Biology:Nimravidae

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Nimravidae is an extinct family of carnivorans, sometimes known as false saber-toothed cats, whose fossils are found in North America, Africa, and Eurasia. Not considered to belong to the true cats (family Felidae), the nimravids are generally considered closely related and classified as a distinct family in the suborder Feliformia. Fossils have been dated from the Middle Eocene through the Late Miocene epochs (Bartonian through Messinian stages, 41.03–7 million years ago), spanning about 34.03 million years.[1]

The barbourofelines, classified as a subfamily of the Nimravidae since 1991, were reassigned to their own distinct family in 2004.[2] However, since 2020, the majority of experts consider barbourofelines as nimravids again.[3][1][4][5][6]

Taxonomy

The family Nimravidae was named by American paleontologist Edward Drinker Cope in 1880,[7] with the type genus as Nimravus. 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).[8]

Nimravids are placed in tribes 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). These nimravids are referred to as "false saber-tooths". The barbourofelids were for a while no longer included in Nimravidae, following elevation to family as sister clade to the true cats (family Felidae).[9][10] However, majority of recent studies have returned them to Nimravidae, with one study suggesting they are part of Nimravinae.[1][4][11][5][6]

Not only did nimravids exhibit diverse dentition, but they also showed the same diversity in size and morphology as cats. Nimravids such as Barbourofelis morrisi, Eusmilus sicarius, and Hoplophoneus occidentalis were leopard-sized,[12][13][14] while some, such as Albanosmilus jourdani and E. adelos, were the size jaguars to small lions.[1][15] The largest nimravids, Quercylurus and Barbourofelis fricki, were able to reach even larger sizes, weighing 200 kg (440 lb) and 328 kg (723 lb) respectively.[16][1] Dinaelurus had the short face, rounded skull, and smaller canines of the modern cheetah,[1] and some such as E. bidentatus and Nanosmilus, were only the size of a small bobcat.[17][18]

Family †Nimravidae
Tribe Image Genus Species
†Dinailurictis (Helbing, 1922)
  • †D. bonali
175px †Dinictis (Leidy, 1854)
  • †D. felina
†Eofelis (Kretzoi, 1938)
  • †E. edwardsii
  • †E. giganteus
175px †Maofelis[19] (Averianov et al., 2016)
  • †M. cantonensis
†Pangurban[20] (Poust et al, 2022)
  • †P. egiae
175px †Pogonodon (Cope, 1880)
  • †P. davisi
  • †P. platycopis
†Quercylurus (Ginsburg 1979)
  • †Q. major
†Nimravini †Dinaelurus (Eaton, 1922)
  • †D. crassus
175px †Nimravus (Cope, 1879)
  • †N. brachyops
  • †N. intermedius
†Hoplophoneini 175px †Hoplophoneus[21] (Cope, 1874)

 (Subgenus: †Eusmilus[21] (Eaton, 1922)) 

  •  †H. bidentatus 
  •  †H. cerebralis 
  • †H. dakotensis
  • †H. mentalis
  • †H. occidentalis
  • †H. oharrai
  • †H. primaevus
  •  †H. sicarius 
  • †H. villebramarensis
†Nanosmilus (Martin, 1992)
  • †N. kurteni

Phylogeny

The phylogenetic relationships of Nimravidae are shown in the following cladogram:[19][21][22]

A 2021 study divides Nimravidae into Hoplophoninae and Nimravinae, the latter including the bulk of species in addition to barbourofelins.

Phylogeny of Nimravidae from the 2022 description of Pangurban:[20]

Nimravidae
 

Maofelis cantonensis

 
 
 

MA-PHQ 348

 
 
Nimravinae
 
 

Dinictis felina

 
 
 

Pogonodon davisi

 
 

Pogonodon platycopis

 
 
 
 
 
 

Dinaelurus crassus

 
 
 

Nimravus brachyops

 
 

Nimravus intermedius

 
 
 
 
 

Eofelis edwardsii

 
 
 

Dinailurictis bonali

 
 

Quercylurus major

 
 
 
 
 
Hoplophoneini
 

Pangurban egiae

 
 
 

Hoplophoneus oharrai

 
 
 
 

