Amphicyonidae

{{Short description|Extinct family of carnivores}} {{About|the extinct family of mammals|the breeds of domestic dog|Karelian Bear Dog|and|Tahltan Bear Dog|dog bears|Hemicyoninae}}{{Confused|Amphicynodontidae|Arctocyonidae}}{{Automatic taxobox | fossil_range = Middle Eocene – Late Miocene {{Fossil range|42|5.3}} | name = Amphicyonids | image = Amphicyon_ingens.JPG | image_caption = Skeleton of ''Amphicyon'' | display_parents = 2 | taxon = Amphicyonidae | authority = Haeckel, 1866 | subdivision_ranks = Subfamilies | subdivision = †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 Ma), spread to Europe by the late Eocene (35 Ma), and further spread to Asia and Africa by the early Miocene (23 Ma). Amphicyonids were among the first lineages of carnivorans to reach large body sizes. The family had largely disappeared worldwide by the late Miocene (9-7 Ma), with the latest recorded species at the end of the Miocene in Africa. Amphicyonids are colloquially referred to as "'''bear-dogs'''".<ref name="Wang-2008">{{Cite book|last=Wang|first=Xiaoming and Richard H. Tedford|title=Dogs; their Fossil Relatives and Evolutionary History|publisher=Columbia University Press|year=2008|isbn=978-0-231-13528-3|pages=10–11}}</ref>

==Taxonomy== thumb|left|Restoration of ''Amphicyon ingens'' 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.<ref>{{Cite journal |issn=0003-0082 |year=2001 |pages=1–20 |doi=10.1206/0003-0082(2001)331<0001:SOAFNA>2.0.CO;2 |title=Small Oligocene Amphicyonids from North America (Paradaphoenus, Mammalia, Carnivora) |last1=Hunt |first1=Robert M. Jr.|journal=American Museum Novitates |issue=3331 |s2cid=198160461 |url=https://www.biodiversitylibrary.org/item/293579 |access-date=2023-07-06 |archive-date=2023-09-04 |archive-url=https://web.archive.org/web/20230904202543/https://www.biodiversitylibrary.org/item/293579 |url-status=live }}</ref><ref name="Wang-2008" /> 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.<ref name="Tomiya-2016" /><ref name="Hunt-2004">{{cite book |last=Hunt |first=Robert M. Jr. |date=2004 |title=Cenozoic Carnivores and Global Climate |chapter=Global Climate and the Evolution of Large Mammalian Carnivores during the Later Cenozoic in North America |chapter-url=http://digitallibrary.amnh.org/dspace/bitstream/2246/453/8/B285a11.pdf |journal=Bulletin of the American Museum of Natural History |volume= |issue=285 |pages=139–285 |doi= 10.1206/0003-0090(2004)285<0139:C>2.0.CO;2|s2cid=86236545 |archive-url=https://web.archive.org/web/20070720132104/http://digitallibrary.amnh.org/dspace/bitstream/2246/453/8/B285a11.pdf |archive-date=July 20, 2007}}</ref><ref>{{Cite journal|issn=0272-4634 |year=2007 |volume=27 |issue=1 |pages=145–159 |doi=10.1671/0272-4634(2007)27[145:CACFWM]2.0.CO;2 |title=Creodonta and Carnivora from Wadi Moghra, Egypt |last1=Morlo |first1=Michael |last2=Miller |first2=Ellen R. |last3=El-Barkooky |first3=Ahmed N. |journal=Journal of Vertebrate Paleontology |s2cid=86235694 }}</ref>

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 {{convert|5|kg|abbr=on}} and as large as {{convert|100|to|773|kg|abbr=on}}<ref>Sorkin, B. 2008: A biomechanical constraint on body mass in terrestrial mammalian predators. ''Lethaia'', Vol. 41, pp. 333–347.</ref> and evolved from wolf-like to bear-like body forms.<ref>Jacobs, Louis L. Jacobs; Scott, Kathleen Marie: Evolution of Tertiary Mammals of North America: Terrestrial carnivores, Cambridge University Press, 1998</ref>

