Pecora Escarpment USGS
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Pecora is an
infraorder Order () is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between Family_(biology), family and Class_(biology), class. In biological classification, the order is a taxonomic rank used in the classific ...
of even-toed hoofed
mammals A mammal () is a vertebrate animal of the class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three middle e ...
with
ruminant Ruminants are herbivorous grazing or browsing artiodactyls belonging to the suborder Ruminantia that are able to acquire nutrients from plant-based food by fermenting it in a specialized stomach prior to digestion, principally through microb ...
digestion. Most members of Pecora have cranial appendages projecting from their
frontal bones In the human skull, the frontal bone or sincipital bone is an unpaired bone which 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 ...
; only two extant genera lack them, ''
Hydropotes The water deer (''Hydropotes inermis'') is a small deer species native to Korea and China. Its prominent tusks, similar to those of musk deer, have led to both subspecies being colloquially named vampire deer in English-speaking areas to which t ...
'' and ''
Moschus Moschus () was an ancient Greek bucolic poet and student of the Alexandrian grammarian Aristarchus of Samothrace. He was born at Syracuse, Magna Graecia, and flourished about 150 BC. Aside from his poetry, he was known for his grammatical work, ...
''. The name "Pecora" comes from the Latin word , which means "cattle".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 Ruminants are herbivorous grazing or browsing artiodactyls belonging to the suborder Ruminantia that are able to acquire nutrients from plant-based food by Enteric fermentation, fermenting it in a specialized stomach prior to digestion, principa ...
appeared in the
Early Eocene In the geologic timescale the Ypresian is the oldest age (geology), age or lowest stage (stratigraphy), stratigraphic stage of the Eocene. It spans the time between , is preceded by the Thanetian Age (part of the Paleocene) and is followed by th ...
and were small, likely
omnivorous An omnivore () is an animal that regularly consumes significant quantities of both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize ...
, forest-dwellers. Molecular dating studies estimate that
Ruminantia Ruminants are herbivorous grazing or browsing artiodactyls belonging to the suborder Ruminantia that are able to acquire nutrients from plant-based food by Enteric fermentation, fermenting it in a specialized stomach prior to digestion, principa ...
split into the two sister clades Pecora and
Tragulina Tragulina (also known as Traguliformes) is an infraorder of even-toed ungulates. Only the chevrotains survive to the present, including the genera ''Tragulus'' (the mouse deer) and '' Hyemoschus'', all within the family Tragulidae. Taxonomy and ...
around 45 million years ago, during the
Eocene The Eocene ( ) is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene'' comes ...
. However, it was not until 15 million years later, at around 30 million years ago during the
Oligocene The Oligocene ( ) is a geologic epoch (geology), epoch of the Paleogene Geologic time scale, Period that extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that defin ...
, that the
evolutionary radiation An evolutionary radiation is an increase in taxonomic diversity that is caused by elevated rates of speciation, that may or may not be associated with an increase in morphological disparity. A significantly large and diverse radiation within ...
of Pecora began and the five
families Family (from ) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictability, structure, and safety as ...
appeared (
Bovidae The Bovidae comprise the family (biology), biological family of cloven-hoofed, ruminant mammals that includes Bos, cattle, bison, Bubalina, buffalo, antelopes (including Caprinae, goat-antelopes), Ovis, sheep and Capra (genus), goats. A member o ...
,
Cervidae A deer (: deer) or true deer is a hoofed ruminant ungulate of the family Cervidae (informally the deer family). Cervidae is divided into subfamilies Cervinae (which includes, among others, muntjac, elk (wapiti), red deer, and fallow deer) a ...
,
Moschidae Moschidae is a family of pecoran even-toed ungulates, containing the musk deer (''Moschus'') and its extinct relatives. They are characterized by long "saber teeth" instead of horns, antlers or ossicones, modest size (''Moschus'' only reaches ...
,
Giraffidae The Giraffidae are a family (biology), family of ruminant artiodactyl mammals that share a recent common ancestor with deer and bovids. This family, once a diverse group spread throughout Eurasia and Africa, presently comprises only two extant ge ...
, and
Antilocapridae The Antilocapridae are a family of ruminant artiodactyls endemic to North America. Their closest extant relatives are the giraffids. Only one species, the pronghorn (''Antilocapra americana''), is living today; all other members of the family ...
