Bathysuchus Tooth
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''Bathysuchus'' ("deep water crocodile") is an
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
genus of
teleosaurid Teleosauridae is a family of extinct typically marine crocodylomorphs similar to the modern gharial that lived during the Jurassic period. Teleosaurids were thalattosuchians closely related to the fully aquatic metriorhynchoids, but were less ad ...
thalattosuchia Thalattosuchia is a clade of marine crocodylomorphs from the Early Jurassic to the Early Cretaceous that had a cosmopolitan distribution. They are colloquially referred to as marine crocodiles or sea crocodiles, though they are not members of Cro ...
n from
Late Jurassic The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 163.5 ± 1.0 to 145.0 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic strata.Owen 1987. In European lithostratigraphy, the name ...
(
Kimmeridgian In the geologic timescale, the Kimmeridgian is an age in the Late Jurassic Epoch and a stage in the Upper Jurassic Series. It spans the time between 157.3 ± 1.0 Ma and 152.1 ± 0.9 Ma (million years ago). The Kimmeridgian follows the Oxfordian ...
) deep water marine deposits in
England England is a country that is part of the United Kingdom. It shares land borders with Wales to its west and Scotland to its north. The Irish Sea lies northwest and the Celtic Sea to the southwest. It is separated from continental Europe b ...
and
France France (), officially the French Republic ( ), is a country primarily located in Western Europe. It also comprises of Overseas France, overseas regions and territories in the Americas and the Atlantic Ocean, Atlantic, Pacific Ocean, Pac ...
. ''Bathysuchus'' displays features that suggest it was more
pelagic The pelagic zone consists of the water column of the open ocean, and can be further divided into regions by depth (as illustrated on the right). The word ''pelagic'' is derived . The pelagic zone can be thought of as an imaginary cylinder or wa ...
than other teleosaurids (along with its close relatives within the Aeolodontinae), including smoother skull bones and reduced armour plating, similar to the fully marine metriorhynchids. This was possibly an adaptation to
rising sea levels Rising may refer to: * Rising, a stage in baking - see Proofing (baking technique) *Elevation * Short for Uprising, a rebellion Film and TV * "Rising" (''Stargate Atlantis''), the series premiere of the science fiction television program ''Starga ...
during the Kimmeridgian, as its earlier relatives such as ''
Teleosaurus ''Teleosaurus'' (from el, τέλειος , 'perfect' and el, σαῦρος , 'lizard') is an extinct genus of teleosaurid crocodyliform found in the Middle Jurassic Calcaire de Caen Formation of France. It was approximately in length and w ...
'' were suited for shallow coasts and lagoon environments.


Description

''Bathysuchus'' is a very long-snouted (longirostrine) teleosaurid, currently known only from several snouts, the back of the skull, teeth, and a few
osteoderms Osteoderms are bony deposits forming scales, plates, or other structures based in the dermis. Osteoderms are found in many groups of extant and extinct reptiles and amphibians, including lizards, crocodilians, frogs, temnospondyls (extinct ...
. It is distinguished from other derived teleosaurids in the following characteristics: strongly ventrally deflected anterior margin of the
premaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammal has b ...
; five premaxillary
alveoli Alveolus (; pl. alveoli, adj. alveolar) is a general anatomical term for a concave cavity or pit. Uses in anatomy and zoology * Pulmonary alveolus, an air sac in the lungs ** Alveolar cell or pneumocyte ** Alveolar duct ** Alveolar macrophage * ...
, the caudal-most being considerably reduced in size; anterodorsally oriented external nares; conical teeth bearing carinae which are only visible on the apical third of the crown. Other salient diagnostic characters can be found in the tooth count, shape of external nares (which is roughly '8'-shaped) and strong deflection of the premaxilla down and outwards. The osteoderms of ''Bathysuchus'' are unlike those of other teleosaurids. Teleosaurid osteoderms are typically heavily ornamented with irregular or tear-drop shaped pits radiating in a starburst pattern, while those of ''Bathysuchus'' have small circular pits arranged in alternating rows (shared with ''Aeolodon''). All but the single caudal osteoderm also lack keels. The osteoderms are also generally smaller and thinner than typical of teleosaurids.