Hoplophoneus occidentalis

 
 

Hoplophoneus primaevus

 
 
 
 

Nanosmilus kurteni

 
 
 

Eusmilus dakotensis

 
 
 

Eusmilus sicarius

 
 
 

Eusmilus adelos

 
 
 

Eusmilus cerebralis

 
 
 

Eusmilus bidentatus

 
 

Eusmilus villebramarensis

 
 
 
 
 
 
 
 
 
 
 
 
 

Evolution

Restoration of Dinictis and Protoceras by Charles R. Knight

The ancestors of nimravids and cats diverged from a common ancestor soon after the Caniformia–Feliformia split, in the middle Eocene about 50 million years ago (Mya), with a minimum constraint of 43 Mya.

Some of the first nimravids, Maofelis and Pangurban, appeared in the middle of the Eocene epoch, about 40 Mya, in Asia and North America respectively.[23] 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 were transforming to scrub and open woodland. This climatic trend continued in the Oligocene, and nimravids evidently flourished in this environment. North America and Asia were connected and shared much related fauna.[24] Europe in the Oligocene was more of an archipelago than a continent, though some land bridges must have existed, for nimravids also spread there. Barbourofelins probably evolved from Nimravinae dispersing into Africa during the Oligocene. The presence of large hyaenodonts prevented them from reaching a large size but were able to carve a niche due to their dental morphology. Eventually, they dispersed from Africa into Eurasia and later into North America.[1]

Extinction

Both Hoplophoninae and Nimravinae died out during the Oligocene epoch, with the last taxa going extinct 29.5 and 25.9 million years ago respectively. Their extinction probably coincided with the expansion of grasslands and possibly competition with amphicyonids.[25] Their extinction started the infamous cat gap, a 7 million year period when no cat-like predators present in North America.[26]

Barbourofelins went extinct around 7 million years ago, during the Late Miocene, for unknown reasons.[1] Antón Mauricio suggested competition with machairodonts such as Machairodus and Nimravides, may have contributed to their extinction, as barbourofelins were widely successful despite the wider expansion of grasslands.[26] However, Paul Barret has contested this hypothesis because of the limited temporal overlap between both clades.[1] In addition, Albanosmilus, the last genus to go extinct in Eurasia, was also able to coexist and compete with machairodonts Amphimachairodus and Machairodus in some localities for over a million years.[27][28][29] Other experts suggested it was more likely they went extinct because of the faunal overturn during the Late Miocene due to the wider expansion of grasslands.[1][30][28]

Morphology

Most nimravids had muscular, low-slung, cat-like bodies, with shorter legs and tails than are typical of cats. Unlike extant Feliformia, the nimravids had a different bone structure in the small bones of the ear. The middle ear of true cats is housed in an external structure called an auditory bulla, which is separated by a septum into two chambers. Nimravid remains show ossified bullae with no septum, or no trace at all of the entire bulla. They are assumed to have had a cartilaginous housing of the ear mechanism.[31] Nimravid feet were short, indicating they walked in a plantigrade or semiplantigrade posture, i.e., on the flat of the feet rather than the toes, like modern cats.[26]

Although some nimravids physically resembled the saber-toothed cats, such as Smilodon, they were not closely related,[32] but evolved a similar form through parallel evolution. They had synapomorphies with the barbourofelins in the cranium, mandible, dentition, and postcranium.[33] They also had a downward-projecting flange on the front of the mandible as long as the canine teeth, a feature that also convergently evolved in the saber-toothed sparassodont Thylacosmilus. While most nimravids were thought to have been ambush predators,[34][35][36][37] some such as Eusmilus adelos and Pogonodon davisi were recovered as pounce-pursuit predators.[38] Albanosmilus whitfordi and Dinaelurus are thought to have been cursorial predators, although no post cranial remains have been found for Dinaelurus.[39][40][1]

A 2021 study has shown that a sizeable number of species developed feline-like morphologies in addition to saber-toothed taxa.[4]

