User:Jts1882/Cat evolution

The Evolution of the Felidae begins in the emergence of cat-like carnivores …

The "false sabre toothed cats", the Barbourofelidae and Nimravidae, are not true cats, but are closely related and together with Felidae and other cat-like carnivores (hyaenas, viverrids and mongooses) make up the feliform carnivores.

The first true cats of the Family Felidae emerged during the Oligocene, about 25 million years ago, with the appearance of Proailurus and Pseudaelurus. The latter species complex was ancestral to two main lines of felids, the cats in the extant subfamilies, Pantherinae and Felinae, and a group of extinct cats, which are assigned to the subfamily Machairodontinae. The machairodonts included the saber-toothed cats such as the Smilodon.

The conical toothed cats, which include all modern cats, originated in Asia and spread across continents by crossing land bridges. Testing of mitochondrial and nuclear DNA revealed that the ancient cats evolved into eight main lineages that diverged in the course of at least 10 migrations (in both directions) from continent to continent via the Bering land bridge and the Isthmus of Panama, with the Panthera genus being the oldest and the Felis genus being the youngest. About 60% of the modern cat species are estimated to have developed within the last million years.

=Phylogeny and overview of cat evolution=

Carnivora and Feliforma (cladograms)

 * Text from Cat gap

All modern carnivorans, including cats, evolved from miacoids, which existed from approximately 66 to 33 million years ago. Miacoid species gave rise to Proailurus (meaning "first cat"; also called "Leman's Dawn Cat"), which appeared about 30 million years ago, and is generally considered the first "true cat". There were other cat-like species before Proailurus, but not within the family of the order Carnivora.

Following the appearance of the dawn cat, there is little in the fossil record for 10 million years to suggest that cats would prosper. In fact, although Proailurus persisted for at least 14 million years, there are so few felid fossils towards the end of the dawn cat's reign that paleontologists refer to this as the "cat gap". The turning point for cats came about with the appearance of a new genus of felids, Pseudaelurus

The increase in disparity through the early Miocene occurs during a time when few feliform fossils have been found in North America. The hypercarnivorous nimravid feliforms were extinct in North America after 26 Ma and felids did not arrive in North America until the Middle Miocene with the appearance of Pseudaelurus. Pseudaelurus crossed over to North America by way of the Bering land bridge from surviving populations in Asia 18.5 million years ago. All modern-day cats are descended from Pseudaelurus.

Nimravids were saber-toothed cat-like animals of the family Nimravidae. Although not "true cats" in the family Felidae, Nimravidae are considered to be a sister taxon to felids. They are basal feliforms, but their exact placement within Carnivora is still uncertain. Physically, Nimravidae resembled Smilodon (which would not evolve until many millions of years later). Nimravidae also became extinct in North America during the "cat gap."

Early cats

 * First felid. Proailurus
 * Three lineages of Pseudaelurus (cladogram)

Proailurus
Proailurus (meaning "first cat"; also known as Leman’s dawn cat) is an extinct carnivoran felid that lived in Europe and Asia approximately 25 million years ago in the Late Oligocene and Miocene. Fossils have been found in Mongolia, Germany, and Spain. One recent phylogeny places it as a basal member of the Feliformia, the suborder that includes mongooses, civets, hyenas, and cats; but other studies suggest that it instead was a felid (a true cat).

Proailurus was a compact and small animal, just a little larger than the domestic cat, weighing about 20 lb (9 kg). It had a long tail, large eyes and sharp claws and teeth, with similar proportions to the modern viverrids. Its claws would have been retractable to some extent. Like the viverrids, Proailurus was at least partially arboreal.

Proailurus was a likely ancestor of Pseudaelurus, which lived 20-10 million years ago, and probably gave rise to the major felid lines, including the extinct machairodontines and the extant felines and pantherines, although the phylogeny of the cats is still not precisely known.