=== Skull === [[File:Daphoenus vetus skull.jpg|thumb|Skull diagram of ''Daphoenus vetus'']] 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.<ref name="Hunt-1998">{{Cite book |author=Hunt, R. M. Jr. |title=Evolution of Tertiary Mammals of North America, volume 1: Terrestrial carnivores, ungulates, and ungulatelike mammals |publisher=Cambridge University Press |year=1998 |isbn=978-0-521-35519-3 |editor1=Janis, Christine M. |location=Cambridge, England |pages=196–227 |chapter=Amphicyonidae |editor2=Scott, Kathleen M. |editor3=Jacobs, Louis L. |chapter-url=https://digitalcommons.unl.edu/cgi/viewcontent.cgi?params=/context/museummammalogy/article/1275/&path_info=Hunt_ETMNA_1998_Amphicyonidae.pdf}}</ref><ref name="Viranti-1996">{{Cite journal |last=Viranti |first=Suvi |date=1996 |title=European Miocene Amphicyonidae – taxonomy, systematics, and ecology |url=https://www.researchgate.net/publication/293537457 |journal=Acta Zoologica Fennica |volume=204 |pages=1–61}}</ref>

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.<ref name="Hunt-1974">{{Cite journal |last=Hunt |first=Robert M. |date=1974 |title=The auditory bulla in carnivora: An anatomical basis for reappraisal of carnivore evolution |url=https://onlinelibrary.wiley.com/doi/10.1002/jmor.1051430103 |journal=Journal of Morphology |language=en |volume=143 |issue=1 |pages=21–75 |doi=10.1002/jmor.1051430103 |pmid=4826105 |bibcode=1974JMorp.143...21H |issn=0362-2525|url-access=subscription }}</ref><ref name="Hunt-1998" /> 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.<ref name="Hunt-1974" /><ref name="Viranti-1996" />

=== Teeth === [[File:Amphicyon.JPG|left|thumb|Cranium and mandible of ''Amphicyon'', Paläontologisches Museum München]] 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.<ref name="Viranti-1996" /><ref name="Solé-2022">{{Cite journal |last1=Solé |first1=Floréal |last2=Lesport |first2=Jean-François |last3=Heitz |first3=Antoine |last4=Mennecart |first4=Bastien |date=2022-06-15 |title=A new gigantic carnivore (Carnivora, Amphicyonidae) from the late middle Miocene of France |journal=PeerJ |language=en |volume=10 |article-number=e13457 |doi=10.7717/peerj.13457 |doi-access=free |pmid=35726261 |pmc=9206431 |issn=2167-8359}}</ref>

At the start of their evolution, amphicyonids retained the typical placental dental formula of {{DentalFormula|upper=3.1.4.3|lower=3.1.4.3}}, but each subfamily follows their own trend in modifying their teeth.<ref name="Hunt-1998" /><ref name="Viranti-1996" /> 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.<ref name="Hunt-1998" /><ref name="Viranti-1996" /> Thaumastocyonines were the most specialized for hypercarnivory, emphasizing massive blade-like carnassials at the expense of the rest of their postcanine teeth.<ref name="Morales-2019" /><ref name="Solé-2022" />

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.<ref name="Morlo-2019">{{cite journal |last1=Morlo |first1=Michael |last2=Bastl |first2=Katharina |last3=Habersetzer |first3=Jörg |last4=Engel |first4=Thomas |last5=Lischewsky |first5=Bastian |last6=Lutz |first6=Herbert |last7=von Berg |first7=Axel |last8=Rabenstein |first8=Renate |last9=Nagel |first9=Doris |date=3 September 2019 |title=The apex of amphicyonid hypercarnivory: solving the riddle of Agnotherium antiquum Kaup, 1833 (Mammalia, Carnivora) |journal=Journal of Vertebrate Paleontology |volume=39 |issue=5 |article-number=e1705848 |bibcode=2019JVPal..39E5848M |doi=10.1080/02724634.2019.1705848 |s2cid=214431583}}</ref> This period is suggested to be short and would have left the animal somewhat vulnerable.<ref name="Morlo-2019"/>

