Amphicyonidae

Last updated

Amphicyonids
Temporal range:
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O
S
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C
P
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Middle Eocene – Late Miocene
Amphicyon ingens.JPG
Skeleton of Amphicyon
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Suborder: Caniformia
Superfamily:† Amphicyonoidea
Family:† Amphicyonidae
Haeckel, 1866
Subfamilies

†Amphicyoninae
†Daphoeninae
†Haplocyoninae
†Temnocyoninae
†Thaumastocyoninae

Contents

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 Amphicyon-ingens reconstruction.jpg
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 Amphicyonidae.png
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 subfamilySubfamily Amphicyoninae Subfamily Haplocyoninae
(Eurasia) [14] [15] 		Subfamily Daphoeninae
(North America)		Subfamily Temnocyoninae
(North America) [16] 		Subfamily Thaumastocyoninae [17]

Related Research Articles

<span class="mw-page-title-main">Nimravidae</span> Extinct family of mammals in the order Carnivora

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, the nimravids are generally considered closely related and classified as a distinct family in the suborder Feliformia. Fossils have been dated from the Middle Eocene through the Late Miocene epochs, spanning about 33.2 million years.

<i>Miacis</i> Extinct genus of carnivores

Miacis is an extinct genus of placental mammals from clade Carnivoraformes, that lived in North America from early to middle Eocene.

<i>Amphicyon</i> Extinct genus of carnivores

Amphicyon is an extinct genus of large carnivorans belonging to the family Amphicyonidae, subfamily Amphicyoninae, from the Miocene epoch. Members of this family received their vernacular name for possessing bear-like and dog-like features. They ranged over North America, Europe, Asia, and Africa.

<span class="mw-page-title-main">Barbourofelidae</span> Extinct family of feliform carnivorans

Barbourofelidae is an extinct family of carnivorans of the suborder Feliformia, sometimes known as false saber-toothed cats, that lived in North America, Eurasia and Africa during the Miocene epoch and existed for about 7.9 million years. Thought to be an independent lineage from the Nimravidae and Machairodontinae, which had all attained elongated canines, recent research argues that it may be a subfamily of the Nimravidae, extending its biochronological range into the Miocene, although this issue is not yet fully resolved.

<span class="mw-page-title-main">Ursoidea</span> Superfamily of mammals

Ursoidea is a superfamily of arctoid carnivoran mammals that includes the families Subparictidae, Amphicynodontidae, and Ursidae which the last family includes the extant lineages of bears as well the extinct Hemicyoninae and Ursavinae. The interrelationships of ursoids has had slight arrangements. In the past it was thought the extinct Amphicyonidae were stem-bears based on morphological analysis of the ear region, though the most recent publications on early amphicyonids suggests they were basal caniforms. The amphicynodontids are sometimes classified as either a subfamily of bears, a paraphyletic assemblage of early bears, or even stem-pinnipeds. The subparictids were previously classified as amphicynodontine/ids. The hemicyonines have been occasionally reclassified as a separate family.

Xinyuictis is an extinct genus of placental mammals from clade Carnivoraformes, that lived in Asia from early to late Eocene.

<i>Agnotherium</i> Extinct genus of carnivores

Agnotherium is a genus of large sized carnivoran mammals, belonging to the Amphicyonidae, which has been found in Western Europe, and possibly China and Northern Africa, and lived during the Late Miocene epoch. Despite only being known from fragmentary remains, the genus notable for hypercarnivorous adaptions, which have been said to represent the "apex" among its family.

<i>Cynelos</i> Extinct genus of carnivores

Cynelos is a large extinct genus of amphicyonids which inhabited North America, Europe, and Africa from the Early Miocene subepoch to the Late Miocene subepoch 20.4–13.7 Mya, existing for approximately 6 million years.

Gobicyon is an extinct genus of large-sized carnivoran mammals, belonging to the Amphicyonidae, that was discovered in China, Mongolia, and Serbia, and lived during the Middle Miocene epoch. Despite only being known from rather fragmentary remains, recent discoveries showcase that it was an aberrant member of the subfamily Haplocyoninae, with adaptions towards bone-crushing similar to those of a hyaena.

<i>Ysengrinia</i> Extinct genus of mammals known as bear dogs

Ysengrinia is an extinct genus of carnivoran in the family Amphicyonidae (beardogs), which lived in Europe, Asia, and North America during the Early Miocene. It was also reported from Egypt and Namibia, but this material has been reassigned to other genera of beardogs.

