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Mesonychia

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Mesonychia
Temporal range:
Early Paleocene—Late Eocene, 63–34 Ma Suspected Late Cretaceous origins[1][2]
Harpagolestes macrocephalus (Mesonychidae) skull reconstruction
Hapalodectes hetangensis (Hapalodectidae) skull
Scientific classification Edit this classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Clade: Scrotifera
Grandorder: Ferungulata
Clade: Pan-Euungulata
Order: †Mesonychia
Van Valen, 1969
Families

†Hapalodectidae
†Mesonychidae

Synonyms

Acreodi (Eberle and McKenna, 2002)[3]

Mesonychia ("middle claws") is an extinct order of small to large-sized omnivorous to carnivorous hoofed mammals related to ungulates. Originally, it was hypothesized that mesonychians were a group of ungulates,[4][5] however recent analysis now found them to be outside of the group entirely. Instead were a basal order of hoofed mammals within Pan-Euungulata.[6][7][1] The order consisted of two families, Hapalodectidae and the more diverse and widespread Mesonychidae.[1] It is believed that mesonychians evolved during the Late Cretaceous, at least 66.7 Ma.[1][5]

Mesonychids, as well as mesonychians as a whole, first appeared in fossil record in the Early Paleocene of East Asia with Dissacus rotundus, Yantanglestes, Hukoutherium, and Dissaccussium.[8][1] Dissacus would later disperse into North America during the Early Paleocene,[9] before dispersing into Northwestern Europe by the end of it.[10] Dissacus was a jackal-sized predator,[11][10] but taxon of a closely related or identical genus, Ankalagon, from the early to middle Paleocene of New Mexico, were far larger, growing to the size of a bear.[12] A later genus, Pachyaena, entered North America by the earliest Eocene, where it evolved into species that were at least as large.[13] Mesonychids were largest predatory mammals in North America and Europe during the Late Paleocene to Middle Eocene.[14][10] In Asia, the record of their history suggests they grew gradually larger and more predatory over time, then shifted to scavenging and bone-crushing lifestyle.[15]

During the Middle Eocene, mesonychians saw a decline in diversity across their distribution.[16][10][17][1] Originally, it was suggested mesonychids, such as Mongolestes, survived into the Early Oligocene.[18][19] This has proven to be questionable as the Ulan Gochu Formation of Mongolia has been revised to the late Middle to earliest Late Eocene.[20] The latest occurrence of the order was within the Ergilin Dzo Formation.[21]

Taxonomy

[edit]

Classification

[edit]
Cladogram showing the position of the Mesonychia

Cope (1975) first classified mesonychids as carnivorans due to the trochelar face of the astragalus is completely grooved.[22] However, in a later paper, Cope reclassified mesonychids as creodonts.[23] In their 1966 paper, Van Valen removed mesonychids from creodonts and placed them within the order Condylarthra.[24] Szalay and Gould (1966) would support this classification within their paper.[18] In 1969, Van Valen suggested mesonychids were a separate order of mammals closely related to Artiodactyla, known as Mesonychia.[25] The order is sometimes referred to by its older name Acreodi.[26][27]

Ting and Li (1987) and Zhou et al. (1995) suggested Hapalodectidae as a separate family of mesonychians,[28][29] a classification supported by phylogenetic modeling.[1][29] Some genera may need revision to clarify the actual number of species or remove ambiguity about genera (such as Dissacus and Ankalagon).[30][1][10] In 2023, Solé and colleagues reclassified nearly European species of Dissacus within the newly revived Hyaenodictis. Additionally, the North American species, "Dissacus" willwoodensis, was also reclassified as Hyaenodictis. Within the same study, they noted the close relation between Ankalagon and Dissacus navajovius, which suggests the former may be a large species of Dissacus.[1]

Relationship with Triisodontidae

[edit]