=== Postcrania === [[File:Daphoenodon superbus restoration & skeleton.png|thumb|Life restoration and skeletal diagram of ''Daphoenodon superbus'']] 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,<ref name="Viranti-1996" /> thaumastocyonines,<ref name="Morales-2021" /> and most daphoenines.<ref name="Hunt-1998" /> 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).<ref name="Viranti-1996" />

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,<ref name="Hunt-2011" /> haplocyonines,<ref>{{Cite journal |last1=Peigné |first1=Stéphane |last2=Heizmann |first2=Elmar P.J. |year=2003 |title=The Amphicyonidae (Mammalia: Carnivora) from Ulm-Westtangente (MN 2, Early Miocene), Baden-Württemberg, Germany – Systematics and ecomorphology |url=https://www.researchgate.net/publication/256846674 |journal=Stuttgarter Beiträge zur Naturkunde, Ser. B |volume=343 |pages=1–133}}</ref> and some species of the large daphoenine ''Daphoenodon''.<ref>{{Cite journal |last=Hunt |first=Robert M. |date=2009 |title=Long-Legged Pursuit Carnivorans (Amphicyonidae, Daphoeninae) from the Early Miocene of North America |url=https://digitalcommons.unl.edu/cgi/viewcontent.cgi?article=1553&context=geosciencefacpub |journal=Bulletin of the American Museum of Natural History |volume=318 |pages=1–95 |doi=10.1206/593.1}}</ref><ref name="Hunt-1998" />

==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. New World amphicyonids include the oldest known amphicyonid, ''Daphoenus'' (40–27 Ma).<ref name="Tomiya-2016">{{Cite journal |url=http://rsos.royalsocietypublishing.org/content/3/10/160518 |doi=10.1098/rsos.160518 |doi-access=free|title=Whence the beardogs? Reappraisal of the Middle to Late Eocene 'Miacis' from Texas, USA, and the origin of Amphicyonidae (Mammalia, Carnivora) |year=2016 |last1=Tomiya |first1=Susumu |last2=Tseng |first2=Zhijie Jack |journal=Royal Society Open Science |volume=3 |issue=10 |article-number=160518 |pmid=27853569 |pmc=5098994 |bibcode=2016RSOS....360518T |access-date=2016-10-13 |archive-date=2016-10-14 |archive-url=https://web.archive.org/web/20161014060307/http://rsos.royalsocietypublishing.org/content/3/10/160518 |url-status=live }}</ref> During the Arikareean in North America, amphicyonids had a competitive advantage over nimravids as the Oligocene progressed due to their greater cursoriality compared to the latter, helping them to persist as the latter disappeared from the continent as terrestrial habitats became more open.<ref>{{Cite journal |last=Castellanos |first=Miguel |date=11 June 2025 |title=Hunting types in North American Eocene–Oligocene carnivores and implications for the 'cat-gap' |url=https://link.springer.com/article/10.1007/s10914-025-09767-2 |journal=Journal of Mammalian Evolution |language=en |volume=32 |issue=2 |article-number=25 |doi=10.1007/s10914-025-09767-2 |issn=1064-7554 |access-date=29 August 2025 |via=Springer Nature Link}}</ref>

