The Felidae:
Cats and Kin After the flood
Chad Arment (2025)
Felines are well-recognized within the creation model as a mammalian family (the Felidae) which shows extensive diversification after the Flood. As only one pair of every unclean terrestrial kind was on the Ark, all living cat species are descended from a single ancestral pair on the Ark. Extensive intrafamilial hybridization, the cognitum for the group, and statistical baraminological studies (incorporating all three subfamilies, including the extinct sabertooth cats) all support familial membership within a single baramin (Robinson and Cavanaugh 1998; Sanders and Wise 2003; Pendragon and Winkler 2011; Lightner 2012; Thompson and Wood 2018).
As with many other baramins, we cannot assume that a given family is the entirety of the baramin. Discontinuity can be noted, but there are many other examples of extensive diversification with familial discontinuity where the only logical interpretation of the data is a single baramin (e.g. lemurs, sloths, etc.). The family level is best considered a minimal baraminic boundary for post-Flood diversification with most Cenozoic terrestrial vertebrates. The baramin itself may be superfamilial or multifamilial in regard to the known fossil record. Wise (2009), for example, proposed the Felidae was part of a baramin encompassing the Feliformia. That would suggest that the original Ark pair was civet- or genet-like, more ‘primitive’ or ‘basal’ in morphology, while modern cats are a more specialized offshoot. In any case, diversification after the Flood was rapid and extensive (Wood 2013), with the various feline clades distinguishing themselves during the post-Flood period equivalent to ‘Miocene’ stratigraphy prior to the Ice Age. (Within an Upper Cenozoic Flood Boundary creation model, strata up through the Cretaceous at least are primarily Flood deposit, while Cenozoic layers beginning somewhere in the Paleogene are a roughly chronological record of the post-Flood period up through the Pleistocene Ice Age and into the modern era.) I suspect the Feliformia may be a single baraminic lineage, but here I will just consider the Felidae as a probable monobaramin.
Feline Hybridization
It is always worth looking at the known hybrids of the Felidae, simply because so many hybrids have been reported.
Captive hybridization:
Felis catus (domestic cat) x Caracal caracal (caracal) (Hartwell 1993-2015)
Felis catus x Felis chaus (jungle cat) (Gray 1954; JFMS 2017)
Felis catus x Felis margarita (sand cat) (Hartwell 1993-2015)
Felis catus x Felis nigripes (black-footed cat) (Hartwell 1993-2015)
Felis catus x Leopardus geoffroyi (Geoffroy’s cat) (JFMS 2017; Rodriguez et al. 2021)
Felis catus x Leopardus tigrina (oncilla) (Hartwell 1993-2015)
Felis catus x Leopardus wiedii (margay) (JFMS 2017)
Felis catus x Lynx rufus (bobcat) (Van Gelder 1977)*
Felis catus x Prionailurus bengalensis (Asian leopard cat) (Davis et al. 2015)
Felis catus x Prionailurus viverrinus (fishing cat) (JFMS 2017)
Felis catus x Leptailurus serval (serval) (Davis et al. 2015)
Felis lybica (African wildcat) x Felis silvestris (European wildcat) (Gray 1954)
Leptailurus serval x Caracal caracal (Hartwell 1993-2015)
Leopardus pardalis (ocelot) x Leopardus wiedii (JFMS 2017)
Leopardus pardalis x Lynx rufus (Hartwell 1993-2015)
Leopardus pardalis x Puma concolor (cougar) (Dubost and Royère 1993)
Panthera leo (lion) x Panthera tigris (tiger) (Van Gelder 1977)
Panthera leo x Panthera pardus (leopard) (Van Gelder 1977)
Panthera leo x Panthera onca (jaguar) (Van Gelder 1977)
Panthera pardus x Panthera onca (Van Gelder 1977)
Panthera pardus x Panthera tigris (Van Gelder 1977)
Panthera pardus x Puma concolor (Gray 1954)
Natural hybridization:
Felis catus x Felis bieti (Chinese mountain cat) (Yu et al. 2021)
Felis catus x Felis lybica (Gray 1954; Tiesmeyer et al. 2020)
Felis catus x Felis silvestris (Gray 1954; Tiesmeyer et al. 2020)
Leopardus tigrina (tigrina) x Leopardus colocolo (pampas cat) (Trigo et al. 2013)
Leopardus tigrina x Leopardus geoffroyi (Trigo et al. 2013)
Leopardus tigrina x Leopardus pardalis (Ruiz-García et al. 2018)
Leopardus tigrina x Leopardus wiedii (Ruiz-García et al. 2018)
Lynx canadensis (Canada lynx) x Lynx rufus (Schwartz et al. 2004)
* Most prior claims of a domestic cat-bobcat hybrid have not stood up to genetic scrutiny. Currently, there is one hybrid cat breeder online (F1Bobcat.com) who states he has F1 hybrids that have been evaluated by the UC-Davis Veterinary Genetics Laboratory and confirmed. No specific paperwork is shown on his site, but the felines in question are shown in video and do appear to evince traits from both sides of the alleged parentage.