When Scott (1894) described Triisodontidae, he noted the family was possibly closely related to mesonychids.[31] Triisodontines were considered to have been a subfamily of mesonychids by Matthew (1907),[32] although they were later moved to Arctocyonidae.[33] However, McKenna and Bell (1997) would reclassify them as a family of mesonychians.[27] Szalay (1969) suggested triisodontines were ancestral to mesonychids as the latter dentition was noted to have been an improvement over triisodontines.[34] Phylogenetic analysis by Tabuce et al. (2011) recovered mesonychians as a monophyletic group within Acreodi evolving from triisodontids-like mammals.[35]

Sarah et al. (2015) recovered mesonychians as basal ungulates most closely related to the "arctocyonids" Mimotricentes, Deuterogonodon and Chriacus. Within the abstract, they found triisodontids to be a paraphyletic stem to mesonychians.[36] However, majority of phylogenetic analyses have recovered mesonychians outside of ungulates entirely, with triisodontids being more closely related to ungulates than to mesonychians.[6][7][1]

Relationship with whales

[edit]

Mesonychians possess unusual triangular molar teeth that are similar to those of Cetacea (whales and dolphins), especially those of the archaeocetes, as well as having similar skull anatomies and other morphologic traits. For this reason, scientists had long believed that mesonychians were the direct ancestor of Cetacea, but the discovery of well-preserved hind limbs of archaic cetaceans, as well as more recent phylogenetic analyses[37][38][39] now indicate cetaceans are more closely related to hippopotamids and other artiodactyls than they are to mesonychians, and this result is consistent with many molecular studies.[40] The similarity in dentition and skull may be the result of primitive ungulate structures in related groups independently evolving to meet similar needs as predators; some researchers have suggested that the absence of a first toe and a reduced metatarsal are basal features (synapomorphies) indicating that mesonychians, perissodactyls, and artiodactyls are sister groups.[30]

Most paleontologists now doubt that whales are descended from mesonychians.[4][41][6][7] Some experts have hypothesized mesonychians were basal ungulates, and that cetaceans are descended from advanced ungulates (Artiodactyla), either deriving from, or sharing a common ancestor with, anthracotheres (the semiaquatic ancestors of hippos).[4] However, mesonychians being recovered within ungulates in cladistic analyses only surfaces following the deletion of Andrewsarchus, which has been recovered as a mesonychid within the cladogram.[41][42][7] One possible conclusion is that Andrewsarchus has been incorrectly classified. The current uncertainty may, in part, reflect the fragmentary nature of the remains of some crucial fossil taxa, such as Andrewsarchus.[41] Current analysis recovered Andrewsarchus as a basal Cetancodontamorpha being closely related to entelodonts, cetaceans, and hippopotamuses.[6][7] Spaulding et al. (2009) only recovered mesonychians being closely related to cetaceans, following the deletion of Carnivora, "Creodonta", "Lipotyphia", and Raoellidae.[7]

Cladogram recovered by Spaulding et al. (2009), which recovered the order outside of ungulates entirely:[7]

Evolution

[edit]

Mesonychians were believed to have evolved during the Late Cretaceous period, at least 66.7 Ma,[1][5] with both families were thought to have originate in the Late Cretaceous based on phylogenetic analysis.[1] The earliest known mesonychians fossils were dated to 63 Ma.[5] In East Asia, mesonychids were known from East Asia during the Early Paleocene and were represented by Dissacus rotundus, Yantanglestes, Hukoutherium, and Dissaccussium.[8][1] Within North America, Dissacus appeared during the late Torrejonian and was contemporary with the "triisodontid" Triisodon, with Ankalagon appearing during the latest Torrejonian.[43] The presence of mesonychids in Asia and in North America suggests mesonychids initial radiation began in the Early Paleocene.[1] Hapalodectids first appeared in the fossil record during the Middle Paleocene in Asia.[1][44] Before the end of the Paleocene, mesonychids would disperse into Europe with the appearance of Dissacus europaeus.[10] Hapalodectids would disperse into North America during the Paleocene-Eocene thermal maximum, which saw the appearance of Hapalodectes anthracinus.[44]

Extinction

[edit]

