Suchosaurus

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Suchosaurus
Temporal range: Early Cretaceous (Valanginian to Barremian), 135–125  Ma
Suchosaurus cultridens holotype tooth.jpg
Holotype tooth of S. cultridens
(NHMUK PV R36536)
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Clade: Dinosauria
Clade: Saurischia
Clade: Theropoda
Family:† Spinosauridae
Subfamily:† Baryonychinae
Genus:† Suchosaurus
Owen, 1841
Type species
†Suchosaurus cultridens
Owen, 1841
Other species
  • †Suchosaurus girardi Sauvage, 1897
Synonyms
List of synonyms

Suchosaurus (meaning "crocodile lizard") is a dubious genus of large theropod dinosaur that lived during the Early Cretaceous in what is now Europe. The type species, S. cultridens, was originally described in 1841 by Richard Owen based on a chimeric assemblage of fossil teeth and vertebrae discovered in the Tilgate Forest, of Sussex, England, in sediments of the Wealden Supergroup. The second species, S. girardi, was established in 1897 by Henri Émile Sauvage from a tooth and fragmentary jaw material recovered from the Papo Seco Formation in Portugal. Initially interpreted as a crocodilian for nearly two centuries, Suchosaurus was only formally reidentified as a spinosaurid following a 2003 publication by Angela Milner, who also considered it as a possible senior synonym of Baryonyx . This proposal was followed by several authors until 2011, when Octávio Mateus and colleagues regarded the genus as dubious due to the non-diagnostic nature of the assigned fossil material. Named only one year before Owen introduced the term Dinosauria in 1842, Suchosaurus ranks among the earliest dinosaurs described in the history of paleontology and represents the first named spinosaurid, although it was not recognized as such at the time of its original descriptions.

Contents

Although known from very limited fossil material, Suchosaurus is estimated to have reached a length of between 8.6–10 metres (28–33 ft), with a minimum body mass of around one tonne. Like other spinosaurids, Suchosaurus was probably a large bipedal carnivore with well-built forelimbs and elongated, crocodile-like skulls. The teeth of Suchosaurus, which constitute the main fossils documenting this taxon, are slightly recurved and display a conical to subconical cross-section. The crown also bears numerous longitudinal flutes. The holotype of S. cultridens appears to lack serrations, whereas at least one tooth attributed to S. girardi bears serrations on its anterior carina. Although now considered as a dubious spinosaurid, Suchosaurus is generally assigned to the subfamily Baryonychinae due to its dental characteristics, which it shares with other genera such as Baryonyx and Suchomimus . Like other members of this group, its teeth were most likely adapted for a piscivorous diet. Based on the fossil record associated with the taxon, the animal lived and hunted in fluvial environments alongside numerous other dinosaurs, as well as pterosaurs, crocodylomorphs, turtles, plesiosaurs, fishes, various invertebrates, and even early mammals.

Research history

Map showing spinosaurid localities of southeast England; 3 is the Tilgate Forest quarry where Suchosaurus was found Wealden Supergroup spinosaurids map.png
Map showing spinosaurid localities of southeast England; 3 is the Tilgate Forest quarry where Suchosaurus was found

The taxonomic history of Suchosaurus begins in 1822, when British palaeontologist Gideon Mantell and his partner Mary Ann mentioned, in their book The Fossils of the South Downs, teeth that had been discovered by Mantell in a quarry in Tilgate Forest, near Cuckfield in Sussex, England. Based on their morphology, characterized by well-defined lateral ridges, Mantell's mentor William Clift, then curator of the Hunterian Museum, London, suggested that these fossils might belong either to a crocodile or to a monitor lizard, the couple favoring the former interpretation. [1] [2] [3] In June 1823, Scottish geologist Charles Lyell brought the fossils described by the Mantell couple to Paris, France, so that they could be examined by French naturalist Georges Cuvier. [2] In his work published the following year, Cuvier endorsed the initial 1822 interpretation, although he described most of the teeth only very briefly. Nevertheless, he illustrated four of the Tilgate fossil teeth discovered by Mantell. [4] In 1827, Mantell redescribed these fossils in greater detail, distinguishing them as belonging to two types of crocodilians, separating the blunt- crowned teeth from the more slender and recurved ones, which he considered comparable to those of gharials. Although he did not erect any scientific name to formalize this observation, he attributed the teeth of the second category to a crocodilian that he referred to by the common name "gavial of Tilgate Forest". [5]

