Amphicyon

Amphicyon ("ambiguous dog") is an extinct genus of large carnivorous bone-crushing mammals, popularly known as bear dogs, of the family Amphicyonidae, subfamily Amphicyoninae, from the Burdigalian Epoch until the late Pliocene. They ranged over North America, Europe, Asia, and Africa from 16.9–2.6 Ma ago, existing approximately 14.3 million years.[2]

Amphicyon Temporal range: Burdigalian–Pliocene PreꞒ Ꞓ O S D C P T J K Pg N Early Miocene to Late Pliocene
A. ingens, American Museum of Natural History
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	Carnivora
Family:	†Amphicyonidae
Subfamily:	†Amphicyoninae
Genus:	†Amphicyon Lartet, 1836
Type species
†Amphicyon major[1] Blainville, 1841
Species
A. frendens Matthew, 1924 A. galushai Hunt, 2003 A. giganteus Kaup, 1884 A. ingens Matthew, 1924 A. laugnacensis Ginsburg, 1989 A. longiramus White, 1942 A. lydekkeri (Pilgrim, 1910) A. major Blainville, 1841 A. palaeindicus Lydekker, 1876

Morphology

Restoration of A. ingens

Amphicyon was the typical bear-dog amphicyonid with morphology similar to both bears and dogs. With its robust build and maximum length of 2.5 m (8 ft), the largest species looked more like a bear than a dog. It had a large heavy tail, thick neck, robust limbs and teeth like a wolf. It was probably an omnivore with a lifestyle comparable to that of the brown bear. The Amphicyon was very large for predators of its time but this advantage eventually became a disadvantage because its large body mass was too large to take faster prey.

A. major has been estimated to have has a body mass of 630 kg (1,400 lb),[3] while A. ingens has been estimated around 600 kilograms (1,300 lb)[4] making it one of the largest known amphicyonids.

Behaviour

Amphicyon is believed to have been an omnivore, but to have tended to eat more meat than plants or other foods. It is widely believed that Amphicyon lived on its own, unlike wolves. It is thought to have targeted slow or injured large prey like the Chalicotherium to feed its large appetite. However in a new study published in 2020, examination of the relative grinding area of the molars of Amphicyon indicates that it was purely carnivorous. Both Amphicyon and its relative, Ischyrocyon possessed skeletal features that are characteristic of both ambush and pursuit living predators. As such, it is believed that Amphicyon probably pursued prey for longer distances but at slower speeds than living ambush predators do. Upon catching up to its prey, Amphicyon probably grabbed its victims with its powerfully muscled forelimbs before killing them by tearing into the prey's ribcage or neck using its large, strong canines set in its narrow snout.[5]

Fossil distribution

The earliest occurrences of Amphicyon in North America are from the early to mid-Miocene, found in the Runningwater Formation in Sioux County, Nebraska, and from the lower part of the Troublesome Formation, Colorado (A. galushai, A. frendens, and A. ingens). Although other large amphicyonids from the Miocene of North America have been placed in Amphicyon, many of these carnivores are now placed in other amphicyonid genera. The Amphicyon lineage in the New World is restricted to the above three species (18.8–14.2 Mya). Particularly rich samples of the large North American species of Amphicyon have been found in the Sheep Creek Formation (A. frendens) and Olcott Formation (A. ingens) of central Sioux County, northwest Nebraska.[1] Amphicyon has also been found in France and Spain[6] in Europe. Amphicyon's youngest range is on the Indian subcontinent, where it disappeared only in the late Pliocene.

Amphicyon range based on fossil finds

Species

A. major jaw

Amphicyon major lived from 16.9–9.0 Ma, approximately 7.9 million years.[7] Specimens have been found in across Europe and in western Turkey.[7] The species was named by De Blainville in 1841.[7] A. major was large in size, comparable to a modern lion or tiger.[8] The estimated mass of A. major is around 180 kg (397 lb) with the functions derived for limb bones and craniodental measurements.[9]

Jaws, Paläontologische Museum München

Amphicyon giganteus was a widespread European species that lived during the early Burdigalian to early Langhian, approximately from 20.4–15.9 Mya,[10] with possible material from Namibia.[11] The species was first described in 1884 by Kaup.[12] A specimen of Iberotherium rexmanueli zbyszewskii with teeth marks from A. giganteus was found in Portugal. It is unknown if the young Iberotherium was attacked or the carcass found and scavenged. The find was described by paleontologists Antunesa et al. in 2006.[13]

Amphicyon galushai represents the first occurrence of Amphicyon in North America, from approximately 18.8–17.5 Mya during the early Hemingfordian. Described by Robert M. Hunt Jr. in 2003, it is mostly known from fossils found in the Runningwater Formation of western Nebraska, a complete adult skull, a partial juvenile skull, 3 mandibles and teeth and postcranial elemenents representing least 15 individuals. In addition there is a skull fragment from the Troublesome Formation of Colorado.[1] It is considered ancestral to the late Hemingfordian species, A. frendens.