The disappearance of Dissacus was contemporary with the appearance of Pachyaena and Palaeonictis, although competition is unlikely to have been the reason for the extinction of Dissacus europaeus.[10] Pachyaena would go extinct during the Intra-European Faunal Turnover roughly 55 Ma, which was likely related to climatic cooling.[10] Following its extinction, European Pachyaena was replaced by European Hyaenodictis.[10][1] The extinction of European mesonychids was contemporary with the Ypresian-Lutetian Mammal Turnover event, which coincided with the cooling period following the Early Eocene Climatic Optimum.[45] Following the extinction of European mesonychids and oxyaenids, hyaenodonts saw a diversification in species and body sizes.[46][45][47][48]

Characteristics

[edit]

Mesonychians have often been reconstructed as resembling wolves albeit superficially, but they would have appeared very different in life. With a short lower spine stiffened by revolute joints, they would have run with stiff backs like modern ungulates rather than bounding or loping with flexible spines like modern carnivorans. While later mesonychians evolved a suite of limb adaptations for running similar to those in both wolves and deer, their legs remained comparatively thick.[30] They would have resembled no group of living animals. Early mesonychians probably walked on the flats of their feet (plantigrade), while later ones walked on their toes (digitigrade). These later mesonychians had hooves, one on each toe, with four toes on each foot. The foot was compressed for efficient running with the axis between the third and fourth toes (paraxonic); it would have looked something like a hoofed paw.[49]

Mesonyx model reconstruction at the Natural History Museum, London

Mesonychians varied in size; some species were as small as a fox, although some species approached the size of moderately sized bears.[49] Some members of the group are known only from skulls and jaws, or have fragmentary postcranial remains.[49][1] But where skeletons are known, they indicate that mesonychians had large heads with strong jaw muscles, relatively long necks, and robust bodies with robust limbs that could run effectively but not rotate the hand or reach out to the side. An unrelated early group of mammalian predators, the creodonts, also had unusually large heads and limbs that traded flexibility for efficiency in running; large head size may be connected to inability to use the feet and claws to help catch and process food, as many modern carnivorans do. Some mesonychians are reconstructed as predatory (comparable to canids), others as scavengers or carnivore-scavengers with bone-crushing adaptations to their teeth (comparable to the large hyenas), and some as omnivorous (comparable to pigs, humans, or black bears). They may not have included hypercarnivores (comparable to felids); their teeth were not as effective at cutting meat as later groups of large mammalian predators.[50] Although some experts consider Mesonyx to be a hypercarnivore due to the metaconids of the lower molars are completely reduced.[18]

There is evidence to suggest that some genera were sexually dimorphic.[51] These "wolves on hooves" were probably one of the more important predator groups in the late Paleocene and Eocene ecosystems of Europe (which was an archipelago at the time), Asia (which was an island continent), and North America.[52] Mesonychian dentition consisted of molars modified to generate vertical shear, thin blade-like lower molars, and carnassial notches, but no true carnassials.[50] The largest species are considered to have been scavengers.[15]