). The appearance of many Pecoran fossils during the
Miocene The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...
suggests that its rapid diversification may correspond to the climate change events of that epoch, as this time period was marked by much of Earth's forest habitats being replaced by grasslands due to widespread cooling and drying. It is likely that the antelopes, giraffids, and pronghorns evolved in an open environment while the cervids, including the caribou, evolved in a woodland habitat. 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 Order () is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between Family_(biology), family and Class_(biology), class. In biological classification, the order is a taxonomic rank used in the classific ...
within the larger
suborder Order () 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 recognized ...
Ruminantia Ruminants are herbivorous grazing or browsing artiodactyls belonging to the suborder Ruminantia that are able to acquire nutrients from plant-based food by Enteric fermentation, fermenting it in a specialized stomach prior to digestion, principa ...
, and is the
sister A sister is a woman or a girl who shares parents or a parent with another individual; a female sibling. The male counterpart is a brother. Although the term typically refers to a familial relationship, it is sometimes used endearingly to ref ...
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
to the
infraorder Order () is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between Family_(biology), family and Class_(biology), class. In biological classification, the order is a taxonomic rank used in the classific ...
Tragulina Tragulina (also known as Traguliformes) is an infraorder of even-toed ungulates. Only the chevrotains survive to the present, including the genera ''Tragulus'' (the mouse deer) and '' Hyemoschus'', all within the family Tragulidae. Taxonomy and ...
(of which
Tragulidae Chevrotains, or mouse-deer, are small, even-toed ungulates that make up the family Tragulidae, and are the only living members of the infraorder Tragulina. The 10 extant species are placed in three genera, but several species also are ...
is the only surviving
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 ...
). Pecora's placement within
Artiodactyla Artiodactyls are placental mammals belonging to the order (biology), order Artiodactyla ( , ). Typically, they are ungulates which bear weight equally on two (an even number) of their five toes (the third and fourth, often in the form of a hoof ...
can be represented in the following
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 ...
:(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 Moschidae is a family of pecoran even-toed ungulates, containing the musk deer (''Moschus'') and its extinct relatives. They are characterized by long "saber teeth" instead of horns, antlers or ossicones, modest size (''Moschus'' only reaches ...
(musk deer) was
sister A sister is a woman or a girl who shares parents or a parent with another individual; a female sibling. The male counterpart is a brother. Although the term typically refers to a familial relationship, it is sometimes used endearingly to ref ...
to
Cervidae A deer (: deer) or true deer is a hoofed ruminant ungulate of the family Cervidae (informally the deer family). Cervidae is divided into subfamilies Cervinae (which includes, among others, muntjac, elk (wapiti), red deer, and fallow deer) a ...
. However, a 2003
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 ...
study by Alexandre Hassanin (of
National Museum of Natural History, France The French National Museum of Natural History ( ; abbr. MNHN) is the national natural history museum of France and a of higher education part of Sorbonne University. The main museum, with four galleries, is located in Paris, France, within the ...
) and colleagues, based on
mitochondria A mitochondrion () is an organelle found in the cells of most eukaryotes, such as animals, plants and fungi. Mitochondria have a double membrane structure and use aerobic respiration to generate adenosine triphosphate (ATP), which is us ...
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 Moschidae is a family of pecoran even-toed ungulates, containing the musk deer (''Moschus'') and its extinct relatives. They are characterized by long "saber teeth" instead of horns, antlers or ossicones, modest size (''Moschus'' only reaches ...
and
Bovidae The Bovidae comprise the family (biology), biological family of cloven-hoofed, ruminant mammals that includes Bos, cattle, bison, Bubalina, buffalo, antelopes (including Caprinae, goat-antelopes), Ovis, sheep and Capra (genus), goats. A member o ...
form a
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
sister to
Cervidae A deer (: deer) or true deer is a hoofed ruminant ungulate of the family Cervidae (informally the deer family). Cervidae is divided into subfamilies Cervinae (which includes, among others, muntjac, elk (wapiti), red deer, and fallow deer) a ...
. According to the study, Cervidae diverged from the Bovidae-Moschidae clade 27 to 28 million years ago. The following
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 ...
is based on the 2003 study. Infraorder Pecora ("horned ruminants", "higher ruminants") *
Incertae sedis or is a term used for a taxonomy (biology), taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertainty ...