Discovery

''Bathysuchus'' was originally described as a new species of ''Teleosaurus'', ''T. megarhinus'', by
John Whitaker Hulke John Whitaker Hulke FRCS FRS FGS (6 November 1830 – 19 February 1895) was a British surgeon, geologist and fossil collector. He was the son of a physician in Deal, who became a Huxleyite despite being deeply religious. Hulke became Huxley's ...
in 1871 on the basis of NHMUK PV OR 43086, an incomplete snout from the
Kimmeridge Clay Formation The Kimmeridge Clay is a sedimentary deposit of fossiliferous marine clay which is of Late Jurassic to lowermost Cretaceous age and occurs in southern and eastern England and in the North Sea. This rock formation is the major source rock for North ...
of
Kimmeridge Bay Kimmeridge Bay () is a bay on the Isle of Purbeck, a peninsula on the English Channel coast in Dorset, England, close to and southeast of the village of Kimmeridge, on the Smedmore Estate. The area is renowned for its fossils, with The Etche ...
in
Dorset Dorset ( ; archaically: Dorsetshire , ) is a county in South West England on the English Channel coast. The ceremonial county comprises the unitary authority areas of Bournemouth, Christchurch and Poole and Dorset (unitary authority), Dors ...
, England. In his 1888 catalog of extinct reptiles and amphibians preserved in the Natural History Museum,
Richard Lydekker Richard Lydekker (; 25 July 1849 – 16 April 1915) was an English naturalist, geologist and writer of numerous books on natural history. Biography Richard Lydekker was born at Tavistock Square in London. His father was Gerard Wolfe Lydekker, ...
referred it to ''
Steneosaurus ''Steneosaurus'' (from el, στενός , 'narrow' and el, σαῦρος , 'lizard') is a dubious genus of teleosaurid crocodyliform from the Middle or Late Jurassic ( Callovian or early Oxfordian) of France. The genus has been used as a w ...
'', as ''S. megarhinus''. Delair (1958) did not accept the attribution of ''T. megarhinus'' to ''Steneosaurus'', and a 2005 cladistic recovered the species as closer to ''Teleosaurus'' than to other species traditionally assigned to ''Steneosaurus''.Mueller-Töwe, I. J. 2005. Phylogenetic relationships of the Thalattosuchia. Zitteliana A45: 211-213. A preliminary report from 2015 indicated that ''"Steneosaurus" megarhinus'' is distinct from all other Middle-Late Jurassic teleosaurids and deserved its own genus.Foffa D, Young MT, Brusatte SL, Steel L. (2015) New specimen and revision of the late Jurassic teleosaurid 'Steneosaurus’ megarhinus. PeerJ PrePrints 3:e1351v1 https://doi.org/10.7287/peerj.preprints.1351v1 This study was followed up in 2019 by a re-description of all the specimens attributed to the species, which were assigned to a new distinct genus, ''Bathysuchus megarhinus''. The generic name ''Bathysuchus'' comes from the
Ancient Greek Ancient Greek includes the forms of the Greek language used in ancient Greece and the ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Dark Ages (), the Archaic peri ...
''βαθύς'' (''bathus'') for 'deep' and ''tσoῦχος'' (''soûkhos'') for 'crocodile', with the intended translation of "deep water crocodile", referring to its inferred lifestyle. Three specimens are known for ''Bathysuchus'': the holotype, NHMUK PV OR 43086, and DORCM G..5067i-v are both incomplete snouts from Kimmeridge Bay, Dorset, part of the deep water
Kimmeridge Clay Formation The Kimmeridge Clay is a sedimentary deposit of fossiliferous marine clay which is of Late Jurassic to lowermost Cretaceous age and occurs in southern and eastern England and in the North Sea. This rock formation is the major source rock for North ...
. An additional skull from the slightly older Aulacostephanus eudoxus Sub-Boreal ammonite Zone of “La Crouzette”, Francoulès, is known from Quercy, France.