References

[edit]
  1. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 Solé, Floréal; Fournier, Morgane; Ladevèze, Sandrine; et al. (2023). "New postcranial elements of mesonychid mammals from the Ypresian of France: New hypotheses for the radiation and evolution of the mesonychids in Europe". Journal of Mammalian Evolution. 30 (2): 371–401. doi:10.1007/s10914-023-09651-x.
  2. ↑ Gingerich, Philip D.; Uhen, Mark D. (1998). "Likelihood estimation of the time of origin of Cetacea and the time of divergence of Cetacea and Artiodactyla". Palaeontologia Electronica. doi:10.26879/98008.
  3. ↑ Eberle, Jaelyn J; McKenna, Malcolm C (June 2002). "Early Eocene Leptictida, Pantolesta, Creodonta, Carnivora, and Mesonychidae (Mammalia) from the Eureka Sound Group, Ellesmere Island, Nunavut". Canadian Journal of Earth Sciences. 39 (6): 899–910. Bibcode:2002CaJES..39..899E. doi:10.1139/e02-001. ISSN 0008-4077.
  4. 1 2 3 Geisler, Jonathan H.; Theodor, Jessica M. (2009). "Hippopotamus and whale phylogeny". Nature. 458 (7236): E1–4, discussion E5. Bibcode:2009Natur.458....1G. doi:10.1038/nature07776. PMID 19295550. S2CID 4320261.
  5. 1 2 3 4 Gingerich, Philip D.; Uhen, Mark D. (1998). "Likelihood estimation of the time of origin of Cetacea and the time of divergence of Cetacea and Artiodactyla". Palaeontologia Electronica. doi:10.26879/98008.
  6. 1 2 3 4 Yu, Yang; Gao, Hongyan; Li, Qiang; Ni, Xijun (2023-01-01). "A new entelodont (Artiodactyla, Mammalia) from the late Eocene of China and its phylogenetic implications". Journal of Systematic Palaeontology. 21 (1). Bibcode:2023JSPal..2189436Y. doi:10.1080/14772019.2023.2189436. ISSN 1477-2019.
  7. 1 2 3 4 5 6 7 Spaulding, Michelle; O'Leary, Maureen A.; Gatesy, John (2009-09-23). "Relationships of Cetacea (Artiodactyla) Among Mammals: Increased Taxon Sampling Alters Interpretations of Key Fossils and Character Evolution". PLOS ONE. 4 (9) e7062. Bibcode:2009PLoSO...4.7062S. doi:10.1371/journal.pone.0007062. ISSN 1932-6203. PMC 2740860. PMID 19774069.
  8. 1 2 Missiaen, Pieter. "亚洲早古近纪哺乳动物生物年代学与生物地理学的新认识."古脊椎动物学报 49.1 (2011). paper
  9. ↑ Toosey, William J.; Williamson, Thomas E.; Shelley, Sarah L.; Brusatte, Stephen L. (2024-11-11). "The osteology of Triisodon crassicuspis (Cope, 1882): New insights into the enigmatic "archaic" placental mammal group "Triisodontidae"". PLOS ONE. 19 (11) e0311187. doi:10.1371/journal.pone.0311187. ISSN 1932-6203. PMC 11554371.
  10. 1 2 3 4 5 6 7 8 9 Solé, Floréal; Godinot, Marc; Laurent, Yves; Galoyer, Alain; Smith, Thierry (2018-09-01). "The European Mesonychid Mammals: Phylogeny, Ecology, Biogeography, and Biochronology". Journal of Mammalian Evolution. 25 (3): 339–379. doi:10.1007/s10914-016-9371-8. ISSN 1573-7055. S2CID 254701971.
  11. ↑ Solé, Floréal; Morse, Paul E.; Bloch, Jonathan I.; Gingerich, Philip D.; Smith, Thierry (July 2021). "New specimens of the mesonychid Dissacus praenuntius from the early Eocene of Wyoming and evaluation of body size through the PETM in North America". Geobios. 66–67: 103–118. doi:10.1016/j.geobios.2021.02.005.
  12. ↑ O'Leary, Maureen A.; Lucas, Spencer G.; Williamson, Thomas E. (2000). "A new specimen of Ankalagon (Mammalia, Mesonychia) and evidence of sexual dimorphism in mesonychians". Journal of Vertebrate Paleontology. 20 (2): 387–93. doi:10.1671/0272-4634(2000)020[0387:ANSOAM]2.0.CO;2. JSTOR 4524103.