1878 lithograph of the holotype tooth of S. cultridens Suchosaurus.jpg
1878 lithograph of the holotype tooth of S. cultridens

In 1841, British palaeontologist Richard Owen established, within the genus Crocodilus , a subgenus and species he named Crocodilus (Suchosaurus) cultridens, based on the fossils discovered by Mantell. [6] The name Suchosaurus comes from the Ancient Greek σοῦχος (souchos, "crocodile"), and σαῦρος (saûros, "lizard"), reflecting its then-accepted affinities with crocodilians. [7] The specific name comes from the Latin culter, "dagger", and dens, "tooth", in reference to the taxon's dental morphology. [8] In his work published the following year (the same work in which he first coined the term Dinosauria), Owen retained Suchosaurus at the rank of subgenus and also referred to it two fossil vertebrae collected from the original locality by Mantell. He further considered the expression "gavial of Tilgate Forest" to be the vernacular name of this taxon. [9] The holotype, cited in the scientific literature since the Cuvier's work, consists of a single 3 centimetres (1.2 inches) tooth broken at the upper third of the crown. Like many fossils discovered at Tilgate, this tooth is now housed in the palaeontological collections of the Natural History Museum, London. The original specimen number was BMNH R36536, [2] but it was later re-catalogued as NHMUK PV R36536. [10] In 1878, Owen elevated Suchosaurus as a distinct genus and illustrated the vertebrae in question for the first time, while noting that they had not been found in direct association with the holotype tooth. [11] In the second volume of his 1884 work, when he figured again the holotype tooth of S. cultridens, he named it as S. laevidens. [12] This designation was probably a lapsus calami , as he did not use this specific epithet elsewhere in the same publication. [2] In 1888, British naturalist Richard Lydekker determined that the two vertebrae previously attributed to the genus actually belonged to ornithischian dinosaurs. He assigned the first to the family Iguanodontidae and suggested that the second might be referred to the ankylosaur Hylaeosaurus . Lydekker also regarded S. laevidens as a junior synonym of S. cultridens, [13] a position likewise adopted in 1890 by British palaeontologists Arthur Smith Woodward and Charles Davies Sherborn. [14]

Type mandible and tooth of S. girardi Suchosaurus girardi.jpg
Type mandible and tooth of S. girardi

In 1897, French palaeontologist Henri-Émile Sauvage erected the second species, Suchosaurus girardi, on the basis of two jaw fragments and a single tooth discovered by Swiss geologist Paul Choffat at Boca do Chapim, in Portugal. The specific epithet honors the Portuguese naturalist Albert Girard. [15] The two jaw fragments are currently numbered as MG 324 at the Museu Geológico  [ pt ] in Lisbon. [16] [17] In 2007, French palaeontologist Éric Buffetaut noted that the associated single tooth had not been relocated in the institution and therefore regarded it as lost. However, he identified a third mandibular fragment, probably complementary to one of the first two and previously undescribed. [18] In a conference abstract published in 2013, Portuguese palaeontologist Elisabete Malafaia and her colleagues reported the rediscovery of the isolated tooth in the palaeontological collections of the National Museum of Natural History and Science, Lisbon. Now catalogued as MNHN/UL.I.F2.176, the tooth is among the fossils that were saved from a fire that destroyed a significant part of the museum in 1978. [17]

After some fossil teeth collected in Sussex and the Isle of Wight were routinely labelled as Suchosaurus, the taxon was only rarely mentioned in the scientific literature after the end of the nineteenth century, probably due to the very limited nature of the fossil material. [2] One of the few notable mentions during the following century comes from Spain, where the palaeontologist José Royo y Gómez  [ es ] in 1927 attributed to this genus fossils discovered in the deposits of Morella, in Castellón. [19] However, these specimens were never described in detail afterwards and appear to have been lost during the Spanish Civil War. [2] They were nevertheless briefly mentioned in 1960 by the Spanish palaeontologist Josep Ramon Bataller  [ ca ]. [20]

Reconstructed skeletal mount of Baryonyx at the National Museum of Nature and Science, Tokyo. Based on its dental characteristics, Suchosaurus has been reinterpreted as a spinosaurid, and a possible synonymy with the latter has even been suggested. Baryonyx walkeri mount NMNS.jpg
Reconstructed skeletal mount of Baryonyx at the National Museum of Nature and Science, Tokyo. Based on its dental characteristics, Suchosaurus has been reinterpreted as a spinosaurid, and a possible synonymy with the latter has even been suggested.