References

  1. ↑ 1.00 1.01 1.02 1.03 1.04 1.05 1.06 1.07 1.08 1.09 1.10 Cite error: Invalid <ref> tag; no text was provided for refs named Barrett21
  2. ↑ Morlo, Michael; Peigné, Stéphane; Nagel, Doris (January 2004). "A new species of Prosansanosmilus: implications for the systematic relationships of the family Barbourofelidae new rank (Carnivora, Mammalia).". Zoological Journal of the Linnean Society 140 (1): 43. doi:10.1111/j.1096-3642.2004.00087.x. 
  3. ↑ Wang, Xiaoming; White, Stuart C.; Guan, Jian (2 May 2020). "A new genus and species of sabretooth, Oriensmilus liupanensis (Barbourofelinae, Nimravidae, Carnivora), from the middle Miocene of China suggests barbourofelines are nimravids, not felids". Journal of Systematic Palaeontology 18 (9): 783–803. doi:10.1080/14772019.2019.1691066. https://escholarship.org/uc/item/0g62362j. 
  4. ↑ 4.0 4.1 4.2 Barrett, P. Z.; Hopkins, W. S. B.; Price, S. A. (2021). "How many sabertooths? Reevaluating the number of carnivoran sabertooth lineages with total-evidence Bayesian techniques and a novel origin of the Miocene Nimravidae". Journal of Vertebrate Paleontology 41 (1). doi:10.1080/02724634.2021.1923523. 
  5. ↑ 5.0 5.1 Werdelin, Lars (November 2021). "African Barbourofelinae (Mammalia, Nimravidae): a critical review". Historical Biology 34 (2): 1347–1355. doi:10.1080/08912963.2021.1998034. https://www.researchgate.net/publication/356101795. 
  6. ↑ 6.0 6.1 Jasinski, Steven E.Expression error: Unrecognized word "et". (November 2022). "New Carnivoran (Mammalia: Carnivora) specimens from the Siwaliks of Pakistan and India and their faunal and evolutionary implications". Historical Biology 35 (11): 2217–2252. doi:10.1080/08912963.2022.2138376. https://www.researchgate.net/publication/365095742. 
  7. ↑ Cope, E. D. (1889). "Synopsis of the Families of Vertebrata". The American Naturalist 23: 1–29. 
  8. ↑ Flynn, John J.; Galiano, Henry (1982). "Phylogeny of early Tertiary Carnivora, with a description of a new species of Protictis from the middle Eocene of Northwestern Wyoming". American Museum Novitates. 
  9. ↑ Morlo, Michael; Peigné, Stéphane; Nagel, Doris (January 2004). "A new species of Prosansanosmilus: implications for the systematic relationships of the family Barbourofelidae new rank (Carnivora, Mammalia)". Zoological Journal of the Linnean Society 140 (1): 43. doi:10.1111/j.1096-3642.2004.00087.x. 
  10. ↑ Werdelin, L.; Yamaguchi, N.; Johnson, W. E.; O'Brien, S. J. (2010). "Phylogeny and evolution of cats (Felidae)". in Macdonald, D. W.; Loveridge, A. J.. Biology and Conservation of Wild Felids. Oxford, UK: Oxford University Press. pp. 59–82. ISBN 978-0-19-923445-5. https://www.researchgate.net/publication/266755142. 
  11. ↑ Barret, Paul Z.; Hopkins, Samantha (August 14, 2024). "Mosaic evolution underlies feliform morphological disparity". The Royal Society Publishing 291 (2028). doi:10.1098/rspb.2024.0756. https://royalsocietypublishing.org/doi/10.1098/rspb.2024.0756. 
  12. ↑ Turner, Alan (1997). The Big Cats and their Fossil Relatives: an illustrated guide. New York: Columbia University Press. pp. 234. ISBN 978-0-231-10228-5. https://archive.org/details/bigcatstheirfoss00turn. 
  13. ↑ Antón, Mauricio (March 3, 2013). "Meet an early "extreme" sabertooth: Eusmilus sicarius". https://chasingsabretooths.wordpress.com/2013/05/03/meet-an-early-extreme-sabertooth-eusmilus-sicarius/. 