** Family †
Dromomerycidae Dromomerycidae is an extinct family of Neogene ruminants belonging to the infraorder Pecora. Dromomerycids, known by two subfamilies, were endemic to North America from the later early Miocene up to the early Pliocene (~5 Ma), leaving no descenda ...
** Family
Antilocapridae The Antilocapridae are a family of ruminant artiodactyls endemic to North America. Their closest extant relatives are the giraffids. Only one species, the pronghorn (''Antilocapra americana''), is living today; all other members of the family ...
(
pronghorn 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 ...
) * Parvorder Bovimorpha ** Superfamily Cervoidea *** Family
Cervidae A deer (: deer) or true deer is a hoofed ruminant ungulate of the family Cervidae (informally the deer family). Cervidae is divided into subfamilies Cervinae (which includes, among others, muntjac, elk (wapiti), red deer, and fallow deer) a ...
(
deer A deer (: deer) or true deer is a hoofed ruminant ungulate of the family Cervidae (informally the deer family). Cervidae is divided into subfamilies Cervinae (which includes, among others, muntjac, elk (wapiti), red deer, and fallow deer) ...
) ** Superfamily
Bovoidea Bovoidea is a superfamily of pecoran ruminants containing the Bovidae and Moschidae Moschidae is a family of pecoran even-toed ungulates, containing the musk deer (''Moschus'') and its extinct relatives. They are characterized by long "sabe ...
*** Family
Moschidae Moschidae is a family of pecoran even-toed ungulates, containing the musk deer (''Moschus'') and its extinct relatives. They are characterized by long "saber teeth" instead of horns, antlers or ossicones, modest size (''Moschus'' only reaches ...
(
musk deer Musk deer can refer to any one, or all eight, of the species that make up ''Moschus'', the only extant genus of the family (biology), family Moschidae. Despite being commonly called deer, they are not true deer belonging to the family Cervidae, b ...
) *** Family
Bovidae The Bovidae comprise the family (biology), biological family of cloven-hoofed, ruminant mammals that includes Bos, cattle, bison, Bubalina, buffalo, antelopes (including Caprinae, goat-antelopes), Ovis, sheep and Capra (genus), goats. A member o ...
(
cattle Cattle (''Bos taurus'') are large, domesticated, bovid ungulates widely kept as livestock. They are prominent modern members of the subfamily Bovinae and the most widespread species of the genus '' Bos''. Mature female cattle are calle ...
,
goat The goat or domestic goat (''Capra hircus'') is a species of Caprinae, goat-antelope that is mostly kept as livestock. It was domesticated from the wild goat (''C. aegagrus'') of Southwest Asia and Eastern Europe. The goat is a member of the ...
s,
sheep Sheep (: sheep) or domestic sheep (''Ovis aries'') are a domesticated, ruminant mammal typically kept as livestock. Although the term ''sheep'' can apply to other species in the genus '' Ovis'', in everyday usage it almost always refers to d ...
, and
antelope The term antelope refers to numerous extant or recently extinct species of the ruminant artiodactyl family Bovidae that are indigenous to most of Africa, India, the Middle East, Central Asia, and a small area of Eastern Europe. Antelopes do ...
s) * Parvorder
Giraffomorpha Giraffomorpha is a clade of pecoran ruminants containing the superfamilies Palaeomerycoidea (Palaeomerycidae) and Giraffoidea (Giraffidae, Prolibytheriidae and Climacoceratidae), of which the giraffe and okapi of the Giraffidae are the only e ...
** Family †
Palaeomerycidae The Palaeomerycidae is an extinct family of Neogene ruminants belonging to the infraorder Pecora. Palaeomerycids lived in Europe and Asia exclusively during the Miocene, coevolving with cervids, bovids, moschids, and tragulids there as part o ...
** Superfamily
Giraffoidea Giraffoidea is a superfamily that includes the families Climacoceratidae, Prolibytheriidae, and Giraffidae. The only extant members in the superfamily are the giraffes and okapi. The Climacoceratidae are also placed in the superfamily, but wer ...
*** Family
Giraffidae The Giraffidae are a family (biology), family of ruminant artiodactyl mammals that share a recent common ancestor with deer and bovids. This family, once a diverse group spread throughout Eurasia and Africa, presently comprises only two extant ge ...
