Biology:Amphicyonidae

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Short description: Extinct family of carnivores

Amphicyonids
Temporal range:
Middle Eocene – Late Miocene
Amphicyon ingens.JPG
Skeleton of Amphicyon
Scientific classification e
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Suborder: Caniformia
Superfamily: Amphicyonoidea
Family: Amphicyonidae
Haeckel, 1866
Subfamilies

Amphicyoninae
Daphoeninae
†Haplocyoninae
Temnocyoninae
Thaumastocyoninae

Amphicyonidae is an extinct family of terrestrial carnivorans belonging to the suborder Caniformia. They first appeared in North America in the middle Eocene (around 45 mya), spread to Europe by the late Eocene (35 mya), and further spread to Asia and Africa by the early Miocene (23 mya). They had largely disappeared worldwide by the late Miocene (5 mya), with the latest recorded species at the end of the Miocene in Africa. They were among the first carnivorans to evolve large body size. Amphicyonids are colloquially referred to as "bear-dogs".[1]

Taxonomy

Restoration of Amphicyon ingens

The family was erected by Haeckel in 1866 (also attributed to Trouessart 1885). Their exact position has long been disputed. Some early paleontologists defined them as members of the family Canidae, but the modern consensus is that they form their own family. Some researchers have defined it as the sister clade to ursids (bears), based on morphological analysis of the ear region.[2][1] However, cladistic analysis and reclassification of several species of early carnivore as amphicyonids has strongly suggested that they may be basal caniforms, from lineages older than the origin of both bears and dogs.[3][4][5]

Description

Amphicyonids ranged in size from as small as 5 kg (11 lb) and as large as 100 to 773 kg (220 to 1,704 lb)[6] and evolved from wolf-like to bear-like body forms.[7]

Early amphicyonids, such as Daphoenodon, possessed a digitigrade posture and locomotion (walking on their toes), while many of the later and larger species were plantigrade or semiplantigrade.[8] The amphicyonids were obligate carnivores, unlike the Canidae, which are hypercarnivores or mesocarnivores.[9]

There is often some confusion with the similar looking (and similarly named) "dog-bears", a more derived group of caniforms that is sometimes classified as a family (Hemicyonidae), but is more often considered a primitive subfamily of ursids (Hemicyoninae).

Evolution

Dymaxion map of Amphicyonidae

It has long been uncertain where amphicyonids originated. It was thought that they may have crossed from Europe to North America during the Miocene epoch, but recent research suggests a possible North American origin from the miacids Miacis cognitus and M. australis (now renamed as the genera Gustafsonia and Angelarctocyon, respectively). As these are of North American origin, but appear to be early amphicyonids, it may be that the Amphicyonidae actually originates in North America.[3]

Other New World amphicyonids include the oldest known amphicyonid, Daphoenus (37–16 Mya).

Amphicyonids began to decline in the late Miocene, and disappeared by the end of the epoch. The exact reasons for this are unclear. The most recent known amphicyonid remains are teeth known from the Dhok Pathan horizon, northern Pakistan , dating to 7.4-5.3 mya.[10] The species is classically named Arctamphicyon lydekkeri, which may actually be synonymous with a species of Amphicyon.[11]

Fossils of juvenile Agnotherium, Ischyrocyon, and Magericyon all show an unusual type of tooth eruption in which there is a vulnerable stage at about two or three years of age where the subadult animal has no functional molar or carnassial teeth, the only functional cheek teeth being several milk premolars.[12] This period was suggested to be "presumably short" but would have made it very difficult for the animal to process food.[13] This type of tooth replacement is not seen in similar carnivorans like ursids or canids, and may have been one factor in the extinction of the Amphicyonidae.

