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Planocraniidae 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 ...
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
eusuchia Eusuchia is a clade of crocodylomorphs that first appeared in the Early Cretaceous with ''Hylaeochampsa''. Along with Dyrosauridae and Sebecosuchia, they were the only crocodyliformes who survived the K-T extinction. Since the other two clades d ...
n
crocodyliform Crocodyliformes is a clade of crurotarsan archosaurs, the group often traditionally referred to as "crocodilians". They are the first members of Crocodylomorpha to possess many of the features that define later relatives. They are the only pseudo ...
s known from the
Paleogene The Paleogene ( ; British English, also spelled Palaeogene or Palæogene; informally Lower Tertiary or Early Tertiary) is a geologic period, geologic period and system that spans 43 million years from the end of the Cretaceous Period million yea ...
of
Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an area ...
,
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 ...
and
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 ...
. The family was coined by Li in 1976, and contains three
genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...
, ''
Boverisuchus ''Boverisuchus'' is an extinct genus of planocraniid crocodyliforms known from the middle Eocene (Lutetian stage) of Germany and western North America. It grew to approximately in length. History The type species ''Boverisuchus magnifrons'' wa ...
'', ''
Duerosuchus ''Duerosuchus'' is an extinct genus of crocodilian. Remains have been found from Corrales del Vino in Zamora, Spain, and are middle Eocene in age (about 40 million years old). ''Duerosuchus'' is known from a single skull that is incomplete but ...
'' and ''
Planocrania ''Planocrania'' is an extinct genus of eusuchian crocodyliforms from what is now China. Two species are currently known to belong to the genus. History The type species, ''Planocrania datangensis'', was named in 1976 from material found from Nan ...
''. Planocraniids were highly specialized crocodyliforms that were adapted to living on land. They had extensive body armor, long legs, and blunt claws resembling hooves, and are sometimes informally called "hoofed crocodiles".


Classification

Prior to 2013, the term Pristichampsidae/Pristichampsinae was used for this group. However, the
type specimen In biology, a type is a particular wiktionary:en:specimen, specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally attached. In other words, a type is an example that serves to a ...
of ''
Pristichampsus ''Pristichampsus'' ("saw crocodile") is a non-diagnostic extinct genus of crocodylian from France and possibly also Kazakhstan that is part of the monotypic Pristichampsidae family. As the type species, ''Pristichampsus rollinatii'', was based o ...
'' was found to be undiagnostic, and considered to be a ''
nomen dubium In binomial nomenclature, a ''nomen dubium'' (Latin for "doubtful name", plural ''nomina dubia'') is a scientific name that is of unknown or doubtful application. Zoology In case of a ''nomen dubium'' it may be impossible to determine whether a s ...
''. As such, Brochu (2013) transferred the other species placed in ''Pristichampsus'' to ''
Boverisuchus ''Boverisuchus'' is an extinct genus of planocraniid crocodyliforms known from the middle Eocene (Lutetian stage) of Germany and western North America. It grew to approximately in length. History The type species ''Boverisuchus magnifrons'' wa ...
'', and resurrected Planocraniidae to replace Pristichampsidae/Pristichampsinae as the name for the
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
. Brochu
cladistically Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived chara ...
defined Planocraniidae as ''
Planocrania hengdongensis ''Planocrania'' is an extinct genus of eusuchian crocodyliforms from what is now China. Two species are currently known to belong to the genus. History The type species, ''Planocrania datangensis'', was named in 1976 from material found from Nan ...
'' and crocodyliforms more closely related to it than to ''
Alligator mississippiensis The American alligator (''Alligator mississippiensis''), sometimes referred to colloquially as a gator or common alligator, is a large crocodilian reptile native to the Southeastern United States. It is one of the two extant species in the gen ...
'' (American alligator), ''
Crocodylus niloticus ''Crocodylus'' is a genus of true crocodiles in the family Crocodylidae. Taxonomy The generic name, ''Crocodylus'', was proposed by Josephus Nicolaus Laurenti in 1768. ''Crocodylus'' contains 13–14 extant (living) species and 5 extinct species ...
'' (Nile crocodile), ''
Gavialis gangeticus The gharial (''Gavialis gangeticus''), also known as gavial or fish-eating crocodile, is a crocodilian in the family Gavialidae and among the longest of all living crocodilians. Mature females are long, and males . Adult males have a distinct b ...
'' (gharial), '' Thoracosaurus macrorhynchus'', ''
Allodaposuchus precedens ''Allodaposuchus'' is an extinct genus of crocodyliforms that lived in what is now southern Europe during the Campanian and Maastrichtian stages of the Late Cretaceous. Although generally classified as a non-crocodylian eusuchian crocodylomorph, ...
'', or ''
Hylaeochampsa vectiana ''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, ...
''.
Phylogenetic 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 ...
analyses based purely on morphological data have generally placed planocraniids in a basal position within the
crocodilia Crocodilia (or Crocodylia, both ) is an order of mostly large, predatory, semiaquatic reptiles, known as crocodilians. They first appeared 95 million years ago in the Late Cretaceous period ( Cenomanian stage) and are the closest living ...
n
crown group In phylogenetics, the crown group or crown assemblage is a collection of species composed of the living representatives of the collection, the most recent common ancestor of the collection, and all descendants of the most recent common ancestor. ...
. Some of these analyses have found that planocraniids lie just outside
Brevirostres Brevirostres is a paraphyletic group of crocodilians that included alligatoroids and crocodyloids. Brevirostres are crocodilians with small snouts, and are distinguished from the long-snouted gharials. It is defined phylogenetically as the last ...
, a group that includes
alligator An alligator is a large reptile in the Crocodilia order in the genus ''Alligator'' of the family Alligatoridae. The two extant species are the American alligator (''A. mississippiensis'') and the Chinese alligator (''A. sinensis''). Additiona ...
s,
caiman A caiman (also cayman as a variant spelling) is an alligatorid belonging to the subfamily Caimaninae, one of two primary lineages within the Alligatoridae family, the other being alligators. Caimans inhabit Mexico, Central and South America fro ...
s, and
crocodile Crocodiles (family (biology), family Crocodylidae) or true crocodiles are large semiaquatic reptiles that live throughout the tropics in Africa, Asia, the Americas and Australia. The term crocodile is sometimes used even more loosely to inclu ...
s but not
gharial The gharial (''Gavialis gangeticus''), also known as gavial or fish-eating crocodile, is a crocodilian in the family Gavialidae and among the longest of all living crocodilians. Mature females are long, and males . Adult males have a distinct b ...
s. However, molecular studies using
DNA sequencing DNA sequencing is the process of determining the nucleic acid sequence – the order of nucleotides in DNA. It includes any method or technology that is used to determine the order of the four bases: adenine, guanine, cytosine, and thymine. Th ...
have found the group Brevirostres to be invalid upon finding that crocodiles and
gavialid Gavialidae is a family of large semiaquatic crocodilians with elongated, narrow snouts. Gavialidae consists of two living species, the gharial (''Gavialis gangeticus'') and the false gharial (''Tomistoma schlegelii''), both occurring in Asia. Man ...
s are more closely related than alligators. A 2018
tip dating Tip dating is a technique used in molecular dating that allows the inference of time-calibrated phylogenetic trees. Its defining feature is that it uses the ages of the samples to provide time information for the analysis, in contrast with traditio ...
study by Lee & Yates using both
molecular A molecule is a group of two or more atoms held together by attractive forces known as chemical bonds; depending on context, the term may or may not include ions which satisfy this criterion. In quantum physics, organic chemistry, and bioche ...
, morphological and
stratigraphic Stratigraphy is a branch of geology concerned with the study of rock layers (strata) and layering (stratification). It is primarily used in the study of sedimentary and layered volcanic rocks. Stratigraphy has three related subfields: lithostrati ...
data instead recovered the planocraniids outside crown group
Crocodylia Crocodilia (or Crocodylia, both ) is an order of mostly large, predatory, semiaquatic reptiles, known as crocodilians. They first appeared 95 million years ago in the Late Cretaceous period (Cenomanian stage) and are the closest living ...
. Below is a
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 ...
from that study: In 2021, Rio & Mannion conducted a new
phylogenetic 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 ...
study using a heavily modified morphological data set, and also noted the lack of consensus and difficulty in placing Planocraniidae. In their study, they recovered Planocraniidae within Crocodylia, as the
sister group 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 ...
to
Longirostres Longirostres is a clade of crocodilians that includes the crocodiles and the gavialids, to the exclusion of the alligatoroids. Named in 2003 by Harshman ''et al.'', Longirostres is a crown group defined phylogenetically as including the last com ...
, as shown in the cladogram below:


