Miracinonyx (colloquially known as the "American cheetah" or the "New World cheetah") is an extinct genus of felids belonging to the subfamily Felinae that was endemic to North America from the Pleistocene epoch (about 2.5 million to 16,000 years ago) and morphologically similar to the modern cheetah (Acinonyx jubatus), although its apparent similar ecological niches have been considered questionable due to anatomical morphologies of the former that would have limited its ability to act as a specialized pursuit predator.[1][2] The genus was originally known from fragments of skeletons, but nearly complete skeletons have been recovered from Natural Trap Cave in northern Wyoming.[3]
Miracinonyx was originally established as a subgenus of the genus Acinonyx, with M. trumani assigned to as the type species.[4]M. studeri was originally included as a second species, but it is more often listed as a junior synonym of M. inexpectatus. M. trumani was exclusive to the Rancholabrean age, while M. inexpectatus ranged from the late Blancan to Irvingtonian ages of North America.[1][5]
Discovery and naming
The first fossils attributed to Miracinonyx were several isolated teeth from Port Kennedy Bone Cave from Pennsylvania, dating back to the Irvingtonian age. The American paleontologist Edward Drinker Cope originally considered these to be related to the spotted hyena, and described the material as Crocuta inexpectata in 1895.[6] However, in his posthumous 1899 publication, he reclassified the specimens as Uncia inexpectata, considering them closely related to the snow leopard.[7]
More fossil material was recovered from deposits of similar age in Maryland and Arkansas, where in 1941 American paleontologist George Gaylord Simpson found them all belonging to the same animal, one he considered to be more closely related to the cougar (Puma concolor), naming the animal as Felis (Puma) inexpectata.[8]
The type species, then known as Felis trumani, was described in 1969 by Phil C. Orr based on a complete skull from the Late Pleistocene found in Crypt Cave, Nevada.[9] In 1979, Daniel B. Adams coined Miracinonyx as a new subgenus of the genus Acinonyx, assigning F. trumani to as the type species: Acinonyx (Miracinonyx) trumani. The subgeneric name, Miracinonyx, is derived from the Latin word mirus ('surprising') and the genus Acinonyx which includes the modern cheetah.[4]
The other species, then known as Felis studeri, was described by Donald E. Savage in 1960 based on a more complete material consisting of extensive postcranial and cranial material of the animal.[10] A 1976 review of fossil pumas by Björn Kurtén considered F. inexpectata and F. studeri to be the same species, with F. inexpectata having priority.[11] In 1979, Adams also included F. studeri as a second species of Miracinonyx, assigned to as Acinonyx (Miracinonyx) studeri; while he recognized Kurtén's proposed synonymy, Adams considered F. inexpectata as a nomen dubium due to the holotype being a fragmentary isolated 4th premolar that prevents definitive classification.[4] In 1990, Blaire Van Valkenburgh and colleagues elevated Miracinonyx to the genus level, and supported the synonymy of "F." studeri and "F." inexpectata, resulting in a new combination, M. inexpectatus.[1]
Taxonomy and evolution
Research into the American cheetah has been contradictory. It was originally believed to be an early cougar representative, before being reclassified in the 1970s as a close relative of the cheetah.[4] This suggested that the ancestors of the cheetah diverged from the Puma lineage in the Americas and migrated back to the Old World, a claim repeated as recently as Johnson et al. (2006),[12] and by Dobrynin et al. (2015)[13] However, other research by Barnett and Faurby, through examining mitochondrial DNA and reanalyzing morphology, has suggested reversing the reclassification: the American cheetah developed cheetah-like characteristics through parallel evolution, but it is most closely related to Puma and not to the modern cheetah of Africa and Asia.[1][14][15] Moreover, Faurby notes that no Acinonyx fossils have been found in North America, and no Miracinonyx fossils elsewhere. However, O'Brien et al. (2016) posit that the supposed homoplasy between the genera is controversial, as it is asserted that is not necessarily any conclusive anatomical or genetic basis for dismissing a homologous relationship between Acinonyx and Miracinonyx.[16] The veracity of the origin of the modern cheetah is also debated; however, Miracinonyx is believed to have evolved from cougar-like ancestors, regardless of whether in the Old World or the New World.[17]
The cougar and M. trumani are believed to have split from a cougar-like ancestor around three million years ago;[14] where M. inexpectatus fits in is unclear, although it is probably a more primitive version of M. trumani.[18]
Below is the phylogeny from Chimento & Dondas (2017) when describing the earliest known fossil record of cougars in South America. As shown here, they found Miracinonyx to be a sister taxon to Puma (though in their paper they considered the former genus to be an extinct subgenus of the latter).[19]