  13. ↑ O'Leary, Maureen A.; Rose, Kenneth D. (1995). "Postcranial Skeleton of the Early Eocene Mesonychid Pachyaena (Mammalia: Mesonychia)". Journal of Vertebrate Paleontology. 15 (2): 401–430. Bibcode:1995JVPal..15..401O. doi:10.1080/02724634.1995.10011238. ISSN 0272-4634. JSTOR 4523639.
  14. ↑ O'Leary, Maureen A.; Rose, Kenneth D. (1995). "Postcranial Skeleton of the Early Eocene Mesonychid Pachyaena (Mammalia: Mesonychia)". Journal of Vertebrate Paleontology. 15 (2): 401–430. Bibcode:1995JVPal..15..401O. doi:10.1080/02724634.1995.10011238. ISSN 0272-4634. JSTOR 4523639.
  15. 1 2 Xun Jin (2012). "New mesonychid (Mammalia) material from the Lower Paleogene of the Erlian Basin, Nei Mongol, China" (PDF). Vertebrata PalAsiatica. 50 (3): 245–257.
  16. ↑ Van, Valkenburgh B. (1999). "Major patterns in the history of carnivorous mammals". Annual Review of Earth and Planetary Sciences. 27: 463–493. Bibcode:1999AREPS..27..463V. doi:10.1146/annurev.earth.27.1.463.
  17. ↑ Jiangzuo, Qigao; Lyras, Georgios; Grohe, Camille; Werdelin, Lars; Niu, Kecheng; Huang, Dongting; Li, Shijie; Jiang, Hao; Fu, Jiao; Wan, Yang; Liu, Jinyi; Wang, Shi-Qi; Deng, Tao (November 2025). "A new ecomorph of Nimravidae, and the early macrocarnivorous niche exploration in Carnivora". Proceedings. Biological Sciences. 292 (2059) 20251686. doi:10.1098/rspb.2025.1686. ISSN 1471-2954. PMC 12646760. PMID 41290163.
  18. 1 2 3 Szalay, Frederick; Gould, S. J. (1966). "Asiatic Mesonychidae (Mammalia, Condylarthra)". Bulletin of the American Museum of Natural History. 132 (2): 127–174.
  19. ↑ Jin, Xun (2005). "Mesonychids from Lushi Basin, Henan Province, China" (PDF). Vertebrata PalAsiatica. 43 (2): 151–164.
  20. ↑ Bai, Bin; Li, Qian; Zhou, Xin-Ying; Wang, Xiao-Yang; Xu, Ran-Cheng; Zhang, Xin-Yue; Quan, Shuo-Shuo; Meng, Jin; Wang, Yuan-Qing (2025). "Litho- and Biostratigraphy of the East Mesa in Shara Murun Region of the Erlian Basin, Inner Mongolia, China, and the subdivision of the Ulangochuian Asian Land Mammal Age". American Museum Novitates. 2025 (4034). doi:10.1206/4034.1. ISSN 0003-0082.
  21. ↑ Tsubamoto, Takehisa, et al. "Fossil evidence of a mesonychid mammal from the upper Eocene Ergilin Dzo Formation, Mongolia." Paleontological Research 16.2 (2012): 171-174.
  22. ↑ Cope, ED (1875). "On the supposed Carnivora of the Eocene of the Rocky Mountains". Proceedings of the Academy of Natural Sciences of Philadelphia. 27: 444--449.
  23. ↑ Cope, E. D. (March–December 1880). "On the Genera of the Creodonta". Proceedings of the American Philosophical Society. 19 (107): 76–82. JSTOR 982610.
  24. ↑ Van Valen, Leigh M. (1966). "Deltatheridia, a new order of mammals". Bulletin of the American Museum of Natural History. 132 (1): 1–126. hdl:2246/1126.
  25. ↑ Van Valen, Leigh (1969). "The Multiple Origins of the Placental Carnivores". Evolution. 23 (1): 118–130. doi:10.2307/2406488. JSTOR 2406488.
  26. ↑ Eberle, Jaelyn J; McKenna, Malcolm C (June 2002). "Early Eocene Leptictida, Pantolesta, Creodonta, Carnivora, and Mesonychidae (Mammalia) from the Eureka Sound Group, Ellesmere Island, Nunavut". Canadian Journal of Earth Sciences. 39 (6): 899–910. Bibcode:2002CaJES..39..899E. doi:10.1139/e02-001. ISSN 0008-4077.
  27. 1 2 Rose, Kenneth D. (2006). "Cete and Artiodactyla". The beginning of the Age of Mammals. Baltimore: Johns Hopkins University Press. ISBN 978-0-8018-9221-9.
