Bathysuchus
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''Bathysuchus'' ("deep water crocodile") is an
extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
genus of teleosaurid
thalattosuchia Thalattosuchia is a clade of mostly 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 member ...
n from
Late Jurassic The Late Jurassic is the third Epoch (geology), epoch of the Jurassic Period, and it spans the geologic time scale, geologic time from 161.5 ± 1.0 to 143.1 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic stratum, strata.Owen ...
(
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 154.8 ±0.8 Ma and 149.2 ±0.7 Ma (million years ago). The Kimmeridgian follows the Oxfordian ...
) deep water marine deposits in
England England is a Countries of the United Kingdom, country that is part of the United Kingdom. It is located on the island of Great Britain, of which it covers about 62%, and List of islands of England, more than 100 smaller adjacent islands. It ...
and
France France, officially the French Republic, is a country located primarily in Western Europe. Overseas France, Its overseas regions and territories include French Guiana in South America, Saint Pierre and Miquelon in the Atlantic Ocean#North Atlan ...
. ''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. The word ''pelagic'' is derived . The pelagic zone can be thought of as an imaginary cylinder or water column between the sur ...
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 The sea level has been rising from the end of the last ice age, which was around 20,000 years ago. Between 1901 and 2018, the average sea level rose by , with an increase of per year since the 1970s. This was faster than the sea level had e ...
during the Kimmeridgian, as its earlier relatives such as ''
Teleosaurus ''Teleosaurus'' (from , 'perfect' and , 'lizard') is an extinct genus of teleosaurid crocodyliform found in the Middle Jurassic Calcaire de Caen Formation of France. It was approximately in length. The holotype is MNHN AC 8746, a quarter of ...
'' 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 taxon, extant and extinct reptiles and amphibians, including lizards, crocodilians, frogs, Temnospondyli, ...
. 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 mammals h ...
; 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 * M ...
, 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' ...
in 1871 on the basis of NHMUK PV OR 43086, an incomplete snout from the
Kimmeridge Clay Formation The Kimmeridge Clay is a sedimentary rock, 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 (geology), form ...
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 ( ; Archaism, archaically: Dorsetshire , ) is a Ceremonial counties of England, ceremonial county in South West England. It is bordered by Somerset to the north-west, Wiltshire to the north and the north-east, Hampshire to the east, t ...
, 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 a British naturalist, geologist and writer of numerous books on natural history. He was known for his contributions to zoology, paleontology, and biogeography. He worked extensively in cata ...
referred it to ''
Steneosaurus ''Steneosaurus'' (from , 'narrow' and , '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 wastebasket taxon for thalattosuchian ...
'', 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 classical antiquity, ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Greek ...
''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 rock, 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 (geology), form ...
. 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 () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical data ...
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 Machimosauridae is an extinct family of teleosauroid thalattosuchian crocodyliforms. Text was copied from this source, which is available under Creative Commons Attribution 4.0 International License The family was first identified in 2016, w ...
,
Teleosauroidea Teleosauroidea is an extinct superfamily of thalattosuchian crocodyliforms living from the Early Jurassic to the Early Cretaceous. It is phylogenetically defined by Mark T. Young and colleagues in 2024 in the ''PhyloCode'' as "the largest clade ...
and Teleosaurinae). ''Bathysuchus'' was found to be a member of 'clade T' most closely related to '' Sericodon'', 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 ...
, and then '' Aeolodon''; Foffa ''et al.'' (2019) initially believed that ''Bathysuchus'' was more closely 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 metriorhynchids and
ophthalmosaurid Ophthalmosauridae is an extinct family of thunnosaur ichthyosaurs from the Middle Jurassic to the early Late Cretaceous (Bajocian - Cenomanian) worldwide. Almost all ichthyosaurs from the Middle Jurassic onwards belong to the family, until the e ...
ichthyosaurs Ichthyosauria is an taxonomy (biology), order of large extinction, extinct marine reptiles sometimes referred to as "ichthyosaurs", although the term is also used for wider clades in which the order resides. Ichthyosaurians thrived during much of ...
. 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 Ocean The Tethys Ocean ( ; ), also called the Tethys Sea or the Neo-Tethys, was a prehistoric ocean during much of the Mesozoic Era and early-mid Cenozoic Era. It was the predecessor to the modern Indian Ocean, the Mediterranean Sea, and the Eurasia ...
and the margins of continental Europe.


References

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