It has been proposed that Miracinonyx is an instance of parallel evolution with the cheetah of the Acinonyx genus. Recent studies, however, suggest that it was not as specialized in chasing as the cheetah, as it retained retractable claws and could supinate its forelimbs to grapple with prey, which may have diminished its ability to run fast compared to the true cheetah.[1][20] Instead, it was more closely related to the cougar; M. trumani might have employed a hunting behavior without modern analogue, which may not have relied on speed as much as a cheetah. M. trumani also had a brain similar to a cougar's, albeit with cheetah-like adaptations in the auditory and visual cortexes, and therefore was not as cognitively prepared for the predatory behavior of true cheetahs. Moreover, it had underdeveloped paranasal sinuses which prevents the brain from overheating in Acinonyx during high-speed chases, though this may reflect the colder temperatures of Pleistocene North America compared to Africa and Iran.[21] Additionally, the injuries that led to the death of a sub-adult Miracinonyx according to a 2022 research article by John-Paul Michael Hodnett et al suggest that Miracinonyx felids, like extant cats such as the puma, regularly fought one another; instances of conspecific aggression among Acinonyx cheetahs are relatively rare.[2][5][22]Miracinonyx species were also larger than a modern cheetah and similar in size to a modern North American cougar. The body mass was typically around 70kg (150lb), with a head-and-body length of 170cm (67in), tail length around 92cm (36in), and shoulder height of 85cm (33in).[23] Large specimens could have weighed more than 95kg (209lb).[24]Miracinonyx most likely preyed on mountain goats (Oreamnus americanus), horses (Equus sp.) and especially pronghorns (Antilocapra americana).
It is often assumed that pronghorns evolved to evade Miracinonyx, but this is still debated. Recent research has suggested that the pronghorn's speed may have been shaped by multiple ecological pressures beyond a single predator, including prehistoric wolves, North American lions, and environmental factors such as open grasslands.[25] In the Neogene period (or more specifically the Miocene epoch), in North America, grassland environments, especially those of C4 carbon fixations, were replacing forested lands in North America as well as C3 plants, causing extensive changes in vegetational structure. As a result, artiodactyls and perissodactyls generally evolved with hypsodont dentition for eating C4 grasses and distally longer limbs for easier movement within open environments. In comparison, true pursuit predators didn't even exist until the Plio-Pleistocene, a more recent development of carnivorans compared to the ungulates that already were built for speed some 20 million years before. This general analysis, however, does not preclude the pronghorn's unusually high speed being a specific adaption to predation pressure from cheetahs, as isotopic analysis does indicate pronghorn were their preferred prey, making up an estimated 40% of their diet.[26][27][28]
M. inexpectatus was more similar to the cougar in build than M. trumani.[5][17] Fossil remains of M. inexpectatus found in Hamilton Cave in West Virginia show that this creature lived with and competed with other large cats like jaguars (Panthera onca) and the saber-toothed cat (Smilodon sp.).[1][33]M. trumani was more cursorial and cheetah-like than M. inexpectatus,[5][17] but unlike cheetahs, both species retained fully retractable claws.[1][2] This genus likely hunted in a way with no modern analogues as its elbow-joint morphology was intermediate between A. jubatus and P. concolor.[2]
Fossils of M. inexpectatus have also been found in Florida,[53][54][55][56]Texas,[57] Colorado,[42]Georgia,[44] South Carolina,[44][58] Pennsylvania, West Virginia,[1] Maryland,[59] and California.[60][61] Some fossil remains from sites in California, including Rancho La Brea and the McKittrick Tar Seeps, were originally misidentified as Puma concolor, but have since been reclassified as Miracinonyx trumani. This revision suggests a broader geographic and ecological range for the species in late Pleistocene California.[62] In 2022, the skeletal remains of a M. inexpectatus were retrieved from a cave in southwestern Virginia.[63]
M. inexpectatus lived between the late Blancan and Irvingtonian ages (Early to Middle Pleistocene), while M. trumani was exclusive to the Rancholabrean age (late Middle to Late Pleistocene),[1] with the youngest specimen assigned to as cf. M. trumani recovered from the Haystack Cave. Based on the shaft fragments of an unidentified mammal from stratum levels 3 and 4, the most reliable radiocarbon dating of this site is estimated at 19,659-16,604 calibrated years BP (14,935 ± 610 uncalibrated years BP),[42][64] which has been suggested to be the latest occurrence of M. trumani.[65]
↑Cope, Edward D. (1895). "The fossil vertebrata from the fissure at Port Kennedy, Pa". Proceedings of the Academy of Natural Sciences of Philadelphia. 47: 446–450. JSTOR4061990.