Some domestic cats have been bred to resemble wild cats, such as the ‘ocicat,’ the ‘pixie-bob,’ or the ‘jaguarundi curl,’ but have no non-domestic DNA. Today, hybrids are a conservation concern, particularly with rarer species that hybridize with feral domestic cats.
For creationists, hybridization is strong evidence for the baraminic unity of living felines. There are several genera which are not directly connected through hybridization (Acinonyx, Catopuma, Herpailurus, Neofelis, Otocolobus, Pardofelis), but the overall genetic and morphological evidence argues for their inclusion. Within early feline speciation after the Flood, hybridization appears to have aided maintenance of genetic diversity and adaptability (Li et al. 2016; Figueiró et al. 2017).
The Earliest Fossil Cats
The fossil origin of felines remains tentatively speculative, as early cats are often sparse in the fossil record and comparable traits for certain genera may not yet be known. Further revisions of known fossils and systematic positions are possible.
The oldest ‘true cat’ fossil is considered to be Proailurus (Werdelin et al. 2010), which appears from Upper Oligocene into Miocene strata in Europe (and a doubtful specimen from Mongolia). It was one of several early aeluroid carnivores, others of which may have been ancestral to different feliform groups. Proailurus was likely the founding stock for Pseudaelurus from the Miocene of Europe and Asia, and possibly several other very similar Miocene genera (many of which were once considered Pseudaelurus). These genera are considered ‘Pseudaelurus-grade’ felines, as a distinct monophyletic clade is unlikely (Werdelin et al. 2010).
During the Miocene radiation, European genera included Pseudaelurus, Styriofelis, Leptofelis, Magerifelis, Pristifelis, and Sivaelurus (Jiangzuo et al. 2021; Salesa et al. 2023). Leptofelis was similar to the lynx-sized Styriofelis, but differed from most early felids by exhibiting adaptations for a more terrestrial lifestyle (Salesa et al. 2017). Magerifelis was about the size of a small caracal or bobcat, but likely more robust (Salesa et al. 2023), and may have preyed on larger animals than its contemporaries. Most other differentiation among the European genera here comes down to craniodental traits (Salesa et al. 2012). In Asia, genera included Pseudaelurus (with some species requiring further review), Sivaelurus, and Leptofelis (Jiangzuo et al. 2021). The leopard-sized Hyperailurictis was the first ‘true cat’ in North America. An unnamed early Miocene felid from Ginn Quarry, Nebraska, exhibits certain ‘intermediate’ traits between Proailurus and Hyperailurictis, and may be a primitive form of the latter or may end up a new genus (Lyras et al. 2019). Africa had its own distinct group of felines: the small Asilifelis, the cougar-sized Diamantofelis, the wildcat-sized Namafelis, and Katifelis (Jiangzuo et al. 2021).
This Miocene diversification contributed significantly to the separate trajectories of felid subfamilies. The Pseudaelurus lineage was closely related to the Machairodontinae (sabertooth cats), Styriofelis to the Felinae (small cats), and Miopanthera to the Pantherinae (big cats). North American Hyperailurictis gave rise to Nimravides (Werdelin et al. 2010), which convergently paralleled other sabertooth cats with elongated canines, but did not survive past the Miocene.