In 1986, British palaeontologists Alan Charig and Angela Milner described the theropod dinosaur Baryonyx walkeri on the basis of a relatively well-preserved partial skeleton discovered near the Ockley, in Surrey. [21] [22] Its initial descriptions highlighted distinctive dental characters, which led some authors to assign several isolated teeth uncovered in the Wealden Supergroup of England to this genus. [23] [2] [3] On the basis of these same morphological criteria, Milner suggested in 2003 that some teeth previously referred to Megalosaurus and Suchosaurus might instead belong to Baryonyx. [24] In his 2007 study, Buffetaut considered the teeth of S. girardi very similar to those of Baryonyx (and S. cultridens) except for the stronger development of the tooth crown flutes (or "ribs"; lengthwise ridges), suggesting that the remains belonged to the same genus. Buffetaut agreed with Milner that the teeth of S. cultridens were almost identical to those of B. walkeri, but with a ribbier surface. The former taxon might be a senior synonym of the latter (since it was published first), depending on whether the differences were within a taxon or between different ones. According to Buffetaut, since the holotype specimen of S. cultridens is a single tooth and that of B. walkeri is a skeleton, it would be more practical to retain the newer name. [18] [25] [2]

In 2010, in a review devoted to the taxonomic history of spinosaurids prior to the original description of the type genus Spinosaurus by the German palaeontologist Ernst Stromer in 1915, Buffetaut stated that it is very likely that the fossils attributed to Suchosaurus by Royo y Gómez belong to the baryonychines, in light of the discovery of additional fossil teeth from this subgroup at Morella. [2] The discovery of these same teeth had previously been reported in a conference abstract by the Spanish palaeontologist José Ignacio Canudo and his colleagues in 2004. [26] In 2011, Portuguese paleontologist Octávio Mateus and colleagues agreed that Suchosaurus was closely related to Baryonyx, but considered both species in the former genus nomina dubia (dubious names) since their holotype specimens were not considered diagnostic (lacking distinguishing features) and could not be definitely equated with other taxa. Nevertheless, since its formal reidentification in the early twenty-first century, Suchosaurus has been recognized as the very first spinosaurid to have been named, as well as one of the earliest dinosaurs ever described in the history of palaeontology, although it was not initially identified as such. [16] Nevertheless, since its formal reidentification in the early 21th century, Suchosaurus has been recognized as the very first spinosaurid to have been named, as well as one of the earliest dinosaurs described in the history of palaeontology, although it was not initially identified as such. [24] [25] [2] [3] [10] [8] Furthermore, the illustration of the holotype tooth published by Cuvier in 1824 is now regarded as the first printed depiction of a spinosaurid fossil, and as one of the oldest dinosaur figures in general, even though these groups had neither been defined nor named at the time of its publication. [2]

Description

Hypothetical life restoration of S. cultridens Life reconstruction of Suchosaurus cultridens.png
Hypothetical life restoration of S. cultridens

Although known only from very fragmentary fossil remains, Suchosaurus would likely have had an overall morphology broadly similar to that of many other spinosaurids, i. e. large bipedal carnivore with robust forelimbs and an elongated skull resembling that of crocodiles. [16] [3] [27] In 2011, American vertebrate palaeontologist Thomas R. Holtz Jr. tentatively estimated Suchosaurus at around 10 metres (33 ft) in length and weighing between 1 and 4 tonnes (1.1 and 4.4 short tons). [28] [29] And in 2016, Spanish palaeontologists Rubén Molina-Pérez and Asier Larramendi estimated S. cultridens at approximately 8.6 m (28 ft) long, 2.15 m (7.1 ft) tall at the hips and weighing 1.4 t (1.5 short tons). [30] When Mantell described in 1827 the assemblage of teeth of the "gavial of Tilgate Forest", now recognized as chimeric, [2] he stated that the largest of these teeth would have belonged to an animal of comparable length, estimated between 6–9 metres (20–30 ft) in length. [5]

