Baryonychines
Temporal range: Early Cretaceous, Possible Santonian record
Skeletal diagram of genera Suchomimus and Baryonyx
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Clade: Saurischia
Clade: Theropoda
Family: †Spinosauridae
Subfamily: †Baryonychinae
Charig & Milner, 1986
Type species
†Baryonyx walkeri
Charig & Milner, 1986
Subgroups

Baryonychinae is an extinct clade or subfamily of spinosaurids from the Early Cretaceous (Valanginian-Albian) of Britain, Portugal, and Niger. In 2023, it consisted of seven genera: Ceratosuchops, Cristatusaurus, Protathlitis, Riparovenator, Suchomimus, Suchosaurus, and Baryonyx, the nominal genus. The clade was named by Charig & Milner in 1986 and defined by Sereno et al. in 1998 and Holtz et al. in 2004 as all taxa more closely related to Baryonyx walkeri than to Spinosaurus aegyptiacus.[1]

Baryonychines were large, bipedal predators with elongated, crocodile-like skulls and lower jaw tips fanning out into rosettes bearing conical, often unserrated, teeth, and a distinct premaxillary notch. They possessed robust forelimbs supporting three-fingered hands with an enlarged first digit claw, to which the subfamily name indirectly refers. Members of this group, unlike the more derived Spinosaurinae, sported only low sails or none at all.

History of discovery

In 1820, paleontologist Gideon Mantell discovered numerous fossil teeth from the Wadhurst Clay Formation of Britain.[2] These were in 1841 named Suchosaurus cultridens by paleontologist Richard Owen, and were identified as a crocodilian.[3] A second species, Suchosaurus girardi, was named in 1897 by Henry-Emile Sauvage from the Papo Seco Formation of Portugal.[4] It was not until the description of Baryonyx in 1986 that these remains were identified as spinosaurid teeth and Suchosaurus was placed in the Spinosauridae.

The second described representative of the subfamily was unearthed in 1983 by fossil collector William John Walker, within the Smokejacks Pit, Weald Clay Formation, Surrey, England. This initiated the involvement of the Natural History Museum of London, discovering a 65% complete skeleton: NHMUK VP R9951. In 1986, the specimen was published and described by Alan J. Charig and Angela C. Milner as Baryonyx walkeri,[5] with a more detailed monograph published in 1997.[6][7] Teeth, hand bones, and vertebrae attributed to the genus were later discovered in 1998 and 2004.[8] The same year, Spinosaurinae and Baryonychinae were cladistically defined by Holtz and colleagues.[1]

In 1973, paleontologist Philippe Taquet discovered specimen MNHN GDF 266 consisting of two premaxillae, a partial maxilla, and a dentary, along with several similar remains from Gadoufaoua, Elrhaz Formation, Niger. They were in 1998 described as the holotype and paratypes of Cristatusaurus lapparenti,[9] although after several inconclusive debates on whether or not the specimen represents the then newly described Baryonyx.[5][6]

In 1997, Paul Sereno and colleagues discovered a ~67% complete skeleton, MNN GDF500, in Gadoufaoua. The next year, Sereno et al. described the specimen as the new baryonychine Suchomimus tenerensis.[10] The species was also the subject of synonymy disputes over Cristatusaurus and Baryonyx throughout the 1990s and 2000s.[11][12]

From 2013 to 2020, several spinosaurid fragments were discovered from the Wessex Formation in Britain. In 2021, Barker et al. described these specimens, IWCMS 2014.95.5, IWCMS 2021.30, IWCMS 2014.95.1-3, IWCMS 2014.95.4, IWCMS 2014.95.6, IWCMS 2014.96.1, 2; 2020.448.1, 2, and IWCMS 2014.96.3, as the two new genera Ceratosuchops inferodios and Riparovenator milnerae. The study defined a new subclade within Baryonychinae: Ceratosuchopsini, defined as all taxa more closely related to Ceratosuchops inferodios than to Baryonyx walkeri. According to the performed analysis and clade definition, the group contains Ceratosuchops, Riparovenator, and Suchomimus.[13][14]