Within the creation model, we can see that this early post-Flood period likely saw the development of the first true cat, which rapidly diversified and spread into various regions of multiple continents. Different lineages developed distinctive traits as genetic groups became more isolated, yet hybridization and gene flow may have initially allowed traits to be exchanged between lineages. Some lineages survived, while others were dead ends. Today, we still see eight different clades from two subfamilies among the living felines: 1) pantherine cats, 2) Asian golden cats 3) caracal-serval, 4) ocelot, 5) lynx, 6) puma-cheetah, 7) Asian leopard cat, and 8) Felis (Johnson et al. 2006; O’Brien and Johnson 2007).
The rise of the Felidae (likely from earlier aeluroid carnivores)
in context with the Miocene basal cats
The emergence of modern feline genera.
(Some appearances may be late due to poor fossilization.)
The Pantherinae (Neofelis,
Panthera)
The pantherine lineage is believed to have diverged early within the Miocene radiation (Johnson et al. 2006). Miopanthera is from middle Miocene deposits (though according to Hemmer [2023a], Sivaelurus has priority as the genus). It is related to the pantherine lineage, but cannot be confirmed yet as ancestral (Geraads and Peigne 2017). A late Miocene-early Pliocene fossil pantherine from the Himalayas was first considered a very early Panthera (Tseng et al. 2014), but cranio-dental differences led to the formation of a new genus, Palaeopanthera (Hemmer 2023a). This genus appears to have led to the rise of two subsequent still-extant lineages: true Panthera and Neofelis. De Bonis et al. (2023) described the genus Pachypanthera from late Miocene sand pits in Thailand, noting it was a large predator well-differentiated from other Felidae, that might offer support for an Asian origin of Panthera.
Neofelis (the clouded leopards) is a sister lineage to Panthera, believed to have split in the late Miocene to early Pliocene (Bursell et al. 2022), but shares functionally similar saber teeth with those of ‘less derived’ machairodonts or sabertooth cats (Christiansen 2006). The genetic separation between Neofelis and sabertooths suggests that the elongated canines are convergent adaptations due to some selective pressure, likely predatory behavior (Harano and Kutsukake 2018). Neofelis has an Asian mainland species, Neofelis nebulosa, and a Sundaland island species, Neofelis diardi. The latter appears to be more derived in saber tooth development (Christiansen 2008).
The earliest true Panthera is Panthera principialis from the late Pliocene of Africa, indicating an earlier dispersal from Asia. Panthera principialis was a lion-sized cat, but not a true lion, though it may have been ancestral to lions and leopards (Hemmer 2023a). Panthera appears to have split with one group in Asia developing into Panthera palaeosinensis (along with snow leopards and tigers), and one in Africa splitting into lions and leopards (Hemmer 2023a). Panthera shawi from early Pleistocene South Africa may have dispersed separately from the Old World into the New World, begetting the ancestral lineage of the modern jaguar, Panthera onca.
Panthera palaeosinensis (which likely includes Panthera zdanskyi [Jiangzuo et al. 2023a]), appears to be a basal pantherine, closely related to the snow leopard, possibly related to the ancestor of the tiger (Hemmer 2023a). The snow leopard and tiger are sister lineages (Davis et al. 2010). The snow leopard lineage begins with Panthera pyrenaica (sometimes Panthera uncia pyrenaica) of Europe, with other snow leopards, Panthera uncia, showing up in late Pleistocene Europe and Asia (Hemmer 2023c; Jiangzuo et al. 2025). A trajectory towards the modern snow leopard morphology can be seen over time, as the lineage adapted to a colder, high-altitude, mountainous environment. Despite low genetic diversity, the snow leopard lineage has managed to purge strong deleterious mutations (Yang et al. 2025).
Panthera zdanskyi from early Pleistocene China has been argued to be an early tiger (Mazák et al. 2011), but clarification on its relationship to Panthera palaeosinensis is required. Early tigers, Panthera tigris, had an extensive range in Asia, from the eastern shores of the Black Sea to the Tibetan Plateau to the Russian Far East, from India to southeast Asia all the way to the Philippines. The current range is far more contracted and discontinuous, and the modern tiger genome appears to be primarily one that survived from the late Pleistocene, with other genetic lineages disappearing (Hu et al. 2022). China served as a late Pleistocene refugium and ‘melting pot’ for several lineages, with later range expansion leading to modern subspecies (which demonstrate little genetic admixture) (Sun et al. 2023).
