevski et al., 2018), Siamosuchus (Lauprasert et al., 2007), as well as other undetermined goniopholidids (Kuzmin et al., 2013;Pu ertolas-Pascual et al., 2015). In general terms, this gastral osteoderm morphology observed in LPUFS 5854 (Fig. 5) and in some goniopholidids, differs from other neosuchians such as thalattosuchians (e.g., € Osi et al., 2018), dyrosaurids , and some tethysuchian pholidosaurids (e.g., Broin, 2002;Jouve and Jalil, 2020). ...
... Interestingly, the nearly trapezoidal morphology of LPUFS 5854 is compatible with some of the osteoderms of the peripheral mesoeucrocodilian shield (Martin et al., 2016), differing from the dorsal osteoderms in both shape and the absence of a sagittal keel. However, several fragmented and non-diagnostic dermal plates have been reported as goniopholidids over the past years, including some dubious South American occurrences (Andrade et al., 2011). ...
The Feliz Deserto Formation (Berriasian–Valanginian, Sergipe-Alagoas Basin, NE Brazil) preserved some of the earliest South American fossil records of the rifting stages which resulted in the Gondwana supercontinent break-up during the Lower Cretaceous. Recently, the first spinosaurid theropod record of this formation was described, based on a tooth recovered from Canafístula 01 fossil locality in Sergipe State. We add herein twenty-seven isolated specimens to the fossil record of the Lower Cretaceous Feliz Deserto Formation. The new material includes seven isolated spinosaurid theropod teeth of the spinosaurine clade, as well as an indeterminate theropod preungual pedal phalanx. In addition, we describe an isolated crocodyliform osteoderm, as well as eighteen isolated teeth, some of which were taxonomically identified in three distinct morphotypes of neosuchian crocodyliforms. These findings expand the Gondwanan fossil record of both spinosaurine theropods and neosuchian crocodyliforms. Despite the fragmented nature of the specimens, these new fossils allowed the characterization of their general taphonomic features with low fluvial transport of bioclast prior to the burial. The depositional paleoenvironment of the Canafístula 01 locality is compatible with the deltaic system unit, that characterizes part of the Feliz Deserto Formation during the Lower Cretaceous. These fossil findings exemplify the co-occurrence of spinosaurids and more than one taxon of crocodyliforms in the deltaic-lacustrine paleoenvironment represented by the Feliz Deserto Formation. These new occurrences reinforce the fossiliferous potential of the Canafístula 01 locality, especially related to the paleovertebrates from the Lower Cretaceous of Brazil.
... Although their dentition is generally poorly preserved and described only superficially, the tooth crowns of some species are quite similar to SGO.PV.1160. Most notably, those of the Lower Cretaceous representatives of the genus Pholidosaurus von Meyer 1841, known from several European localities, have been described as slender, conical, slightly recurved, bearing enamel ornamentation formed by apicobasal ridges and either lacking carinae or possessing such weakly developed ones that they are difficult to differentiate from the rest of the ridges (Andrews 1913;Martin et al. 2016). Oceanosuchus boecensis (Hua et al. 2007) from the lower Cenomanian of western France and Chalawan thailandicus (Buffetaut and Ingavat 1980) from the Jurassic of northeastern Thailand are other pholidosaurids whose tooth crowns also bear apicobasal enamel ridges (Buffetaut andIngavat 1980, 1984;Hua et al. 2007). ...
... Their dentition consists of conical tooth crowns with varying degrees of robustness and lingual curvature, and striated/fluted enamel surfaces. However, their teeth, as those of spinosaurids, are invariably bicarinate (Andrade et al. 2011;Puértolas-Pascual et al. 2015;Martin et al. 2016;Ristevski et al. 2018). Additionally, the distribution of these neosuchians was restricted to Laurasia (Allen 2012;Halliday et al. 2015;Puértolas-Pascual et al. 2015;Martin et al. 2016;Ristevski et al. 2018), therefore we consider it safe to conclude that SGO.PV.1160 does not belong to a goniopholidid either. ...
