''Planocrania'' 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 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 ...

eusuchia Eusuchia is a clade of neosuchian Crocodylomorpha, crocodylomorphs that first appeared in the Early Cretaceous, which includes modern Crocodilia, crocodilians. Along with Dyrosauridae and Sebecosuchia, they were the only crocodyliformes who survi ...

crocodyliform Crocodyliformes is a clade of Crurotarsi, 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 ...

s from what is now

China China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...

. Two species are currently known to belong to the genus.

History

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 ...

, ''Planocrania datangensis'', was named in 1976 from material found from

Nanxiong Nanxiong (), historically Namyung, Namhung, and Nanhsiung, is a county-level city of northern Guangdong province, People's Republic of China, bordering Jiangxi to the north, east and southeast. It is under the administration of Guangdong Province. ...

Guangdong Province ) means "wide" or "vast", and has been associated with the region since the creation of Guang Prefecture in AD 226. The name "''Guang''" ultimately came from Guangxin ( zh, labels=no, first=t, t= , s=广信), an outpost established in Han dynasty ...

, and the new genus ''Planocrania'' was placed in a newly erected

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 ...

Planocraniidae Planocraniidae is an extinct family of eusuchian crocodyliforms known from the Paleogene of Asia, Europe and North America. The family was coined by Li in 1976, and contains three genera, '' Boverisuchus'', '' Duerosuchus'' and '' Planocrania' ...

. A second species, ''Planocrania hengdongensis'', is known from

Hengdong County Hengdong County () is a county located in the east of Hengyang prefecture-level city, Hunan province, China. It was established from the eastern part of Hengshan County in 1966 and named after the ''east of Mount Heng''. The county has an area ...

Hunan Province Hunan is an inland province in Central China. Located in the middle reaches of the Yangtze watershed, it borders the province-level divisions of Hubei to the north, Jiangxi to the east, Guangdong and Guangxi to the south, and Guizhou and Chon ...

and was described in 1984. ''Planocrania'' was subsequently included in the family Pristichampsidae along with the genus ''

Pristichampsus ''Pristichampsus'' (from , 'saw' and , 'crocodile') is a non-diagnostic and potentially dubious extinct genus of crocodylian from France and possibly also Kazakhstan that is part of the monotypic Pristichampsidae family. As the type species, ' ...

''. Both genera were previously assigned to the

subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end botanical subfamily names with "-oideae", and zo ...

Pristichampsinae within the family

Crocodylidae Crocodiles (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 more loosely to include all extant membe ...

before they were placed in their own family, the Pristichampsidae. However, in a 2013

phylogenetic 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 dat ...

analysis, the two Asian species of ''Planocrania'' were found to be most closely related to the North American and European ''

Boverisuchus ''Boverisuchus'' is an extinct genus of planocraniid crocodyliforms known from the early to middle Eocene (Ypresian to Lutetian stages) of Germany and western North America. It was a relatively small crocodyliform with an estimated total length ...

'', and the family

was reinstated to contain these genera and replace Pristichampsidae.

Phylogeny

Phylogenetic analyses based purely on morphological data have generally placed planocraniids in a basal position within the

crocodilia Crocodilia () is an 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 crocodylomorph pseudosuchia ...

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 group of crocodilians that, as originally formulated, included alligatoroids and crocodyloids, but not gavialoids. Results of molecular phylogenetic analysis uniformly draw gavialoids and crocodyloids into a close relationshi ...

, a group that includes

alligator An alligator, or colloquially gator, is a large reptile in the genus ''Alligator'' of the Family (biology), family Alligatoridae in the Order (biology), order Crocodilia. The two Extant taxon, extant species are the American alligator (''A. mis ...

caiman A caiman ( (also spelled cayman) from Taíno language, Taíno ''kaiman'') is an alligatorid belonging to the subfamily Caimaninae, one of two primary lineages within the Alligatoridae family (biology), family, the other being alligators. ...

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 more loosely to include ...

s but not

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 ...

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, thymine, cytosine, and guanine. The ...

have found the group Brevirostres to be invalid upon finding that crocodiles and gavialids 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 that are held together by attractive forces known as chemical bonds; depending on context, the term may or may not include ions that satisfy this criterion. In quantum physics, organic chemistry, ...

, 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: lithost ...

data instead recovered the planocraniids outside crown group

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 ...

. Below is a

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

from that study: In 2021, Rio & Mannion conducted a new

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 ...

Longirostres Longirostres is a clade of crocodilians that includes the crocodiles and the gavialids, to the exclusion of the alligatoroids. Defined in 2003 by Harshman ''et al.'', Longirostres is a crown group defined phylogenetically as including the last ...

, as shown in the cladogram below:

References

Neosuchia Prehistoric pseudosuchian genera Paleocene crocodylomorphs Paleocene reptiles of Asia Fossil taxa described in 1976 {{paleo-archosaur-stub