Lions and leopards are a sister lineage, with jaguars closely related (Davis et al. 2010). The earliest fossil leopard, Panthera pardus, is from the Pliocene Laetoli site in Tanzania (Stein and Hayssen 2013), and additional early fossils support an African origin (Pajimans et al. 2018). Leopards spread into Asia and Europe during the middle to late Pleistocene, even reaching the United Kingdom (Diedrich 2013; Marciszak et al. 2022). (The ‘leopard’ fossil situation in middle and possibly late Pleistocene Europe is a bit convoluted, as climatic changes and species turnover may have brought three similar species [leopard, snow leopard, European puma] into the same regions over and over again, though this is still under investigation [Hemmer 2023c; Madurell-Malapeira 2025].) The modern leopard is highly adaptable, and is found in a wide range of habitats and climates in Africa, Arabia, India, central Asia, southeast Asia, and the Russian Far East (Uphyrkina et al. 2001).
The lion, Panthera leo, originated in Africa, with an early fossil appearance at the late Pliocene Laetoli site in Tanzania (Haas et al. 2005). An early dispersion of lions into Eurasia led to the development of first the early Pleistocene steppe lion, Panthera fossilis, then the late Pleistocene cave lion, Panthera spelaea (Sabol et al. 2022; Madurell-Malapeira 2025). Panthera fossilis was very large and occupied a temperate climate, preying on large megafauna, while the smaller Panthera spelaea became cold-adapted. Panthera spelaea crossed the Bering Strait and occupied the Alaskan Beringia, with the North American lion, Panthera atrox, possibly diverging from that population in the late Pleistocene. Panthera atrox became genetically isolated from its cave lion founding stock (Barnett et al. 2009). (An alternative theory has been proposed by Sotnikova and Foronova [2014] based on a steppe lion from western Siberia and certain shared ‘primitive’ traits, that Panthera atrox developed directly from Panthera fossilis. Or, as proposed by Salis et al. [2022], there were multiple waves of dispersal of Panthera spelaea across the Bering Strait, with Panthera atrox diverging from an earlier wave, thus genetically distinct from later Beringean lions.) Panthera atrox fossils have been found throughout western North America, with a few sites known east of the Mississippi, as well as into Mexico and possibly South America (Bravo-Cuevas et al. 2016; Chimento and Agnolin 2017). Both cave lions and North American lions disappeared near the end of the Pleistocene (Barnett et al. 2016). A very large Pleistocene lion (the ‘Natodomeri lion’) was found in Kenya (Manthi et al. 2018), with some affinity to Panthera fossilis, suggesting that that species may have briefly immigrated back into Africa (Sherani and Sherani 2025). While lions moved into India and other parts of western and central Asia during the Pleistocene, the arrival of modern Asian lions is still under debate, with some arguing for a more recent Holocene immigration, along with the cheetah, after the Pleistocene megafauna disappeared (Schnitzler and Hermann 2019).
Panthera gombaszoegensis of Pleistocene Eurasia, has traditionally been placed within the jaguar lineage (Hemmer et al. 2010), though some argue it is closer to the tigers (Chatar et al. 2022). The recent discovery of the species in northeastern China from the middle Pleistocene, however, fills a biogeographical gap that supports dispersal across the Bering Strait (Jiangzuo et al. 2023a). Panthera onca, the modern jaguar, has certain derived dental traits from adaptation to hard-shelled prey, which do not show up in Old World fossils (Jiangzuo and Liu 2020). Jaguar fossils show up in North America during the early Pleistocene, and in South America during the middle to late Pleistocene (Hemmer et al. 2010). Certain fossils indicate the presence of a much larger jaguar form that likely disappeared alongside many other large predators during the Quaternary megafaunal extinction (Ruiz-Ramoni et al. 2020).