... However, their teeth, as those of spinosaurids, are invariably bicarinate (Andrade et al. 2011;Puértolas-Pascual et al. 2015;Martin et al. 2016;Ristevski et al. 2018). Additionally, the distribution of these neosuchians was restricted to Laurasia (Allen 2012;Halliday et al. 2015;Puértolas-Pascual et al. 2015;Martin et al. 2016;Ristevski et al. 2018), therefore we consider it safe to conclude that SGO.PV.1160 does not belong to a goniopholidid either. ...
We describe the first crocodyliform tooth found in continental deposits from the Quebrada Monardes Formation (Lower Cretaceous) at Cerro La Isla, Atacama Region, northern Chile. Though the fossil (SGO.PV.1160) is poorly preserved, the enamel surfaces clearly bear tightly-packed apicobasal ridges. Comparisons of SGO.PV.1160 with the dental morphology of taxa that it may belong to suggest a hitherto unknown pholidosaurid or, more likely, notosuchian. Therefore, the tooth crown is here provisionally identified as belonging to an indeterminate mesoeucrocodylian, pending future discoveries of more complete material. This find increases our knowledge of the fauna in the Lower Cretaceous deposits of Cerro La Isla, as well as the importance of the fossil site, since pholidosaurids have not been reported from Chile, while if it proves to be notosuchian, it may represent one of the earliest known taxa of the group in South America, and the entire world.
... It is absent in most "advanced ziphosuchians" (Pol & Leardi, 2015); for example, Mariliasuchus (de Andrade & Bertini, 2008), Notosuchus (Fiorelli & Calvo, 2008), sphagesaurids (Pol et al., 2014), and sebecosuchians (Busbey, 1986;Kellner et al., 2014) but present in baurusuchids (Montefeltro et al., 2020). The muscle was also relegated to the medial and ventral mandible in many advanced neosuchians including goniopholids (Martin et al., 2020;Martin, Delfino, & Smith, 2016), pholidosaurs (Martin, Raslan-Loubatié, & Mazin, 2016), and paralligatorids (Turner, 2015). Basal eusuchians also lack the lateral insertion, including Bernissartia (Martin et al., 2020). ...
Jaw muscles are key features of the vertebrate feeding apparatus. The jaw musculature is housed in the skull whose morphology reflects a compromise between multiple functions, including feeding, housing sensory structures, and defense, and the skull constrains jaw muscle geometry. Thus, jaw muscle anatomy may be suboptimally oriented for the production of bite force. Crocodylians are a group of vertebrates that generate the highest bite forces ever measured with a flat skull suited to their aquatic ambush predatory style. However, basal members of the crocodylian line (e.g., Prestosuchus) were terrestrial predators with plesiomorphically tall skulls, and thus the origin of modern crocodylians involved a substantial reorganization of the feeding apparatus and its jaw muscles. Here, we reconstruct jaw muscles across a phylogenetic range of crocodylians and fossil suchians to investigate the impact of skull flattening on muscle anatomy. We used imaging data to create 3D models of extant and fossil suchians that demonstrate the evolution of the crocodylian skull, using osteological correlates to reconstruct muscle attachment sites. We found that jaw muscle anatomy in early fossil suchians reflected the ancestral archosaur condition but experienced progressive shifts in the lineage leading to Metasuchia. In early fossil suchians, musculus adductor mandibulae posterior and musculus pterygoideus (mPT) were of comparable size, but by Metasuchia, the jaw musculature is dominated by mPT. As predicted, we found that taxa with flatter skulls have less efficient muscle orientations for the production of high bite force. This study highlights the diversity and evolution of jaw muscles in one of the great transformations in vertebrate evolution.
... The shoulder girdle of Confractosuchus closely resembles Anteophthalmosuchus hooleyi (see fig. 16 in (Martin Delfino and Smith, 2016)). Anteophthalmosuchus shoulder girdle morphology was reported to be similar to Isisfordia duncani (scapula only see fig. 2 in Salisbury et al. (2006) and Susisuchus anatoceps (schematic outline, see fig. 3 in Salisbury et al. (2003); however, the published images of both these susisuchids were insufficient to draw such similarities to Confractosuchus. ...
