Pholidosauridae is an extinct

family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...

of aquatic

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 crocodylomorphs more closely related to ''Crocodylus niloticus' ...

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 M ...

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. During Mesozoic and early Cenozoic times, cro ...

Fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...

s have been found in

Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...

(

Denmark ) , song = ( en, "King Christian stood by the lofty mast") , song_type = National and royal anthem , image_map = EU-Denmark.svg , map_caption = , subdivision_type = Sovereign state , subdivision_name = Danish Realm, Kingdom of Denmark ...

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 ...

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 ...

Germany Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe after Russia, and the most populous member state of the European Union. Germany is situated betwe ...

Spain , image_flag = Bandera de España.svg , image_coat = Escudo de España (mazonado).svg , national_motto = ''Plus ultra'' (Latin)(English: "Further Beyond") , national_anthem = (English: "Royal March") , i ...

and

Sweden Sweden, formally the Kingdom of Sweden,The United Nations Group of Experts on Geographical Names states that the country's formal name is the Kingdom of SwedenUNGEGN World Geographical Names, Sweden./ref> is a Nordic country located on ...

Africa Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...

(

Algeria ) , image_map = Algeria (centered orthographic projection).svg , map_caption = , image_map2 = , capital = Algiers , coordinates = , largest_city = capital , relig ...

Niger ) , official_languages = , languages_type = National languagesMali Mali (; ), officially the Republic of Mali,, , ff, 𞤈𞤫𞤲𞥆𞤣𞤢𞥄𞤲𞤣𞤭 𞤃𞤢𞥄𞤤𞤭, Renndaandi Maali, italics=no, ar, جمهورية مالي, Jumhūriyyāt Mālī is a landlocked country in West Africa. Mali ...

Morocco Morocco (),, ) officially the Kingdom of Morocco, is the westernmost country in the Maghreb region of North Africa. It overlooks the Mediterranean Sea to the north and the Atlantic Ocean to the west, and has land borders with Algeria to ...

and

Tunisia ) , image_map = Tunisia location (orthographic projection).svg , map_caption = Location of Tunisia in northern Africa , image_map2 = , capital = Tunis , largest_city = capital , ...

North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...

(

Canada Canada is a country in North America. Its ten provinces and three territories extend from the Atlantic Ocean to the Pacific Ocean and northward into the Arctic Ocean, covering over , making it the world's second-largest country by tot ...

and the

United States The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 states, a federal district, five major unincorporated territorie ...

) and

South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the southe ...

(

Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin America. At and with over 217 million people, Brazil is the world's fifth-largest country by area ...

and

Uruguay Uruguay (; ), officially the Oriental Republic of Uruguay ( es, República Oriental del Uruguay), is a country in South America. It shares borders with Argentina to its west and southwest and Brazil to its north and northeast; while bordering ...

). The pholidosaurids first appeared in the fossil record during the

Bathonian In the geologic timescale the Bathonian is an age and stage of the Middle Jurassic. It lasted from approximately 168.3 Ma to around 166.1 Ma (million years ago). The Bathonian Age succeeds the Bajocian Age and precedes the Callovian Age. Strat ...

stage of the

Middle Jurassic The Middle Jurassic is the second epoch of the Jurassic Period. It lasted from about 174.1 to 163.5 million years ago. Fossils of land-dwelling animals, such as dinosaurs, from the Middle Jurassic are relatively rare, but geological formations co ...

. Jouve & Jalil (2020) described postcranial material of a pholidosaurid from the

Paleocene The Paleocene, ( ) or Palaeocene, is a geological epoch (geology), epoch that lasted from about 66 to 56 mya (unit), million years ago (mya). It is the first epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), E ...

(

Danian The Danian is the oldest age or lowest stage of the Paleocene Epoch or Series, of the Paleogene Period or System, and of the Cenozoic Era or Erathem. The beginning of the Danian (and the end of the preceding Maastrichtian) is at the Cretaceous� ...

) of

Ouled Abdoun Basin The Oulad Abdoun Basin (also known as the Ouled Abdoun Basin or Khouribga Basin) is a phosphate sedimentary basin located in Morocco, near the city of Khouribga. It is the largest in Morocco, comprising 44% of Morocco's phosphate reserves, and a ...

(Morocco), representing the most recent record of the family. The authors also reinterpreted putative

Maastrichtian The Maastrichtian () is, in the ICS geologic timescale, the latest age (uppermost stage) of the Late Cretaceous Epoch or Upper Cretaceous Series, the Cretaceous Period or System, and of the Mesozoic Era or Erathem. It spanned the interval from ...

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

Sabinosuchus ''Sabinosuchus'' (meaning "Sabinas crocodile") is a genus of Mesoeucrocodylian, from the Maastrichtian Escondido Formation of Coahuila, Mexico, with ''Sabinosuchus coahuilensis'' as the type species. First described as a putative dyrosauridae, dy ...