Pseudaelurus
Pseudaelurus is a prehistoric cat that lived in Europe, Asia and North America in the Miocene between approximately 20 to 8 million years ago. It is an ancestor of today's felines and pantherines as well as the extinct machairodont saber-tooths, and is a successor to Proailurus. It originated from Eurasia and was the first felid to reach North America, when it entered the continent at about 18.5 Ma ending a 'cat-gap' of 7 million years. The slender proportions of the animal, together with its short, viverrid-like legs, suggest that it may have been an agile climber of trees.

Traditionally all all the Pseudaelurus species from Europe, Asia and North America have been assigned to a single genus, even though the paraphyletic nature of the group has often being noted and several authorities have split Pseudaelurus into separate genera or subgenera, including Hyperailurictis, Styriofelis, Miopanthera and Schizailurus (see Werdelin et al 2010). Pseudaelurus thus represents an evolutionary grade, with several different subgroups containing the ancestors to later felids.

Werdelin el al (2010) suggested dividing Pseudaelurus to represent three separate lineages, each giving rise to a different descendant group of felids. The North American species were assigned to Hyperailurictis, which is ancestral to the North American genus Nimravides. Three European species – P. turnauensis, P. loreti, P. romieviensis – were assigned to Styriofelis (junior synonyms Miopanthera and Schizailurus), which forms the sister taxon to all modern felids, the Felinae sensu lato (i.e. including the pantherine cats). The remaining Eurasian Pseudaelurus species form a sister group to the sabre-toothed cats, the Machairodontinae.

The following cladogram is based on Piras et al (2013) and illustrates the three more derived lineages that evolved from "Pseudaelurus" species

Sabre-toothed cats

 * Traditional tribes
 * Eumachairodonta

The Machairodontinae originated in the early or middle Miocene of Africa. The early felid Pseudaelurus quadridentatus showed a trend towards elongated upper canines, and is believed to be at the base of the machairodontine evolution. The earliest known machairodont genus is the middle Miocene Miomachairodus from Africa and Turkey. Until the late Miocene, machairodontines co-existed at several places together with barbourofelids, archaic large carnivores that also bore long sabre-teeth.

Traditionally, three different tribes of machairodontines were recognized, the Smilodontini with typical dirk-toothed forms, such as Megantereon and Smilodon, the Machairodontini or Homotherini with scimitar-toothed cats, such as Machairodus or Homotherium, and the Metailurini, containing genera such as Dinofelis and Metailurus. However, some have recently regrouped the Metailurini within the other felid subfamily, the Felinae, along with all modern cats. The last machairodontine genera, Smilodon and Homotherium, did not disappear until late in the Pleistocene, roughly 10,000 years ago in the Americas.

The name 'saber-toothed tigers' is misleading. Machairodonts were not even in the same subfamily as tigers, there is no evidence that they had tiger-like coat patterns, and this broad group of animals certainly did not all live or hunt in the same manner as the modern tiger. DNA analysis published in 2005 confirmed and clarified cladistic analysis in showing that the Machairodontinae diverged early from the ancestors of modern cats and are not closely related to any living feline species.

Saber-tooths also coexisted in many places with conical-toothed cats. In Africa and Eurasia, sabertooth cats competed with several pantherines and cheetahs until the early or middle Pleistocene. Homotherium survived in northern Europe even until the late Pleistocene. In the Americas, they coexisted with the cougar, American lion, American cheetah, and jaguar until the late Pleistocene. Saber-toothed and conical-toothed cats competed with each other for food resources, until the last of the former became extinct. All recent felids have more or less conical-shaped upper canines.