Classification

Family Amphicyonidae

Not assigned to a subfamily Subfamily Amphicyoninae Subfamily Haplocyoninae
(Eurasia)[14][15]
Subfamily Daphoeninae
(North America)
Subfamily Temnocyoninae
(North America)[16]
Subfamily Thaumastocyoninae[17]
  • Afrocyon[25]
    • A. burolleti
  • Amphicyon
    • A. astrei
    • A. carnutense?
    • A. cooperi
    • A. frendens
    • A. galushai
    • A. giganteus?
    • A. gutmanni
    • A. ingens
    • A. lactorensis
    • A. laugnacensis
    • A. longiramus
    • A. lyddekeri?
    • A. major (type)
    • A. palaeindicus?
    • A. pannonicus
    • A. shahbazi
    • A. ulungurensis
    • A. zhanxiangi
  • Askazansoria[21][26]
    • A. mavrini
  • Bonisicyon[27]
    • B. illacabo
  • Cynelos[28]
    • C. caroniavorus
    • C. crassidens
    • C. helbingo
    • C. idoneus
    • C. jitu
    • C. jourdan
    • C. lemanensis
    • C. macrodon
    • C. minor
    • C. pivetaui
    • C. rugosidens
    • C. schlosseri
    • C. sinapius
  • Cynodictis[29][19]
    • C. cayluxensis
    • "C. compressidens"
    • C. crassus
    • C. exilis
    • C. ferox
    • C. intermedius
    • C. lacustris
    • C. longirostris
    • "C. palmidens"
    • C. peignei
  • Dehmicyon
    • D. schlosseri
  • Goupilictis
    • G. minor
  • Hecubides[28]
    • H. euryodon
  • Ictiocyon
    • I. socialis
  • Ischyrocyon
    • I. gidleyi
  • Janvierocyon
    • J. pontignensis
  • Maemohcyon[30][21]
    • M. potisati
  • Magericyon
    • M. anceps
    • M. castellanus
  • Myacyon[31]
    • M. dojambir
    • M. kiptalami
    • M. peignei
  • Mogharacyon[28]
    • M. anubisi
  • Namibiocyon[32]
    • N. ginsburgi
  • Paludocyon[33]
    • P. bohemicus
  • Pliocyon
    • P. medius
    • P. ossifragus
    • P. robustus
  • Pseudocyon
    • P. sansaniensis
    • P. steinheimensis?
    • P. styriacus
  • Pseudarctos
    • P. bavaricus
  • Pseudocyonopsis[21][19]
    • P. ambiguus
    • P. antiquus
    • P. landesquei
    • P. quercensis
  • Tartarocyon
    • T. cazanavei
  • Adilophontes
    • A. brachykolos
  • Brachyrhyncocyon
    • B. dodgei
    • B. intermedius
    • B. montanus
  • Daphoenictis
    • D. tedfordi
  • Daphoenodon
    • D. falkenbachi
    • D. notionastes
    • D. robustum
    • D. periculosus
    • D. skinneri
    • D. superbus
  • Daphoenus
    • D. felinus
    • D. hartshornianus
    • D. lambei
    • D. nebrascensis
    • D. socialis
    • D. transversus
    • D. vetus
  • Delotrochanter
    • D. major
    • D. oryktes
    • D. petersoni
  • Mammacyon
    • M. ferocior
    • M. obtusidens
  • Rudiocyon
    • R. amplidens
  • Temnocyon
    • T. altigenis
    • T. ferox
    • T. fingeruti
    • T. macrogenys
    • T. percussor
    • T. subferox
  • Agnotherium
    • A. antiquus
  • Ammitocyon
    • A. kainos
  • Crassidia
    • C. intermedia
  • Peignecyon
    • P. felinoides
  • Thaumastocyon
    • T. bourgeoisi
    • T. dirus
  • Tomocyon[34]
    • T. grivense
  • Ysengrinia
    • "Y. americanus"
    • Y. depereti
    • Y. geraniana
    • Y. tolosana
    • "Y. valentiana"

