Pecora Ridge
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Pecora is an infraorder of even-toed hoofed mammals with ruminant digestion. Most members of Pecora have cranial appendages projecting from their
frontal bones 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, part ...
; only two extant genera lack them, '' Hydropotes'' and '' Moschus''. The name “Pecora” comes from the Latin word ''pecus'', which means “horned livestock”.Bubenik, A. Epigenetical, Morphological, Physiological, and Behavioral Aspects of Evolution of Horns, Pronghorns, and Antlers. in ''Horns, Pronghorns, and Antlers''. G. Bubenik and A. Bubenik eds. Springer-Verlag. New York. 1990 Although most pecorans have cranial appendages, only some of these are properly called “horns”, and many scientists agree that these appendages did not arise from a common ancestor, but instead evolved independently on at least two occasions.Janis, C., K. Scott. The Interrelationships of Higher Ruminant Families with Special Emphasis on the Members of the Cervoidea. ''American Museum Novitates''. 2893: 1-85. 1987. http://digitallibrary.amnh.org/dspace/handle/2246/5180 Likewise, while Pecora as a group is supported by both molecular and morphological studies, morphological support for interrelationships between pecoran families is disputed.


Evolutionary history

The first fossil ruminants appeared in the
Early Eocene In the geologic timescale the Ypresian is the oldest age or lowest stratigraphic stage of the Eocene. It spans the time between , is preceded by the Thanetian Age (part of the Paleocene) and is followed by the Eocene Lutetian Age. The Ypresian i ...
and were small, likely omnivorous, forest-dwellers. Artiodactyls with cranial appendages first occur in the early Miocene. The appearance of Pecora during the Miocene suggests that its rapid diversification may correspond to the climate change events of that epoch. It is likely that the antelopes, giraffids, and pronghorns evolved in an open environment while the cervids, including the caribou, evolved in
woodland habitat.
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The type of gallop in Pecorian species is shown to be closely related to their environment and anatomy, where light Pecorian species use both flexed and extended suspensions in their fast gallops. The white-tail and mule-deer have been observed to primarily use the extended suspension, since in this phase of their gallop they leap over bushes and logs that are present in their brush environment. However, heavy Pecorian species do not use extended suspensions as most have backs that slope downward with shorter hind legs.


Taxonomy and Classification

Pecora is an infraorder within the larger
suborder Order ( la, ordo) is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between family and class. In biological classification, the order is a taxonomic rank used in the classification of organisms and ...
Ruminantia, and is the sister
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, ...
to the infraorder Tragulina (of which Tragulidae is the only surviving family). Pecora's placement within Artiodactyla can be represented in the following cladogram:(see e.g. Fig S10) Current attempts to determine the relationships among pecoran families (as well as all artiodactyls) rely on molecular studies, as little consensus exists in morphological studies. Different families within Pecora are recognized as valid by different groups of scientists.and sources therein, pp. 4–5 Until the beginning of the 21st century it was understood that the family Moschidae (musk deer) was sister to Cervidae. However, a 2003 phylogenetic study by Alexandre Hassanin (of National Museum of Natural History, France) and colleagues, based on
mitochondria A mitochondrion (; ) is an organelle found in the Cell (biology), cells of most Eukaryotes, such as animals, plants and Fungus, fungi. Mitochondria have a double lipid bilayer, membrane structure and use aerobic respiration to generate adenosi ...
l and
nuclear Nuclear may refer to: Physics Relating to the nucleus of the atom: * Nuclear engineering *Nuclear physics *Nuclear power *Nuclear reactor *Nuclear weapon *Nuclear medicine *Radiation therapy *Nuclear warfare Mathematics *Nuclear space *Nuclear ...
analyses, revealed that Moschidae and Bovidae form a
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, ...
sister to Cervidae. According to the study, Cervidae diverged from the Bovidae-Moschidae clade 27 to 28 million years ago. The following cladogram is based on the 2003 study. Infraorder Pecora ("horned ruminants", "higher ruminants") * Family †
Gelocidae The Gelocidae are an extinct group of hornless ruminants, ruminantia that are estimated to have lived during the Eocene and Oligocene epochs, from 36 MYA (unit), MYA to 6 MYA. The family generally includes extinct hornless ruminants which do not ...
* Family †
Palaeomerycidae The Palaeomerycidae are an extinct family of ruminants in the order Artiodactyla. Palaeomerycids lived in North America, Europe, Africa and Asia from 33 to 4.9 million years ago (from the Late Eocene to Pliocene epochs), existing for about 28 mi ...
* Family Cervidae ( deer) * Family Moschidae (
musk deer Musk deer can refer to any one, or all seven, of the species that make up ''Moschus'', the only extant genus of the family Moschidae. Despite being commonly called deer, they are not true deer belonging to the family Cervidae, but rather their fa ...
) * Family Bovidae ( cattle, goats, sheep, and
antelope The term antelope is used to refer to many species of even-toed ruminant that are indigenous to various regions in Africa and Eurasia. Antelope comprise a wastebasket taxon defined as any of numerous Old World grazing and browsing hoofed mammals ...
s) * Superfamily Giraffoidea ** Family Antilocapridae ( pronghorn) ** Family Giraffidae (
giraffe The giraffe is a large African hoofed mammal belonging to the genus ''Giraffa''. It is the tallest living terrestrial animal and the largest ruminant on Earth. Traditionally, giraffes were thought to be one species, ''Giraffa camelopardalis ...
, okapi) ** Family † Climacoceratidae


