Planocraniidae

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Planocraniidae
Temporal range: Paleocene - Eocene, 61.6–40  Ma [1]
Boverisuchus magnifrons white background.jpg
Skeleton of Boverisuchus magnifrons
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Clade: Archosauria
Clade: Pseudosuchia
Clade: Crocodylomorpha
Clade: Crocodyliformes
Clade: Neosuchia
Clade: Eusuchia
Family: Planocraniidae
Li, 1976
Type genus
Planocrania
Li, 1976
Genera

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 . [2] [3] 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". [4]

Contents

Classification

Prior to 2013, the term Pristichampsidae/Pristichampsinae was used for this group. However, the type specimen of Pristichampsus was found to be undiagnostic, and considered to be a nomen dubium . [2] As such, Brochu (2013) transferred the other species placed in Pristichampsus to Boverisuchus , and resurrected Planocraniidae to replace Pristichampsidae/Pristichampsinae as the name for the clade. [2] Brochu cladistically defined Planocraniidae as Planocrania hengdongensis and crocodyliforms more closely related to it than to Alligator mississippiensis (American alligator), Crocodylus niloticus (Nile crocodile), Gavialis gangeticus (gharial), Thoracosaurus macrorhynchus , Allodaposuchus precedens , or Hylaeochampsa vectiana .

Phylogenetic analyses based purely on morphological data have generally placed planocraniids in a basal position within the crocodilian crown group. Some of these analyses have found that planocraniids lie just outside Brevirostres, a group that includes alligators, caimans, and crocodiles but not gharials. [2] [5] [6] However, molecular studies using DNA sequencing have found the group Brevirostres to be invalid upon finding that crocodiles and gavialids are more closely related than alligators. [7] [8] [9] [10] [11]

A 2018 tip dating study by Lee & Yates using both molecular, morphological and stratigraphic data instead recovered the planocraniids outside crown group Crocodylia. Below is a cladogram from that study: [10]

Goniopholis

Bernissartia

Shamosuchus

Eusuchia

Acynodon

Hylaeochampsa

Allodaposuchus

Lohuecosuchus

Borealosuchus

"Thoracosaurs"

Eothoracosaurus

Thoracosaurus

Argochampsa

Eogavialis

Eosuchus

Planocraniidae

Planocrania hengdongensis

Planocrania datangensis

Boverisuchus

Crocodylia

In 2021, Rio & Mannion conducted a new phylogenetic 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 to Longirostres, as shown in the cladogram below: [1]

Eusuchia

Hylaeochampsidae

Allodaposuchidae

Borealosuchus

Crocodylia

Alligatoroidea

Planocraniidae

Planocrania datangensis

Planocrania hengdongensis

Boverisuchus magnifrons

Boverisuchus vorax

Longirostres

Gavialoidea

Crocodyloidea

Description

Skull of holotype of the extinct crocodile "Weigeltisuchus geiseltalensis" KUHN, 1938 (specimen no. GMH Leo X 8001; subsequently "Pristichampsus geiseltalensis", now referred to Boverisuchus magnifrons KUHN, 1938) Skull of Pristichampsus geiseltalensis.JPG
Skull of holotype of the extinct crocodile “Weigeltisuchus geiseltalensis” KUHN, 1938 (specimen no. GMH Leo X 8001; subsequently “ Pristichampsus geiseltalensis ”, now referred to Boverisuchus magnifrons KUHN, 1938)

Planocraniids were land-living (terrestrial) crocodyliforms with longer legs than living crocodilians. They grew to a maximum size of 2 to 3 metres (6.6 to 9.8 ft) in length. [12] Nearly complete skeletons of Boverisuchus indicate that planocraniids were more heavily armored than living crocodilians, with bony plates called osteoderms 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, [13] suggesting that planocraniids may have been unguligrade, [14] walking on the tips of their toes like mammalian ungulates. 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. [12]

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. [12]

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 bone at the tip of the upper jaw and the maxilla 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. [12]

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 clades existing. [15]

Related Research Articles

<span class="mw-page-title-main">Gavialidae</span> Family of gharial crocodylians

Gavialidae is a family of large semiaquatic crocodilians with elongated, narrow snouts. Gavialidae consists of two living species, the gharial and the false gharial, both occurring in Asia. Many extinct members are known from a broader range, including the recently extinct Hanyusuchus. Gavialids are generally regarded as lacking the jaw strength to capture the large mammalian prey favoured by crocodiles and alligators of similar size so their thin snout is best used to catch fish, however the false gharial has been found to have a generalist diet with mature adults preying upon larger vertebrates, such as ungulates.

<i>Ceratosuchus</i> Extinct genus of reptiles

Ceratosuchus is an extinct genus of alligatorine crocodylian from latest Paleocene rocks of Colorado's Piceance Basin and earliest Eocene rocks of Wyoming's Bighorn Basin in North America, a slice of time known as the Clarkforkian North American Land Mammal Age. Like its modern relatives, Ceratosuchus was a swamp-dwelling predator. It is named for the pair of flattened, triangular bony plates that extend from the back of its head.

<i>Boverisuchus</i> Extinct genus of reptiles

Boverisuchus is an extinct genus of planocraniid crocodyliforms known from the middle Eocene of Germany and western North America. It was a relatively small crocodyliform with an estimated total length of approximately 2.2–3.6 metres (7.2–11.8 ft).