References

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  2. Hunt Jr., Robert M. (2001). "Small Oligocene Amphicyonids from North America (Paradaphoenus, Mammalia, Carnivora)". American Museum Novitates (3331): 1–20. doi:10.1206/0003-0082(2001)331<0001:SOAFNA>2.0.CO;2. ISSN 0003-0082. https://www.biodiversitylibrary.org/item/293579. Retrieved 2023-07-06. 
  3. 3.0 3.1 Tomiya, Susumu; Tseng, Zhijie Jack (2016). "Whence the beardogs? Reappraisal of the Middle to Late Eocene 'Miacis' from Texas, USA, and the origin of Amphicyonidae (Mammalia, Carnivora)". Royal Society Open Science 3 (10): 160518. doi:10.1098/rsos.160518. PMID 27853569. PMC 5098994. Bibcode2016RSOS....360518T. http://rsos.royalsocietypublishing.org/content/3/10/160518. Retrieved 2016-10-13. 
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  5. Morlo, Michael; Miller, Ellen R.; El-Barkooky, Ahmed N. (2007). "Creodonta and Carnivora from Wadi Moghra, Egypt". Journal of Vertebrate Paleontology 27 (1): 145–159. doi:10.1671/0272-4634(2007)27[145:CACFWM2.0.CO;2]. ISSN 0272-4634. 
  6. Sorkin, B. 2008: A biomechanical constraint on body mass in terrestrial mammalian predators. Lethaia, Vol. 41, pp. 333–347.
  7. Jacobs, Louis L. Jacobs; Scott, Kathleen Marie: Evolution of Tertiary Mammals of North America: Terrestrial carnivores, Cambridge University Press, 1998
  8. Wang, Xiaoming and Tedford, Richard H. Dogs: Their Fossil Relatives and Evolutionary History. New York: Columbia University Press, 2008. p10-11, 29
  9. Hunt, R. M. Jr. (1998). "Amphicyonidae". Evolution of Tertiary Mammals of North America, volume 1: Terrestrial carnivores, ungulates, and ungulatelike mammals. Cambridge, England: Cambridge University Press. pp. 196–227. ISBN 978-0-521-35519-3. https://books.google.com/books?id=I-RgojcDyWYC&q=R.+M.+Hunt.+1998.+Amphicyonidae.&pg=PA196. 
  10. Sehgal, R. K. and A. C. Nanda (2002). "Age of the fossiliferous Siwalik sediments exposed in the vicinity of Nurpur, District Kangra, Himachal Pradesh". Current Science 82 (4): 392–395. 
  11. Stéphane Peigné (2006). "A new amphicyonid (Mammalia, Carnivora, Amphicyonidae) from the late middle Miocene of northern Thailand and a review of the amphicyonine record in Asia". Thailand Journal of Asian Earth Sciences 26 (5): 519–532. doi:10.1016/j.jseaes.2004.11.003. Bibcode2006JAESc..26..519P. 
  12. Morlo, Michael; Bastl, Katharina; Habersetzer, Jörg; Engel, Thomas; Lischewsky, Bastian; Lutz, Herbert; von Berg, Axel; Rabenstein, Renate et al. (3 September 2019). "The apex of amphicyonid hypercarnivory: solving the riddle of Agnotherium antiquum Kaup, 1833 (Mammalia, Carnivora)". Journal of Vertebrate Paleontology 39 (5): e1705848. doi:10.1080/02724634.2019.1705848. Bibcode2019JVPal..39E5848M. 
  13. Morlo, Michael; Bastl, Katharina; Habersetzer, Jörg; Engel, Thomas; Lischewsky, Bastian; Lutz, Herbert; von Berg, Axel; Rabenstein, Renate et al. (3 September 2019). "The apex of amphicyonid hypercarnivory: solving the riddle of Agnotherium antiquum Kaup, 1833 (Mammalia, Carnivora)". Journal of Vertebrate Paleontology 39 (5): e1705848. doi:10.1080/02724634.2019.1705848. Bibcode2019JVPal..39E5848M. 