Evolutionary history and origin of phenotype
Phylogeny of Machairodontines with the three out-groups, Proailurus, Pseudaelurus, and all modern, conical-toothed cats, with a brief description of each genus:

Until the recent discovery of the Late Miocene fossil depository known as Batallones-1 in the 1990s, specimens of Smilodontini and Homotheriini ancestors were rare and fragmentary, so the evolutionary history of the saber-toothed phenotype, a phenotype affecting craniomandibular, cervical forelimb and forelimb anatomy, was largely unknown. Prior to the excavation of Batallones-1, the predominating hypothesis was that the highly derived saber-toothed phenotype arose rapidly through pleiotropic evolution. Batollnes-1 unearthed new specimens of Promegantereon ogygia, a Smilodontini ancestor, and Machairodus aphanistus, a Homotheriini ancestor, shedding light on evolutionary history. (Though the Smilodontini ancestor was originally assigned to the genus Paramachairodus, it was later revised to the genus Promegantereon). The leopard-sized P. ogygia (living 9.0 Ma) inhabited Spain (and perhaps additional territory), and its most studied descendants, the members of the tiger-sized genus Smilodon, lived up to 10,000 years ago in the Americas. The lion-sized M. aphanistus (living 15.0 Ma) roamed Eurasia, as did its most studied descendants, members of the lion-sized genus Homotherium (living 3.0-5.0 Ma).

The current hypothesis for the evolution of the saber-toothed phenotype, made possible by Batollnes-1, is that this phenotype arose gradually over time through mosaic evolution. Although the exact cause is uncertain, current findings have supported the hypothesis that a need for the rapid killing of prey was the principle pressure driving the development of the phenotype over evolutionary time. As indicated by high instances of broken teeth, the biotic environment of saber-toothed cats was one marked by intense competition.

Broken teeth indicate the frequency at which teeth contact bone. Increased teeth-bone contact suggests either increased consumption of carcasses, rapid consumption of prey, or increased aggression over kills – all three of which point to decreased prey availability, heightening competition between predators. Such a competitive environment would favor the faster killing of prey, because if prey is taken away before consumption (such as by out-competing) the energetic cost of capturing that prey is not reimbursed, and, if this occurs often enough in the lifetime of a predator, death by exhaustion or starvation would result. The earliest adaptations improving the speed at which prey was killed are present in the skull and mandible of P. ogygia and of M. aphanistus, and in the cervical vertebrae and forelimb of P. ogygia. They provide further morphological evidence for the importance of speed in the evolution of the saber-toothed phenotype.

Evolutionary Relationships

 * copied from Machairodontini (June 28, 2017

Previous analysis of the saber-toothed cats used Linnaean classification based the morphology of the upper canines, which suggested that there were four tribes within the subfamily of Machairodontinae. These tribes include Machairodontini, Homotherini, Smildontini, and Matailurini seen in the cladogram of Machairodontinae. Then upon finding further fossils, Machairodontini was assigned to the Homotherini tribe. The Homotherini includes species such as Machairodus, (M. aphanistus and M. giganteus). These species are characterized by their scimitar teeth that make up the large upper canines. The Smildontini tribe includes species such as Paramachairodus, Megantereon, and Smilodon. These species have been characterized based on their longer, dagger-like teeth. The other tribe Matailurini includes species such as Matailurus and Dinofelis. These have been classified by having teeth different than the previous two clades, the long serrated teeth and the long, dagger-like teeth. Unfortunately, there are few postcranial skeletons of saber-toothed cats preserved, so available phylogenetic characters are largely cranial. New research published in 2013 took a different approach to elucidating the evolutionary relationships of saber-toothed cats. This approach used cladistic phylogeny instead of the previous Linnaean classification. The results concluded the original four tribes could not be recovered by parsimony analysis because many of the synapomorphies of the saber toothed cats are based on other bones that the prominent, upper canines.

Workers now regocnize a clade for true saber toothed cats called Eumachairodontia that includes species from all of the previously proposed tribes: Megantereon, Smilodon, Amphimachairodus, Homotherium and Xenosmilus. The synapomorphy for the “true saber-toothed” clade Eumachairodontia is the hypertrophied, greatly flattened upper canines. The true machairodontines have a synapomorphy of flattened, small, lower canines and other bone variations such as the small upper first molar compared to basal, pre-saber toothed cats large, transversely situated upper first molar and large upper third premolar parastyle.

