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Nimravidae is an
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...
of
carnivora Carnivora is a Clade, monophyletic order of Placentalia, placental mammals consisting of the most recent common ancestor of all felidae, cat-like and canidae, dog-like animals, and all descendants of that ancestor. Members of this group are f ...
ns, sometimes known as false saber-toothed cats, whose
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
s are found in
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and th ...
and
Eurasia Eurasia (, ) is the largest continental area on Earth, comprising all of Europe and Asia. Primarily in the Northern and Eastern Hemispheres, it spans from the British Isles and the Iberian Peninsula in the west to the Japanese archipelag ...
. Not considered to belong to the true cats (family
Felidae Felidae () is the family of mammals in the order Carnivora colloquially referred to as cats, and constitutes a clade. A member of this family is also called a felid (). The term "cat" refers both to felids in general and specifically to the dom ...
), the nimravids are generally considered closely related and classified as a distinct family in the
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 ...
Feliformia Feliformia is a suborder within the order Carnivora consisting of "cat-like" carnivorans, including cats (large and small), hyenas, mongooses, viverrids, and related taxa. Feliformia stands in contrast to the other suborder of Carnivora, Canifor ...
. Fossils have been dated from the Middle
Eocene The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', ...
through the Late
Miocene The Miocene ( ) is the first 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 means "less recen ...
epochs (
Bartonian The Bartonian is, in the ICS's geologic time scale, a stage or age in the middle Eocene Epoch or Series. The Bartonian Age spans the time between . It is preceded by the Lutetian and is followed by the Priabonian Age. Stratigraphic defini ...
through
Tortonian The Tortonian is in the geologic time scale an age or stage of the late Miocene that spans the time between 11.608 ± 0.005 Ma and 7.246 ± 0.005 Ma (million years ago). It follows the Serravallian and is followed by the Messinian. The Torto ...
stages, 40.4–7.2 million years ago), spanning about . The
barbourofelid Barbourofelidae is an extinct family of carnivorans of the suborder Feliformia, sometimes known as false saber-toothed cats, that lived in North America, Eurasia and Africa during the Miocene epoch (16.9—9.0 million years ago) and existed for a ...
s, which were formerly classified as a
subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus Genus ( plural genera ) is a taxonomic rank used in the biological classifica ...
of the Nimravidae, were reassigned to their own distinct family
Barbourofelidae Barbourofelidae is an extinct family of carnivorans of the suborder Feliformia, sometimes known as false saber-toothed cats, that lived in North America, Eurasia and Africa during the Miocene epoch (16.9—9.0 million years ago) and existed for a ...
in 2004. However, some recent studies suggest the barbourofelids are a branch of the nimravids, suggesting that this debate might not be settled yet.