  14. 14.0 14.1 Jiangzuo, Q. et al. (September 2019). "New material of Gobicyon (Carnivora, Amphicyonidae, Haplocyoninae) from northern China and a review of Aktaucyonini evolution". Papers in Palaeontology 7 (1): 307–327. doi:10.1002/spp2.1283. 
  15. Morales, J. et al. (May 2021). "Ammitocyon kainos gen. et sp. nov., a chimerical amphicyonid (Mammalia, Carnivora) from the late Miocene carnivore traps of Cerro de los Batallones (Madrid, Spain)". Journal of Systematic Palaeontology 19 (5): 393–415. doi:10.1080/14772019.2021.1910868. https://ddd.uab.cat/record/241083. Retrieved 2023-02-18. 
  16. Hunt, Robert (2011-01-01). "Evolution of large carnivores during the mid-Cenozoic of North America: The Temnocyonine Radiation (Mammalia, Amphicyonidae)". Papers in the Earth and Atmospheric Sciences. https://digitalcommons.unl.edu/geosciencefacpub/542. Retrieved 2023-04-07. 
  17. Morales, J.; Fejfar, O.; Heizmann, E.; Wagner, J.; Valenciano, A.; Bella, J. (2019). "A new Thaumastocyoninae (Amphicyonidae, Carnivora) from the early Miocene of Tuchořice, the Czech Republic". Fossil Imprint 75 (3–4): 397–411. doi:10.2478/if-2019-0025. https://ddd.uab.cat/pub/artpub/2019/220418/fosimp_a2019m12v75n3-4p397-411.pdf. 
  18. Berger, Jean-Pierre (June 1998). "'Rochette' (Upper Oligocene, Swiss Molasse): a strange example of a fossil assemblage". Review of Palaeobotany and Palynology 101 (1–4): 95–110. doi:10.1016/S0034-6667(97)00071-7. Bibcode1998RPaPa.101...95B. 
  19. 19.0 19.1 19.2 19.3 19.4 Fournier, Morgane; Ladevèze, Sadrine; Le Verger, Kévin; Fischer, Valentin; Speijer, Robert P.; Solé, Floréal (2020). "On the morphology of the astragalus and calcaneus of the amphicyonids (Carnivora, Mammalia) from the Paleogene of Europe: implications for the ecology of the European bear-dogs". Geodiversitas 42 (18): 305–325. doi:10.5252/geodiversitas2020v42a18. 
  20. Ginsburg, Léonard (2002). "Un Amphicyonidae (Carnivora, Mammalia) nouveau du Miocène moyen de Vieux-Collonges (Rhône)" (in french). Symbioses 7: 55–57. 
  21. 21.0 21.1 21.2 21.3 Peign, Stphane; Salesa, Manuel J.; Antn, Mauricio; Morales, Jorge (2008). "A New Amphicyonine (Carnivora: Amphicyonidae) from the Upper Miocene of Batallones-1, Madrid, Spain" (in en). Palaeontology 51 (4): 943–965. doi:10.1111/j.1475-4983.2008.00788.x. Bibcode2008Palgy..51..943P. https://onlinelibrary.wiley.com/doi/10.1111/j.1475-4983.2008.00788.x. Retrieved 2023-05-01. 
  22. Solé, Floréal; Fischer, Valentin; Denayer, Julien; Speijer, Robert P.; Fournier, Morgane; Le Verger, Kévin; Ladevèze, Sandrine; Folie, Annelise et al. (2020). "The upper Eocene-Oligocene carnivorous mammals from the Quercy Phosphorites (France) housed in Belgian collections". Geologica Belgica 24 (1–2): 1–16. doi:10.20341/gb.2020.006. 
  23. de Bonis, Louis (2020). "New genus of amphicyonid carnivoran (Mammalia, Carnivora, Amphicyonidae) from the phosphorites of Quercy (France)". Fossil Imprint 76 (1): 201–208. doi:10.37520/fi.2020.013. 
  24. Wang, Xiaoming; Hong-jiang, Wang; Jiangzuo, Qigao (2016). "New record of a haplocyonine amphicyonid in early Miocene of Nei Mongol fills a long-suspected geographic hiatus". Vertebrata PalAsiatica 54 (1): 21–35. 
