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''Fruitachampsa'' is a
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 ...
of shartegosuchid
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 ...
from the
Upper Jurassic The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 163.5 ± 1.0 to 145.0 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic strata.Owen 1987. In European lithostratigraphy, the name ...
Morrison Formation The Morrison Formation is a distinctive sequence of Late Jurassic, Upper Jurassic sedimentary rock found in the western United States which has been the most fertile source of dinosaur fossils in North America. It is composed of mudstone, sandsto ...
of
Colorado Colorado (, other variants) is a state in the Mountain West subregion of the Western United States. It encompasses most of the Southern Rocky Mountains, as well as the northeastern portion of the Colorado Plateau and the western edge of t ...
. It is known from multiple specimens that show it to have been a relatively long-limbed terrestrial quadrupedal predator less than long, with a short face and a prominent pair of canine-like teeth in the lower jaw. Before it was formally described in 2011, it was also known as the "
Fruita The City of Fruita is a home rule municipality located in western Mesa County, Colorado, United States. The city population was 13,395 at the 2020 United States Census. Fruita is a part of the Grand Junction, CO Metropolitan Statistical Area a ...
form". Its
type species In 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 species that contains the biological type specimen ...
is ''F. callisoni''.


Discovery

Fossils of ''Fruitachampsa'' have been found from the Fruita Paleontological Area (FPA) in
Fruita, Colorado The City of Fruita is a home rule municipality located in western Mesa County, Colorado, United States. The city population was 13,395 at the 2020 United States Census. Fruita is a part of the Grand Junction, CO Metropolitan Statistical Area a ...
. The deposits in the FPA belong to the
Morrison Formation The Morrison Formation is a distinctive sequence of Late Jurassic, Upper Jurassic sedimentary rock found in the western United States which has been the most fertile source of dinosaur fossils in North America. It is composed of mudstone, sandsto ...
, which is
Late Jurassic The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 163.5 ± 1.0 to 145.0 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic strata.Owen 1987. In European lithostratigraphy, the name ...
in age. The first remains of ''Fruitachampsa'' were found by paleontologists James Clark and George Callison in 1975, who discovered a diverse assemblage of microvertebrates at the FPA. More complete material of ''Fruitachampsa'' was found in 1976, 1977, and 1979. ''Fruitachampsa'' has been found alongside another more basal crocodylomorph, ''
Macelognathus ''Macelognathus'' is an extinct genus of sphenosuchian crocodylomorph from the Late Jurassic. Originally it was believed be a turtle and later a dinosaur. It lived in what is now Wyoming, in North America.Moodie, R.L. 1908. The relationship of ...
''. The name ''Fruitachampsa'' has existed since the late 1980s. Because the name was not published with a formal description, ''Fruitachampsa'' was considered a ''
nomen nudum In taxonomy, a ''nomen nudum'' ('naked name'; plural ''nomina nuda'') is a designation which looks exactly like a scientific name of an organism, and may have originally been intended to be one, but it has not been published with an adequate descr ...
'' or "naked name." ''Fruitachampsa callisoni'' was formally described by James Clark in 2011, with the specific name honoring George Callison.