panthera leo
panthera tigris
panthera pardus
panthera onca
panthera uncia
neofelis nebulosa
Felinae: The Asian Golden Cat Lineage (Catopuma,
Pardofelis)
This lineage differentiated genetically very early (Johnson et al. 2006), but there is little fossil material. The Asian golden cat (Catopuma temminckii) is found in the Malay Peninsula down to Sumatra. Fossils are known from late Pleistocene or early Holocene Sumatra (Patel et al. 2016) and late Pleistocene Vietnam (Louys 2014). The bay cat (Catopuma badia) is the Asian golden cat’s sister species, and found only on the island of Borneo. The bay cat is both endangered and rarely seen (Kitchener et al. 2004). The marbled cat (Pardofelis marmorata) is a forest cat found in southeast Asia, south to Borneo, and west into the Himalayas. Pardofelis, as a basal member of the Felinae, has been linked in the past to the pantherine felines (Werdelin et al. 2010). It does share a similar coat pattern, proportionally long tail, and flexible ankle joints with the clouded leopard, Neofelis, but how much of this might be convergent due to a similar arboreal lifestyle, is unknown (Yuan et al. 2023).
Catopuma temminckii
catopuma badia
pardofelis marmorata
Felinae: The Asian Leopard Cat
Lineage (Otocolobus, Prionailurus)
Prionailurus is an Asian genus of small felines exemplified by the widespread leopard cat, Prionailurus bengalensis. That species can be found from India and Pakistan, through China and southeast Asia, to the Russian Far East and Japan. There is some evidence for an attempt to domesticate this species within China during the Middle-Late Yangshao period (Vigne et al. 2016). The Sunda leopard cat, Prionailurus javanensis, is an island species found in Sumatra, Borneo, and certain other islands of Indonesia and the Philippines. The flat-headed cat, Prionailurus planiceps, is found in Borneo, Sumatra, and the mainland of Thailand and Malaysia. The fishing cat, Prionailurus viverrinus, is found discontinuously through a broad range of southern Asia, such as India, Sri Lanka, Pakistan, Myanmar, and Thailand. The rusty-spotted cat, Prionailurus rubiginosus, is found in India, Sri Lanka, and Nepal. Despite a close genetic relationship, these species are often disparate in behavior and habitat, allowing them to occupy distinct niches (Silva et al. 2020).
The fossil record for Prionailurus bengalensis includes early Pleistocene Java, middle Pleistocene Laos and China, and late Pleistocene Java, Borneo, and China, while Prionailurus planiceps is known from Pleistocene Borneo (Louys 2014). A particularly small species, Prionailurus kurteni, has been described from the middle Pleistocene of southern China (Jiangzuo et al. 2024b).
Nuclear DNA suggests that the Pallas’s cat, Otocolobus manul, nests alongside Prionailurus (Johnson et al. 2006), while mDNA suggests a sister relationship with Felis, with that pair grouping alongside Prionailurus (Li et al. 2016). This species is widespread throughout Central Asia.
prionailurus bengalensis
prionailurus bengalensis
prionailurus planiceps
prionailurus viverrinus
prionailurus rubiginosus
otocolobus manul
Felinae: The Caracal Lineage (Caracal,
Leptailurus)
Both Caracal and Leptailurus are known back to Pliocene deposits in Africa (Turner 1987; PBDB), though those early fossils may not represent modern species. Today, the caracal (Caracal caracal) can be found throughout different regions of Africa and the Middle East, into India and Central Asia, preferring drier regions with vegetative cover. The African golden cat (Caracal aurata) can be found in the rainforests of Central and West Africa. The serval (Leptailurus serval) is found in West, Central, and Southern Africa, preferring thickets, wetlands, and grasslands. A Pliocene fossil cat in Spain, Caracal depereti (Morales et al. 2003), requires further review.
caracal caracal
caracal aurata
leptailurus serval
Felinae: The Lynx Lineage (Lynx)
The fossil Lynx issiodorensis is considered the likely ancestral species to all living Lynx species (Werdelin 1981). Lynx issiodorensis is known from Pliocene deposits in Africa, Europe, and Asia, but by the Pleistocene had disappeared from Africa (PBDB; Werdelin 1981). There has been debate over whether the species issiodorensis might be better placed under the genus Caracal, but recent analysis demonstrated its affinity with Lynx (Cuccu et al. 2023). Felis rexroadensis from Mio-Pliocene North America has been suggested as an early member of the lynx lineage, but is disputed by other authors (MacFadden and Galiano 1981; Werdelin 1985).