<span class="mw-page-title-main">Hemicyoninae</span> Extinct subfamily of bears

Hemicyoninae is an extinct subfamily of Ursidae, often called dog bears. They were bear-like carnivorans living in Europe, North America, Africa and Asia during the Oligocene through Miocene epochs 33.9–5.3 Ma, existing for approximately 28.6 million years. They are sometimes classified as a separate family.

<i>Gustafsonia</i> Extinct genus of carnivores

Gustafsonia is an extinct genus of carnivoran belonging to the family Amphicyonidae. The type species, Gustafsonia cognita, was described in 1986 by Eric Paul Gustafson, who originally interpreted it as a miacid and named it Miacis cognitus. It was subsequently considered to be the only species of the diverse genus Miacis that belonged to the crown-group Carnivora, within the Caniformia, and it was ultimately assigned to the family Amphicyonidae. The type specimen or holotype was discovered in Reeve's bonebed, western Texas, in the Chambers Tuff Formation in 1986. The University of Texas holds this specimen. It is the only confirmed fossil of this species.

<span class="mw-page-title-main">Hyaenodonta</span> Extinct order of mammals

Hyaenodonta is an extinct order of hypercarnivorous placental mammals of clade Pan-Carnivora from mirorder Ferae. Hyaenodonts were important mammalian predators that arose during the early Paleocene in Europe and persisted well into the late Miocene.

<i>Magericyon</i> Extinct genus of carnivores

Magericyon is an extinct genus of Amphicyonid ("bear-dog") that lived during the Miocene 10-9 Ma in what is now Spain.

<span class="mw-page-title-main">Dissopsalini</span> Extinct tribe of mammals

Dissopsalini is an extinct tribe of teratodontid hyaenodonts. Fossil remains of these mammals are known from early to late Miocene deposits in Asia and Africa.

Ammitocyon is a genus of large sized carnivoran mammals, belonging to the Amphicyonidae, that lived during the Late Miocene in what is now Spain. It is notable for its extreme adaptations towards hypercarnivory, its extremely robust skeleton, and was one of the last surviving members of its family.

<span class="mw-page-title-main">Thaumastocyoninae</span> Extinct subfamily of amphicyonids

Thaumastocyoninae is an extinct subfamily of amphicyonids, large terrestrial carnivores, which inhabited what is now Europe during the Miocene epoch. The subfamily was erected by Hürzeler (1940), and is defined by the complete suppression of m1 metaconid, reduction of the premolars, except the p4, which is reinforced, and the oblique abrasion of the teeth, and the possession of hypercarnivorous tendencies. Thaumastocyonines are poorly known, with only about 65 dental specimens, most of those isolated teeth, being known as of 2020, although more complete remains have recently been discovered.

Myacyon is an extinct genus of large sized carnivoran mammals, belonging to the family Amphicyonidae, that lived in Africa 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. 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 or Namibiocyon.

Bonisicyon is an extinct genus of carnivoran mammals, belonging to the family Amphicyonidae. It is the last-surviving member of its family, living in East Africa during the end of the Miocene epoch. Known only from a damaged mandible and isolated teeth, its closer taxonomic affinities are unclear. It is notable for both its small size, and its unique dentition.

Namibiocyon is an extinct genus of carnivoran mammals, belonging to the family Amphicyonidae, that lived in Namibia during the Early Miocene epoch. Before the erection of this taxon in 2022, the type and only species, N. ginsburgi, had been assigned to a variety of other genera. It is notable for its adaptions toward hypercarnivory.