Conical-toothed cats

 * The eight felid lineages

Traditionally, the conical-toothed cats are divided into three subfamilies based on phenotypical features: the Felinae, the Pantherinae, and the Acinonychinae (cheetahs).

Molecular phylogenetic analysis suggests that living (extant) felids fall into eight lineages (clades). The placement of the cheetah within the Puma lineage invalidates the traditional subfamily Acinonychinae, and recent sources use only two subfamilies for extant genera. The eight lineages divide between these as follows:


 * Subfamily Pantherinae:
 * Lineage 1 (Panthera lineage): Panthera, Neofelis
 * Subfamily Felinae:
 * Lineage 2 (Bay cat lineage): Pardofelis, Catopuma
 * Lineage 3 (Caracal lineage): Leptailurus, Caracal (including Profelis)
 * Lineage 4 (Ocelot cat lineage): Leopardus
 * Lineage 5 (Lynx lineage): Lynx
 * Lineage 6 (Puma lineage): Puma, Acinonyx
 * Lineage 7 (Leopard cat lineage): Prionailurus, Otocolobus
 * Lineage 8 (Domestic cat lineage): Felis

The last four lineages are more related to each other than to any of the first four, so form a clade within the Felinae subfamily of family Felidae.

Phylogeny
The phylogenetic relationships of extant felids are shown in the following cladogram, based on the molecular phylogenetic analysis of Johnson et al. (2006). The lineages, genera and species are as used in that study.

=Palaeobiology and Biogeography (evolutionary time line)=

First cats
Climate cooling during miocene First felids occur after the ‘Grande Coupure’ marking the Eocene/Oligocene boundary (c. 33.9 Ma;

Barbourofelidae lineage (from 32 mya to about 10mya) (Eurasian and Africa) Haplogale, Stenogale and Proailurus lineage

Proailurus, the first true cat, is found in Europe about 30 mya. Arboreal. A similar felid with primative featires was found in NA around 17mya. Could be first NA felid and ancestor of NA Pseudaelurus grade felids or Hyperailurictis.

Addition species described by Peigne? (not in PaleoDB)

Pseudaelurus grade cats appear about 20 mya. Climate change and open landscapes.

Four species of Pseudaelurus are known from Europe. (first three are styriofelis) In the order of increasing size they are: P. turnauensis (= P. transitorius), (earliest and smallest, 20mya) P. lorteti, in MN 4 (18–17 Ma). P. romieviensis, in MN 4 (18–17 Ma). P. quadridentatus (in MN 4 (18–17 Ma).

P. lorteti and P. romieviensis become extinct at the end of the Middle Miocene (c. 11.6 Ma), However, P. quadridentatus and P. turnauensis survive into the Late Miocene (MN 9, c. 11.2–9.5 Ma), contemporary with Miomachairodus pseudailuroides, the earliest record of Machairodontinae, which was found in Turkey (Schmidt-Kittler 1976; Viranta and Werdelin 2003 via Werdelin 2010)

In North America
NA Pseudaelurus P. validus (stratigraphic range c. 17.5–16.5 Ma), [Lynx to Puma-sized] P. Skinneri (c. 17.5–17.1 Ma), P. Intrepidus (c. 17.1–13.3 Ma), P. Stouti (c. 15.2–12.7 Ma), and P. Marshi (c. 16.4–12.7 Ma).

Migrations to NA (after Qiu 2003):

Migration to NA around 20 Ma, probably from 21 Ma to 19–18 Ma: Proailurus-like felid?

Nimravides in North America 14mya

Four species of Nimravides are known: N. Thinobates (c. 11.0–9.6 Ma), N. Pedionomus (c. 12.0–11.5 Ma), N. Hibbardi (c. 7.0–6.4 Ma), and N. Galiani (c. 11.6–10.7 Ma).

Sabre-toothed felids
Ginsburgsmilus:Africa during the early Miocene. There is only one known specimen of Ginsburgsmilus napakensis,[2] dated to 20-19 mya.