  28. ↑ Ting, Suyin; Li, Chuankuei (1987). "The skull of Hapalodectes (?Acreodi, Mammalia), with notes on some Chinese Paleocene mesonychids" (PDF). Vertebrata PalAsiatica (in Chinese and English). 25: 161–186. Archived (PDF) from the original on 2020-11-02.
  29. 1 2 Zhou, Xiaoyuan; Zhai, Renjie; Gingerich, Philip D.; Chen, Liezu (1995). "Skull of a New Mesonychid (Mammalia, Mesonychia) from the Late Paleocene of China". Journal of Vertebrate Paleontology. 15 (2): 387–400. doi:10.1080/02724634.1995.10011237.
  30. 1 2 3 tetrapodzoology (15 August 2009). "Mesonyx and the other mesonychid mesonychians (mesonychians part IV)". scienceblogs.com. Retrieved 2019-05-16.
  31. ↑ Scott, W. B. (1894). "A Revision of the North American Creodonta with Notes on Some Genera Which Have Been Referred to That Group". Proceedings of the Academy of Natural Sciences of Philadelphia. 44: 291–323.
  32. ↑ Osborn, Henry F.; Matthew, William D. (1909). "Cenozoic mammal horizons of western North America, with faunal lists of the Tertiary Mammalia of the West". US Geol Surv Bull. 361: 91–120. doi:10.3133/b361.
  33. ↑ Matthew, W.D. (1937). "Paleocene faunas of the San Juan Basin, New Mexico". Transactions of the American Philosophical Society. 30 (3532). American Philosophical Society: 1–510. Bibcode:1937Natur.140...46C. doi:10.2307/1005521. JSTOR 1005521.
  34. ↑ Szalay, Frederick S. (1969). "Origin and Evolution of Function of the Mesonychid Condylarth Feeding Mechanism". Evolution. 23 (4): 703–720. doi:10.1111/j.1558-5646.1969.tb03552.x. JSTOR 2406864.
  35. ↑ Rodolphe Tabuce, Julien Clavel and Miguel Telles Antunes (2011). "A structural intermediate between triisodontids and mesonychians (Mammalia, Acreodi) from the earliest Eocene of Portugal". Naturwissenschaften. 98 (2): 145–155. Bibcode:2011NW.....98..145T. doi:10.1007/s00114-010-0747-y. PMID 21181109. S2CID 22526630.
  36. ↑ Sarah L. Shelley, Thomas E. Williamson, Stephen L. Brusatte, Resolving the higher-level phylogenetic relationships of “Triisodontidae” (‘Condylarthra’) within Placentalia, October 2015, Society of Vertebrate Paleontology (abstract)
  37. ↑ Geisler, Jonathan H.; Uhen, Mark D. (2003). "Morphological support for a close relationship between hippos and whales". Journal of Vertebrate Paleontology. 23 (4): 991–6. doi:10.1671/32. JSTOR 4524409. S2CID 59143599.
  38. ↑ Geisler, Jonathan H.; Uhen, Mark D. (2005). "Phylogenetic Relationships of Extinct Cetartiodactyls: Results of Simultaneous Analyses of Molecular, Morphological, and Stratigraphic Data". Journal of Mammalian Evolution. 12 (1–2): 145–60. doi:10.1007/s10914-005-4963-8. S2CID 34683201.
  39. ↑ Boisserie, J.-R.; Lihoreau, F.; Brunet, M. (2005). "The position of Hippopotamidae within Cetartiodactyla". Proceedings of the National Academy of Sciences. 102 (5): 1537–41. Bibcode:2005PNAS..102.1537B. doi:10.1073/pnas.0409518102. JSTOR 3374466. PMC 547867. PMID 15677331.
  40. ↑ Gatesy, J.; Hayashi, C.; Cronin, M. A.; Arctander, P. (1996). "Evidence from milk casein genes that cetaceans are close relatives of hippopotamid artiodactyls". Molecular Biology and Evolution. 13 (7): 954–63. doi:10.1093/oxfordjournals.molbev.a025663. PMID 8752004.
  41. 1 2 3 Thewissen, J. G. M.; Cooper, Lisa Noelle; Clementz, Mark T.; Bajpai, Sunil; Tiwari, B. N. (2009). "Thewissen et al. Reply". Nature. 458 (7236): E5. Bibcode:2009Natur.458....5T. doi:10.1038/nature07775. S2CID 4431497.