Description

Planocraniids were land-living (
terrestrial Terrestrial refers to things related to land or the planet Earth. Terrestrial may also refer to: * Terrestrial animal, an animal that lives on land opposed to living in water, or sometimes an animal that lives on or near the ground, as opposed to ...
) crocodyliforms with longer legs than living crocodilians. They grew to a maximum size of in length. Nearly complete skeletons of ''Boverisuchus'' indicate that planocraniids were more heavily armored than living crocodilians, with bony plates called
osteoderm 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 amp ...
s tightly interlocking along the back, completely encasing the tail, and extending down the legs. The claws were blunt and have been described as hoof-like in shape, suggesting that planocraniids may have been unguligrade, walking on the tips of their toes like mammalian
ungulate Ungulates ( ) are members of the diverse clade Ungulata which primarily consists of large mammals with hooves. These include odd-toed ungulates such as horses, rhinoceroses, and tapirs; and even-toed ungulates such as cattle, pigs, giraffes, cam ...
s. The areas on the leg bones where muscles attach were in different positions in planocraniids than they are in living crocodilians, possibly as an adaptation to walking on land. While most crocodilians have flattened skulls, planocraniids had tall and narrow (or laterally compressed) skulls. Their teeth were also laterally compressed and not conical like those of modern crocodilians. The combination of a laterally compressed skull and laterally compressed teeth is called the "ziphodont" condition. The teeth of the upper jaw completely overlapped the teeth of the lower jaw when the mouth was closed, giving planocraniids an alligator-like overbite. Planocraniids had a notch between 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 ...
bone at the tip of the upper jaw and the
maxilla The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The t ...
behind it. Living crocodiles also have this notch, which provides room for the enlarged fourth tooth of the lower jaw when the mouth is closed. In planocraniids the fourth tooth was small and did not fit into the notch.


Evolution

The evolution of Planocraniidae occurred after the K-T extinction when niches were open. Because of the lack of competition, the Planocraniidae evolved to hunt on land which would have been almost impossible with the
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 ...
clades existing.


References

{{Taxonbar, from=Q15076991 Neosuchians Paleocene crocodylomorphs Terrestrial crocodylomorphs Paleocene first appearances Eocene extinctions Eocene crocodylomorphs Eocene reptiles of Asia Eocene reptiles of Europe Eocene reptiles of North America Prehistoric reptile families