Description

Most of the body of ''Fruitachampsa'' is known from several partial skeletons and skulls. It is a small crocodyliform under in length. Although small, ''Fruitachampsa'' is large in comparison to other shartegosuchids, with a maximum skull length of . Viewed from above, the skull is square-shaped in the back and narrows toward the snout. The snout is similar in appearance to that of the living caiman genus ''
Paleosuchus ''Paleosuchus'' is a South American genus of reptiles in the subfamily Caimaninae of the family (biology), family Alligatoridae. They are the smallest members of the order Crocodilia in the Americas. The genus contains two extant species and a ye ...
'', although it is not flattened like in modern crocodylians. As in other early crocodylomorphs, a bone called the
palpebral An eyelid is a thin fold of skin that covers and protects an eye. The levator palpebrae superioris muscle retracts the eyelid, exposing the cornea to the outside, giving vision. This can be either voluntarily or involuntarily. The human eyel ...
extends backwards over the eye socket like an
eyebrow An eyebrow is an area of short hairs above each eye that follows the shape of the lower margin of the brow ridges of some mammals. In humans, eyebrows serve two main functions: first, communication through facial expression, and second, prevent ...
. The teeth at the back of the jaws have flattened tops with ridges running along their length. Unlike other crocodyliforms, ''Fruitachampsa'' lacks mandibular or antorbital fenestrae (holes in the lower jaw and in front of the eye sockets). ''Fruitachampsa'' differs from other shartegosuchids in that it has procoelous vertebrae with concave anterior surfaces and convex posterior surfaces. As a terrestrial crocodyliform, ''Fruitachampsa'' has long and extremely slender limbs. 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 cover the back in two overlapping rows, and are much thinner than those of living crocodylians. Most specimens of ''Fruitachampsa'' probably belong to adult individuals because the
neural arch The spinal column, a defining synapomorphy shared by nearly all vertebrates,Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic i ...
is fused to the
body Body may refer to: In science * Physical body, an object in physics that represents a large amount, has mass or takes up space * Body (biology), the physical material of an organism * Body plan, the physical features shared by a group of anima ...
of each vertebra, and the sacral ribs are fused to the
sacrum The sacrum (plural: ''sacra'' or ''sacrums''), in human anatomy, is a large, triangular bone at the base of the spine that forms by the fusing of the sacral vertebrae (S1S5) between ages 18 and 30. The sacrum situates at the upper, back part ...
. For most living crocodylians, young individuals have many unfused elements in their vertebral column. Other features such as a deeply pitted
skull roof The skull roof, or the roofing bones of the skull, are a set of bones covering the brain, eyes and nostrils in bony fishes and all land-living vertebrates. The bones are derived from dermal bone and are part of the dermatocranium. In comparati ...
and the fusing of the
frontal bone The frontal bone is a bone in the human skull. The bone consists of two portions.''Gray's Anatomy'' (1918) These are the vertically oriented squamous part, and the horizontally oriented orbital part, making up the bony part of the forehead, par ...
s in some individuals also suggest that the known material of ''Fruitachampsa'' represent mature individuals. Therefore, it is unlikely that ''Fruitachampsa'' grew much larger in body size than the specimens indicate.


Classification

Before it was formally described, ''Fruitachampsa'' was included in several
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 as the "Fruita form". These analyses usually placed it as a basal member of a group called
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 ...
. When it was named in 2011, a phylogenetic analysis of ''Fruitachampsa'' grouped it with a family of Asian crocodyliforms called Shartegosuchidae. Shartegosuchids lie outside Mesoeucrocodylia in a more basal position among Crocodyliformes. These non-mesoueucrocodylian crocodyliforms are often called
protosuchia Protosuchia is a group of extinct Mesozoic crocodyliforms. They were small in size (~1 meter in length) and terrestrial. In phylogenetic terms, Protosuchia is considered an informal group because it is a grade of basal crocodyliforms, not a true ...
ns, although Protosuchia is rarely considered a
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 ...
grouping. ''Fruitachampsa'' was probably placed among more advanced crocodyliforms in earlier analyses because these studies did not include the poorly known Asian shartegosuchids. ''Fruitachampsa'' possesses several features in common with more advanced crocodyliforms, including procoelous vertebrae and a
secondary palate The secondary palate is an anatomical structure that divides the nasal cavity from the oral cavity in many vertebrates. In human embryology, it refers to that portion of the hard palate that is formed by the growth of the two palatine shelves medi ...
. These features are highly variable in crocodyliforms, and probably evolved independently in ''Fruitachampsa''. Below is a cladogram showing the phylogenetic placement of ''Fruitachampsa'' from Clark (2011):


Paleobiology

''Fruitachampsa'' is the only shartegosuchid known from North America, and is one of the youngest members of the group. The presence of ''Fruitachampsa'' in the western United States suggests that shartegosuchids radiated from Asia into North America. During the Late Jurassic, the only route between Asia and North America would have been through Europe. Although shartegosuchids likely inhabited Europe, no fossils have been found from the continent.


References

{{Taxonbar, from=Q3089314 Late Jurassic crocodylomorphs of North America Late Jurassic crocodylomorphs Morrison fauna Terrestrial crocodylomorphs Fossil taxa described in 2011 Prehistoric pseudosuchian genera