The North American bobcat (Lynx rufus) is a distinctive smaller member of the group. Molecular dating suggests it diverged early within the lineage, while morphological comparison suggests a close relationship to the similarly small Lynx hei from early Pleistocene China (Jianzuo et al. 2022). In North America, bobcats first show up in the Blancan stage (early-mid Pliocene to early Pleistocene) (PBDB; Werdelin 1981). This suggests the two lynx lineages likely diverged in Pliocene Eurasia (Jianzuo et al. 2022), driven first by size, followed by other morphological (particularly craniodental) trait change. Lynx issiodorensis is thus not necessarily ancestral to the bobcat lineage, but may rather reflect the sister lynx lineage.
The divergence of the modern lynx species (Iberian lynx, Lynx pardinus; Eurasian lynx, Lynx lynx; Canadian lynx, Lynx canadensis) from Lynx issiodorensis was geographic in nature. Lynx canadensis likely arose from a Pleistocene migration to North America (distinct from the bobcat lineage migration), Lynx lynx arose in early Pleistocene Asia via a local subspecies (moving into Europe in the late Pleistocene), and Lynx pardinus developed from several European subspecies (Mecozzi et al. 2021). Lynx pardinus had a wider distribution initially, in southern Europe, than its current range, now restricted to the southern Iberian Peninsula (possibly the remnant of a glacial refugium [Johnson et al. 2004]). Several of these species demonstrated plasticity in body size over time during the Plio-Pleistocene, which may or may not be related to climate fluctuations (Mecozzi et al. 2021). The Pleistocene Lynx thomasi, in Morocco, likely diverged as an immigrant from Europe, from Lynx pardinus (Geraads 2008).
lynx rufus
lynx canadensis
lynx lynx
lynx pardinus
Felinae: The Cheetah/Puma Lineage
(Acinonyx, Herpailurus, Puma)
This lineage likely first split between cheetah-like and puma-like cats very early after the Miocene radiation, as Acinonyx and Puma are both known from Pliocene and Pleistocene deposits (PBDB).
The cheetahs, Acinonyx, first appeared, though not indisputably, in eastern and southern Africa from the middle to late Pliocene (Van Valkenburgh et al. 2018). These fragmentary bones are not referable to a specific species, but are a bit larger than the modern cheetah. This ancestral cheetah may have given rise to the modern cheetah, Acinonyx jubatus, which shows up in Pleistocene Africa, as well as, separately, the larger ‘giant cheetah’, Acinonyx pardinensis, which was widespread from China and India to much of southern Europe through most of the Pleistocene. A cheetah from Morocco, Acinonyx aicha, was originally thought to be Pliocene but revised to early Pleistocene (Geraads 2008; Cherin et al. 2014). There are a few possible late Pliocene Acinonyx from Mongolia, Russia, and Europe (Cherin et al. 2014), so an African origin for the genus is not uncontested. Pliocene and early Pleistocene skulls often show generalized pantherine-like traits with skulls that are less ‘cheetah-like’ or domed (Cherin et al. 2014; Cherin et al. 2018), but still exhibit a distinctive cheetah tooth morphology (Geraads 2014). Skull variability has led some authors to use Sivapanthera for certain Chinese fossils, but many recent authors synonymize them with Acinonyx (Geraads 2014). Jiangzuo et al. (2024a) has proposed that during the Pleistocene, Acinonyx pardinensis transitioned into the smaller Acinonyx pleistocaenicus, which may be more closely related to the modern cheetah. Historically, Acinonyx jubatus also occupied parts of the Middle East over to India, but has been extirpated there. The modern cheetah genomes exhibit significantly reduced genetic variation, suggesting Pleistocene bottlenecks responsible for the decrease (Dobrynin et al. 2015).
Puma as a genus originated in the Old World, with the Eurasian Puma pardoides, similar in size to the modern cougar. Fossils are known from late Pliocene sites in Georgia, Mongolia, Italy, and Morocco, to the late early Pleistocene in Spain, and are found from Europe to Russia (Hemmer et al. 2004; Cherin et al. 2013; Hemmer 2023b). Hemmer (2023b) has proposed that a fossil feline at the Pliocene Laetoli site in Tanzania is a Puma species, and speculated that introgressive hybridization between it and Panthera species led to a later Pleistocene form, Puma incurva. How early African Puma might affect the origin of Puma in the Old World is still under debate.
