Pseudaelurus

Pseudaelurus is a prehistoric cat that lived in Europe, Asia and North America in the Miocene between approximately twenty and eight million years ago. It is related to today's felines and pantherines as well as the extinct machairodonts (saber-tooths), and is a successor to Proailurus. It originated from Eurasia and was the first cat to reach North America, when it entered the continent at about 18.5 Ma ending a 'cat-gap' of 7 million years.[1][2] The slender proportions of the animal, together with its short, viverrid-like legs, suggest that it may have been an agile climber of trees.[3]

Pseudaelurus Temporal range: Miocene, 20–8 Ma PreꞒ Ꞓ O S D C P T J K Pg N
Pseudaelurus jawbone from the Staatliches Museum für Naturkunde Stuttgart, Germany
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	Carnivora
Suborder:	Feliformia
Family:	Felidae
Genus:	†Pseudaelurus Gervais, 1850
Type species
†Pseudaelurus quadridentatus Blainville, 1843
Species
†P. cuspidatus Wang et al., 1998 †P. guangheesis Cao et al., 1990 †P. intrepidus Leidy, 1858 †P. lorteti Gaillard, 1899 †P. marshi Thorpe, 1922 †P. quadridentatus Blainville, 1843 †P. romieviensis Roman & Viret, 1934 †P. skinneri Rothwell, 2003 †P. stouti Schultz & Martin, 1972 †P. turnauensis Hoernes, 1882 †P. validus Rothwell, 2001
Synonyms
Pseudaelurus Hyperailurictis Styriofelis Miopanthera Schizailurus P. turnauensis Pseudaelurus transitorius

Species and distribution

Europe

The first and most primitive species Pseudaelurus turnauensis (= Pseudaelurus transitorius) from the early Miocene was about the size of a domestic cat and probably evolved directly from the Oligocene Proailurus. The European Pseudaelurus turnauensis gave rise to additional species. Pseudaelurus lorteti was lynx-sized and the even larger Pseudaelurus quadridentatus weighed about 30 kg (66 lb) and was approximately the size of a cougar. The latter showed a trend towards slightly enlarged upper canines, indicating that it may have given rise to the later saber-toothed machairodontines. Unexpectedly, the early Pseudaelurus turnauensis survived until 8 million years ago, at which time it is recorded in Germany, whereas the more derived Pseudaelurus lorteti and Pseudaelurus quadridentatus died out 2 million years earlier.[4] A fourth European species is described as Pseudaelurus romieviensis, smaller than P. quadridentatus but larger than P. lorteti.[2]

Asia

P. turnauensis is also recorded in Saudi Arabia from the early Miocene. P. lorteti is also recognized from the middle Miocene of Asia (Jiangsu, China), where two additional species, Pseudaelurus guangheensis from the middle Miocene of Gansu (China) and Pseudaelurus cuspidatus from the middle Miocene of Xinjiang (China) are reported.[2]

North America

Main article: Hyperailurictis

A total of five Pseudaelurus-grade species are known from North America. The species P. validus and P. skinneri are both from the late Hemingfordian, while P. intrepidus, P. marshi, and P. stouti (originally described as Lynx stouti) are all from the Early and Late Barstovian. Of those, P. validus, P. intrepidus, and P. marshi are all relatively large species, overlapping in size with each other and with the Eurasian species P. quadridentatus. P. stouti was smaller and more slender in form, and P. skinneri was also a smaller species, overlapping in size with P. lorteti, P. cuspidatus, and P. guagnheensis.[5][2]

Taxonomy and phylogeny

Location of Pseudaelurus grade fossils based on Paleobiology Database.[6]

• †Pseudaelurus
• †Styriofelis
• †Hyperailurictis
• now assigned to †Afrosmilus within the family †Barbourofelidae

Traditionally all the Pseudaelurus-grade 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. Several authorities have split Pseudaelurus into separate genera or subgenera, including Hyperailurictis, Styriofelis, Miopanthera and Schizailurus. These different groups of Pseudaelurus-grade felids are often considered to have given rise to later felid lineages.

The genus Styriofelis was originally proposed in 1929 by Kretzoi for the species Pseudaelurus turnaeunsis.[7] Kretzoi also proposed the genus Hyperailurictis for the North American species Pseudaelurus intrepidus,[8] and Miopanthera for Ps. lorteti.[9] In 1964, Beaumont elaborated on Kretzoi's proposal and split Pseudaelurus into three separate genera: Pseudaelurus for the European Ps. quadridentatus, Schizailurus for Ps. lorteti, and Hyperailurictis for Ps. intrepidus.[10]

Taxonomic history

In 1843, the paleontologist de Blainville published a description of a felid cranium and lower jaw fragment from Sansan, France. He assigned these fossils to a new species, Felis quadridentata. The cranium was later reassigned to another species, but in 1850 the lower jaw fragment was assigned to a new genus by Paul Gervais as Pseudaelurus quadridentatus, due to having certain primitive features.[11]