Prosansanosmilus ... Europe during the Miocene epoch (16.9—16.0 mya)

Barbourofelids (16.9—9.0 Ma) first appear in the fossil record in the Early Miocene of Africa. By the end of the Early Miocene, a land bridge had opened between Africa and Eurasia, allowing for a faunal exchange between the two continents. Barbourofelids migrated at least three times from Africa to Europe (Morlo 2006).[6] While the genus Sansanosmilus evolved in Europe, barbourofelids also migrated through Eurasia and reached North America by the late Miocene, represented there solely by the genus Barbourofelis. (wikipedia: Barbourofelidae)

Sansanosmilus? endemic to Europe and Asia, which lived during the Miocene, 13.6—11.1 mya (none in NA) Migration to NA around 13 mya: Sansanosmilus? (Qiu-2003) ancestor of Barbourofelis

First machairodontinae .. lineages and locations [In Europe 13-14mya]

Miomachaerodus pseudailuroides in Turkey [MN 7/8 and MN 9 (c. 12.5–9.5 Ma). ] =Machaerodus aphanistus (Ginsberg 1981)

Anton-2004: Machairodus should be restricted in content to Vallesian (c. 11.2–9.0 Ma) forms, while Turolian (c. 9.0–5.3 Ma) forms should be referred to Amphimachairodus (Fig. 2.2, letter F).

Amphimachairodus lineage in Eurasia 10mya

Machairodus migrated to North America (some time in the Vallesian, (11.6—9.0 Ma)), ancestral to M. coloradensis (c. 9.0–5.3 Ma)

Paramachaerodus lineage Eurasia 11–10 Paramachaerodus and derivates Eurasia 9

Metailurini Eurasia 9–8

?Migration to NA around 7-8 mya: Machairodus. (Qiu-2003)

Homotheriini in Eurasia 6mya Derived Homotheriini in Africa 5mya North American Homotheriini ca 4–3

Smilodontini Eurasia, Africa, and North America 5–4

Migration to NA around 4 mya: Homotherium, Lynx (Qiu-2003) Migration to Eurasia from NA around 4 mya: Meganteron (Qiu-2003)

Splitting into lineages – location and spread
Data from Li et al 2006; average divergence times from Table S3]


 * Panthera lineage: diverged 11.5mya/ radiate 5.7 (Neofelis sepatates 5.7 mys, division of Panthera at 3.7mya
 * Small cats into seven lineages
 * Caracal lineage (3 species) diverghes at 10.7mya
 * Leptailurus (serval) 6.3mya serval splits off
 * Caracal (Caracal, African golden cat)
 * Ocelot lineage (8 species) 9.8 mya/3.1mya
 * Leopardus (northern and southern tiger cats, guina, Geoffrooy’s cat, Andean cat, Pampas cat, Margay, Ocelot)
 * 8.8mya lynx and bay cat group diverges from Puma-leopard cat-domestic cat line
 * Splits at 8.1mya (bay cat from lynx lineage
 * Bay cat lineage: (3 species)
 * Pardofelis (marbled cat) diverges at 5.5.mya
 * Catopuma (Bay cat, Asian golden cat)
 * Lynx lineage (4 species) – modern species from 3.5mya (bobcat from lynxes which split at 1.6mya)
 * Lynx (bobcat, Canada, Eurasian and Iberian lynxes)
 * Puma lineage (3 species) diverges at 8.2mya (from leopard cat-domestic cat lineage)
 * Acinonyx (Cheetah): splits off at 5mya
 * Puma genus, splits 3.7mya into
 * Herpailurus (Jaguarundi)
 * Puma (Puma)
 * 7..3mya final split between leopard cat and domestic cat lineahes
 * Leopard cat lineage (five species) – splits at 6mya (Otocolobus and Prionailurus)
 * Otocolobus (Pallas cat)
 * Prionailurus (Leopard cat, fishing cat, flat headed cat, rusty spotted cat) divereges at 3.8mya
 * Domestic cat lineage (5 species): last common ancestor of living species 4.2mya
 * Felis (wildcat (including domestic cat), Chinese mountain cat, sand cat, black-footed cat, jungle cat)