References

  1. 1 2 Wang, Xiaoming and Richard H. Tedford (2008). Dogs; their Fossil Relatives and Evolutionary History. Columbia University Press. pp. 10–11. ISBN   9780231135283.
  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. S2CID   198160461. Archived from the original on 2023-09-04. Retrieved 2023-07-06.
  3. 1 2 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. Bibcode:2016RSOS....360518T. doi:10.1098/rsos.160518. PMC   5098994 . PMID   27853569. Archived from the original on 2016-10-14. Retrieved 2016-10-13.
  4. ↑ Hunt, Robert M. Jr. (2004). "Global Climate and the Evolution of Large Mammalian Carnivores during the Later Cenozoic in North America" (PDF). Cenozoic Carnivores and Global Climate. pp. 139–285. doi:10.1206/0003-0090(2004)285<0139:C>2.0.CO;2. S2CID   86236545. Archived from the original (PDF) on July 20, 2007.{{cite book}}: |journal= ignored (help)
  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:CACFWM]2.0.CO;2. ISSN   0272-4634. S2CID   86235694.
  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". In Janis, Christine M.; Scott, Kathleen M.; Jacobs, Louis L. (eds.). 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.
  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. JSTOR   24106648.
  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. Bibcode:2006JAESc..26..519P. doi:10.1016/j.jseaes.2004.11.003.
  12. ↑ Morlo, Michael; Bastl, Katharina; Habersetzer, Jörg; Engel, Thomas; Lischewsky, Bastian; Lutz, Herbert; von Berg, Axel; Rabenstein, Renate; Nagel, Doris (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. Bibcode:2019JVPal..39E5848M. doi:10.1080/02724634.2019.1705848. S2CID   214431583.
  13. ↑ Morlo, Michael; Bastl, Katharina; Habersetzer, Jörg; Engel, Thomas; Lischewsky, Bastian; Lutz, Herbert; von Berg, Axel; Rabenstein, Renate; Nagel, Doris (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. Bibcode:2019JVPal..39E5848M. doi:10.1080/02724634.2019.1705848. S2CID   214431583.
  14. 1 2 Jiangzuo, Q.; Wang, S.; 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. S2CID   204264754.
  15. ↑ Morales, J.; Abella, 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. S2CID   235363054. Archived from the original on 2023-04-06. 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. Archived from the original on 2023-03-26. 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" (PDF). Fossil Imprint. 75 (3–4): 397–411. doi:10.2478/if-2019-0025. S2CID   210921722. Archived (PDF) from the original on 2022-08-14.
  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. Bibcode:1998RPaPa.101...95B. doi:10.1016/S0034-6667(97)00071-7.
  19. 1 2 3 4 5 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 . S2CID   220714559.
  20. ↑ Ginsburg, Léonard (2002). "Un Amphicyonidae (Carnivora, Mammalia) nouveau du Miocène moyen de Vieux-Collonges (Rhône)". Symbioses (in French). 7: 55–57.
  21. 1 2 3 4 Peign, Stphane; Salesa, Manuel J.; Antn, Mauricio; Morales, Jorge (2008). "A New Amphicyonine (Carnivora: Amphicyonidae) from the Upper Miocene of Batallones-1, Madrid, Spain". Palaeontology. 51 (4): 943–965. Bibcode:2008Palgy..51..943P. doi: 10.1111/j.1475-4983.2008.00788.x . S2CID   55334897. Archived from the original on 2023-04-09. Retrieved 2023-05-01.
  22. ↑ Solé, Floréal; Fischer, Valentin; Denayer, Julien; Speijer, Robert P.; Fournier, Morgane; Le Verger, Kévin; Ladevèze, Sandrine; Folie, Annelise; Smith, Thierry (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 . S2CID   224860287.
  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 . S2CID   229196919.
  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. hdl:10261/137042. ISSN   1698-6180. Archived from the original on 2022-11-27. Retrieved 2023-05-01.
  26. ↑ Grohé, Camille; de Bonis, Louis; Chaimanee, Yaowalak; Chavasseau, Olivier; Rugbumrung, Mana; Yamee, Chotima; Suraprasit, Kantapon; Gibert, Corentin; Surault, Jérôme; Blondel, Cécile; Jaeger, Jean-Jacques (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. S2CID   219296152. Archived from the original on 2023-05-26. Retrieved 2023-05-18.
  27. ↑ Werdelin, Lars; Simpson, Scott W. (2009). "The last amphicyonid (Mammalia, Carnivora) in Africa". Geodiversitas. 31 (4): 775–787. doi:10.5252/g2009n4a775. ISSN   1280-9659. S2CID   73677723.
  28. 1 2 3 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" (PDF). Communications of the Geological Survey of Namibia. 24: 1–16. Archived (PDF) from the original on 2023-04-05. Retrieved 2023-05-01– via Google Scholar.
  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" (PDF). Geodiversitas. 42 (16): 239–255. doi:10.5252/geodiversitas2020v42a16. S2CID   220281575. Archived (PDF) from the original on 2023-09-04. 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. Bibcode:2018JVPal..38E9857J. doi:10.1080/02724634.2018.1539857. ISSN   0272-4634. S2CID   91998860. Archived from the original on 2023-12-19. 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 . S2CID   146620949.
  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" (PDF). Communications of the Geological Survey of Namibia. 24: 1–16. Archived (PDF) from the original on 2023-04-05. Retrieved 2023-05-01– via Google Scholar.
  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 . S2CID   245032640. Archived from the original on 2023-04-18. 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. S2CID   85904748.
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