Myacyon
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''Myacyon'' is an extinct genus of large sized carnivoran mammals, belonging to the family
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 appear in Asia, ...
(“bear dogs”), that lived in
Africa Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...
during the Miocene epoch. Due to the limited scope and fragmentary nature of the severely damaged holotype, as well as the illustrations in its descriptions, which have been called inadequate, usage of this genus poses serious issues. However, it is notable for being one of the last surviving members of its family and its adaptions to
hypercarnivory A hypercarnivore is an animal which has a diet that is more than 70% meat, either via active predation or by scavenging. The remaining non-meat diet may consist of non-animal foods such as fungi, fruits or other plant material. Some extant exampl ...
. Its relationships to other amphicyonids are obscure, and it is not closely related to '' Bonisicyon'', the other late surviving African genus, although it has been proposed that it descends from a species of ''
Cynelos '' Cynelos'' is a large extinct genus of bear dogs which inhabited North America, Europe, and Africa from the Early Miocene subepoch to the Late Miocene subepoch 20.4—13.7 Annum, Mya, existing for approximately . Species * ''C. caroniavorus'' ...
''.


History and species

The genus was erected in 1992 on the basis of a severely damaged and fragmented distal half of a right
mandible In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
containing m1-m2, and a not yet erupted m3, found at the locality Oued Mya in
Algeria ) , image_map = Algeria (centered orthographic projection).svg , map_caption = , image_map2 = , capital = Algiers , coordinates = , largest_city = capital , relig ...
. In 2005, the species ''
Agnotherium ''Agnotherium'' is a genus of large sized carnivoran mammals, belonging to the Amphicyonidae ("bear dogs"), which has been found in Western Europe, and possibly Northern Africa, and lived during the Late Miocene epoch. Despite only being known f ...
kiptalami'' was described based on a snout broken off behind the second molar (KNM BN 488). A revision of African amphicyonids by Morales et al. in 2016 moved this species, as well as all African material referred to ''Agnotherium'' to the genus ''Myacyon''. However, a redescription of ''Agnotherium'' in 2020 tentatively included material from
Tunisia ) , image_map = Tunisia location (orthographic projection).svg , map_caption = Location of Tunisia in northern Africa , image_map2 = , capital = Tunis , largest_city = capital , ...
in this genus instead of ''Myacyon'', although they note that the lack of definitive upper teeth belonging to ''Agnotherium'' does not allow them to confidently assign these remains. This material consists of a right maxilla fragment (NOM T-370), a proximal left radius fragment (NOM T-179) and a metatarsal V (NOM T-2269).Morlo M, Bastl K, Habersetzer J, Engel T, Lischewsky B, Lutz H, von Berg A, Rabenstein R, Nagel D. 2020
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.
Morales et al. further referred teeth (KNM-LT 23049, KNM-LT 23073 and KNM-LT 23051) of a large amphicyonid from
Lothagam Lothagam is a geological formation located in Kenya, near the southwestern shores of Lake Turkana, from Kanapoi. It is located between the Kerio and Lomunyenkuparet Rivers on an uplifted fault block. Lothagam has deposits dating to the Miocene-P ...
, which represents one of the youngest occurrences of the family worldwide,Werdelin, Lars (2003). Mio-Pliocene Carnivora from Lothagam, Kenya. In
''Lothagam: The Dawn of Humanity in Eastern Africa''
pp. 261–328. New York. Columbia University Press. .
to ''Myacyon'' cf. ''kiptalami''. Additionally, they proposed that a m2 from Beni Mellal,
Morocco Morocco (),, ) officially the Kingdom of Morocco, is the westernmost country in the Maghreb region of North Africa. It overlooks the Mediterranean Sea to the north and the Atlantic Ocean to the west, and has land borders with Algeria to ...
represents a new species. The species ''?Myacyon peignei'' was described in 2019 based on remains found in
Kenya ) , national_anthem = "Ee Mungu Nguvu Yetu"() , image_map = , map_caption = , image_map2 = , capital = Nairobi , coordinates = , largest_city = Nairobi , ...
and tentatively assigned to the genus. It is only known from teeth—the holotype is a P4 (KNM-FT 3611), while the other remains are a left m1 (KNM-FT 3379) and right m1 (KNM-FT 3399). The genus name references the
Wadi Wadi ( ar, وَادِي, wādī), alternatively ''wād'' ( ar, وَاد), North African Arabic Oued, is the Arabic term traditionally referring to a valley. In some instances, it may refer to a wet (ephemeral) riverbed that contains water onl ...
Mya, which is located close to the site where the remains were found. The species name ''dojambir'' is the Arabic word for “December”, the month in which the remains were discovered, whereas the names of the other two species honor Kiptalam Cheboi and Stéphane Peigné, respectively.