=== Extinction === Amphicyonids began to decline in the late Miocene,<ref name="Solé-2022" /> with amphicyonids going extinct in North America and Eurasia around 9 and 7 Ma, respectively.<ref name="Hunt-1998" /><ref name="Solé-2022" /> The most recent securely dated amphicyonid remains belong to the genus ''Bonisicyon'' from the latest Miocene (6.5–5.3 Ma) of Lothagam.<ref name="Werdelin-2009">{{Cite journal |last1=Werdelin |first1=Lars |last2=Simpson |first2=Scott W. |date=2009 |title=The last amphicyonid (Mammalia, Carnivora) in Africa |url=http://www.bioone.org/doi/abs/10.5252/g2009n4a775 |journal=Geodiversitas |language=en |volume=31 |issue=4 |pages=775–787 |doi=10.5252/g2009n4a775 |bibcode=2009Geodv..31..775W |issn=1280-9659 |s2cid=73677723}}</ref> The extinction of amphicyonids have been hypothesized by experts.<ref name="Solé-2022" />

Viranti (1996) goes over potential hypothesizes to the decline and extinction of European amphicyonids. First, they discussed the possibility that the extinction of amphicyonids In Europe was the result of environmental changes. Secondly, they discuss on how much the decline of prey coincides with the extinction of the amphicyonids. They found evidence that the extinction of amphicyonids in Western Europe was contemporary with climatic changes and the reduction of forests.<ref name="Viranti-1996" />

Viranti (1996) also found no evidence of hypercarnivorous amphicyonid diversity being directly connected perissodactyls evolution, as perissodactyls increased in diversity after the extinction of the hypercarnivorous anphicyonids. Lastly, they discussed the possibility of the extinction of actively hunting amphicyonids with bone crushing capabilities was the result of competing with hyenas (including percrocutines), machairodonts, large mustelids, and ursids. They argued while the majority of amphicyonids went extinct prior to the appearance of the hyaenine ''Adcrocuta'', the last amphicyonid species may have been outcompeted by ''Adcrocuta''. They also suggested that machairodonts were likely competitors to hypercarnivorous amphicyonids. On the other hand, amphicyonids may not have competed with percrocutines and mustelids due to their rarity and/or absence MN9 localities. With ursids being able to co-occur with amphicyonids without outcompeting them.<ref name="Viranti-1996" />

Valkenburgh (1999) hypothesized that the decline and extinction of amphicyonids may have been due to the arrival of agriotherini bears and were replaced by taxa more specialized in carnivory (e.g. felids and hyenas) and omnivory (bears).<ref name="Van Valkenburgh-1999">{{Cite journal |last=Van Valkenburgh |first=B. |year=1999 |title=Major patterns in the history of carnivorous mammals |journal=Annual Review of Earth and Planetary Sciences |volume=27 |pages=463–493 |bibcode=1999AREPS..27..463V |doi=10.1146/annurev.earth.27.1.463}}</ref> Werdelin and Simpson (2009) suggested the extinction of ''Bonisicyon'' may have been the result of the inability to compete with ''Eucyon''.<ref name="Werdelin-2009" />