The dentition of Suchosaurus shows some notable differences between the two species. However, the teeth display only the basic characteristics typical of baryonychines and do not exhibit any diagnostic traits allowing distinction at the generic level, which has led the taxon to be regarded as a nomen dubium. [16] Like Baryonyx, the holotype tooth of S. cultridens is round in cross-section, slightly recurved, and bears numerous flutes on the crown. It differs, however, in having more pronounced fluting and an apparent absence of serrations on the carinae (the anterior and posterior cutting edges of a tooth). Because the carina is strongly worn, it remains difficult to determine whether the tooth was truly devoid of serrations or whether their absence simply results from this wear. The teeth of S. girardi are ziphodont , being subcircular in cross-section and bearing seven denticles per millimetre, a number comparable to that observed in Baryonyx. The roots, very long and slender, exceed half the length of the crown. The teeth of S. girardi are just as strongly fluted as the holotype of S. cultridens, showing eight longitudinal ridges on their inner surface and an enamel texture that is microscopically wrinkled. Nevertheless, unlike the type species, at least one tooth of S. girardi appears to possess a serrated anterior carina. Little information has been reported about the rare fossil jaw fragments of the holotype of S. girardi because of their very incomplete preservation, but they are understood to represent the right portion of the dentary (the largest bone of the lower jaw in diapsids). [18] [16]

Palaeobiology

Though few skull material has been discovered for Suchosaurus, [18] [16] it is known that spinosaurid skulls resembled those of crocodiles; they were long, low, narrow and expanded at their front ends into a terminal rosette-like shape, with a robust secondary palate on the roof of the mouth that made them more resistant to stress and bending. In contrast, the primitive and typical condition for theropods was a tall, broader and wedge-like snout with a less developed secondary palate. The skull adaptations of spinosaurids converged with those of crocodilians; early members of the latter group had skulls similar to typical non-avian (or non-bird) theropods, later developing elongated snouts, conical teeth, and secondary palates. These adaptations may have been the result of a dietary change from terrestrial prey to fish. [31] [32] Most theropod dinosaurs have recurved, blade-like teeth with serrated carinae for slicing through flesh, whereas spinosaurid teeth evolved to be straighter, more conical, and have small or nonexistent serrations. Such dentition is seen in living piscivorous predators such as gharials, as it is better suited for piercing and maintaining grip on slippery aquatic prey so it can be swallowed whole, rather than torn apart. [27] [31] [33]

Palaeoecology

England

The original stratigraphic position of the holotype tooth of S. cultridens does not appear to be unanimously agreed upon among geologists. The specimen comes from the Grinstead Clay, a unit dated to the late Valanginian of the Early Cretaceous, whose status varies according to authors: it is considered either as a distinct geological formation or a member of the Tunbridge Wells Sand Formation. In any case, it originates from the fossil record of the Wealden Supergroup of England, a geological group divided into numerous formations with ages that are both older and younger. [10] During the Early Cretaceous, the Weald area of Surrey, Sussex, and Kent was partly covered by the large, fresh-to-brackish water Wealden Lake. Two large rivers drained the northern area (where London now stands), flowing into the lake through a river delta; the Anglo-Paris Basin was in the south. Its climate was sub-tropical, similar to the present Mediterranean region. [34] [35]

Portugal

S. girardi is known from the Papo Seco Formation, which dates to the early Barremian age of the Early Cretaceous epoch. [18] [16] [17] The formation's depositional environment exhibits a shift from shallow marine to continental, with stratigraphically higher layers representing an estuarine setting. [36] [37] Many other fossil taxa have been found in localities of the Papo Seco Formation. [18] [17] The only formally identified contemporaneous theropod is the spinosaurid Iberospinus (whose holotype had previously been interpreted as belonging to Baryonyx [16] ), [36] although fragmentary bones also indicate the presence of representatives of other lineages, including a possible dromaeosaurid [38] and a coelurosaurian. [39] Two theropod footprint types have been found, one that is smaller with narrow digits, belonging to an indeterminate theropod, and another referrable to the ichnogenus Megalosauripus , likely produced by a carnosaurian theropod such as a spinosaurid. [40] The other dinosaurs present in the formation are sauropods and ornithopods. Sauropod material includes the remains of indeterminate sauropods, eusauropods, titanosauriforms, and titanosaurs. [41] The only formally described ornithopod genus is Cariocecus , [37] but other evidence of their presence includes fragmentary remains of unnamed taxa, consisting of isolated teeth and bones (e.g., incomplete vertebrae and limb bones). [42] A probable ornithopod footprint has also been described from the formation. [43] Non-dinosaurian vertebrate fossils from the Papo Seco Formation include teeth referred to ornithocheirid and ctenochasmatoid pterosaurs, cf. Anteophthalmosuchus sp. (a goniopholidid crocodyliform), and cf. Lepidotes sp. (a semionotiform actinopterygian) and carapace fragments and a partial radius of pancheloniids (sea turtles). [44] [38] Invertebrate fossils comprise aquatic bivalves ( Eomiodon , Nipponomaia , and ostreidans) and gastropods ( Chemnitzia , Natica sp., Turritella ). [45]