  42. ↑ O'Leary, Maureen A.; Gatesy, John (2008). "Impact of increased character sampling on the phylogeny of Cetartiodactyla (Mammalia): Combined analysis including fossils". Cladistics. 24 (4): 397–442. doi:10.1111/j.1096-0031.2007.00187.x. PMID 34879630. S2CID 85141801.
  43. ↑ Toosey, William J.; Williamson, Thomas E.; Shelley, Sarah L.; Brusatte, Stephen L. (2024-11-11). "The osteology of Triisodon crassicuspis (Cope, 1882): New insights into the enigmatic "archaic" placental mammal group "Triisodontidae"". PLOS ONE. 19 (11) e0311187. doi:10.1371/journal.pone.0311187. ISSN 1932-6203. PMC 11554371.
  44. 1 2 Solé, Floréal; Bast, Eric D.; Yang, Jian; Li, Cheng-Sen; Smith, Thierry (2017). "The first species of Hapalodectes (Mesonychia, Mammalia) from the middle Paleocene of China (Qianshan Basin, Anhui Province) sheds light on the initial radiation of hapalodectids". Paleontology. 60 (3): 433–449. doi:10.5061/dryad.v4q0r.
  45. 1 2 Solé, Floréal; Fischer, Valentin; Le Verger, Kévin; Mennecart, Bastien; Speijer, Robert P.; Peigné, Stéphane; Smith, Thierry (2022). "Evolution of European carnivorous mammal assemblages through the Paleogene". Biological Journal of the Linnean Society. 135 (4): 734–753. doi:10.1093/biolinnean/blac002.
  46. ↑ Solé, Floréal; Mennecart, Bastien (2019). "A large hyaenodont from the Lutetian of Switzerland expands the body mass range of the European mammalian predators during the Eocene". Acta Palaeontologica Polonica. 64 (2). doi:10.4202/app.00581.2018. Retrieved 10 September 2024.
  47. ↑ Solé, Floréal; Falconnet, Jocelyn; Yves, Laurent (24 July 2014). "New proviverrines (Hyaenodontida) from the early Eocene of Europe; phylogeny and ecological evolution of the Proviverrinae: Evolution of the Proviverrinae". Zoological Journal of the Linnean Society. 171 (4): 878–917. doi:10.1111/zoj.12155. Retrieved 7 November 2024 – via Wiley Online Library.
  48. ↑ Solé, Floreal; Morlo, M.; Schaal, T.; Lehmann, T. (2021). "New hyaenodonts (Mammalia) from the late Ypresian locality of Prémontré (France) support a radiation of the hyaenodonts in Europe already at the end of the early Eocene". Geobios. 66–67: 119–141. Bibcode:2021Geobi..66..119S. doi:10.1016/j.geobios.2021.02.004. ISSN 0016-6995. S2CID 234848856.
  49. 1 2 3 O'Leary, Maureen A.; Rose, Kenneth D. (1995). "Postcranial Skeleton of the Early Eocene Mesonychid Pachyaena (Mammalia: Mesonychia)". Journal of Vertebrate Paleontology. 15 (2): 401–430. doi:10.1080/02724634.1995.10011238. ISSN 0272-4634. JSTOR 4523639.
  50. 1 2 Zhou, X.; Sanders, W. J.; Gingerich, P. D. (1992). "Functional and Behavioral Implications of Vertebral Structure in Pachyaena ossifraga (Mammalia, Mesonychia)" (PDF). Contributions from the Museum of Paleontology the University of Michigan. 28: 289–319.
  51. ↑ O'Leary, Maureen A.; Lucas, Spencer G.; Williamson, Thomas E. (2000-06-27). "A new specimen of Ankalagon (Mammalia, Mesonychia) and evidence of sexual dimorphism in mesonychians". Journal of Vertebrate Paleontology. 20 (2): 387–393. doi:10.1671/0272-4634(2000)020[0387:ANSOAM]2.0.CO;2. ISSN 0272-4634.
  52. ↑ Van, Valkenburgh B. (1999). "Major patterns in the history of carnivorous mammals". Annual Review of Earth and Planetary Sciences. 27: 463–493. Bibcode:1999AREPS..27..463V. doi:10.1146/annurev.earth.27.1.463.
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