(
giraffe The giraffe is a large Fauna of Africa, African even-toed ungulate, hoofed mammal belonging to the genus ''Giraffa.'' It is the Largest mammals#Even-toed Ungulates (Artiodactyla), tallest living terrestrial animal and the largest ruminant on ...
s and
okapi The okapi (; ''Okapia johnstoni''), also known as the forest giraffe, Congolese giraffe and zebra giraffe, is an artiodactyl mammal that is endemic to the northeast Democratic Republic of the Congo in central Africa. However, non-invasive gen ...
) *** Family †
Climacoceratidae Climacoceratidae is a family of superficially deer-like artiodactyl ungulates which lived in the Miocene epoch in Africa. They are close to the ancestry of giraffes, with some genera, such as '' Prolibytherium'', originally classified as giraffes ...


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 Tragulina (also known as Traguliformes) is an infraorder of even-toed ungulates. Only the chevrotains survive to the present, including the genera ''Tragulus'' (the mouse deer) and '' Hyemoschus'', all within the family Tragulidae. Taxonomy and ...
: an
astragalus Astragalus may refer to: * ''Astragalus'' (plant), a large genus of herbs and small shrubs *Astragalus (bone) The talus (; Latin for ankle or ankle bone; : tali), talus bone, astragalus (), or ankle bone is one of the group of foot bones known ...
with parallel sides, a loss of the trapezium, and differences in parts of the skull such as the
petrosal bone The petrous part of the temporal bone is pyramid-shaped and is wedged in at the base of the skull between the sphenoid and occipital bones. Directed medially, forward, and a little upward, it presents a base, an apex, three surfaces, and three a ...
. The distinguishing features of most pecoran families are cranial appendages. Most modern pecorans (with the exception of the
Moschidae Moschidae is a family of pecoran even-toed ungulates, containing the musk deer (''Moschus'') and its extinct relatives. They are characterized by long "saber teeth" instead of horns, antlers or ossicones, modest size (''Moschus'' only reaches ...
) 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 Keratin () is one of a family of structural fibrous proteins also known as ''scleroproteins''. It is the key structural material making up Scale (anatomy), scales, hair, Nail (anatomy), nails, feathers, horn (anatomy), horns, claws, Hoof, hoove ...
. They are indicative of Bovidae. Horns develop in the
periosteum The periosteum is a membrane that covers the outer surface of all bones, except at the articular surfaces (i.e. the parts within a joint space) of long bones. (At the joints of long bones the bone's outer surface is lined with "articular cartila ...
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. *
Antler Antlers are extensions of an animal's skull found in members of the Cervidae (deer) Family (biology), family. Antlers are a single structure composed of bone, cartilage, fibrous tissue, skin, nerves, and blood vessels. They are generally fo ...
s are bony structures that are shed and replaced each year in members of the family
Cervidae A deer (: deer) or true deer is a hoofed ruminant ungulate of the family Cervidae (informally the deer family). Cervidae is divided into subfamilies Cervinae (which includes, among others, muntjac, elk (wapiti), red deer, and fallow deer) a ...
. 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 commonly as the whitetail and the Virginia deer, is a medium-sized species of deer native to North, Central and South America. It is the most widely-distributed mainland ungulate ...
''), or palmate, as in the moose (''
Alces alces The moose (: 'moose'; used in North America) or elk (: 'elk' or 'elks'; used in Eurasia) (''Alces alces'') is the world's tallest, Largest cervids, largest and heaviest extant taxon, extant species of deer and the monotypic taxon, only specie ...
''). *
Ossicone 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 Horn (anatomy), horns and antlers ...
s 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 The Giraffidae are a family (biology), family of ruminant artiodactyl mammals that share a recent common ancestor with deer and bovids. This family, once a diverse group spread throughout Eurasia and Africa, presently comprises only two extant ge ...
and closely related extinct clades, represented in modern animals by the giraffe (''
Giraffa camelopardalis The northern giraffe (''Giraffa camelopardalis''), also known as three-horned giraffe, is the type species of giraffe, ''G. camelopardalis'', and is native to North Africa, although alternative taxonomic hypotheses have proposed the northern gir ...
'') and the okapi ('' Okapia johnstoni''). *
Pronghorn 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 ...
s 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'').


References


External links

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