''Pholidosaurus'' is an extinct

genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...

neosuchia Neosuchia is a clade within Mesoeucrocodylia that includes all modern extant crocodilians and their closest fossil relatives. It is defined as the most inclusive clade containing all Crocodylomorpha, crocodylomorphs more closely related to ''Croc ...

crocodylomorph Crocodylomorpha is a group of pseudosuchian archosaurs that includes the crocodilians and their extinct relatives. They were the only members of Pseudosuchia to survive the end-Triassic extinction. Extinct crocodylomorphs were considerably mor ...

. It is the

type genus In biological taxonomy, the type genus (''genus typica'') is the genus which defines a biological family and the root of the family name. Zoological nomenclature According to the International Code of Zoological Nomenclature, "The name-bearin ...

of the

family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...

Pholidosauridae Pholidosauridae is an extinct family of aquatic neosuchian mesoeucrocodylian crocodylomorphs. Fossils have been found in Europe (Denmark, England, France, Germany, Spain and Sweden), Africa (Algeria, Niger, Mali, Morocco and Tunisia), North Amer ...

. Fossils have been found in northwestern

Germany Germany, officially the Federal Republic of Germany, is a country in Central Europe. It lies between the Baltic Sea and the North Sea to the north and the Alps to the south. Its sixteen States of Germany, constituent states have a total popu ...

. The genus is known to have existed during the

Berriasian In the geological timescale, the Berriasian is an age/ stage of the Early/Lower Cretaceous. It is the oldest subdivision in the entire Cretaceous. It has been taken to span the time between 143.1 ±0.6 Ma and 137.05 ± 0.2 (million years ago) ...

Albian The Albian is both an age (geology), age of the geologic timescale and a stage (stratigraphy), stage in the stratigraphic column. It is the youngest or uppermost subdivision of the Early Cretaceous, Early/Lower Cretaceous epoch (geology), Epoch/s ...

stages of the

Early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name) is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 143.1 ...

. Fossil material found from the Annero and Jydegård Formations in

Skåne Scania ( ), also known by its native name of Skåne (), is the southernmost of the historical provinces of Sweden, provinces () of Sweden. Located in the south tip of the geographical region of Götaland, the province is roughly conterminous w ...

Sweden Sweden, formally the Kingdom of Sweden, is a Nordic countries, Nordic country located on the Scandinavian Peninsula in Northern Europe. It borders Norway to the west and north, and Finland to the east. At , Sweden is the largest Nordic count ...

and on the island of

Bornholm Bornholm () is a List of islands of Denmark, Danish island in the Baltic Sea, to the east of the rest of Denmark, south of Sweden, northeast of Germany and north of Poland. Strategically located, Bornholm has been fought over for centuries. I ...

Denmark Denmark is a Nordic countries, Nordic country in Northern Europe. It is the metropole and most populous constituent of the Kingdom of Denmark,, . also known as the Danish Realm, a constitutionally unitary state that includes the Autonomous a ...

, have been referred to as a mesoeucrocodylian, and possibly represent the genus ''Pholidosaurus''.

Description

An early description of the genus by Lydekker (1888) mentioned that the

orbit In celestial mechanics, an orbit (also known as orbital revolution) is the curved trajectory of an object such as the trajectory of a planet around a star, or of a natural satellite around a planet, or of an artificial satellite around an ...

is slightly smaller than the supratemporal fossa, the

nasals In phonetics, a nasal, also called a nasal occlusive or nasal stop in contrast with an oral stop or nasalized consonant, is an occlusive consonant produced with a lowered velum, allowing air to escape freely through the nose. The vast majorit ...

reach 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 ...

e, and the

vomer The vomer (; ) is one of the unpaired facial bones of the skull. It is located in the midsagittal line, and articulates with the sphenoid, the ethmoid, the left and right palatine bones, and the left and right maxillary bones. The vomer forms ...

appears on the

palate The palate () is the roof of the mouth in humans and other mammals. It separates the oral cavity from the nasal cavity. A similar structure is found in crocodilians, but in most other tetrapods, the oral and nasal cavities are not truly sep ...

. It is similar in appearance to and about as large as the modern

gharial The gharial (''Gavialis gangeticus''), also known as gavial or fish-eating crocodile, is a crocodilian in the family (biology), family Gavialidae and among the longest of all living crocodilians. Mature females are long, and males . Adult males ...

Species

The

type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...

of ''Pholidosaurus'' is ''P. schaumburgensis'', named in 1841 from the Wealden of

Bückeburg Bückeburg (; Northern Low Saxon: ''Bückeborg'') is a town in Lower Saxony, Germany, on the border with North Rhine Westphalia. It is located in the district of Schaumburg close to the northern slopes of the Weserbergland ridge. Bückeburg ha ...