'' as a pholidosaurid, and argued that at least two independent pholidosaurid lineages reached the Maastrichtian, among which one survived the

Cretaceous–Paleogene extinction event The Cretaceous–Paleogene (K–Pg) extinction event (also known as the Cretaceous–Tertiary extinction) was a sudden mass extinction of three-quarters of the plant and animal species on Earth, approximately 66 million years ago. With the ...

. Before the publication of this study it was thought that the family became extinct during the Late

Turonian The Turonian is, in the ICS' geologic timescale, the second age in the Late Cretaceous Epoch, or a stage in the Upper Cretaceous Series. It spans the time between 93.9 ± 0.8 Ma and 89.8 ± 1 Ma (million years ago). The Turonian is preceded by t ...

stage of the

Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', the ...

. ''

Sarcosuchus ''Sarcosuchus'' (; ) is an extinct genus of crocodyliform and distant relative of living crocodilians that lived during the Early Cretaceous, from the late Hauterivian to the early Albian, 133 to 112 million years ago of what is now Africa and ...

'' is one of the best known pholidosaurs. It is believed to have attained lengths of up to 9.5 m and weighed up to 4.3 metric tons. Related to ''Sarcosuchus'', ''

Chalawan thailandicus ''Chalawan'' (from th, ชาละวัน ) is an extinct genus of pholidosaurid mesoeucrocodylian known from the Late Jurassic or Early Cretaceous Phu Kradung Formation of Nong Bua Lamphu Province, northeastern Thailand. It contains a sing ...

'' could reached more than in length. One

genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...

, '' Suchosaurus'', once thought to be a pholidosaur, has since been shown to be a

spinosaurid The Spinosauridae (or spinosaurids) are a clade or family of tetanuran theropod dinosaurs comprising ten to seventeen known genera. They came into prominence during the Cretaceous period. Spinosaurid fossils have been recovered worldwide, includi ...

theropod Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally c ...

dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...

(''incertae sedis'' within

Baryonychinae Baryonychinae is an extinct clade or subfamily of spinosaurids from the Early Cretaceous (Valanginian-Albian) of Britain, Portugal, and Niger. In 2021, it consisted of six genera: ''Ceratosuchops'', ''Cristatusaurus'', ''Riparovenator'', ''Suchomi ...

; possibly a

junior synonym The Botanical and Zoological Codes of nomenclature treat the concept of synonymy differently. * In botanical nomenclature, a synonym is a scientific name that applies to a taxon that (now) goes by a different scientific name. For example, Linna ...

of ''

Baryonyx ''Baryonyx'' () is a genus of theropod dinosaur which lived in the Barremian stage of the Early Cretaceous period, about 130–125 million years ago. The first skeleton was discovered in 1983 in the Smokejack Clay Pit, of Surrey, England, in se ...

''). The

Cenomanian The Cenomanian is, in the ICS' geological timescale, the oldest or earliest age of the Late Cretaceous Epoch or the lowest stage of the Upper Cretaceous Series. An age is a unit of geochronology; it is a unit of time; the stage is a unit in the s ...

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 Saskatche ...

'' was the Pholidosaurid species that appeared to be the most common during the

Phylogeny

Pholidosauridae is usually considered to be most closely related to the

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

. However, the relationship between these families is not fully understood. Pholidosauridae might be

monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...

paraphyletic In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...

or even a

polyphyletic A polyphyletic group is an assemblage of organisms or other evolving elements that is of mixed evolutionary origin. The term is often applied to groups that share similar features known as homoplasies, which are explained as a result of converg ...

in relation to Dyrosauridae. For example, Fortier, Perea & Schultz (2011) found the family to be monophyletic, and include to main lineages: the ''Elosuchus''–''Meridiosaurus'' lineage and the ''Pholidosaurus'' lineage. The

cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...

below shows their phylogenetic analysis, which is based on an expanded version of Pol and Gasparini (2009) analysis. de Andrade ''et al.'' (2011) recovered a

traditional Pholidosauridae. In their analysis the "''Elosuchus'' lineage" was found to be basal to the "''Pholidosaurus'' lineage"+Dyrosauridae. They used the name Elosuchidae for the ''Elosuchus'' lineage and restricted Pholidosauridae to its type genus. The following cladogram simplified after their analysis.

References

External links

Pholidosauridae at the Paleobiology Database
* Mikko's Phylogeny Archive o

{{Taxonbar, from=Q137740 Jurassic crocodylomorphs Early Cretaceous crocodylomorphs Late Cretaceous crocodylomorphs Middle Jurassic first appearances Late Cretaceous extinctions Prehistoric reptile families