In 1858, Joseph Leidy described the species Felis intrepidus, from North America, and reassigned the species as Pseudaelurus intrepidus in 1869. After that discovery, another eight species of Pseudaelurus would be described in North America, but only five are still considered valid.[2]

In 1872, Henri Filhol described the species Pseudaelurus edwardsi from France, but the species was reassigned to the nimravid genus Eofelis in 2000.[12]

In 1882, a second species from Europe was described as Pseudaelurus turauensis, and a third species, Pseudaelurus lorteti, in 1899. The fourth European species, Pseudaelurus romieviensis, was described in 1934. In addition, the species Pseudaelurus transitorius was described in 1892, but most later authors considered it a synonym of P. turnaeunsis.[2]

In 1914, fossils from Africa were described and assigned to the species Pseudaelurus africanus. However, the species was later reassigned to first the genus Metailurus and finally to Afrosmilus.[13]

In Asia, the first description of Pseudaelurus was in 1910, when a fragmentary fossil was assigned to Pseudaelurus chinjiensis; however, it was reassigned in 1915 to the new genus Sivaelurus.[14] The next appearance of Pseudaelurus-grade felids in Asia wasn't until 1986, when a lower jaw fragment and some dental fragments were assigned to the species Pseudaelurus lorteti. In 1990, the species Pseudaelurus guangheensis was described.[15] In 1998, a second Asian species, Pseudaelurus cuspidatus, was also described.[16] Both of the Asian species are known only from fragmentary fossils.[2]

In 1998, while measuring fragmentary fossils from the Hsanda-Gol locality in Mongolia, Robert Hunt referred a lower jaw fragment to Proailurus sp.; while this was reassigned to the nimravid genus Eofelis in 1999 instead, a 2004 review of felid material from other localities in Mongolia suggested that it could belong to Pseudaelurus cuspidatus instead, on basis of having similar features. However, the Hsanda-Gol specimen is dated back to the Oligocene, while Pseudaelurus cuspidatus is found solely in Miocene-aged localities. The same paper also described a pair of fragments (a lower jaw fragment and a metapodial) and attributed them to Pseudaelurus sp.[14]

In 2010, a review of the Felidae as a whole suggested that Pseudaelurus be split into three genera: Hyperailurictis for the five North American species, Styriofelis for two of the European species (P. lorteti and P. turnaunensis), and Pseudaelurus sensu stricto for P. quadridentatus. The status of P. romieviensis, P. guangheensis, and P. cuspidatus was given as uncertain. In addition, Miopanthera and Schizailurus were recognized as junior synonyms of Styriofelis.[10]

In 2012, a new species Styriofelis vallesiensis was described based on a specimen found in Spain.[17] However, a review of the species in 2017 concluded that Leptofelis was sufficiently different from other Styriofelis species as to require a separate genus. It was thus reassigned to the new genus Leptofelis as Leptofelis vallesiensis.[18]

In 2017, a review of the species Felis pamiri, which was named based on a snout fragment from Turkey and dated to the late Miocene, concluded that Felis pamiri and Pseudaelurus lorteti were likely closely related to each other, and ancestral to the Panthera lineage. Both species were reassigned to the genus Miopanthera as Miopanthera lorteti and Miopanthera pamiri.[19]

Phylogeny

The following cladogram is based on Piras et al (2013)[20] and illustrates the three more derived lineages that evolved from "Pseudaelurus" species.

Felidae

†Proailurus  Proailurinae   †Proailurus bourbonnensis     †Proailurus lemanensis     †Proailurus major      "Pseudaelurus"    Pseudaelurus lineage  †Pseudaelurus    †Pseudaelurus quadridentatus     †Pseudaelurus cuspidatus     †Pseudaelurus guangheesis       †Machairodontinae      †Hyperailurictis  Hyperailurictis lineage   †Hyperailurictis intrepidus     †Hyperailurictis marshi     †Hyperailurictis stouti     †Hyperailurictis validus     †Hyperailurictis skinneri      †Sivaelurus  †Sivaelurus chinjiensis     Styriofelis lineage  †Styriofelis    †Styriofelis turnauensis     †Styriofelis romieviensis      Felinae    Felinae      †Miopanthera    †Miopanthera lorteti     †Miopanthera pamiri       Pantherinae     sensu lato   (grade)