In 2023, Santos-Cubedo et al. described a new genus and species of baryonychine, Protathlitis cinctorrensis, based on a partial skeleton, recovered from the Arcillas de Morella Formation of Castellón, Spain.[15]

Numerous undescribed specimens have been found as well, such as XMDFEC V0010, described in 2010 by Hone, Xu and Wang; a theropod tooth from the Majiacun Formation of China dated to ~86-85 million years ago. They interpreted the tooth as belonging to a probable baryonychine, which would expand the temporal range of Baryonychinae, and Megalosauroidea as a whole, well into the Late Cretaceous.[16][17] However, this tooth lacks spinosaurid synapomorphies.[18] Other undescribed specimens include UT-JAW2 from Libya[19] and various remains from Spain, such as Baryonychinae indet.[20] from Vallipón, Castellote, Spain, LAD0-2 from Spain[21] CMP-2 from Cantera del Mas de la Parreta 1,[22] from Castilla y Leon,[23] from the El Castellar Formation,[24] Tenadas del Jabali,[25] and from Mas de Curolles.[26]

Description

Reconstructed forelimb and hand of Suchomimus, Museum of Ancient Life, Utah

Even though baryonychines were on average smaller than the more advanced spinosaurines, they were still decently large compared to theropods in general. The smallest members, Ceratosuchops and Riparovenator, are estimated at 8–9 m (26–29 ft) and ~1.4-2 tons, while the largest member, Suchomimus, is estimated to measure 9.5–11 m (31 –36 ft) in length and 3–4.7 tons in weight.[27][7][28] Members of this family, like other spinosaurids, sported robust forelimbs with large, three-clawed hands.[29][30] However, unlike the more derived spinosaurines, these animals possessed small sails, as in Suchomimus, Riparovenator, and Ceratosuchops (for the latter two, sails were assumed judging by their phylogenetic position);[10][13] some with only the vertebrae of the sacral region being elongated, or none at all, as in Baryonyx.[6]

Skull

Closeup of the teeth of Suchomimus

Like most other spinosaurids, baryonychines had a very elongated skull compared to other theropods.[29] Furthermore, even in comparison to the spinosaurines their skulls were long. Perhaps the most proportionally-lengthened skull belongs to Suchomimus. In its very crocodilian skull, there is little to no concavity or convexity from the front (premaxillae) to the back (parietals) of the skull, unlike in spinosaurines and most other theropods.[10] Baryonychines possess reduced antorbital fenestrae in comparison to other theropods,[31] with most of the front snout being solid bone formed by the premaxillae and maxillae. The tips of the premaxillae were expanded into a "terminal rosette" holding enlarged, recurved teeth.[32] Behind these expansions, baryonychines featured a subnarial gap complex where dentary teeth fit into, and, further posterior, a dentary gap that the large anteriormost maxillary teeth indented.[29] These animals also bore reduced, narrow premaxillary crests.[33]

Classification

The subfamily Baryonychinae was first implicitly named in 1986 by Alan J. Charig and Angela Milner when they named the family "Baryonychidae" to include Baryonyx. Those who name families are considered the nominal authors of the subfamilies also. The family Baryonychidae was invalidated when Baryonyx was found to be a spinosaurid.[34] Milner stated that it was likely that Suchosaurus belonged to this subfamily as well.

In 1998, the newly described Cristatusaurus was agreed to be very closely related to, if not identical to, Baryonyx by Charig & Milner, 1986 & 1997,[5][6] Sereno, 1998,[10] and Rauhut, 2003.[12] Later in 1998, Sereno et al. described the genus Suchomimus, and placed it in Baryonychinae along with Baryonyx. They defined the clade's distinguishing characteristics as "numerous small-sized, serrated teeth in the dentary behind the terminal rosette and deeply-keeled anterior dorsal vertebrae."[10] By 2002, the conclusion was that the subfamily contained the genera Baryonyx, Cristatusaurus, Suchomimus, and Suchosaurus.[35] The clade was phylogenetically defined by Holtz et al. as all taxa more closely related to Baryonyx walkeri than to Spinosaurus aegyptiacus.[1]

In the 2012 description of Ichthyovenator, Allain et al. found it to belong to this subfamily,[36] although almost all subsequent studies have found otherwise, placing it in Spinosaurinae.[13][37][38]