Puma dispersed from the Old World to the New World across the Bering Strait, likely sometime in the Pliocene. It is uncertain whether the late Pliocene-early Pleistocene ‘Felis’ lacustris could be part of the Puma lineage (Kurtén and Anderson 1980; Werdelin 1985). In North America, a branch split off into the cheetah-like Miracinonyx (Barnett et al. 2005). When the Puma lineage crossed into Central and South America (likely part of the Great American Biotic Interchange), it appears to have rapidly split into Puma concolor and Puma pumoides. Puma pumoides is the lineage that developed into the jaguarundi, Herpailurus yagouaroundi. It is possible, based on molecular evidence, that the lineage split just prior to entering South America, with the pumoides branch dispersing more quickly south than the concolor branch (Ruiz-García et al. 2022). Puma pumoides is known from upper Pliocene Argentina (Chimento et al. 2014). A Puma concolor skull is known from early-middle Pleistocene Argentina (Chimento and Dondas 2018). A fossil cat calcaneum (heel) from late Pliocene–early Pleistocene Argentina shows intermediate (cursorial) features between a modern puma and the cheetah-like cats (Ercoli et al. 2019). Puma concolor, the modern cougar, immigrated north back to North America. Puma concolor fossils begin to appear in North America only during the late Pleistocene (Culver et al. 2000). Jaguarundi also moved north, as far as the southern U.S., but with only modern records, as alleged Pleistocene fossils have been dismissed (Ray 1964; Werdelin 1985).
The Plio-Pleistocene ‘cheetah-like’ cats, Miracinonyx, were somewhat convergent to the African cheetah, and it is no surprise that they are part of the same radiating lineage (Van Valkenburgh et al. 1990). Still, the more cursorial species, Miracinonyx trumani, did not have the extreme adaptations found in the modern cheetah and likely used a predatory method not comparable in modern cats (Figueirido et al. 2023)—suggesting that the theory that Miracinonyx was a responsible factor in the co-adaptational development of high-speed pronghorns, is implausible.
acinonyx jubatus
puma concolor
herpailurus yagouaroundi
herpailurus yagouaroundi
Felinae: The Ocelot Lineage (Leopardus)
The small, mostly spotted (though not always) wild cats of South America are now placed in the genus Leopardus, after phylogenetic reviews indicated that several previously described genera should be synonymized. Traditionally, these included the ocelot, margay, oncilla, pampas cat, kodkod, Geoffroy’s cat, and Andean mountain cat, but several recent phylogenetic revisions have split, elevated, and merged species, so there are approximately 14 species currently recognized, with continued gene flow between several species (Ruiz-García et al. 2018; Santos et al. 2018; Nascimento et al. 2021; Trindade et al. 2021; Lescroart et al. 2023; Ruiz-García et al. 2023).
Leopardus vorohuensis has been described from the early Pleistocene strata of Argentina (Berta 1983; Prevosti 2006; Prevosti et al. 2021), and is likely closely related to the colocolo or pampas cat, Leopardus colocola. Fossils of L. colocola are known from late Pleistocene Argentina and late Pleistocene to Holocene Tierra del Fuego, Chile (Prevosti 2006). Fossils of the ocelot, Leopardus pardalis, are known from the late Pleistocene of the southern U.S., Mexico, Brazil, and Argentina (Prevosti et al. 2021). The margay, Leopardus wiedii, is known from late Pleistocene-Holocene fossils in the southern U.S. (Prevosti et al. 2021).
This lineage’s forebears (or forecats) migrated across the Bering Strait from Asia into North America, then later into South America (Johnson et al. 2006). The identity of the felines in this ‘ghost lineage’ in North America are unknown, though several ‘Felis’ fossils have been suggested, several of which appear as early as the late Miocene. The ocelot lineage was distinct prior to the formation of the isthmus of Panama (part of the Miocene Asian radiation), but molecular dating of the diversification into modern species is roughly compatible with occurring after the lineage entered South America (Werdelin et al. 2010). The Great American Biotic Interchange occurred with the formation of this land bridge in the late Pliocene period. Some research suggests that the ocelot and closely-related species developed first, with other species diverging later (Ruiz-García et al. 2022; Lescroart et al. 2023). If all Leopardus speciation occurred within South America, some species, like the ocelot, expanded their range north back over the land bridge, and re-entered the southern U.S. during the late Pleistocene (Prevosti et al. 2021).