Fossil distribution

''M. dojambir -'' * Oued Mya 1, Algeria, ca. 11.2-9 Ma ''M. kiptalami -'' * Member D,
Ngorora Formation The Ngorora Formation is a geological formation in Kenya preserving fossils dating to the Miocene. The uppermost member of the formation shows sign of a faunal turnover that occurred around 11 to 10.5 million years ago, coinciding with faunal cha ...
, Kenya, ca. 12 Ma * Kipsaraman, Muruyur Formation, Kenya, ca. 14.5 Ma * Hondeklip Bay, South Africa, ca. 12 Ma ''M.''
cf. The abbreviation ''cf.'' (short for the la, confer/conferatur, both meaning "compare") is used in writing to refer the reader to other material to make a comparison with the topic being discussed. Style guides recommend that ''cf.'' be used onl ...
''kiptalami -'' * Bled Douarah, Beglia Formation, Tunisia, 13-11 Ma * Samburu Hills, Kenya, 9.5 Ma * Lower Nawata Formation,
Lothagam Lothagam is a geological formation located in Kenya, near the southwestern shores of Lake Turkana, from Kanapoi. It is located between the Kerio and Lomunyenkuparet Rivers on an uplifted fault block. Lothagam has deposits dating to the Miocene-P ...
, Kenya, 7.4 ± 0.1 – 6.5 ± 0.1 MaStorrs, G. W. (2003). Numerical Age Control for the Miocene-Pliocene Succession at Lothagam, a Hominoid-bearing Sequence in the Northern Kenya Rift. In
''Lothagam: The Dawn of Humanity in Eastern Africa''
pp. 43–64. New York. Columbia University Press. .
''M. peignei -'' *
Fort Ternan Fort Ternan is a small town in Kericho County, Kenya, located 50 kilometres east of Kisumu and five kilometres east of Koru. It is named after Col. Trevor Ternan C.M.G.D.S.O. Fort Ternan is located at the western border of the former Rift Valley Pr ...
, Kenya, early
Serravallian The Serravallian is, in the geologic timescale, an List of time periods, age or a stage (stratigraphy), stage in the middle Miocene epoch (geology), Epoch/series (stratigraphy), Series, which spans the time between 13.82 annum, Ma and 11.63 Ma (m ...
, 13.8-13.7 Ma ''Myacyon'' sp. - * Beni Mellal, Morocco, ca. 14 Ma


