Ferae

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Ferae
Temporal range: 78.9–0 Ma[1]
Late Cretaceous - present
Order Carnivora.jpg
High diversity of carnivorans
Pangolin borneo.jpg
Low species diversity in Pholidotes: a pangolin
Scientific classification e
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Clade: Ferungulata
Clade: Ferae
Linnaeus, 1758[2]
Subgroups

Possible members:[3]

Ferae (/ˈfɪər/ FEER-ee, Latin[ˈfɛrae̯]) is a clade of mammals, consisting of the orders Carnivora (wolves, cats, seals, and so on ; over 260 species around the globe) and Pholidota (eight species of pangolins in tropical Africa and Asia). An alternate name, Ostentoria, has also been proposed for a grouping of the Carnivora and Pholidota.[4] The last common ancestor of extant Ferae is supposed to have diversified ca. 78.9 million years ago.[1] Several extinct orders, relatives of the Pholidota, such as Creodonts,[3] are members of Ferae as well.

Phylogeny of extant taxa[edit]

Position of pangolins[edit]

Pangolins were long thought to be the closest relatives of Xenarthra (armadillos, anteaters, and sloths), contributing to the polyphyletic group of Edentata. Research based on immunodiffusion technique[5] and comparison of protein and DNA sequences[6][7][8] revealed the close relationships between pangolins and carnivorans. Living pangolins and carnivorans also share few unusual derived morphological and anatomical traits, such as the ossified tentorium cerebelli and the fusion of the scaphoid and lunate bones in the wrist.[9]

Sister groups to Ferae[edit]

According to recent studies (reflected in the diagram below), the closest relatives of Ferae are Perissodactyla (horses, tapirs, and rhinos) and Cetartiodactyla (which combines Artiodactyla—camels, pigs, ruminants and hippos—with Cetacea—whales and dolphins).[10][11] An alternate phylogeny (less supported) holds that the closest relatives to the Ferae are the Perissodactyla and Chiroptera (bats), not Cetartiodactyla.[12] Ferae together with Perissodactyla has been called Zooamata. Ferae, Perissodactyla, and Chiroptera together has been called Pegasoferae.

Phylogenetic position of the Ferae in the context of the order-level cladogram of Boreoeutheria.
 Boreoeutheria 

 Euarchontoglires
 (primates, colugos, treeshrews, rodents, rabbits)  Lepus timidus - 1700-1880 - Print - Iconographia Zoologica -(white background).jpg

 Laurasiatheria 

 Eulipotyphla
 (hedgehogs, shrews, moles, solenodons)  Erinaceus europaeus - 1700-1880 - Print - Iconographia Zoologica - Special Collections University of Amsterdam -(white background).jpg

 Scrotifera 

 Chiroptera
 (bats and flying foxes)  Flying fox at botanical gardens in Sydney (cropped and flipped).jpg

 Fereuungulata 
 Ferae 

 Pholidota
 (pangolins)  Pangolin Hardwicke (white background).jpg

 Carnivora
 (cats, hyenas, dogs, bears, seals, etc.)  Dogs, jackals, wolves, and foxes (Plate XI).jpg

 Euungulata 

 Perissodactyla
  (horses, tapirs, rhinos, etc.) Equus quagga (white background).jpg

 Cetartiodactyla
 (camels, pigs, ruminants, hippos, whales, etc.)  The deer of all lands (1898) Hangul white background.png

The cladogram has been reconstructed from mitochondrial and nuclear DNA and protein characters.

Fossil members[edit]

Position of Creodonta[edit]

Various enigmatic Palaeocene mammals have been found to belong to Ferae. While there has been strong support in the inclusion of creodonts into Ferae, they were usually recovered as sister taxon to Carnivoramorpha (carnivorans and their stem-relatives).[13] The phylogenetic analysis of hundreds of morphological characters of Paleocene placentals found instead that creodonts might be the sister group to Pholidotamorpha (pangolins and their stem-relatives).[3]

Phylogenetic hypothesis for the position of Creodonta within Ferae.
 Ferungulata 

Euungulata Equus quagga (white background).jpg

 Ferae 
 Pholidotamorpha 

Pholidota (sensu stricto) Pangolin Hardwicke (white background).jpg

Palaeanodonta Metacheiromys DB152-2.jpg

 (Pholidota sensu lato) 
 †Oxyaenodonta 

Oxyaenidae Patriofelis ferox by R. B. Horsfall (coloured).png

 †Hyaenodonta 

Altacreodus

Tinerhodon

Hyaenodonta (sensu stricto) Hyaenodon horridus by R. B. Horsfall (coloured).jpg

 sensu lato 
 Carnivoramorpha 
 †Viverravoidea 

Viverravidae

 ? 

