# Amphicyonidae

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".

## Taxonomy

The family was erected by Haeckel in 1866 (also attributed to Trouessart 1885). Their exact position has long been disputed. Early paleontologists usually defined them as members of Canidae (the dog family) or Ursidae (the bear family), but the modern consensus is that they form their own family. Some researchers have defined it as the sister clade to ursids, based on morphological analysis of the ear region. However, cladistic analysis and reclassification of several species of early carnivore as amphicyonids has strongly suggested that they may be basal caniforms, a lineage older than the origin of both bears and dogs.
Amphicyonids should not be confused with the similar looking (and similarly nicknamed) "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). They should also not be confused with Amphicynodontidae (another family of extinct caniforms which were related to bears or pinnipeds) or Arctocyonidae (a family of "condylarths" which literally translates to "bear-dogs").

## 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) and evolved from wolf-like to bear-like body forms.

### Skull

Amphicyonids tended to have relatively large skulls, with the snout shorter than the rear portion of the cranium. In some large members of the family, such as Amphicyon, the back of the skull develops a sharp sagittal crest which defines attachment points for large jaw muscles.
Amphicyonids had a relatively rudimentary form of auditory bulla, a bony sheath which encases the middle ear cavity. The bulla is small, mostly formed by the crescent-shaped ectotympanic bone below the middle ear. The entotympanics only make a minor contribution whenever they are ossified, which only becomes commonplace in Miocene amphicyonids. In these regards, amphicyonids are similar to living bears, otters, walruses, eared seals, and the red panda. The bulla also helps to distinguish the evolutionary trajectory of amphicyonids: early bears such as Cephalogale have large bullae which are reduced through the course of their evolution, while dogs start out with large bullae which persist through their entire existence. Amphicyonids differ from both dogs and bears in that they start with a small bulla which gradually becomes more strongly developed later in their evolution.

### Teeth

Like most carnivorans, amphicyonid teeth were adapted for carnivory, with large canines near the front and shearing carnassials at the back of the jaw. Amphicyonids were typically mesocarnivorous (majority meat-eating, like dogs) or hypercarnivorous (entirely meat-eating, like cats), and some were adapted for tough abrasive food. Only two small Miocene amphicyonines, Pseudarctos and Ictiocyon, show any evidence for a hypocarnivorous (majority plant-eating) diet.
At the start of their evolution, amphicyonids retained the typical placental dental formula of 3.1.4.33.1.4.3, but each subfamily follows their own trend in modifying their teeth. Daphoenines, for example, have dog-like teeth, with substantial premolars and reduced second and third molars. Temnocyonines and haplocyonines take this approach even further, with massive crushing premolars akin to hyenas. Amphicyonines follow the opposite path, reducing most premolars and greatly enlarging and strengthening the carnassials and second molar. Bears also have large molars, but their teeth are modified into wide rectangular forms for grinding plant material. Amphicyonids did not pursue the same adaptations; their upper molars always maintain a roughly triangular profile for shearing and crushing meat. Thaumastocyonines were the most specialized for hypercarnivory, emphasizing massive blade-like carnassials at the expense of the rest of their postcanine teeth.
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. This period is suggested to be short and would have left the animal somewhat vulnerable.

### Postcrania

Many amphicyonids had cat-like bodies, with a long tail and relatively short, strong limbs suitable for stalking and pouncing on their prey. Later and larger species tended to be plantigrade or semiplantigrade, walking with most or all of the surface of the foot against the ground like bears. This was the norm for amphicyonines, thaumastocyonines, and most daphoenines. It is entirely possible that the largest amphicyonids were capable of both bear-style hunting (chasing down and mauling their prey with teeth and claws) and cat-style hunting (a quick ambush where the prey is killed with a bite to the neck).
Many amphicyonid lineages instead adopted a digitigrade posture and locomotion (walking on their toes) and long legs specialized for running with a primarily front-to-back arc of movement. These cursorial wolf- or hyena-like forms included temnocyonines, haplocyonines, and some species of the large daphoenine Daphoenodon.

## Evolution

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.
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. The species is classically named Arctamphicyon lydekkeri, which may actually be synonymous with a species of Amphicyon.

## Ecology

Amphicyonids are suggested to have ranged in ecology from omnivores to hypercarnivores, with some amphicyonids suggested to have engaged in bone-crushing like some modern hyenas. At least some amphicyonids are suggested to have been solitary hunters.

## Classification

Family Amphicyonidae
| Not assigned to a subfamily | Subfamily Amphicyoninae | Subfamily Haplocyoninae (Eurasia) | Subfamily Daphoeninae (North America) | Subfamily Temnocyoninae (North America) | Subfamily Thaumastocyoninae |
| Amphicyanis Angelarctocyon A. australis (formerly Miacis australis) Brachycyon B. reyi B. palaeolycos B. gaudryi Gustafsonia G. cognita (formerly Miacis cognitus) Guangxicyon G. sinoamericanus Harpagocyon H. inusitatus Harpagophagus Meiniogale M. getti Sarcocyon (Possible affinities with the Haplocyoninae) S. ferox Storchictis S. miacinus Symplectocyon S. praecursor Vishnucyon V. chinjiensis | Afrocyon 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 A. mavrini Bonisicyon B. illacabo Cynelos 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 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 H. euryodon Ictiocyon I. socialis Ischyrocyon I. gidleyi Janvierocyon J. pontignensis Maemohcyon M. potisati Magericyon M. anceps M. castellanus Myacyon M. dojambir M. kiptalami M. peignei Mogharacyon M. anubisi Namibiocyon N. ginsburgi Paludocyon P. bohemicus Pliocyon P. medius P. ossifragus P. robustus Pseudocyon P. sansaniensis P. steinheimensis? P. styriacus Pseudarctos P. bavaricus Pseudocyonopsis P. ambiguus P. antiquus P. landesquei P. quercensis Tartarocyon T. cazanavei | Aktaucyon A. brachifacialis Gobicyon G. acutus G. macrognathus G. serbiae G. yei G. zhegalloi Haplocyon H. elegans H. dombrowskii H. crucians Haplocyonoides H. mordax H. serbiae H. ponticus Haplocyonopsis H. crassidens | Adilophontes A. brachykolos Brachyrhynchocyon 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 Paradaphoenus P. cuspigerus P. minimus P. tooheyi | 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 T. grivense Ysengrinia Y. americanus Y. depereti Y. geraniana Y. tolosana Y. valentiana |