Up until 2021, with Cristatusaurus and Suchosaurus being considered too incomplete and dubious, only the baryonychines Suchomimus and Baryonyx have been included in phylogenetic analyses, nearly always finding them to be sister genera in Baryonychinae, such as in the analysis performed by Arden et al. in 2018, shown below.[37]

Spinosauridae

Praia das Aguncheiras taxon

Baryonychinae
Baryonyx walkeri
Suchomimus tenerensis
Spinosaurinae
Siamosaurus suteethorni

Eumeralla taxon

Ichthyovenator laosensis
Irritator challengeri
Oxalaia quilombensis
Spinosaurini

Gara Samani taxon

Sigilmassasaurus brevicollis
Spinosaurus aegyptiacus

In 2021, Chris Barker, Hone, Darren Naish, Andrea Cau, Lockwood, Foster, Clarkin, Schneider, and Gostling described two new spinosaurid species, Ceratosuchops inferodios and Riparovenator milnerae, and placed them well-supportedly in Baryonychinae. They placed them within the newly created tribe Ceratosuchopsini alongside Suchomimus. Barker et al. diagnosed three autapomorphies to distinguish the clade: "1. postorbital facet of frontal dorsoventrally thick (height more than 40% of length) and excavated by a deep, longitudinal slot; 2. well-defined and strongly curved anterior margins of supratemporal fossa; 3. occipital surface of the basisphenoid collateral oval scars excavated." Members of this clade range in length from 7.7 to 9.5 m (25.3 to 31 ft).[39] The results of their Bayesian analysis appear below:[13]

Megalosauridae

Spinosauridae

Vallibonavenatrix

Baryonychinae
Iberospinus
Baryonyx
Ceratosuchopsini
Suchomimus
Riparovenator
Ceratosuchops
Spinosaurinae
Camarillasaurus
Ichthyovenator
Irritator
Spinosaurini
Sigilmassasaurus

"Spinosaurus B" (=?cf. Sigilmassasaurus/Spinosaurus sp.)

MSNM-V4047 (=?cf. Sigilmassasaurus/Spinosaurus sp.)

FSAC-KK11888 (Spinosaurus sp.)

Spinosaurus holotype

Paleobiology

Feeding

Baryonychine teeth are small and recurved with little to no serrations, resembling those of crocodiles. These are considered adaptations for piscivory, as numerous recurved teeth aid in holding a struggling slippery animal within the jaws and down the throat; rather than the serrated teeth in most other theropods which are generalized for cutting and ripping flesh.[40] Vullo et al, 2016 likened the cranial evolution and adaptations to piscivory in spinosaurids to those of the Muraenesocidae, a modern family of predatory eels with a similarly evolved skull.

Baryonyx as one of the most complete representatives of the group shows evidence of a generalist behavior. One recorded instance is the holotype of Baryonyx found with both fish as well as a juvenile iguanodontid contents within the stomach region.[41] Another instance is pointed out by a 2016 study by the Belgian palaeontologist Christophe Hendrickx and colleagues. They found that adult spinosaurs could displace their mandibular rami (halves of the lower jaw) sideways when the jaw was depressed, which allowed the pharynx (opening that connects the mouth to the oesophagus) to be widened. This jaw-articulation is similar to that seen in pterosaurs and living pelicans, and would likewise have allowed spinosaurids to swallow large prey such as fish and other animals.

They also reported that the possible Portuguese Iberospinus[42] (formerly seen as Baryonyx) fossils were found associated with isolated Iguanodon teeth, and listed it along with other such associations as support for opportunistic feeding behaviour in spinosaurs.[33]