  25. Morales, Jorge; Pickford, M.; Valenciano, Alberto (2016). "Systematics of African Amphicyonidae, with descriptions of new material from Napak (Uganda) and Grillental (Namibia)". Journal of Iberian Geology 42 (2): 131–150. doi:10.13039/501100003329. ISSN 1698-6180. https://digital.csic.es/handle/10261/137042. Retrieved 2023-05-01. 
  26. Grohé, Camille; de Bonis, Louis; Chaimanee, Yaowalak; Chavasseau, Olivier; Rugbumrung, Mana; Yamee, Chotima; Suraprasit, Kantapon; Gibert, Corentin et al. (2020). "The Late Middle Miocene Mae Moh Basin of Northern Thailand: The Richest Neogene Assemblage of Carnivora from Southeast Asia and a Paleobiogeographic Analysis of Miocene Asian Carnivorans". American Museum Novitates (3952): 1–57. doi:10.1206/3952.1. https://www.biodiversitylibrary.org/bibliography/174894. Retrieved 2023-05-18. 
  27. Werdelin, Lars; Simpson, Scott W. (2009). "The last amphicyonid (Mammalia, Carnivora) in Africa" (in en). Geodiversitas 31 (4): 775–787. doi:10.5252/g2009n4a775. ISSN 1280-9659. http://www.bioone.org/doi/abs/10.5252/g2009n4a775. 
  28. 28.0 28.1 28.2 Morales, Jorge; Pickford, Martin (2022). "The taxonomic status of "Ysengrinia" ginsburgi Morales et al. 1998 (Amphicyonidae, Carnivora) from the basal middle Miocene of Arrisdrift, Namibia.". Communications of the Geological Survey of Namibia 24: 1–16. https://www.mme.gov.na/files/publications/b07_CommsGSN24pp1_16Morales_Pickford2022.pdf. Retrieved 2023-05-01. 
  29. Le Verger, Kévin; Solé, Floréal; Ladevèze, Sandrine (2020). "Description of a new species of Cynodictis Bravard & Pomel, 1850 (Carnivora, Mammalia) from the Quercy Phosphorites with comments on the use of skull morphology for phylogenetics". Geodiversitas 42 (16): 239–255. doi:10.5252/geodiversitas2020v42a16. https://www.zora.uzh.ch/id/eprint/213539/1/g2020v42a16.pdf. Retrieved 2023-07-06. 
  30. Jiangzuo, Qigao; Li, Chunxiao; Wang, Shiqi; Sun, Danhui (2018-11-02). "Amphicyon zhanxiangi, sp. nov., a new amphicyonid (Mammalia, Carnivora) from northern China". Journal of Vertebrate Paleontology 38 (6): e1539857. doi:10.1080/02724634.2018.1539857. ISSN 0272-4634. Bibcode2018JVPal..38E9857J. https://doi.org/10.1080/02724634.2018.1539857. Retrieved 2023-05-09. 
  31. Werdelin, Lars (2019). "Middle Miocene Carnivora and Hyaenodonta from Fort Ternan, western Kenya". Geodiversitas 41 (sp1): 267–283. doi:10.5252/geodiversitas2019v41a6. 
  32. Morales, Jorge; Pickford, Martin (2022). "The taxonomic status of "Ysengrinia" ginsburgi Morales et al. 1998 (Amphicyonidae, Carnivora) from the basal middle Miocene of Arrisdrift, Namibia.". Communications of the Geological Survey of Namibia 24: 1–16. https://www.mme.gov.na/files/publications/b07_CommsGSN24pp1_16Morales_Pickford2022.pdf. Retrieved 2023-05-01. 
  33. Morales; Fejfar; Heizmann; Wagner; Valenciano; Abella (2021). "The Amphicyoninae (Amphicyonidae, Carnivora, Mammalia) Of The Early Miocene From Tuchořice, The Czech Republic". Fossil Imprint 77 (1): 126–144. doi:10.37520/fi.2021.011. https://zenodo.org/record/7167294. Retrieved 2023-05-01. 
  34. Fejfar, O.; Heizmann, E. (October 2015). "An illustrated summary of the lower Miocene carnivores (Mammalia, Carnivora) of Tuchořice, Czech Republic". Historical Biology 28 (1–2): 316–329. doi:10.1080/08912963.2015.1029923. 

External links


Wikidata ☰ Q248422 entry