Morphology and evolution

Most nimravids had muscular, low-slung, cat-like bodies, with shorter legs and tails than are typical of cats. Unlike
extant Extant is the opposite of the word extinct. It may refer to: * Extant hereditary titles * Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English * Extant taxon, a taxon which is not extinct, ...
Feliformia, the nimravids had a different bone structure in the small bones of the ear. The
middle ear The middle ear is the portion of the ear medial to the eardrum, and distal to the oval window of the cochlea (of the inner ear). The mammalian middle ear contains three ossicles, which transfer the vibrations of the eardrum into waves in the ...
of true cats is housed in an external structure called an
auditory bulla The tympanic part of the temporal bone is a curved plate of bone lying below the squamous part of the temporal bone, in front of the mastoid process, and surrounding the external part of the ear canal. It originates as a separate bone (tympanic ...
, which is separated by a
septum In biology, a septum (Latin for ''something that encloses''; plural septa) is a wall, dividing a cavity or structure into smaller ones. A cavity or structure divided in this way may be referred to as septate. Examples Human anatomy * Interat ...
into two chambers. Nimravid remains show
ossified Ossification (also called osteogenesis or bone mineralization) in bone remodeling is the process of laying down new bone material by cells named osteoblasts. It is synonymous with bone tissue formation. There are two processes resulting in t ...
bullae with no septum, or no trace at all of the entire bulla. They are assumed to have had a
cartilaginous Cartilage is a resilient and smooth type of connective tissue. In tetrapods, it covers and protects the ends of long bones at the joints as articular cartilage, and is a structural component of many body parts including the rib cage, the neck a ...
housing of the ear mechanism. Nimravid feet were short, indicating they walked in a
plantigrade 151px, Portion of a human skeleton, showing plantigrade habit In terrestrial animals, plantigrade locomotion means walking with the toes and metatarsals flat on the ground. It is one of three forms of locomotion adopted by terrestrial mammals. ...
or semiplantigrade posture, i.e., on the flat of the feet rather than the toes, like modern cats. Although some nimravids physically resembled the saber-toothed cats, such as ''
Smilodon ''Smilodon'' is a genus of the extinct machairodont subfamily of the felids. It is one of the most famous prehistoric mammals and the best known saber-toothed cat. Although commonly known as the saber-toothed tiger, it was not closely rela ...
'', they were not closely related, but
evolved Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variati ...
a similar form through
parallel evolution Parallel evolution is the similar development of a trait in distinct species that are not closely related, but share a similar original trait in response to similar evolutionary pressure.Zhang, J. and Kumar, S. 1997Detection of convergent and paral ...
. They possessed
synapomorphies In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to ha ...
with the barbourofelids in the
cranium The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, th ...
,
mandible In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...
,
dentition Dentition pertains to the development of teeth and their arrangement in the mouth. In particular, it is the characteristic arrangement, kind, and number of teeth in a given species at a given age. That is, the number, type, and morpho-physiolo ...
, and postcranium. They also had a downward-projecting flange on the front of the mandible as long as the canine teeth, a feature which also convergently evolved in the saber-toothed sparassodont ''
Thylacosmilus ''Thylacosmilus'' is an extinct genus of saber-toothed metatherian mammals that inhabited South America from the Late Miocene to Pliocene epochs. Though ''Thylacosmilus'' looks similar to the " saber-toothed cats", it was not a felid, like the ...
''. The ancestors of nimravids and cats diverged from a common ancestor soon after the
Caniformia Caniformia is a suborder within the order Carnivora consisting of "dog-like" carnivorans. They include dogs (wolves, foxes, etc.), bears, raccoons, and mustelids. The Pinnipedia (seals, walruses and sea lions) are also assigned to this group. ...
Feliformia Feliformia is a suborder within the order Carnivora consisting of "cat-like" carnivorans, including cats (large and small), hyenas, mongooses, viverrids, and related taxa. Feliformia stands in contrast to the other suborder of Carnivora, Canifor ...
split, in the middle
Eocene The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', ...
about 50 million years ago ( Mya), with a minimum constraint of 43 Mya. Recognizable nimravid
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
s date from the late Eocene (37 Mya), from the
Chadronian The Chadronian age within the North American Land Mammal Ages chronology is the North American faunal stage typically set from 38,000,000 to 33,900,000 years BP, a period of . It is usually considered to fall within the Eocene epoch. The Chadroni ...
White River Formation at Flagstaff Rim,
Wyoming Wyoming () is a state in the Mountain West subregion of the Western United States. It is bordered by Montana to the north and northwest, South Dakota and Nebraska to the east, Idaho to the west, Utah to the southwest, and Colorado to t ...
, to the late
Miocene The Miocene ( ) is the first 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 means "less recen ...
(5 Mya). Nimravid diversity appears to have peaked about 28 Mya. A 2021 study has shown that a sizeable amount of species developed feline-like morphologies in addition to saber-toothed taxa.