Amphicyon frendens lived during the late Hemingfordian, 17.5–15.9 Mya,[10] The species was originally described by W. Matthew in 1924 from specimens found in the middle member of the Sheep Creek Formation, Sioux County, Nebraska.[14] A. frendens specimens have since been found at sites in Harney and Malheur Counties, Oregon. A specimen examined by S. Legendre and C. Roth in 1988 yielded an estimated body mass of 135.6 kg (300 lb),[3] similar to that of Ischyrocyon, Amphicyon galushai and its borophagine competitor, Epicyon, with which it coexisted.

Amphicyon ingens lived during the early to middle Barstovian, 15.8–14.0 Mya.[15] The species was originally described by W. Matthew in 1924 from specimens found in the Olcott Formation, Sioux County, Nebraska.[14] Specimens attributed to this species have since been found in California, Colorado, and New Mexico.

Amphicyon palaeindicus is known from the Bugti Hills in Pakistan. It was first described by Richard Lydekker in 1876. The exact age of the fossil sites it was recovered from is unclear, though they seem to range from the late Oligocene to the late Miocene. Its status as an actual species is unclear, as nearly all remains attributed to Amphicyon in the region were attributed to it.

Amphicyon lydekkeri is known from the Dhok Pathan horizon in Pakistan. It was originally described by Pilgrim in 1910 and attributed to its own genus, Arctamphicyon. However, the differences between "Arctamphicyon" and Amphicyon may ultimately be negligible, and it is most likely part of the genus.[16] With the Dhok Pathan deposits dating to the late Pliocene, Amphicyon lydekkeri is the youngest amphicyonid known.

References

1. Hunt, Robert M. (2003). "Intercontinental Migration of Large Mammalian Carnivores: Earliest Occurrence of the Old World Beardog Amphicyon (Carnivora, Amphicyonidae) in North America" (PDF). Bulletin of the American Museum of Natural History. 279: 77–115. doi:10.1206/0003-0090(2003)279<0077:c>2.0.co;2.
2. "Fossilworks: Amphicyon". fossilworks.org.
3. Legendre, S.; Roth, C. (1988). "Correlation of carnassial tooth size and body weight in recent carnivores (Mammalia)". Historical Biology. 1 (1): 85–98. doi:10.1080/08912968809386468.
4. Sorkin, B. (2008-04-10). "A biomechanical constraint on body mass in terrestrial mammalian predators". Lethaia. 41 (4): 333–347. doi:10.1111/j.1502-3931.2007.00091.x.
5. https://www.researchgate.net/publication/40661208_Ecomorphology_of_the_giant_bear-dogs_Amphicyon_and_Ischyrocyon
6. Rafael Fraguas (January 7, 2010). "Animales prehistóricos en el Metro". El País (in Spanish). Madrid.
7. "Fossilworks: Amphicyon major". fossilworks.org.
8. National Geographic Prehistoric Mammals by Alan Turner
9. Figueirido; et al. (2011). "Body mass estimation in amphicyonid carnivoran mammals: A multiple regression approach from the skull and skeleton" (PDF). Acta Palaeontologica Polonica. 56 (2): 225–246. doi:10.4202/app.2010.0005.
10. Hunt, Robert M. (1998). "Amphicyonidae". In Janis, C. M.; Scott, K.M.; Jacobs, L. L (eds.). Evolution of tertiary mammals of North America, volume 1: Terrestrial carnivores, ungulates and ungulatelike mammals. Cambridge, UK: Cambridge University Press. pp. 196–227. ISBN 978-0521355193.
11. Pickford; et al. (1996). "Preliminary results of new excavations at Arrisdrift, middle Miocene of southern Namibia". C. R. Acad. Sci. Paris. II (332): 991–996.
12. Catalogue of the Fossil Mammalia in the British Museum Natural History, Dept. of Geology, Richard Lydekker
13. M. T. Antunes; et al. (2006). "Ichnological evidence of a Miocene rhinoceros bitten by a bear-dog (Amphicyon giganteus)". Annales de Paléontologie. 92: 31–39. doi:10.1016/j.annpal.2005.10.002.
14. W. D. Matthew. 1924. Third contribution to the Snake Creek Fauna. Bulletin of the American Museum of Natural History 50:59-210
15. Sorkin, B. (2006). "Ecomorphology of the giant bear-dogs Amphicyon and Ischyrocyon". Historical Biology. 18 (4): 375–388. doi:10.1080/08912960600618073.
16. Peigné, Stéphane (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.