Ravenictis

Carnivoraformes Dogs, jackals, wolves, and foxes (Plate XI).jpg

 (Carnivora sensu lato) 
 †Creodonta 
[14][15]

Possible members[edit]

In addition various supposedly "hoofed mammals" like the mesonychians and arctocyonids (usually considered as stem-artiodactyls[16]) also belong to the group. In addition, Mesonychians might be the sister group to carnivoramorphs, while arctocyonids are polyphyletic with Arctocyon and Loxolophus sister to pantodonts and periptychids, Goniacodon and Eoconodon sister to the Carnivoramorpha-Mesonychia clade, most other genera allied with creodonts and palaeoryctidans, and Protungulatum not a placental mammal.[3] This enlarged Ferae was also found to be the sister group to Chiroptera.[3]

Below is a phylogeny of the interrelationships within Ferae among the various extant and fossil groups, as well as their sister placement to Chiroptera after Halliday et al. (2015)[3]:

Chiroptera

 Ferae 

Carnivoramorpha

Mesonychia

 †Triisodontidae 

Goniacodon

Eoconodon

Pentacodontidae

Periptychidae

Pantodonta

 †Arctocyonidae 

Arctocyon

Loxolophus

 Pholidotamorpha 

Taeniodonta

Escavadodontidae

Wyolestidae

Nyctitheriidae

Pantolestidae

Palaeanodonta

Pholidota

 †Oxyclaenidae 

Chriacus

Thryptacodon

Claenodon

Anacodon

Oxyclaenus

Palaeoryctidae

Didelphodus

Acmeodon

Gelastops

 †Creodonta 

Oxyaenidae

Hyaenodonta

References[edit]

  1. ^ a b Gaubert, Philippe; Antunes, Agostinho; Meng, Hao; Miao, Lin; Peigné, Stéphane; Justy, Fabienne; Njiokou, Flobert; Dufour, Sylvain; Danquah, Emmanuel; Alahakoon, Jayanthi; Verheyen, Erik (11 May 2018). "The Complete Phylogeny of Pangolins: Scaling Up Resources for the Molecular Tracing of the Most Trafficked Mammals on Earth". Journal of Heredity. 109 (4): 347–359. doi:10.1093/jhered/esx097. ISSN 0022-1503.
  2. ^ "'Ferae' - The Linnean Collections". linnean-online.org. Retrieved 26 February 2020.
  3. ^ a b c d e f Halliday, Thomas J. D.; Upchurch, Paul; Goswami, Anjali (2015). "Resolving the relationships of Paleocene placental mammals" (PDF). Biological Reviews. 92 (1): 521–550. doi:10.1111/brv.12242. ISSN 1464-7931. PMC 6849585. PMID 28075073.
  4. ^ Amrine-Madsen, H.; Koepfli, K.P.; Wayne, R.K.; Springer, M.S. (2003). "A new phylogenetic marker, apolipoprotein B, provides compelling evidence for eutherian relationships". Molecular Phylogenetics and Evolution. 28 (2): 225–240. doi:10.1016/S1055-7903(03)00118-0. PMID 12878460.
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  7. ^ Madsen, Ole; Scally, Mark; Douady, Christophe J.; Kao, Diana J.; DeBry, Ronald W.; Adkins, Ronald; Amrine, Heather M.; Stanhope, Michael J.; de Jong, Wilfried W.; Springer, Mark S. (2001). "Parallel adaptive radiations in two major clades of placental mammals". Nature. 409 (6820): 610–614. doi:10.1038/35054544. ISSN 1476-4687. PMID 11214318.
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  9. ^ Gaudin, Timothy J.; Gaubert, Philippe; Billet, Guillaume; Hautier, Lionel; Ferreira-Cardoso, Sérgio; Wible, John R. (1 January 2020), Challender, Daniel W. S.; Nash, Helen C.; Waterman, Carly (eds.), "Chapter 1 - Evolution and morphology", Pangolins, Biodiversity of World: Conservation from Genes to Landscapes, Academic Press, pp. 5–23, doi:10.1016/b978-0-12-815507-3.00001-0, ISBN 978-0-12-815507-3, retrieved 26 February 2020
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  11. ^ Zhou, X.; et al. (2011). "Phylogenomic analysis resolves the interordinal relationships and rapid diversification of the Laurasiatherian mammals". Systematic Biology. 61 (1): 150–64. doi:10.1093/sysbio/syr089. PMC 3243735. PMID 21900649. Retrieved 3 October 2011. (Advance Access; published online 7 September 2011)
  12. ^ Nishihara, H.; Hasegawa, M; Okada, N (2006). "Pegasoferae, an unexpected mammalian clade revealed by tracking ancient retroposon insertions". Proceedings of the National Academy of Sciences. 103 (26): 9929–34. Bibcode:2006PNAS..103.9929N. doi:10.1073/pnas.0603797103. PMC 1479866. PMID 16785431.
  13. ^ McKenna, M. C. (1975). "Toward a phylogenetic classification of the Mammalia". In Luckett, W. P.; Szalay, F. S. (eds.). Phylogeny of the Primates. New York: Plenum. pp. 21–46.
  14. ^ Solé, F. & Ladevèze, S. (2017). "Evolution of the hypercarnivorous dentition in mammals (Metatheria, Eutheria) and its bearing on the development of tribosphenic molars." Evolution & Development, 19(2), 56–68.
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  16. ^ Smith, De Bast. "Reassessment of the Small 'Arctocyonid' Prolatidens waudruae from the Early Paleocene of Belgium, and Its Phylogenetic Relationships with Ungulate-Like Mammals". Journal of Vertebrate Paleontology. doi:10.1080/039.033.0410 (inactive 22 January 2020). Retrieved 8 August 2013. Cite journal requires |journal= (help)