References

1. Tedford, R. H.; Galusha, T.; Skinner, M. F.; Taylor, B. E.; Fields, R. W.; Macdonald, J. R.; Rensberger, J. M.; Webb, S. D.; Whistler, D.P. (1987). "Faunal succession and biochronology of the Arikareean through Hemphillian interval (late Oligocene through earliest Pliocene epochs) in North America". In Woodburne, M. O. (ed.). Cenozoic mammals of North America: Geochronology and biostratigraphy. Berkeley: University of California Press. pp. 153–210. ISBN 0-520-05392-3.
2. Rothwell, Tom (2003). "Phylogenetic systematics of North American Pseudaelurus (Carnivora: Felidae)" (PDF). American Museum Novitates. 3403: 1–64. doi:10.1206/0003-0082(2003)403<0001:psonap>2.0.co;2. hdl:2246/2829. ISSN 0003-0082.
3. Turner, Alan; Antón, Mauricio (1997). The Big Cats and their fossil relatives. New York: Columbia University Press. ISBN 0-231-10228-3.
4. Augusti, Jordi (2002). Mammoths, Sabertooths and Hominids: 65 Million Years of Mammalian Evolution in Europe. New York: Columbia University Press. ISBN 0-231-11640-3.
5. Rothwell, Tom (2001). "A Partial Skeleton of Pseudaelurus (Carnivora: Felidae) from the Nambé Member of the Tesuque Formation, Española Basin, New Mexico" (PDF). American Museum Novitates. 3343. ISSN 0003-0082.
6. "Pseudaelurus". paleobiodb.org. The data were downloaded from the Paleobiology Database on June 27, 2017 using the taxon name 'Pseudaelurus'. Retrieved June 27, 2017.
7. Kretzoi, M. (1929a). "Feliden-Studien". A Magyar Királyi Földtani Intézet Hazinyomdaja. 24: 1–22.
8. Kretzoi, M. (1929b). "Materialien zur phylogenetischen Klassifikation der Aeluroideen". Int Cong Zool Budapest. 10: 1293–1355.
9. Kretzoi, M. (1938). "Die Raubtiere von Gombaszög nebst einer Übersicht der Gesamtfauna". Ann Mus Nat Hungar. 31: 88–157.
10. Werdelin, Lars; O'Brien, Stephen J.; Johnson, Warren E.; Yamaguchi, Nobuyuki (2010). "Phylogeny and evolution of cats (Felidae)". In Macdonald, D.W.; Loveridge, A.J. (eds.). Biology and Conservation of Wild Felids. Oxford: Oxford University Press.
11. Gervais, Paul (1850). "Zoologie et paléontologie françaises. Nouvelles recherches sur les animaux vertébrés dont on trouve les ossements enfouis dans les sol de le France et sur leur comparaison avec les espèces propres aux autres regions du globe". Zoologie et Paléontologie Françaises. 8: 1–271.
12. Peigné, Stéphane (2000). "A new species of Eofelis (Carnivora: Nimravidae) from the Phosphorites of Quercy, France". Comptes Rendus de l'Académie des Sciences, Série IIA. 330 (9): 653–658. doi:10.1016/s1251-8050(00)00199-3.
13. Morales, Jorge; Pickford, M.; Salesa, Manuel J. (2008). "Creodonta and Carnivora from the Early Miocene of the Northern Sperrgebiet, Namibia". Memoir of the Geological Survey of Namibia. 20: 291–310.
14. Rothwell, Tom (2004). "Chapter 12: New Felid Material from the Ulaan Tologoi Locality, Loh Formation (Early Miocene) of Mongolia". Bulletin of the American Museum of Natural History. 285: 157–165. CiteSeerX 10.1.1.616.5669. doi:10.1206/0003-0090(2004)285<0157:c>2.0.co;2.
15. Cao, Z.; Du, H; Zhao, Q.; Cheng, J (1990). "Discovery of the middle Miocene fossil mammals in Guanghe District, Gansu and their stratigraphic significance". Geoscience. 4: 16–32.
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17. Salesa, Manuel J.; Antón, Mauricio; Morales, Jorge; Peigné, Stéphane (2012). "Systematics and phylogeny of the small felines (Carnivora, Felidae) from the Late Miocene of Europe: a new species of Felinae from the Vallesian of Batallones (MN 10, Madrid, Spain)". Journal of Systematic Palaeontology. 10 (1): 87–102. doi:10.1080/14772019.2011.566584. ISSN 1477-2019.
18. Salesa, Manuel J.; Siliceo, Gema; Antón, Mauricio; Peigné, Stéphane; Morales, Jorge (2017). "Functional and Systematic Implications of the Postcranial Anatomy of a Late Miocene Feline (Carnivora, Felidae) from Batallones-1 (Madrid, Spain)". Journal of Mammalian Evolution. 26: 1–31. doi:10.1007/s10914-017-9414-9.
19. Geraads, Denis; Peigné, Stéphane (2017). "Re-appraisal of 'Felis' pamiri Ozansoy 1959 (Carnivora, Felidae) from the upper Miocene of Turkey: the earliest pantherin cat?". Journal of Mammalian Evolution. 24 (4): 415–425. doi:10.1007/s10914-016-9349-6.
20. Piras, P.; Maiorino, L.; Teresi, L.; Meloro, C.; Lucci, F.; Kotsakis, T.; Raia, P. (2013). "Bite of the Cats: Relationships between Functional Integration and Mechanical Performance as Revealed by Mandible Geometry". Systematic Biology. 62 (6): 878–900. doi:10.1093/sysbio/syt053. ISSN 1063-5157. PMID 23925509.

• Christiansen, Per (2008). "Phylogeny of the great cats (Felidae: Pantherinae), and the influence of fossil taxa and missing characters". Cladistics. 24 (6): 977–992. doi:10.1111/j.1096-0031.2008.00226.x.