References

  1. 1 2 3 Holtz, Thomas R.; Molnar, Ralph E.; Currie, Philip J. (2019). "Basal Tetanurae". In Weishampel, David B.; Dodson, Peter; Osmólska, Halszka (eds.). The Dinosauria, Second Edition. pp. 71–110. doi:10.1525/9780520941434-009. ISBN 978-0-520-94143-4. S2CID 226816827.
  2. ↑ Mantell, G.A., 1822, The fossils of the South Downs or Illustrations of the Geology of Sussex, London, Rupton Relfe
  3. ↑ Owen, Richard (1840). Odontography; or, A treatise on the comparative anatomy of the teeth; their physiological relations, mode of development, and microscopic structure, in the vertebrate animals. H. Baillière. pp. 1–32. OCLC 5294428.
  4. ↑ Sauvage, H. E (1897). Vertébrés fossiles du Portugal: contributions à l'étude des poissons et des reptiles du jurassique et du crétacique [Fossil vertebrates from Portugal: contributions to the study of Jurassic and Cretacic fish and reptiles] (in French). l'Académie royale des Sciences. OCLC 1015745741.
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  6. 1 2 3 4 Charig, A. J.; Milner, A. C. (1997). "Baryonyx walkeri, a fish-eating dinosaur from the Wealden of Surrey". Bulletin of the Natural History Museum of London. 53: 11–70.
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  13. 1 2 3 4 Barker, C.T.; Hone, D.; Naish, D.; Cau, A.; Lockwood, J.; Foster, B.; Clarkin, C.; Schneider, P.; Gostling, N. (2021). "New spinosaurids from the Wessex Formation (Early Cretaceous, UK) and the European origins of Spinosauridae". Scientific Reports. 11 (1): 19340. Bibcode:2021NatSR..1119340B. doi:10.1038/s41598-021-97870-8. PMC 8481559. PMID 34588472.
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  16. ↑ Hone, David; Xu, Xing; Wang, Deyou (January 2010). "中国河南晚白垩世地层一枚可能属于重爪龙亚科(兽脚亚目：棘龙科)的牙齿化石" [A probable Baryonychine (Theropoda: Spinosauridae) tooth from the Upper Cretaceous of Henan Province, China]. Vertebrata PalAsiatica (in Chinese). 48: 19–26.
  17. ↑ Majiacun Formation at Fossilworks.org
  18. ↑ Katsuhiro, Kubota; Yuji, Takakuwa; Yoshikazu, Hasegawa (2017). "Second discovery of a spinosaurid tooth from the Sebayashi Formation (Lower Cretaceous), Kanna Town, Gunma Prefecture, Japan" (PDF). Bulletin of the Gunma Museum of Natural History. 21: 1–6.
  19. ↑ Le Loeuff, Jean; Métais, Eddy; Dutheil, Didier B.; Rubino, Jean Loup; Buffetaut, Eric; Lafont, François; Cavin, Lionel; Moreau, Fabrice; Tong, Haiyan; Blanpied, Christian; Sbeta, Ali (September 2010). "An Early Cretaceous vertebrate assemblage from the Cabao Formation of NW Libya" (PDF). Geological Magazine. 147 (5): 750–759. Bibcode:2010GeoM..147..750L. doi:10.1017/S0016756810000178. S2CID 130450882.
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  28. ↑ Campione, Nicolás E.; Evans, David C.; Brown, Caleb M.; Carrano, Matthew T. (2014). "Body mass estimation in non-avian bipeds using a theoretical conversion to quadruped stylopodial proportions". Methods in Ecology and Evolution. 5 (9): 913–923. doi:10.1111/2041-210X.12226. S2CID 84317234.
  29. 1 2 3 Hone, David William Elliott; Holtz, Thomas Richard (June 2017). "A Century of Spinosaurs - A Review and Revision of the Spinosauridae with Comments on Their Ecology". Acta Geologica Sinica - English Edition. 91 (3): 1120–1132. doi:10.1111/1755-6724.13328. S2CID 90952478.
  30. ↑ S., Paul, Gregory (2016-10-25). The Princeton field guide to dinosaurs (2nd ed.). Princeton, N.J. ISBN 9781400883141. OCLC 954055249.{{cite book}}: CS1 maint: location missing publisher (link) CS1 maint: multiple names: authors list (link)