== 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.<ref name="Solé-2022" /> It was thought that many amphicyonids are suggested to have been solitary hunters.<ref name="Morlo-2019" /><ref name="Sorkin-2006">{{cite journal |last1=Sorkin |first1=B. |year=2006 |title=Ecomorphology of the giant bear-dogs ''Amphicyon'' and ''Ischyrocyon'' |url=https://www.researchgate.net/publication/40661208 |journal=Historical Biology |volume=18 |issue=4 |pages=375–388 |bibcode=2006HBio...18..375S |doi=10.1080/08912960600618073 |s2cid=85341612}}</ref><ref name="Viranti-1996" /> However, in the case of ''Daphoenodon'', five adults and one juvenile were found in three adjacent burrows, suggesting this species were gregarious predators.<ref name="Sorkin-2006" /> == Classification == '''Family Amphicyonidae''' {| class="wikitable" |- style="vertical-align: top;" ! Not assigned to a subfamily ! Subfamily Amphicyoninae ! Subfamily Haplocyoninae (Eurasia)<ref name="Jiangzuo-2019"/><ref name="Morales-2021">{{cite journal |last1= Morales |first1= J. |last2= Abella |first2= J. |display-authors= etal |date= May 2021 |title= ''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= Journal of Systematic Palaeontology |volume= 19 |issue= 5 |pages= 393–415 |doi= 10.1080/14772019.2021.1910868 |bibcode= 2021JSPal..19..393M |s2cid= 235363054 |url= https://ddd.uab.cat/record/241083 |access-date= 2023-02-18 |archive-date= 2023-04-06 |archive-url= https://web.archive.org/web/20230406065951/https://ddd.uab.cat/record/241083 |url-status= live }}</ref> ! Subfamily Daphoeninae (North America) ! Subfamily Temnocyoninae (North America)<ref name="Hunt-2011">{{Cite journal |last=Hunt |first=Robert |date=2011-01-01 |title=Evolution of large carnivores during the mid-Cenozoic of North America: The Temnocyonine Radiation (Mammalia, Amphicyonidae) |url=https://digitalcommons.unl.edu/geosciencefacpub/542 |journal=Bulletin of the American Museum of Natural History |volume=358 |pages=1–153 |doi=10.1206/358.1 |archive-url=https://web.archive.org/web/20230802082331/https://digitalcommons.unl.edu/cgi/viewcontent.cgi?article=1554&context=geosciencefacpub |archive-date=2023-08-02 |access-date=2023-04-07}}</ref> ! Subfamily Thaumastocyoninae<ref name="Morales-2019">{{cite journal |last1=Morales |first1=J. |last2=Fejfar |first2=O. |last3=Heizmann |first3=E. |last4=Wagner |first4=J. |last5=Valenciano |first5=A. |last6=Bella |first6=J. |date=2019 |title=A new Thaumastocyoninae (Amphicyonidae, Carnivora) from the early Miocene of Tuchořice, the Czech Republic |url=https://ddd.uab.cat/pub/artpub/2019/220418/fosimp_a2019m12v75n3-4p397-411.pdf |journal=Fossil Imprint |volume=75 |issue=3–4 |pages=397–411 |doi=10.2478/if-2019-0025 |s2cid=210921722 |archive-date=2022-08-14 |archive-url=https://web.archive.org/web/20220814223253/https://ddd.uab.cat/pub/artpub/2019/220418/fosimp_a2019m12v75n3-4p397-411.pdf |url-status=live }}</ref> |- style="vertical-align: top;" | * '''''Amphicyanis''''' * '''''Angelarctocyon''''' ** ''A. australis'' (formerly ''Miacis australis'') * '''''Brachycyon''''' ** ''B. reyi'' ** ''B. palaeolycos'' ** ''B. gaudryi'' * '''''Gustafsonia''''' ** ''G. cognita'' (formerly ''Miacis cognitus'') * '''''Guangxicyon''''' ** ''G. sinoamericanus'' * '''''Harpagocyon'''''<ref