... A resemblance with Pachycheilosuchus trinquei (Rogers, 2003) can be established despite a slightly different orientation in the figures and only one view supplied of each (see fig. 6A,B in Rogers (2003)). Much like the description of Anteophthalmosuchus hooleyi (Martin, Delfino and Smith, 2016) the coracoid resembles the morphology of eusuchians. ...
Crocodylians are among Earth’s most successful hyper-carnivores, with their crocodyliform ancestors persisting since the Triassic. The diets of extinct crocodyliforms are typically inferred from distinctive bite-marks on fossil bone, which indicate that some species fed on contemporaneous dinosaurs. Nevertheless, the most direct dietary evidence (i.e. preserved gut contents) of these interactions in fossil crocodyliforms has been elusive. Here we report on a new crocodyliform, Confractosuchus sauroktonos gen. et sp. nov., from the Cenomanian (92.5–104 Ma) of Australia, with exceptionally preserved abdominal contents comprising parts of a juvenile ornithopod dinosaur. A phylogenetic analysis recovered Confractosuchus as the sister taxon to a clade comprising susisuchids and hylaeochampsids. The ornithopod remains displayed clear evidence of oral processing, carcass reduction (dismemberment) and bone fragmentation, which are diagnostic hallmarks of some modern crocodylian feeding behaviour. Nevertheless, a macro-generalist feeding strategy for Confractosuchus similar to extant crocodylians is supported by a morphometric analysis of the skull and reveals that dietary versatility accompanied the modular assembly of the modern crocodylian bauplan. Of further interest, these ornithopod bones represent the first skeletal remains of the group from the Winton Formation, previously only known from shed teeth and tracks, and may represent a novel taxon.
... Two ventral pits are present on the ventral surface of the premaxilla. The maxillary depression is present on the laterodorsal surface of the posterior process (figure 1d ), as in the other goniopholidids and pholidosaurids [35]. The number of subfossae in the depression is five (figure 4) as in G. simus [36,37], while it is four in Amphicotylus lucasii and three in G. kiplingi [8] and Anteophthalmosuchus hooleyi [35]. ...
... The maxillary depression is present on the laterodorsal surface of the posterior process (figure 1d ), as in the other goniopholidids and pholidosaurids [35]. The number of subfossae in the depression is five (figure 4) as in G. simus [36,37], while it is four in Amphicotylus lucasii and three in G. kiplingi [8] and Anteophthalmosuchus hooleyi [35]. A neurovascular opening is present at the bottom of each chamber in G. kiplingi [8]; however, it is only present in the second chamber of the left depression of Amphicotylus milesi (GMNH-PV0000229). ...
... A neurovascular opening is present at the bottom of each chamber in G. kiplingi [8]; however, it is only present in the second chamber of the left depression of Amphicotylus milesi (GMNH-PV0000229). The posterior margin of the nasals is divided by the frontal as in G. kiplingi [8] and Anteophthalmosuchus hooleyi [35]. ...
Goniopholididae is a group of basal neosuchian crocodyliforms closely related to Paralligatoridae and Eusuchia that lived during the Jurassic and Early Cretaceous. Goniopholidids have the long, flat snout and secondary palate of modern crocodylians, the acquisition of which is regarded as a key feature in the early evolution of crocodylian body plan and their aquatic adaptation. Here, we report a new species, Amphicotylus milesi , with the description from the best-preserved specimen to date of Goniopholididae from Wyoming, USA. Its posterior extension of the nasopharyngeal passage (pterygoid secondary palate) and the shortening and dorsal deflection of the ceratobranchial suggest that basal neosuchians could raise their gular valve to separate oral and pharyngeal cavities as in modern crocodylians. The anatomy of Amphicotylus milesi sheds light on the acquisition of this new respiratory system in the crocodyliform evolution and their early aquatic adaptation, leading to modern crocodylians.