Description

The dimensions of ''Myacyon dojambir'' approach those of the largest '' Amphicyon giganteus,'' which have been estimated to weigh over 300 kg, whereas the material from the Samburu Hills and Beni Mellal is considerably smaller. ''M. kiptalami'' is as large or slightly larger than a lion, and the remains from Kipsaraman, Bled Douarah and Lothagam suggest a similar size. ?''M. peignei'' is not nearly as large, being smaller than most Early Miocene species of ''Cynelos'', except for ''C. euryodon,'' itself comparable in size to the 42 kg heavy ''C. lemanensis''. It is characterized by its sectorial molars, showing adaptions towards hypercarnivory. The large m1 is unlike that of any other amphicyonid, possessing a shallow carnassial notch. It has an elongated trigonid, a strong protoconid, which possesses a strong and tall trenchant anterior crest and a short talonid, with a robust, crested hypoconid and a far distally located entoconid which is small, poorly developed and lacks a crest. The paraconid is indistinct and separated from the protoconid by a very weak notch, which is visible on the buccal margin of the anterior crest, while the metaconid is reduced and situated slightly posteriorly. The short and oblong m2 lacks a paraconid and only possesses a poorly developed metaconid, which is situated at the level of the trenchant protoconid. Its talonid is short and narrower than the trigonid, possessing a strong hypoconid but lacking an entoconid. The trigonids of both teeth have a strong buccal cingulum. The m3 is present, but probably reduced. The cranial remains of ''M. kiptalami'' superficially resembles those of large
felids Felidae () is the family of mammals in the order Carnivora colloquially referred to as cats, and constitutes a clade. A member of this family is also called a felid (). The term "cat" refers both to felids in general and specifically to the do ...
and
nimravids Nimravidae is an extinct family of carnivorans, sometimes known as false saber-toothed cats, whose fossils are found in North America and Eurasia. Not considered to belong to the true cats (family Felidae), the nimravids are generally considered ...
, with a short snout, enlarged canines and long diastemata. As in lions, the anterior extremities of its nasals flare outwards and slightly downwards, thereby shifting the nasal notches laterally and widening the nostrils. Its maxillo-premaxillary does not touch the nasals until it has reached the orbital line, which is much further back than in canids. Furthermore, the upper border of the stout premaxilla is lower than in dogs, but similarly projects anteriorly, even if it is less pointed, resulting in the incisors lying in an almost straight line, curved incisor battery of dogs. The incisors have steep roots and long axes and occlusal apices, which point sharply towards the rear. As a result of the outbowing of the maxilla in the region of the carnassial and molar, the maxilla is antero-posteriorly shortened. A further adaption towards shortening the snout is the 90° rotation of the P3 along the part of the maxilla in which it is located. It is tucked behind the parastyle of the carnassial, a position usually be filled by a protocone among other amphicyonids. Above it, a large infraorbital foramen is located. It furthermore possesses a large facial fossa, which lies anterior to the orbit and extends anteriorly and ventrally as a curved furrow immediately behind the canine jugum.As a result of these adaptions, Agnotherium had a considerable gape, with canine tips extending 40 mm beneath the occlusal surface of the carnassial and incisors. ''M. peignei'' is only known from a handful tooth, although the development of the protoconid shelf on its P4 is a distinct feature of this taxon.