References

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  37. 1 2 Bertozzo, F.; Camilo, B.; Araújo, R.; Manucci, F.; Kullberg, J. C.; Cerio, D. G.; Carvalho, V. F.; de Marrecas, P.; Figueiredo, S. D.; Godefroit, P. (2025). "Cariocecus bocagei, a new basal hadrosauroid from the Lower Cretaceous of Portugal". Journal of Systematic Palaeontology . 23 (1) 2536347. doi:10.1080/14772019.2025.2536347. ISSN   1477-2019. S2CID   281435871.
  38. 1 2 Domingues Figueiredo, S.; Bachtsevanidou Strantzali, I.; Gomes, M.; Pimenta, R.; Santos, M. (2017). "Preliminary data of new dinosaurs and turtles remains from the basal deposits of the Papo-Seco Formation in Areias do Mastro's Quarry (Cabo Espichel - Portugal)". Arnava (in Portuguese). 6 (1): 117–130. ISSN   2320-0103.
  39. ↑ Domingues Figueiredo, S.; Rosina, P.; Gomes, M. (2018). "Novos Restos De Um Dinossáurio Aviano Do Cretácico Inferior De Portugal (Areias Do Mastro – Sesimbra)" [New Remains of an Avian Dinosaur from the Lower Cretaceous of Portugal (Mastro area – Sesimbra)]. Revista de Geistória e Pré-História. 1 (in Portuguese). 1 (2): 66–69.
  40. ↑ Figueiredo, S.; Carvalho, I.; Cunha, P.; Neto de Carvalho, C.; Díaz-Martínez, I.; Garcias, T. (2025). "Pegadas de terópodes da Formação de Papo Seco (Barremiano inferior, Cabo Espichel, Bacia Lusitânica)" [Theropod tracks from the Papo Seco Formation (lower Barremian, Cabo Espichel, Lusitanian Basin)]. Comunicações Geológicas (in Portuguese). 112: 81–85. doi: 10.34637/E936-4184 .
  41. ↑ Figueiredo, S. D.; Marques, C.; R., I. M.; Cunha, P. P.; de Souza Carvalho, I. (2023). "Sauropod dinosaur remains from the Papo-Seco formation (lower Barremian, Lusitanian Basin, Cabo Espichel, Portugal): Previous and new discoveries". Cretaceous Research . 151 105657. Bibcode:2023CrRes.15105657F. doi:10.1016/j.cretres.2023.105657. S2CID   260414572.
  42. ↑ Domingues Figueiredo, S.; Carvalho, I. d. S.; Pereda-Suberbiola, X.; Proença Cunha, P.; Bachtsevanidou Strantzali, I.; Antunes, V. (2022). "Ornithopod dinosaur remains from the Papo Seco Formation (lower Barremian, Lusitanian Basin, Portugal): a review of old and new finds". Historical Biology . 35 (11): 2181–2192. doi: 10.1080/08912963.2022.2138372 . ISSN   0891-2963. S2CID   253390618.
  43. ↑ Figueiredo, S.; Dinis, P.; Belo, J.; Rosina, P.; Bachtsevanidou Strantzali, I. (2017). "A new record of a possible ornithopod footprint from the Lower Cretaceous of Cabo Espichel (Sesimbra, Portugal)" (PDF). Bollettino della Società Paleontologica Italiana. 56 (2): 217–231. doi:10.4435/BSPI.2017.20. ISSN   0375-7633. S2CID   44179652.
  44. ↑ Figueiredo, S. D.; Rosina, P.; Figuti, L. (2015). "Dinosaurs and other vertebrates from the Papo-Seco Formation (Lower Cretaceous) of southern Portugal". Journal of Iberian Geology . 41 (3): 301–314. doi:10.5209/rev_JIGE.2015.v41.n3.47828. ISSN   1886-7995. S2CID   53970915.
  45. ↑ Figueiredo, S.; Bachtsevanidou Strantzali, I.; Rosina, P.; Gomes, M. (2016). "New Data about the Paleo Environment of the Papo-Seco Formation (Lower Cretaceous) of Southern Portugal" (PDF). Journal of Environmental Science and Engineering. 41 (3): 463–470.
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