Anatomy

Pecorans share characteristics with other artiodactyls, including a four-chambered stomach, and a paraxonic foot, meaning that it supports weight on the third and fourth digits. Several characteristics distinguish Pecora from its sister taxon, Tragulina: an astragalus with parallel sides, a loss of the trapezium, and differences in parts of the skull such as the petrosal bone. The distinguishing features of most pecoran families are cranial appendages. Most modern pecorans (with the exception of the Moschidae) have one of four types of cranial appendages: horns, antlers, ossicones, or pronghorns. * True horns have a bone core that is covered in a permanent sheath of keratin. They are indicative of Bovidae. Horns develop in the periosteum over the frontal bone, and can be curved or straight. Surface features on the keratin sheath (e.g., ridges or twists) are thought to be caused by differential rates of growth around the bone core. * Antlers are bony structures that are shed and replaced each year in members of the family Cervidae. They grow from a permanent outgrowth of the frontal bone called the pedicle. Antlers can be branched, as in the white-tailed deer (''
Odocoileus virginianus The white-tailed deer (''Odocoileus virginianus''), also known as the whitetail or Virginia deer, is a medium-sized deer native to North America, Central America, and South America as far south as Peru and Bolivia. It has also been introduced t ...
''), or palmate, as in the moose ('' Alces alces''). *
Ossicones Ossicones are columnar or conical skin-covered bone structures on the heads of giraffes, male okapi, and some of their extinct relatives. Ossicones are distinguished from the superficially similar structures of horns and antlers by their unique ...
are permanent bone structures that fuse to the frontal or parietal bones during the lifetime of an animal. They are found only in the Giraffidae and closely related extinct clades, represented in modern animals by the giraffe ('' Giraffa camelopardalis'') and the okapi ('' Okapia johnstoni''). *
Pronghorns The pronghorn (, ) (''Antilocapra americana'') is a species of artiodactyl (even-toed, hoofed) mammal indigenous to interior western and central North America. Though not an antelope, it is known colloquially in North America as the American an ...
are similar to horns in that they have keratinous sheaths covering permanent bone cores; however, these sheaths are deciduous and can be shed like antlers. Very little is known about the development of pronghorns, but they are generally presumed to have evolved independently. The only extant animal with pronghorns is the pronghorn antelope (''
Antilocapra americana The pronghorn (, ) (''Antilocapra americana'') is a species of artiodactyl (even-toed, hoofed) mammal indigenous to interior western and central North America. Though not an antelope, it is known colloquially in North America as the American ante ...
'').


References


External links

* {{Taxonbar, from=Q772442 Ruminants Extant Burdigalian first appearances