Taxonomy

The family Nimravidae was named by American paleontologist
Edward Drinker Cope Edward Drinker Cope (July 28, 1840 – April 12, 1897) was an American zoologist, paleontologist, comparative anatomist, herpetologist, and ichthyologist. Born to a wealthy Quaker family, Cope distinguished himself as a child prodigy interes ...
in 1880, with the type genus as ''
Nimravus ''Nimravus'' is an extinct genus of "false" saber-toothed cat in the family Nimravidae, that was endemic to North America during the Oligocene epoch 33.3—26.3 Annum, mya, existing for approximately . Not closely related to true saber-toothed ca ...
.'' The family was assigned to Fissipedia by Cope (1889); to Caniformia by Flynn and Galiano (1982); to Aeluroidea by Carroll (1988); to Feliformia by Bryant (1991); and to Carnivoramorpha, by Wesley-Hunt and Werdelin (2005). Nimravids are placed in
tribes The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide usage of the term in English is in the discipline of anthropology. This definition is contested, in part due to confli ...
by some authors to reflect closer relationships between genera within the family. Some nimravids evolved into large, toothed, cat-like forms with massive flattened upper canines and accompanying mandibular flanges. Some had dentition similar to felids, or modern cats, with smaller canines. Others had moderately increased canines in a more intermediate relationship between the saber-toothed cats and felids. The upper canines were not only shorter, but also more conical, than those of the true saber-toothed cats (
Machairodontinae Machairodontinae is an extinct subfamily of carnivoran mammals of the family Felidae (true cats). They were found in Asia, Africa, North America, South America, and Europe from the Miocene to the Pleistocene, living from about 16 million unti ...
). These nimravids are referred to as "false saber-tooths". Not only did nimravids exhibit diverse dentition, but they also showed the same diversity in size and morphology as cats. Some were
leopard The leopard (''Panthera pardus'') is one of the five extant species in the genus '' Panthera'', a member of the cat family, Felidae. It occurs in a wide range in sub-Saharan Africa, in some parts of Western and Central Asia, Southern Russia, ...
-sized, others the size of today's
lion The lion (''Panthera leo'') is a large cat of the genus '' Panthera'' native to Africa and India. It has a muscular, broad-chested body; short, rounded head; round ears; and a hairy tuft at the end of its tail. It is sexually dimorphic; adu ...
s and
tiger The tiger (''Panthera tigris'') is the largest living Felidae, cat species and a member of the genus ''Panthera''. It is most recognisable for its dark vertical stripes on orange fur with a white underside. An apex predator, it primarily pr ...
s, one had the short face, rounded skull, and smaller canines of the modern
cheetah The cheetah (''Acinonyx jubatus'') is a large cat native to Africa and central Iran. It is the fastest land animal, estimated to be capable of running at with the fastest reliably recorded speeds being , and as such has evolved specialized ...
, and one, '' Nanosmilus'', was only the size of a small
bobcat The bobcat (''Lynx rufus''), also known as the red lynx, is a medium-sized cat native to North America. It ranges from southern Canada through most of the contiguous United States to Oaxaca in Mexico. It is listed as Least Concern on the ...
. The Barbourofelids were for a while no longer included in Nimravidae, following elevation to family as sister clade to the true cats (family
Felidae Felidae () is the family of mammals in the order Carnivora colloquially referred to as cats, and constitutes a clade. A member of this family is also called a felid (). The term "cat" refers both to felids in general and specifically to the dom ...
). However, several recent studies have returned them to Nimravidae, including as part of Nimravinae.


Phylogeny

The phylogenetic relationships of Nimravidae are shown in the following cladogram: A 2021 study divides Nimravidae into Hoplophoninae and Nimravinae, the latter including the bulk of species in addition to barbourofelids.


Natural history

Nimravids appeared in the middle of the
Eocene The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', ...
epoch, about 40 mya, in North America and Asia. The global climate at this time was warm and wet, but was trending cooler and drier toward the late Eocene. The lush forests of the
Eocene The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', ...
were transforming to scrub and open
woodland A woodland () is, in the broad sense, land covered with trees, or in a narrow sense, synonymous with wood (or in the U.S., the '' plurale tantum'' woods), a low-density forest forming open habitats with plenty of sunlight and limited shade (see ...
. This climatic trend continued in the
Oligocene The Oligocene ( ) is a geologic epoch of the Paleogene Period and extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that define the epoch are well identified but t ...
, and nimravids evidently flourished in this environment. North America and Asia were connected and shared much related fauna. Europe in the Oligocene was more of an
archipelago An archipelago ( ), sometimes called an island group or island chain, is a chain, cluster, or collection of islands, or sometimes a sea containing a small number of scattered islands. Examples of archipelagos include: the Indonesian Arch ...
than a continent, though some land bridges must have existed, for nimravids also spread there. In the Miocene, the fossil record suggests that many animals suited for living in forest or woodland were replaced by grazers suited for grassland. This suggests that much of North America and Asia became dominated by savanna. Nimravids disappeared along with the woodlands, but survived in relictual humid forests in Europe to the late Miocene. When conditions ultimately changed there in the late Miocene, the last nimravids disappeared about 9 Mya.


See also

*
Oxyaenidae Oxyaenidae ("sharp hyenas") is a family of extinct carnivorous placental mammals. Traditionally classified in order Creodonta, this group is now classified in its own order Oxyaenodonta ("sharp tooth hyenas") within clade Pan-Carnivora in mirord ...


References


External links

* * {{Taxonbar, from=Q131889 Prehistoric mammal families Bartonian first appearances Tortonian extinctions Taxa named by Edward Drinker Cope Fossil taxa described in 1880