=The big cats=

Pantherine evolution and biogeography

Extinct pantherines

Lions



 * Earliest possible lion fossil ~4mya

Cave lions:


 * Panthera leo spelaea or P. spelaea, the Eurasian or European cave lion
 * Panthera leo fossilis or P. spelaea fossilis or P. fossilis, the Early Middle Pleistocene European cave lion
 * Panthera leo atrox or P. atrox, the American lion or American cave lion

From Lion:

P. leo evolved in Africa between 1 million and 800,000 years ago, before spreading throughout the Holarctic region. It appeared in the fossil record in Europe for the first time 700,000 years ago with the subspecies Panthera leo fossilis at Isernia in Italy. From this lion derived the later cave lion (Panthera leo spelaea), which appeared about 300,000 years ago. Lions died out in northern Eurasia at the end of the last glaciation, about 10,000 years ago; this may have been secondary to the extinction of Pleistocene megafauna.

Tigers
... evolutionary history of the tiger, the expansion and contraction of its range after the Middle Pleistocene.... biogeographic diversification, distribution and evolutionary success of the tiger ... bottleneck and the Toba volcano, with subsequent expansion from a refugium, and the effects of the LGM.


 * Sources on the evolutionary history of the tiger (from Talk:Tiger)
 * Biogeographical change in the tiger Panthera_tigris (Kitchener & Dugmore, 2000)
 * Predicted Pleistocene–Holocene range shifts of the tiger (Panthera tigris) (Cooper et al, 2016)
 * Planning tiger recovery: Understanding intraspecific variation for effective conservation (Wilting et al, 2015)
 * Mitochondrial Phylogeography Illuminates the Origin of the Extinct Caspian Tiger and Its Relationship to the Amur Tiger (Driscoll et al, 2009)
 * The Last Wild Tigers (Matthiessen, 2014)
 * http://www.paleobiodb.org/classic/displayReference?reference_no=34644
 * A taxonomic revision of the tigers (Panthera tigris) of Southeast Asia (Mazak & Groves, 2006. This a study of the island tiger subspecies and proposes to separate tigers into three distinct species based on the Phylogenetic Species Concept. The principle author (Mazak) wrote the ASM Mammalian Species guide for the tiger. The proposal hasn't been followed but is similar to the proposals to reduce the subspecies to two and the article discusses the geographic isolation.
 * Phylogeography and Genetic Ancestry of Tigers (Panthera tigris) (Luo et al, 2004) - Mt-DNA analysis suggests common ancestor of modern tigers around 75kya. Discusses possibility of Toba eruption and retreat to refugium in south China.


 * The Caspian Tiger,(Panthera tigris virgata) genetic roots span over a million years. A nearly cataclysmic event during the late Pleistocene Era, some 10,000 to 12,000 years ago nearly wiped out all tigers. Fortunately, a small remnant of tigers survived.


 * Read more at: https://phys.org/news/2009-01-caspian-tiger-extinct-siberian.html#jCp

The tiger lineage overall goes back more than a million years (see Wilting et al 2015 for references). Then around 75,000 years ago the Toba supereruption caused a global winter and this had a catastrophic effect on tiger populations, reducing them to a refugium in southern China (see Luo et al 2004, Wilting et al, 2015 ), from which the modern tigers expanded and differentiated into subspecies. This refugium explains the basal position of the South Chinese tiger (P. tigris amoyensis) in Mt-DNA analysis. Later, the LCM (10-20kya?) cause a retreat from northern Asia. Afterwards (ca 10kya) the ancestors of the Siberian and Caspian tigers migrated back into the vacated northern areas, either by the Gansu-silk road route (Driscoll et al 2009) or via northern China (Cooper et al, 2016).

=References=