↑Cope, Edward D. (1899). "Vertebrate remains from Port Kennedy bone deposit". Journal of the Academy of Natural Sciences of Philadelphia. 11 (2).
↑Orr, P.C. (1969). "Felis trumani, a new radiocarbon dated cat skull from Crypt Cave, Nevada". Bulletin of the Santa Barbara Society of Natural History. 2: 1–8.
↑Savage, Donald E. (1960). "A Survey of Various Late Cenozoic Vertebrate Faunas of the Panhandle of Texas: Felidae". University of California Publications in Geological Sciences. 36: 317–343.
↑Kurten, B. (1976). "Fossil Puma (Mammalia: Felidae) in North America". Netherlands Journal of Zoology. 26: 502–534.
↑Haaramo, Mikko (2005-11-15). "Felinae – small cats". Felidae. Mikko's phylogeny archive. Archived from the original on 2007-03-27. Retrieved 2007-02-20.
↑Chimento, N.R.; Dondas, A. (2017). "First record of Puma concolor (Mammalia, Felidae) in the Early-Middle Pleistocene of South America". Journal of Mammalian Evolution. 25 (3): 381–389. doi:10.1007/s10914-017-9385-x. hdl:11336/48212. S2CID16249074.
↑Van Valkenburg, Blaire; Pang, Benison; Cherin, Marco; Rook, Lorenzo (2018). "The Cheetah: The Evolutionary History and Paleoecology". In Marker, Laurie; K. Boast, Lorraine; Schmidt-Kuntzel, Anne (eds.). Cheetahs: Biology and Conservation: Biodiversity of the World: Conservation from Genes to Landscapes. Academic Press. pp.25–32. ISBN978-0-12-804120-8.
↑Caro, T.M. (1994). Cheetahs of the Serengeti Plains: Group Living in an Asocial Species. Chicago: University of Chicago Press. p.500. ISBN978-0-226-09433-5.
↑The microtine rodents of the Cheetah Room fauna, Hamilton Cave, West Virginia, and the spontaneous origin of Synaptomys (Report). 1988. doi:10.3133/b1853.
↑Carranza-Castañeda, Oscar; Miller, Wade E. (May 1996). "Hemphillian and Blancan Felids from central Mexico". Journal of Paleontology. 70 (3): 509–518. doi:10.1017/s0022336000038439. ISSN0022-3360.
123Sanders, A.E. (2002). "Additions to the Pleistocene Mammal Faunas of South Carolina, North Carolina, and Georgia". Transactions of the American Philosophical Society. 92 (5): i. doi:10.2307/4144916. JSTOR4144916.
↑Scott, M.; Gary, S. (1987). Late Pliocene (Late Blancan) vertebrates from the St. Petersburg Times site, Pinellas County, Florida, with a brief review of Florida Blancan faunas. Florida Paleontological Society. OCLC182860777.
↑Simpson, G.G. (1929). Pleistocene mammalian fauna of the Seminole Field, Pinellas County, Florida. American Museum of Natural History. OCLC729432.
↑Morgan, Gary S.; Seymour, Kevin L. (1997-10-01). "Fossil history of the panther (Puma concolor) and the cheetah-like cat (Miracinonyx inexpectatus) in Florida". Bulletin of the Florida Museum of Natural History. 40 (2): 177–219. doi:10.58782/flmnh.emto6774. ISSN0071-6154.
↑Slaughter, B.H. (1966). "The Moore Pit Local Fauna; Pleistocene of Texas". Journal of Paleontology. 40 (1): 78–91. JSTOR1301775.
↑Kohn, M.J.; McKay, M.P.; Knight, J.L. (2005). "Dining in the Pleistocene—Who's on the menu?". Geology. 33 (8): 649–652. doi:10.1130/g21476ar.1. S2CID55786870.
↑Steven D. Emslie; David J. Meltzer (2019). "Late Quaternary vertebrates from the Upper Gunnison Basin, Colorado, and small-mammal community resilience to climate change since the last glacial maximum". Quaternary Research. 92 (2): 388–407. Bibcode:2019QuRes..92..388E. doi:10.1017/qua.2019.26. S2CID198418756.
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