Paleoecology

At Oued Mya, Myacyon coexisted with the equid ''
Hipparion ''Hipparion'' (Greek, "pony") is an extinct genus of horse that lived in North America, Asia, Europe, and Africa during the Miocene through Pleistocene ~23 Mya—781,000 years ago. It lived in non-forested, grassy plains, shortgrass prairie or st ...
'' and the rhinoceros ''
Aceratherium ''Aceratherium'' (Greek: "without (a) horn" (keratos), "beast" (therion)) was a genus of rhinoceros of the subfamily Aceratheriinae that lived in Eurasia during the Miocene. Taxonomy ''Aceratherium'' was coined by Kaup (1832) for ''"Rhinoceros ...
'', as well as the giraffid ''
Palaeotragus ''Palaeotragus'' ("ancient goat") is a genus of very large, primitive, okapi-like giraffids from the Miocene of Africa and Eurasia. ''Palaeotragus primaevus'' is the older species, being found in early to mid-Miocene strata, while ''P. germaini' ...
'' and
proboscideans The Proboscidea (; , ) are a taxonomic order of afrotherian mammals containing one living family (Elephantidae) and several extinct families. First described by J. Illiger in 1811, it encompasses the elephants and their close relatives. From ...
. The discoveries at Member D of the Ngorora Formation suggests that it was once covered by open woodland, with fluvial conditions being present. Here ''Myacyon'' was found among the immense
hyaenodont Hyaenodonta ("hyena teeth") is an extinct order of hypercarnivorous placental pan-carnivoran mammals from mirorder Ferae. Hyaenodonts were important mammalian predators that arose during the early Paleocene in Europe and persisted well into the ...
''
Megistotherium ''Megistotherium'' ("greatest beast") is an extinct genus of hyainailourid hyaenodonts from paraphyletic subfamily Hyainailourinae that lived in Africa. Etymology The name of this genus comes and . The name of species ''Megistotherium osteo ...
'' and its smaller relative ''
Dissopsalis ''Dissopsalis'' ("double scissors") is a genus of teratodontine hyaenodonts of the tribe Dissopsalini. The older species, ''D. pyroclasticus'', lived in Kenya during the middle Miocene, while the type species, ''D. carnifex'', lived in Pakista ...
'', the lioness-sized hyaena ''
Percrocuta ''Percrocuta'' is an extinct genus of hyena-like feliform carnivores. It lived in Europe, Asia, and Africa, during the Miocene epoch. Characteristics With a maximum length of 1.50 m (5 ft), ''Percrocuta'' was much bigger than its moder ...
'' as well as a variety of smaller
carnivorans Carnivora is a monophyletic order of placental mammals consisting of the most recent common ancestor of all cat-like and dog-like animals, and all descendants of that ancestor. Members of this group are formally referred to as carnivorans, ...
. A large variety of
herbivores A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...
are known from this site, among them the proboscidean ''
Deinotherium ''Deinotherium'' was a large elephant-like proboscidean that appeared in the Middle Miocene and survived until the Early Pleistocene. Although superficially resembling modern elephants, they had notably more flexible necks, limbs adapted to a mo ...
'',
bovids The Bovidae comprise the biological family of cloven-hoofed, ruminant mammals that includes cattle, bison, buffalo, antelopes, and caprines. A member of this family is called a bovid. With 143 extant species and 300 known extinct species, the ...
, suids and the eary
hippopotamus The hippopotamus ( ; : hippopotamuses or hippopotami; ''Hippopotamus amphibius''), also called the hippo, common hippopotamus, or river hippopotamus, is a large semiaquatic mammal native to sub-Saharan Africa. It is one of only two extan ...
''
Kenyapotamus ''Kenyapotamus'' is a possible ancestor of living hippopotamuses that lived roughly 16 million to 8 million years ago during the Miocene epoch. Its name reflects that its fossils were first found in modern-day Kenya. Although little is known ab ...
''. Fossils of
crocodylians Crocodilia (or Crocodylia, both ) is an order of mostly large, predatory, semiaquatic reptiles, known as crocodilians. They first appeared 95 million years ago in the Late Cretaceous period (Cenomanian stage) and are the closest living ...
, freshwater turtles and fish have also been found. As indicated by palaeosol carbon isotope studies and tooth microwear patterns, Fort Ternan, where ?''M. peignei'' has been found, was covered by open woodland and located adjacent to highlands. The locality shares many taxons with the Ngorora Formation, among them '' Afrochoerodon'', ''
Climacoceras ''Climacoceras'' (from Greek for "ladder horns") is a genus of extinct artiodactyl ungulates that lived in Africa and Europe during the Miocene. The members of ''Climacoceras'' were related to giraffes, and the genus was formerly placed within th ...
, Chilotheridium, Gentrytragus,
Kipsigicerus ''Kipsigicerus'' is an extinct genus of East African antelope from the Middle Miocene. Its closest living relative is the Four-horned Antelope, four-horned-antelope. It was discovered in Fort Ternan, Kenya and was originally described as a specie ...
,
Dorcatherium ''Dorcatherium'' is an extinct genus of tragulid ruminant which existed in Europe, East Africa and the Siwaliks during the Miocene and Pliocene The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5 ...
'' and ''Kenyapotamus.'' The carnivoran assemblage is also similar, including ''Percrocuta'', ''Dissopsalis'' and '' Tugenictis'', with a gigantic hyainailourine as the largest predator. The hominoid ''
Kenyapithecus ''Kenyapithecus wickeri'' is a fossil ape discovered by Louis Leakey in 1961 at a site called Fort Ternan in Kenya. The upper jaw and teeth were dated to 14 million years ago. One theory states that ''Kenyapithecus'' may be the common ancestor ...
'' is among the few genera unique to this site.


References

{{Taxonbar, from=Q116223872 Bear dogs Prehistoric carnivoran genera