<i>Borealosuchus</i> Extinct genus of reptiles

Borealosuchus is an extinct genus of crocodyliforms that lived from the Late Cretaceous to the Eocene in North America. It was named by Christopher Brochu in 1997 for several species that had been assigned to Leidyosuchus. The species assigned to it are: B. sternbergii, the type species, from the Maastrichtian of Colorado, Montana, North Dakota, South Dakota, and Wyoming; B. acutidentatus, from the Paleocene of Saskatchewan; B. formidabilis, from the Paleocene of North Dakota; B. griffithi, from the Paleocene of Alberta; and B. wilsoni, from the Eocene of Wyoming. B. formidabilis is particularly well-known, represented by the remains of many individuals from the Wannagan Creek site in North Dakota.

Argochampsa is an extinct genus of eusuchian crocodylomorph, usually regarded as a gavialoid crocodilian, related to modern gharials. It lived in the Paleocene of Morocco. Described by Hua and Jouve in 2004, the type species is A. krebsi, with the species named for Bernard Krebs. Argochampsa had a long narrow snout, and appears to have been marine in habits.

<span class="mw-page-title-main">Eusuchia</span> Clade of reptiles

Eusuchia is a clade of neosuchian crocodylomorphs that first appeared in the Early Cretaceous, which includes modern crocodilians. Along with Dyrosauridae and Sebecosuchia, they were the only crocodyliformes who survived the K-Pg extinction.

<span class="mw-page-title-main">Alligatoroidea</span> Superfamily of reptiles

Alligatoroidea is one of three superfamilies of crocodylians, the other two being Crocodyloidea and Gavialoidea. Alligatoroidea evolved in the Late Cretaceous period, and consists of the alligators and caimans, as well as extinct members more closely related to the alligators than the two other groups.

Dollosuchoides, colloquially known as the Crocodile of Maransart, is an extinct monospecific genus of gavialoid crocodilian, traditionally regarded as a member of the subfamily Tomistominae. Fossils have been found in the Brussel Formation of Maransart, Belgium and date back to the middle Eocene.

<span class="mw-page-title-main">Crocodyliformes</span> Clade of reptiles

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 pseudosuchians to survive the K-Pg extinction event.

Maroccosuchus zennaroi is an extinct gavialoid crocodylian from the Early Eocene of Morocco, traditionally regarded as a member of the subfamily Tomistominae.

Paratomistoma is an extinct monospecific genus of gavialoid crocodylian. It is based on the holotype specimen CGM 42188, a partial posterior skull and lower jaw discovered at Wadi Hitan, Egypt, in Middle Eocene-age rocks of the Gehannam Formation. The skull is unfused but considered morphologically mature. Paratomistoma was named in 2000 by Christopher Brochu and Philip Gingerich; the type species is P. courti in honor of Nicholas Court, who found CGM 42188. They performed a phylogenetic analysis and found Paratomistoma to be a derived member of Tomistominae, related to the false gharial. It may have been a marine or coastal crocodilian.

<i>Planocrania</i> Extinct genus of reptiles

Planocrania is an extinct genus of eusuchian crocodyliforms from what is now China. Two species are currently known to belong to the genus.

Prodiplocynodon is an extinct genus of basal crocodyloid crocodylian. It is one of the only crocodyloids known from the Cretaceous and existed during the Maastrichtian stage. The only species of Prodiplocynodon is the type species P. langi from the Lance Formation of Wyoming, known only from a single holotype skull lacking the lower jaw.

<span class="mw-page-title-main">Crocodyloidea</span> Superfamily of crocodiles

Crocodyloidea is one of three superfamilies of crocodilians, the other two being Alligatoroidea and Gavialoidea, and it includes the crocodiles. Crocodyloidea may also include the extinct Mekosuchinae, native to Australasia from the Eocene to the Holocene, although this is disputed.

<span class="mw-page-title-main">Gavialoidea</span> Superfamily of large reptiles

Gavialoidea is one of three superfamilies of crocodylians, the other two being Alligatoroidea and Crocodyloidea. Although many extinct species are known, only the gharial Gavialis gangeticus and the false gharial Tomistoma schlegelii are alive today, with Hanyusuchus having become extinct in the last few centuries.

<span class="mw-page-title-main">Brevirostres</span> Taxon of reptiles

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 common ancestor of Alligator mississippiensis and Crocodylus niloticus and all of its descendants. This classification was based on morphological studies primarily focused on analyzing skeletal traits of living and extinct fossil species, and placed the gharials outside the group due to their unique skull structure, and can be shown in the simplified cladogram below:

<span class="mw-page-title-main">Allodaposuchidae</span> Extinct clade of reptiles

Allodaposuchidae is an extinct clade of eusuchians that lived in Europe during the Late Cretaceous (Santonian-Maastrichtian).

Tomistoma cairense is an extinct species of gavialoid crocodilian from the Lutetian stage of the Eocene era. It lived in North East Africa, especially Egypt. Remains of T. cairense have been found in the Mokattam Formation, in Mokattam, Egypt. Tomistoma cairense did not have a Maxilla process within their lacrimal gland, whereas all extant (living) crocodilians do.

Portugalosuchus is an extinct genus of eusuchian crocodyliform that was possibly a basal crocodylian – if so then it would be the oldest known crocodylian to date. The type species is P. azenhae, described in 2018, and it is known from the Late Cretaceous (Cenomanian)-aged Tentugal Formation in Portugal. A 2021 morphological study recovered Portugalosuchus within Crocodylia as a member of Gavialidae closely related to similar "thoracosaurs", while also noting that it might also possibly be outside of Crocodylia completely. A 2022 tip dating analysis incorporating both morphological and DNA data placed Portugalosuchus outside of Crocodylia, as the sister taxon of the family Allodaposuchidae. A cladogram simplified after that analysis is shown below:

<span class="mw-page-title-main">Longirostres</span> Clade of crocodilians

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 common ancestor of Crocodylus niloticus and Gavialis gangeticus and all of its descendants.

References

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