  14. ↑ Antón, Mauricio (2013). Sabertooth. Bloomington, Indiana: University of Indiana Press. p. 104. ISBN 978-0-253-01042-1. 
  15. ↑ Antón, Mauricio (2013). Sabertooth. Bloomington, Indiana: University of Indiana Press. p. 104. ISBN 978-0-253-01042-1. 
  16. ↑ Peigne, Stephane (May 2003). "Systematic review of European Nimravinae (Mammalia, Carnivora, Nimravidae) and the phylogenetic relationships of Palaeogene Nimravidae". Zoologica Scripta 32 (3): 199–229. doi:10.1046/j.1463-6409.2003.00116.x. https://onlinelibrary.wiley.com/doi/abs/10.1046/j.1463-6409.2003.00116.x. 
  17. ↑ Martin, L. D. (1991, January). A new miniature saber-toothed nimravid from the Oligocene of Nebraska. In Annales Zoologici Fennici (pp. 341-348). Finnish Zoological Publishing Board, formed by the Finnish Academy of Sciences, Societas Biologica Fennica Vanamo, Societas pro Fauna et Flora Fennica, and Societas Scientiarum Fennica.
  18. ↑ Antón, Mauricio (2013). Sabertooth. Bloomington, Indiana: University of Indiana Press. pp. 97. ISBN 978-0-253-01042-1. 
  19. ↑ 19.0 19.1 Averianov, Alexander; Obraztsova, Ekaterina; Danilov, Igor; Skutschas, Pavel; Jin, Jianhua (2016-05-10). "First nimravid skull from Asia" (in En). Scientific Reports 6 (1). doi:10.1038/srep25812. PMID 27161785. Bibcode: 2016NatSR...625812A. 
  20. ↑ 20.0 20.1 Cite error: Invalid <ref> tag; no text was provided for refs named Pangurban
  21. ↑ 21.0 21.1 21.2 Barrett, P.Z. (2016). "Taxonomic and systematic revisions to the North American Nimravidae (Mammalia, Carnivora)". PeerJ 4. doi:10.7717/peerj.1658. PMID 26893959. 
  22. ↑ Peigne, Stéphane (May 2003). "Systematic review of European Nimravinae (Mammalia, Carnivora, Nimravidae) and the phylogenetic relationships of Palaeogene Nimravidae" (in en). Zoologica Scripta 32 (3): 199–229. doi:10.1046/j.1463-6409.2003.00116.x. 
  23. ↑ Averianov, Alexander; Obraztsova, Ekaterina; Danilov, Igor; Skutschas, Pavel; Jin, Jianhua (2016). "First nimravid skull from Asia". Scientific Reports 6. doi:10.1038/srep25812. PMID 27161785. Bibcode: 2016NatSR...625812A. 
  24. ↑ Prothero, Donald R. (2006). After the Dinosaurs: The Age of Mammals. Bloomington, Indiana: Indiana University Press. pp. 9, 132–134, 160, 167, 174, 176, 198, 222–233. ISBN 978-0-253-34733-6. 
  25. ↑ Castellanos, Miguel (2025). "Hunting types in North American Eocene–Oligocene carnivores and implications for the ‘cat-gap’". Journal of Mammalian Evolution 32 (2): 1-12. doi:10.1007/s10914-025-09767-2. https://link.springer.com/article/10.1007/s10914-025-09767-2. 
  26. ↑ 26.0 26.1 26.2 Antón, Mauricio (2013). Sabertooth. Bloomington, Indiana: University of Indiana Press. p. 90. ISBN 978-0-253-01042-1. 
  27. ↑ Jiangzuo, Q; Werdelin, L; Sanisidro, O; Yang, Rong; Fu, Jiao; Li, Shijie; Wang, Shiqi; Deng, Tao (April 2023). "Origin of adaptations to openenvironments and social behaviour insabretoothed cats from the northeasternborder of the Tibetan Plateau". Proceedings of the Royal Society B: Biological Sciences 290 (1997): 7–8. doi:10.1098/rspb.2023.0019. PMID 37072045. PMC 10113030. https://www.researchgate.net/publication/370122411. 
  28. ↑ 28.0 28.1 Michael Morlo (2006). "New remains of Barbourofelidae from the Miocene of Southern Germany: implications for the history of barbourid migrations". Beiträge zur Paläontologie, Wien 30: 339–346. https://www.researchgate.net/publication/233791187. 