, Germany. ''P. schaumburgensis'' was named on the basis of a natural mold of part of a thorax discovered in around 1830 from the Berriasian Obernkirchen Sandstein. This mould is known as IMGPGö 741-1. The individual that the mould belonged to is thought to have been around in length. ''Macrorhynchus'' is a

junior synonym In taxonomy, the scientific classification of living organisms, a synonym is an alternative scientific name for the accepted scientific name of a taxon. The botanical and zoological codes of nomenclature treat the concept of synonymy differently. ...

of ''Pholidosaurus''. It was named in 1843 from the same stratigraphic unit and region as ''P. schaumbergensis'', with the type species being ''M. meyeri''. Because ''M. meyeri'' bears a strong resemblance to ''Pholidosaurus schaumburgensis'', it is now regarded as a species of ''Pholidosaurus''. It was reassigned to the genus ''Pholidosaurus'' in 1887 by

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 ...

because of this synonymy, and also because the name ''Macrorhynchus'' was preoccupied by a genus of fish named in 1880. ''P. meyeri'' differs from ''P. schaumburgensis'' in that the bar separating the supratemporal fenestrae is rounded, while in the type species it is rounded.

Misassigned species

''Pholidosaurus decipiens'' was erected for a partial cranium, NHMUK 28432, that was originally assigned to the new genus and species ''Petrosuchus laevidens'' by

Richard Owen Sir Richard Owen (20 July 1804 – 18 December 1892) was an English biologist, comparative anatomy, comparative anatomist and paleontology, palaeontologist. Owen is generally considered to have been an outstanding naturalist with a remarkabl ...

in 1878. ''Petrosuchus laevidens'' was based on this cranium and a mandibular ramus called BMNH 41099, both of which were collected from

Swanage Swanage () is a coastal town and civil parish in the south east of Dorset, England. It is at the eastern end of the Isle of Purbeck and one of its two towns, approximately south of Poole and east of Dorchester, Dorset, Dorchester. In the Unit ...

, England. A later study in 1911 concluded that the material belonged to two different species; NHMUK 28432 was reassigned to ''Pholidosaurus'' and NHMUK 41099 was designated the lectotype of ''Petrosuchus laevidens''. The species name ''decipiens'' was coined in reference to Owen's oversight, and ''Petrosuchus'' is now considered a junior synonym of '' Goniopholis simus''. Another species from England, ''P. purbeckensis'', was originally described as a species of ''

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 ...

'' in 1888. The holotype is an almost complete cranium, referred to as DORCM G97, missing the anterior portion of the rostrum. The skull was found from either Swanage or the

Isle of Purbeck The Isle of Purbeck is a peninsula in Dorset, England. It is bordered by water on three sides: the English Channel to the south and east, where steep cliffs fall to the sea; and by the marshy lands of the River Frome, Dorset, River Frome and Poo ...

(hence the species name), although the exact locality from which the skull originated is not specified by the author of the original description. This material was also once referred to ''Macrorhynchus''. The author of the 1888 description considered ''S. purbeckensis'' an intermediate form between ''Steneosaurus'' and ''

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 ...

''. However, in 2002, a new study showed that ''S. purbeckensis'' was conspecific with ''P. decipiens'', creating the new combination ''Pholidosaurus purbeckensis''. Another species of ''Pholidosaurus'', ''P. laevis'', was named in 1913 from Swanage, based on the partial cranium NHMUK R3414. This has been considered a junior synonym of ''P. purbeckensis'' by both Salisbury ''et al.'' (1999) and Salisbury (2002). In an SVPCA abstract, Smith ''et al.'' (2016) noted that ''Pholidosaurus purbeckensis'' is not congeneric with the type species, and instead is closely related to '' Fortignathus'' and members of

Dyrosauridae Dyrosauridae is a family of extinct neosuchian crocodyliforms that lived from the Campanian to the Eocene. Dyrosaurid fossils are globally distributed, having been found in Africa, Asia, Europe, North America and South America. Over a dozen sp ...

.Thomas J. Smith, Lorna Steel, and Mark T. Young, 2016. Re-evaluation of ''Pholidosaurus purbeckensis'' (Crocodyliformes: Tethysuchia) from the Early Cretaceous of England, with implications for the evolution of Pholidosauridae and Dyrosauridae. ''Symposium of Vertebrate Palaeontology and Comparative Anatomy 2016 Abstract Book''. p. 29.

Classification

Richard Lydekker assigned ''Pholidosaurus'' to the family

Goniopholididae Goniopholididae is an extinct family of moderate-sized semi-aquatic neosuchian crocodyliformes. Their bodyplan and morphology are convergent on living crocodilians. They lived across Laurasia (Asia, Europe and North America) between the Middle J ...

in 1887 along with ''

Hylaeochampsa ''Hylaeochampsa'' is an extinct genus of eusuchian crocodylomorphs. It is known only from a partial skull recovered from Barremian-age rocks of the Lower Cretaceous Vectis Formation (Wealden Group) of the Isle of Wight. This skull, BMNH R 177, ...