name="Berger-1998">{{cite journal |last1= Berger |first1= Jean-Pierre |date= June 1998 |title= 'Rochette' (Upper Oligocene, Swiss Molasse): a strange example of a fossil assemblage |journal= Review of Palaeobotany and Palynology |volume= 101 |issue= 1–4 |pages= 95–110 |doi= 10.1016/S0034-6667(97)00071-7|bibcode= 1998RPaPa.101...95B }}</ref><ref name="Fournier-2020">{{cite journal|last1=Fournier|first1=Morgane|last2=Ladevèze|first2=Sadrine|last3=Le Verger|first3=Kévin|last4=Fischer|first4=Valentin|last5=Speijer|first5=Robert P.|last6=Solé|first6=Floréal|year=2020|title=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|journal=Geodiversitas|volume=42|issue=18|pages=305–325|doi=10.5252/geodiversitas2020v42a18|s2cid=220714559 |doi-access=free |bibcode=2020Geodv..42..305F }}</ref> ** ''H. inusitatus'' * '''''Harpagophagus''''' * '''''Meiniogale'''''<ref>{{cite journal|last=Ginsburg|first=Léonard|year=2002|title=Un Amphicyonidae (Carnivora, Mammalia) nouveau du Miocène moyen de Vieux-Collonges (Rhône)|journal=Symbioses|language=french|volume=7|pages=55–57}}</ref><ref name="Peigné-2008"/> ** ''M. getti'' * '''''Sarcocyon''''' {{Small|(Possible affinities with the Haplocyoninae)}}<ref>{{cite journal|last1=Solé|first1=Floréal|last2=Fischer|first2=Valentin|last3=Denayer|first3=Julien|last4=Speijer|first4=Robert P.|last5=Fournier|first5=Morgane|last6=Le Verger|first6=Kévin|last7=Ladevèze|first7=Sandrine|last8=Folie|first8=Annelise|last9=Smith|first9=Thierry|year=2020|title=The upper Eocene-Oligocene carnivorous mammals from the Quercy Phosphorites (France) housed in Belgian collections|journal=Geologica Belgica|volume=24|issue=1–2|pages=1–16|doi=10.20341/gb.2020.006|s2cid=224860287 |doi-access=free |bibcode=2020GeolB..24....1S }}</ref><ref name="Jiangzuo-2019">{{cite journal |last1= Jiangzuo |first1= Q. |last2= Wang |first2= S. |display-authors=etal |date= September 2019 |title= New material of ''Gobicyon'' (Carnivora, Amphicyonidae, Haplocyoninae) from northern China and a review of Aktaucyonini evolution |journal= Papers in Palaeontology |volume= 7 |issue= 1 |pages= 307–327 |doi= 10.1002/spp2.1283|s2cid= 204264754 }}</ref> ** ''S. ferox'' * '''''Storchictis'''''<ref>{{cite journal|last=de Bonis|first=Louis|year=2020|title=New genus of amphicyonid carnivoran (Mammalia, Carnivora, Amphicyonidae) from the phosphorites of Quercy (France)|journal=Fossil Imprint|volume=76|issue=1|pages=201–208|doi=10.37520/fi.2020.013|s2cid=229196919 |doi-access=free}}</ref> ** ''S. miacinus'' * '''''Symplectocyon'''''<ref name="Fournier-2020"/> ** ''S. praecursor'' * '''''Vishnucyon'''''<ref>{{cite journal|last1=Wang|first1=Xiaoming|last2=Hong-jiang|first2=Wang|last3=Jiangzuo|first3=Qigao|year=2016|title=New record of a haplocyonine amphicyonid in early Miocene of Nei Mongol fills a long-suspected geographic hiatus|journal=Vertebrata PalAsiatica|volume=54|issue=1|pages=21–35}}</ref> ** ''V. chinjiensis'' | * '''''Afrocyon'''''<ref name="Morales-2016">{{Cite journal |last1=Morales |first1=Jorge |last2=Pickford |first2=M. |last3=Valenciano |first3=Alberto |date=2016 |title=Systematics of African Amphicyonidae, with descriptions of new material from Napak (Uganda) and Grillental (Namibia) |journal=Journal of Iberian Geology |volume=42 |issue=2 |pages=131–150 |url=https://digital.csic.es/handle/10261/137042 |doi=10.13039/501100003329 |hdl=10261/137042 |issn=1698-6180 |access-date=2023-05-01 |archive-date=2022-11-27 |archive-url=https://web.archive.org/web/20221127134814/https://digital.csic.es/handle/10261/137042 |url-status=live }}</ref> ** ''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'''''<ref name="Peigné-2008"/><ref>{{cite journal|last1=Grohé|first1=Camille|last2=de Bonis|first2=Louis|last3=Chaimanee|first3=Yaowalak|last4=Chavasseau|first4=Olivier|last5=Rugbumrung|first5=Mana|last6=Yamee|first6=Chotima|last7=Suraprasit|first7=Kantapon|last8=Gibert|first8=Corentin|last9=Surault|first9=Jérôme|last10=Blondel|first10=Cécile|last11=Jaeger|first11=Jean-Jacques|year=2020|title=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|journal=American Museum Novitates|number=3952|pages=1–57|doi=10.1206/3952.1|s2cid=219296152|url=https://www.biodiversitylibrary.org/bibliography/174894|access-date=2023-05-18|archive-date=2023-05-26|archive-url=https://web.archive.org/web/20230526212002/https://www.biodiversitylibrary.org/bibliography/174894|url-status=live}}</ref> ** ''A. mavrini'' * '''''Bonisicyon'''''<ref name="Werdelin-2009"/> ** ''B. illacabo'' * '''''Cynelos'''''<ref name="Morales-2022"/> ** ''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'''''<ref>{{cite journal|last1=Le Verger|first1=Kévin|last2=Solé|first2=Floréal|last3=Ladevèze|first3=Sandrine|year=2020|title=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|journal=Geodiversitas|volume=42|issue=16|pages=239–255|doi=10.5252/geodiversitas2020v42a16|bibcode=2020Geodv..42..239L |s2cid=220281575|url=https://www.zora.uzh.ch/id/eprint/213539/1/g2020v42a16.pdf|access-date=2023-07-06|archive-date=2023-09-04|archive-url=https://web.archive.org/web/20230904202546/https://www.zora.uzh.ch/id/eprint/213539/1/g2020v42a16.pdf|url-status=live}}</ref><ref name="Fournier-2020"/> ** ''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'''''<ref name="Morales-2022"/> ** ''H. euryodon'' * '''''Ictiocyon''''' ** ''I. socialis'' * '''''Ischyrocyon''''' ** ''I. gidleyi'' * '''''Janvierocyon''''' ** ''J. pontignensis'' * '''''Maemohcyon'''''<ref>{{Cite journal |last1=Jiangzuo |first1=Qigao |last2=Li |first2=Chunxiao |last3=Wang |first3=Shiqi |last4=Sun |first4=Danhui |date=2018-11-02 |title=Amphicyon zhanxiangi, sp. nov., a new amphicyonid (Mammalia, Carnivora) from northern China |journal=Journal of Vertebrate Paleontology |volume=38 |issue=6 |article-number=e1539857 |doi=10.1080/02724634.2018.1539857 |bibcode=2018JVPal..38E9857J |s2cid=91998860 |issn=0272-4634 }}</ref><ref name="Peigné-2008"/> ** ''M. potisati'' * '''''Magericyon''''' ** ''M. anceps'' ** ''M. castellanus'' * '''''Myacyon'''''<ref name="Werdelin-2019">{{cite journal |last1= Werdelin |first1= Lars |date= 2019 |title= Middle Miocene Carnivora and Hyaenodonta from Fort Ternan, western Kenya |journal= Geodiversitas |volume= 41 |issue= sp1 |pages= 267–283 |doi= 10.5252/geodiversitas2019v41a6|s2cid= 146620949 |doi-access= free |bibcode= 2019Geodv..41S.267W }}</ref> ** ''M. dojambir'' ** ''M. kiptalami'' ** ''M. peignei'' * '''''Mogharacyon'''''<ref name="Morales-2022">{{Cite journal |last1=Morales |first1=Jorge |last2=Pickford |first2=Martin |date=2022 |title=The taxonomic status of "Ysengrinia" ginsburgi Morales et al. 1998 (Amphicyonidae, Carnivora) from the basal middle Miocene of Arrisdrift, Namibia. |url=https://www.mme.gov.na/files/publications/b07_CommsGSN24pp1_16Morales_Pickford2022.pdf |journal=Communications of the Geological Survey of Namibia |volume=24 |pages=1–16 |via=Google Scholar |access-date=2023-05-01 |archive-date=2023-04-05 |archive-url=https://web.archive.org/web/20230405214443/https://mme.gov.na/files/publications/b07_CommsGSN24pp1_16Morales_Pickford2022.pdf |url-status=live }}</ref> ** ''M. anubisi'' * '''''Namibiocyon'''''<ref name="Morales-2022"/> ** ''N. ginsburgi'' * '''''Paludocyon'''''<ref>{{Cite journal |last1=Morales |last2=Fejfar |last3=Heizmann |last4=Wagner |last5=Valenciano |last6=Abella |date=2021 |title=The Amphicyoninae (Amphicyonidae, Carnivora, Mammalia) Of The Early Miocene From Tuchořice, The Czech Republic |url=https://zenodo.org/record/7167294 |journal=Fossil Imprint |volume=77 |issue=1 |pages=126–144 |doi=10.37520/fi.2021.011 |s2cid=245032640 |doi-access=free |access-date=2023-05-01 |archive-date=2023-04-18 |archive-url=https://web.archive.org/web/20230418063714/https://zenodo.org/record/7167294 |url-status=live }}</ref> ** ''P. bohemicus'' * '''''Pliocyon''''' ** ''P. medius'' ** ''P. ossifragus'' ** ''P. robustus'' * '''''Pseudocyon''''' ** ''P. sansaniensis'' ** ''P. steinheimensis?'' ** ''P. styriacus'' * '''''Pseudarctos''''' ** ''P. bavaricus'' * '''''Pseudocyonopsis'''''<ref name="Peigné-2008">{{Cite journal |last1=Peigné |first1=Stéphane |last2=Salesa |first2=Manuel J. |last3=Antón |first3=Mauricio |last4=Morales |first4=Jorge |title=A New Amphicyonine (Carnivora: Amphicyonidae) from the Upper Miocene of Batallones-1, Madrid, Spain |date=2008 |journal=Palaeontology |language=en |volume=51 |issue=4 |pages=943–965 |doi=10.1111/j.1475-4983.2008.00788.x |bibcode=2008Palgy..51..943P |s2cid=55334897 |doi-access=free }}</ref><ref name="Fournier-2020"/> ** ''P. ambiguus'' ** ''P. antiquus'' ** ''P. landesquei'' ** ''P. quercensis'' * '''''Tartarocyon'''''<ref name="Solé-2022" /> ** ''T. cazanavei'' | * '''''Aktaucyon''''' ** ''A. brachifacialis'' * '''''Gobicyon''''' ** ''G. acutus'' ** ''G. macrognathus'' ** ''G. serbiae'' ** ''G. yei'' ** ''G. zhegalloi'' * '''''Haplocyon'''''<ref name="Fournier-2020"/> ** ''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'''''<ref name="Fejfar-2015">{{cite journal |last1= Fejfar |first1= O. |last2= Heizmann |first2= E. |date= October 2015 |title= An illustrated summary of the lower Miocene carnivores (Mammalia, Carnivora) of Tuchořice, Czech Republic |journal= Historical Biology |volume= 28 |issue= 1–2 |pages= 316–329 |doi= 10.1080/08912963.2015.1029923|s2cid= 85904748 }}</ref> ** ''T. grivense'' * '''''Ysengrinia''''' ** ''Y. americanus'' ** ''Y. depereti'' ** ''Y. geraniana'' ** ''Y. tolosana'' ** ''Y. valentiana'' |}

==References== {{Reflist}}

==External links== * [http://rsos.royalsocietypublishing.org/content/3/10/160518 Whence the beardogs? Reappraisal of the Middle to Late Eocene 'Miacis' from Texas, US, and the origin of Amphicyonidae ]

Category:Amphicyonidae Category:Pliocene extinctions Category:Eocene first appearances Category:Taxa named by Ernst Haeckel Category:Prehistoric mammal families Category:Taxa described in 1886