  29. ↑ Domingo, Laura; Domingo, M. Soledad; Koch, Paul L.; Alberdi, M. Teresa (May 10, 2017). "Carnivoran resource and habitat use in the context of a Late Miocene faunal turnover episode". Palaeontology 60 (4): 461–483. doi:10.1111/pala.12296. https://onlinelibrary.wiley.com/doi/10.1111/pala.12296. 
  30. ↑ Jiangzuo, Qigao; Li, Shijie; Deng, Tao (2022). "Parallelism and lineage replacement of the late Miocene scimitar-toothed cats from the old and New World". iScience 25 (12). doi:10.1016/j.isci.2022.105637. PMID 36505925. PMC 9730133. Bibcode: 2022iSci...25j5637J. https://www.cell.com/iscience/pdf/S2589-0042(22)01909-5.pdf. 
  31. ↑ Turner, Alan (1997). The Big Cats and their Fossil Relatives: an illustrated guide. New York: Columbia University Press. p. 234. ISBN 0-231-10228-3. 
  32. ↑ "Meet the Cat Family". The Sunday Observer (Colombo, Sri Lanka: Associated Newspapers of Ceylon). July 16, 2012. http://archives.sundayobserver.lk/2012/07/15/jun04.asp. 
  33. ↑ Bryant, Harold N. (February 1991). "Phylogenetic Relationships and Systematics of the Nimravidae (Carnivora)". Journal of Mammalogy (Lawrence, Kansas: American Society of Mammalogists) 72 (1): 56–78. doi:10.2307/1381980. 
  34. ↑ Naples, V. L.; Martin, L. D. (June 2008). "Restoration of the superficial facial musculature in nimravids". Zoological Journal of the Linnean Society 130 (1): 55–81. doi:10.1111/j.1096-3642.2000.tb02195.x. 
  35. ↑ Ormsby, Christianne (May 2021). Morphology and Paleoecology of Nimravides galiani (Felidae) and Barbourofelis loveorum (Barbourofelidae) from the Late Miocene of Florida (MS (Master of Science) thesis).
  36. ↑ Morlo, Michael; Gunnell, Gregg F.; Nagel, Doris (2010). "10 - Ecomorphological analysis of carnivore guilds in the Eocene through Miocene of Laurasia" (in English). Carnivoran Evolution. Cambridge University Press. pp. 269–310. doi:10.1017/CBO9781139193436.011. ISBN 978-1-139-19343-6. https://www.academia.edu/24842570. 
  37. ↑ Castellanos, Miguel (2025). "Hunting types in North American Eocene–Oligocene carnivores and implications for the ‘cat-gap’". Journal of Mammalian Evolution 32 (2): 1-12. doi:10.1007/s10914-025-09767-2. https://link.springer.com/article/10.1007/s10914-025-09767-2. 
  38. ↑ Castellanos, Miguel (2025). "Hunting types in North American Eocene–Oligocene carnivores and implications for the ‘cat-gap’". Journal of Mammalian Evolution 32 (2): 1-12. doi:10.1007/s10914-025-09767-2. https://link.springer.com/article/10.1007/s10914-025-09767-2. 
  39. ↑ Bryant, Harold N. (1996). "Nimravidae". in Donald R. Prothero and Robert J. Emry. The Terrestrial Eocene-Oligocene Transition in North America. Cambridge, England: Cambridge University Press. p. 468. ISBN 0-521-43387-8. 
  40. ↑ Andersson, Ki; Werdelin, Lars (December 2003). "The evolution of cursorial carnivores in the Tertiary: Implications of elbow-joint morphology". Proceeding of the Royal Society B 270 (Suppl 2): S163-5. doi:10.1098/rsbl.2003.0070. PMID 14667370. PMC 1809930. https://www.researchgate.net/publication/8967327. 
  • Peigné, Stephane; De Bonis, Louis (August 2003). "Juvenile cranial anatomy of Nimravidae (Mammalia, Carnivora): biological and phylogenetic implications". Zoological Journal of the Linnean Society 138 (4): 477–493. doi:10.1046/j.1096-3642.2003.00066.x. 
  • Data related to Nimravidae at Wikispecies

Template:Nimravidae Wikidata ☰ Q131889 entry



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