'', '' Theriosuchus'', ''

Goniopholis ''Goniopholis'' (meaning "angled scale") is an extinct genus of goniopholidid crocodyliform that lived in Europe and North America during the Late Jurassic and Early Cretaceous. Like other goniopholidids, it resembled living crocodilians, and pro ...

'', and ''Petrosuchus'' because the

vertebra Each vertebra (: vertebrae) is an irregular bone with a complex structure composed of bone and some hyaline cartilage, that make up the vertebral column or spine, of vertebrates. The proportions of the vertebrae differ according to their spina ...

e are amphicoelus and the orbit communicates with the lateral

temporal fossa The temporal fossa is a fossa (shallow depression) on the side of the skull bounded by the temporal lines above, and the zygomatic arch below. Its floor is formed by the outer surfaces of four bones of the skull. The fossa is filled by the te ...

. ''Pholidosaurus'' has often been grouped with other longirostrine, or long-snouted, crocodylomorphs, including

dyrosaurids Dyrosauridae is a family (biology), family of extinct neosuchian crocodyliforms that lived from the Campanian to the Eocene. Dyrosaurid fossils are globally distributed, having been found in Africa, Asia, Europe, North America and South America. ...

and

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 ...

ns. Buckley and Brochu (1999) concluded that ''Pholidosaurus'', ''

Sokotosuchus ''Sokotosuchus'' is an extinct genus of dyrosaurid crocodyliform which existed during the Maastrichtian in western Africa. Fossils of the genus were found in the Dukamaje Formation of Nigeria, and some cranial material has possibly been found in ...

'', Dyrosauridae, and Thalattosuchia formed a longirostrine

clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...

that was the

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 ...

Crocodylia Crocodilia () is an Order (biology), order of semiaquatic, predatory reptiles that are known as crocodilians. They first appeared during the Late Cretaceous and are the closest living relatives of birds. Crocodilians are a type of crocodylomorp ...

. However, Thalattosuchia was traditionally considered a more basal clade of crocodylomorphs, being a more basal lineage of

Mesoeucrocodylia Mesoeucrocodylia is the clade that includes Eusuchia and crocodyliforms formerly placed in the paraphyletic group Mesosuchia. The group appeared during the Early Jurassic, and continues to the present day. Diagnosis It was long known that Me ...

than dyrosaurids or ''Pholidosaurus'', both of which were considered neosuchians. The results of the phylogenetic analysis by Buckley and Brochu (1999) were attributed to the similarity in characters associated with snout elongation seen in these crocodylomorphs, even though these characters may have been independently derived in each group. More recent studies have revealed Thalattosuchia as a more basal clade when dyrosaurids are removed from the data set. More recent studies show that ''Pholidosaurus'' is closely related to the Thalattosuchia, with both taxa closely related to a clade containing ''

Terminonaris ''Terminonaris'' is a genus of extinct pholidosaurid crocodyliforms that lived during the Late Cretaceous (Cenomanian and TuronianWu X-C, Russell AP, & Cumbaa SL. 2001. ''Terminonaris'' (Archosauria: Crocodyliformes): new material from Saskatchew ...

'' and the Dyrosauridae. In a phylogenetic analysis conducted by Sereno ''et al.'' (2001), ''Pholidosaurus'' was placed as a distant sister taxon to the other longirostrine crocodylomorphs, with ''Terminonaris'' and the newly named ''

Sarcosuchus ''Sarcosuchus'' (), from Ancient Greek σάρξ (''sárx''), meaning "flesh", and σούχος (''soúkhos''), meaning "crocodile", is an extinct genus of crocodyliform a that lived during the Early Cretaceous, from the late Hauterivian to the ...

'' being closely related to one another and ''

Dyrosaurus ''Dyrosaurus'' is a genus of extinct crocodylomorph that lived during the early Eocene. The name ''Dyrosaurus'' comes from () the Greek for lizard or reptile, and Dyr for Djebel Dyr (mountain) close to where the type species was discovered. It ...

'' being the next closest taxon to the group. The later phylogenetic analysis of Brochu ''et al.'' (2002) again showed that ''Pholidosaurus'' was closely related to Thalattosuchia. In the study, both taxa formed a clade that was the sister taxon to a clade containing ''Sokotosuchus'' and Dyrosauridae. Jouve ''et al.'' (2006) concluded that ''Pholidosaurus'' was closely related thalattosuchians were also included within the family, which would be considered

paraphyletic Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...

without them. Jouve ''et al.'' (2006), like Buckley and Brochu (1999), attributed this result to phylogenetic problems that exist among longirostrine crocodylomorphs due to similarities in their morphology.

References

External links

''Pholidosaurus''
in the

Paleobiology Database The Paleobiology Database (PBDB) is an online resource for information on the distribution and classification of fossil animals, plants, and microorganisms. History The Paleobiology Database originated in the NCEAS-funded Phanerozoic Marine Pale ...

{{Taxonbar, from=Q3381939 Early Cretaceous reptiles of Europe Early Cretaceous crocodylomorphs of Europe Tethysuchia Prehistoric pseudosuchian genera