... A total of 189 characters and 107 taxa were included. IRSNB R47 and NHMUK PV R 3876 were scored as Anteophthalmosuchus episkator based on Martin et al. (2016) and Ristevski et al. (2018), and this taxon was selected as the outgroup (see Supplemental Data 2). The matrix was analyzed with TNT v. 1.1 (Goloboff et al., 2008), using a traditional search based on 1,000 replications of Wagner trees (using random addition sequences), followed by tree bisection recognition (TBR) as a swapping algorithm, saving 100 trees by replication, and considering 10 random seed values. ...
Duerosuchus piscator is a middle Eocene eusuchian known only from Corrales del Vino (Zamora, Spain). The species was defined based on an incomplete skull, partial lower jaws and two vertebrae from a single individual, and several osteoderms referred to other specimens. A detailed study of these remains allows us to question the attribution of all these remains to the same form. Just the cranial remains are considered as indisputably attributable to it; the validity of this species being supported. The present study provides a detailed description and an amended diagnosis for Duerosuchus piscator, which is included for the first time in a phylogenetic analysis in order to establish its systematic position within Crocodylia. As a result of this study, the Eocene crocodyliform paleobiodiversity in the Duero Basin is recognized as comprising a notosuchian (Iberosuchus macrodon), as well as three crocodylians, each belonging to a clade: the alligatoroid Diplocynodon tormis, a crocodyloid traditionally attributed to the genus Asiatosuchus, and Duerosuchus piscator, which is here identified as a planocraniid, up to now unrecognized in the Iberian fossil record.
... 10A-10D). One osteoderm (Fig. 10A) preserves a portion of the forward projecting spine that originates on the anterolateral corner (Martin, Delfino & Smith, 2016). It is smooth with no ornamentation. ...
... It is smooth with no ornamentation. Martin, Delfino & Smith (2016) describe these processes as inserting into a distinct groove on the posterolateral corner of the anteriorly adjacent osteoderm. This insertion is a smooth, concave surface or fossa when seen in ventral view and can be seen on one of the osteoderms (Fig. 10D). ...
... All ventral osteoderms are smooth on their dorsal surfaces and ornamented with large circular pits on their ventral side. Martin, Delfino & Smith (2016) describe similar hexagonal-shaped ventral osteoderms as the most central osteoderm on the ventral shield of the coelognathosuchian Anteophthalmosuchus. Marginal ventral osteoderms are not hexagonal but four-to five-sided (Martin, Delfino & Smith, 2016), suggesting that osteoderm UA-2016-13-22 (Fig. 10E) is from the margin of the ventral shield and UA-2016-13-22 (Fig. 10F) is a central ventral osteoderm. The hexagonal osteoderm from the Arkansas material is un-sutured, suggesting this individual was a sub-adult. ...
We present a previously discovered but undescribed late Early Cretaceous vertebrate fauna from the Holly Creek Formation of the Trinity Group in Arkansas. The site from the ancient Gulf Coast is dominated by semi-aquatic forms and preserves a diverse aquatic, semi-aquatic, and terrestrial fauna. Fishes include fresh- to brackish-water chondrichthyans and a variety of actinopterygians, including semionotids, an amiid, and a new pycnodontiform, Anomoeodus caddoi sp. nov. Semi-aquatic taxa include lissamphibians, the solemydid turtle Naomichelys , a trionychid turtle, and coelognathosuchian crocodyliforms. Among terrestrial forms are several members of Dinosauria and one or more squamates, one of which, Sciroseps pawhuskai gen. et sp. nov., is described herein. Among Dinosauria, both large and small theropods ( Acrocanthosaurus , Deinonychus , and Richardoestesia ) and titanosauriform sauropods are represented; herein we also report the first occurrence of a nodosaurid ankylosaur from the Trinity Group. The fauna of the Holly Creek Formation is similar to other, widely scattered late Early Cretaceous assemblages across North America and suggests the presence of a low-diversity, broadly distributed continental ecosystem of the Early Cretaceous following the Late Jurassic faunal turnover. This low-diversity ecosystem contrasts sharply with the highly diverse ecosystem which emerged by the Cenomanian. The contrast underpins the importance of vicariance as an evolutionary driver brought on by Sevier tectonics and climatic changes, such as rising sea level and formation of the Western Interior Seaway, impacting the early Late Cretaceous ecosystem.