leopardus pardalis
leopardus geoffroyi
Felinae: The Domestic Cat Lineage
(Felis)
Felis was initially a catch-all taxon for many fossil cats, but most secular arrangements note that the genetic lineage for true Felis appears to only go back to the Pliocene and the fossil record tends to agree (O’Brien and Johnson 2007). There is the small Felis christoli from late Miocene Italy, however (Morales et al. 2003). A ‘Felis sp.’ is known from the Pliocene Kanapoi, Kenya, site (Werdelin et al. 2010). A fossil feline closely related to the European wild cat, Felis silvestris, is known from Pliocene Spain (Morales et al. 2003).
In Europe, Felis lunensis from the Plio-Pleistocene boundary likely developed into the Pleistocene-modern Felis silvestris (Kurtén 1965). The latter appears to have rapidly expanded its range in the late Pleistocene into Africa and the Middle East, leading to the origin of the African wild cat, Felis lybica. Felis catus was domesticated from the African wild cat in Egypt and the Near East (Yamaguchi et al. 2004). The earliest zooarchaeological evidence comes from pre-pottery Neolithic Cyprus, and human-aided transport during and after the Neolithic helped spread cats throughout the Old World, including Europe and Asia (Hu et al. 2014; Ottoni et al. 2017; Haruda et al. 2020; Ottoni and Van Neer 2020).
felis lybica
felis lybica
felis silvestris
felis chaus
The Machairodontinae (sabertooth cats)
This extinct subfamily first showed up in middle Miocene deposits, expanded throughout Africa, Europe, Asia, North America, and South America, then disappeared at the end of the Pleistocene (with Smilodon surviving into the early Holocene in South America). Most exhibited distinctly enlarged canine teeth. Throughout much of this period, sabertooth cats were more numerous and more diverse than their ‘conical-toothed’ felinae and pantherinae cousins (Christiansen 2013). Depending upon the study, 3 to 4 tribes make up the machairodonts. These lineages diverged early, within the Miocene (Christiansen 2013; Jiangzuo, Werdelin, and Sun 2022). Machairodus, for example, was a large Miocene-Pleistocene sabertooth, with a particularly large species, Machairodus lahayishpup, found in Miocene North America (Orcutt and Calede 2021). Homotherium was another widespread genus, with Pliocene-Pleistocene fossils found in Africa, Europe, Asia, and the Americas (Antón et al. 2014). The presence of multiple genera at certain fossil sites suggests distinct predatory strategies among sabertooths in ecologically mixed environments (Jiangzuo et al. 2023b), and of course, sabertooths were in competition with a wide range of other similar-sized carnivores such as ursids, amphicyonids, and felines (Domingo et al. 2016). The machairodonts were not even the only saber-toothed carnivores. Other feliform groups, the Nimravinae and Barbourofelinae, also exhibited saber-like teeth, as did the metatherian Thylacosmilidae.
Why a saber tooth? While there has been discussion of sexual selection, both male and female sabertooth cats have the elongated canines, so that does not appear to be the selective pressure that induced and maintained the elongation. The teeth were too fragile for violent stabbing, but other models of prey capture have been suggested that appear feasible: some sabertooth cats had longer, flexible necks that allowed them to position their bite around the prey’s neck and then rotate their heads to penetrate the throat with their saber teeth in a shearing motion (Antón et al. 2019). Restorations suggest that sabertooth cats with very long canines had exposed saber teeth, not covered in a soft tissue pouch (Antón et al. 2024). The morphological specialization of the saber tooth may have proven less adaptable when larger prey started to disappear during the Pleistocene, spelling the end of the sabertooth cats (Piras et al. 2018).
smilodon populator
Display at the Museum of Zoology of the University of São Paulo, Brazil
(CC BY-SA 4.0, Wilfredor)
hoplophoneus
(Hough 1950)
Homotherium latidens
'Mummy of a juvenile sabre-toothed cat Homotherium latidens from the Upper Pleistocene of Siberia', Lopatin, et al. Nature (14 Nov. 2024)
https://doi.org/10.1038/s41598-024-79546-1
(CC BY 4.0)
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