  31. ↑ Rayfield, Emily J.; Milner, Angela C.; Xuan, Viet Bui; Young, Philippe G. (12 December 2007). "Functional morphology of spinosaur 'crocodile-mimic' dinosaurs". Journal of Vertebrate Paleontology. 27 (4): 892–901. doi:10.1671/0272-4634(2007)27[892:fmoscd]2.0.co;2. S2CID 85854809.
  32. ↑ Sales, M.A.F.; Schultz, C.L. (2017). "Spinosaur taxonomy and evolution of craniodental features: Evidence from Brazil". PLOS ONE. 12 (11): e0187070. Bibcode:2017PLoSO..1287070S. doi:10.1371/journal.pone.0187070. PMC 5673194. PMID 29107966.
  33. 1 2 Hendrickx, Christophe; Mateus, Octávio; Buffetaut, Eric (6 January 2016). "Morphofunctional Analysis of the Quadrate of Spinosauridae (Dinosauria: Theropoda) and the Presence of Spinosaurus and a Second Spinosaurine Taxon in the Cenomanian of North Africa". PLOS ONE. 11 (1): e0144695. Bibcode:2016PLoSO..1144695H. doi:10.1371/journal.pone.0144695. PMC 4703214. PMID 26734729.
  34. ↑ Buffetaut, Eric (November 2007). "The spinosaurid dinosaur Baryonyx (Saurischia, Theropoda) in the Early Cretaceous of Portugal" (PDF). Geological Magazine. 144 (6): 1021–1025. Bibcode:2007GeoM..144.1021B. doi:10.1017/S0016756807003883. S2CID 130212901.
  35. ↑ Buffetaut, Eric; Ouaja, Mohamed (1 September 2002). "A new specimen of Spinosaurus (Dinosauria, Theropoda) from the Lower Cretaceous of Tunisia, with remarks on the evolutionary history of the Spinosauridae" (PDF). Bulletin de la Société Géologique de France. 173 (5): 415–421. doi:10.2113/173.5.415. hdl:2042/216.
  36. ↑ Allain, Ronan; Xaisanavong, Tiengkham; Richir, Philippe; Khentavong, Bounsou (May 2012). "The first definitive Asian spinosaurid (Dinosauria: Theropoda) from the early cretaceous of Laos". Naturwissenschaften. 99 (5): 369–377. Bibcode:2012NW.....99..369A. doi:10.1007/s00114-012-0911-7. PMID 22528021. S2CID 2647367.
  37. 1 2 Arden, T.M.S.; Klein, C.G.; Zouhri, S.; Longrich, N.R. (2018). "Aquatic adaptation in the skull of carnivorous dinosaurs (Theropoda: Spinosauridae) and the evolution of aquatic habits in Spinosaurus". Cretaceous Research. 93: 275–284. doi:10.1016/j.cretres.2018.06.013. S2CID 134735938.
  38. ↑ Malafaia, E.; Miguel Gasulla, J.; Escaso, F.; Narváez, I.; Luis Sanz, J.; Ortega, F. (2019). "A new spinosaurid theropod (Dinosauria: Megalosauroidea) from the late Barremian of Vallibona, Spain: Implications for spinosaurid diversity in the Early Cretaceous of the Iberian Peninsula". Cretaceous Research. 106: 104221. doi:10.1016/j.cretres.2019.104221. S2CID 202189246.
  39. ↑ Paul, G.S., 2010, The Princeton Field Guide to Dinosaurs, Princeton University Press p. 87
  40. ↑ Sues, Hans-Dieter; Frey, Eberhard; Martill, David M.; Scott, Diane M. (19 September 2002). "Irritator challengeri , a spinosaurid (Dinosauria: Theropoda) from the Lower Cretaceous of Brazil". Journal of Vertebrate Paleontology. 22 (3): 535–547. doi:10.1671/0272-4634(2002)022[0535:ICASDT]2.0.CO;2. S2CID 131050889.
  41. ↑ Charig, A. J.; Milner, A. C. (1997). "Baryonyx walkeri, a fish-eating dinosaur from the Wealden of Surrey". Bulletin of the Natural History Museum of London. 53: 11–70.
  42. ↑ Mateus, Octávio; Estraviz-López, Darío (16 February 2022). "A new theropod dinosaur from the early cretaceous (Barremian) of Cabo Espichel, Portugal: Implications for spinosaurid evolution". PLOS ONE. 17 (2): e0262614. Bibcode:2022PLoSO..1762614M. doi:10.1371/journal.pone.0262614. PMC 8849621. PMID 35171930.
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