Classification

''Bathysuchus'' has consistently been recognised as a teleosauroid thalattosuchian, although its affinities to ''Teleosaurus'' and ''Steneosaurus'' were only based on superficial features of the skull. The anatomical distinctiveness of ''Bathysuchus'' was recognised in 2019 by Foffa and colleagues through a thorough re-description of the known specimens. This also allowed them to include it in an updated
phylogenetic analysis In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
of thalattosuchians incorporating the newly recognised traits. Their analysis found Teleosauroidea to be split into two major groups, one containing ''Teleosaurus'' and other generally long-snouted genera that they referred to as 'clade T' (now Aeolodontinae), and another of ''Steneosaurus edwardsi'', the durophagous Machimosaurini, and their closest relatives that they called 'clade S' (now Machimosaurini,
Teleosauroidea Teleosauroidea is an extinct superfamily of thalattosuchian crocodyliforms living from the Early Jurassic to the Early Cretaceous The Early Cretaceous ( geochronological name) or the Lower Cretaceous (chronostratigraphic name), is the earlier o ...
and Teleosaurinae). ''Bathysuchus'' was found to be a member of 'clade T' most closely related to ''
Sericodon ''Sericodon'' is an extinct genus of teleosaurid crocodyliform from the Late Jurassic (Tithonian) of Germany and Switzerland. The genus contains a single species, ''S. jugleri''. ''Sericodon'' was placed in 'Clade T' (Aeolodontinae) and was foun ...
'', its
sister taxon In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and t ...
, and then ''
Aeolodon ''Aeolodon'' is an extinct genus of teleosaurid crocodyliform reptile from the Late Jurassic (Tithonian) of Germany and France that was initially named as a species of ''Crocodylus'' in 1814. Although previously synonymized with ''Steneosaurus'' ...
''; Foffa ''et al.'' (2019) initially believed that ''Bathysuchus'' was more clocely related to ''Aeolodon'' than to ''Sericodon'', but Johnson ''et al.'' (2020) have proven otherwise. Their results are shown simplified below: This simplified cladogram by Johnson ''et al.'' (2020) shows the updated location of ''Bathysuchus'' within Teleosauridae and Aeolodontinae:


Palaeobiology

''Bathysuchus'' was identified as sharing several characteristics with ''Aeolodon'' and metriorhynchids that suggest it was more pelagic than other teleosaurids. Anatomically, ''Bathysuchus'' shares with them the reduced ornamentation of the skull, as well as smaller, less sculpted osteoderms. It is unknown if it also had proportionately smaller limb bones, as known from its close relative ''Aeolodon''. The reduction of the osteoderms in size and complexity is suggested to be related to thermoregulation, as the vascularised osteoderms of crocodylomorphs are thought to radiate heat into the body. In pelagic marine reptiles basking is not as necessary as the ambient water temperatures fluctuate less, and a similar shift may have occurred in the lineage leading to ''Bathysuchus'' and its relative ''Aeolodon'' that allowed them to reduce their osteoderm coverage. The presence of ''Bathysuchus'' fossils in the Kimmeridge Clay is also significant, as this represents a deep water environment on the outer shelf with depths estimated to be around 150—200 metres, and was associated with deep water fauna including large pliosaurs,
geosaurine Geosaurinae is a subfamily of metriorhynchid crocodyliforms from the Middle Jurassic to the Early Cretaceous (Bathonian - Aptian) of Europe, North America and South America. Named by Richard Lydekker, in 1889, it contains the metriorhynchid ...
metriorhynchids and ophthalmosaurid
ichthyosaurs Ichthyosaurs (Ancient Greek for "fish lizard" – and ) are large extinct marine reptiles. Ichthyosaurs belong to the order known as Ichthyosauria or Ichthyopterygia ('fish flippers' – a designation introduced by Sir Richard Owen in 1842, alt ...
. This environment was part of the Jurassic Sub-Boreal Seaway, which was noted to show a decline in teleosaurid diversity across the Middle—Late Jurassic boundary as global sea levels rose. It's possible then that ''Bathysuchus'' and ''Aeolodon'' represent an attempt at shallow-water teleosaurids to adapt to a more pelagic lifestyle as the local environment changed, while other teleosaurids became restricted to the Tethys seaway and the margins of continental Europe.


References

{{Taxonbar, from=Q41168134 Fossil taxa described in 1871 Late Jurassic crocodylomorphs Thalattosuchians Fossil taxa described in 2019 Prehistoric pseudosuchian genera