Hyaenodonta

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Hyaenodonta
Temporal range: Early Paleocene to Late Miocene
61.6–10  Ma
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(Suspected Late Cretaceous origin, but unconfirmed by fossils yet) [1] [2]
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Mirorder: Ferae
Clade: Pan-Carnivora
Order:† Hyaenodonta
Van Valen, 1967 [3]
Subgroups
[see classification]
Synonyms
  • Hyaenodontida (Solé, 2010) [4]
  • Hyaenodontidae (Leidy, 1869)
  • Proviverroidea (Morlo, 2009) [5]

Hyaenodonta ("hyena teeth") is an extinct order of mostly carnivorous placental mammals of clade Pan-Carnivora from mirorder Ferae. [6] [7] Hyaenodonts were important mammalian predators that are believed to have arose either in the Late Cretaceous [8] [9] or Early Paleocene [10] within Europe, and persisted well into the Late Miocene. [11] [12] Hyaenodonts were found across Africa, Eurasia, and North America throughout the early Cenozoic. They occupied a variety of ecosystems and displayed a variety of body shapes, diet, and sizes. Some hyaenodonts, such as Hyaenodon , were cursorial predators.

Contents

The order currently consists of three superfamilies, Hyaenodontoidae, Hyainailouroidea, and Limnocyonoidea, in addition to smaller groups such as sinopines and the Galecyon clade. The hyaenodontoids included Hyaenodontidae and Proviverrinae, while hyainailouroids included Hyainailouridae, Prionogalidae, and Teratodontidae, and limnocyonoids consisted of Limnocyon and their close relatives.

Phylogeny

Classification

Hyaenodonts were considerably more widespread and successful than the oxyaenids, the other clade of mammals originally classified along with the hyaenodonts as part of Creodonta. [13] In 2015 phylogenetic analysis of Paleogene mammals, by Halliday et al., monophyly of Creodonta was supported and was placed in the clade Ferae, closer to Pholidota than to Carnivora. [14] However, order Creodonta is now considered to be a polyphyletic wastebasket taxon containing two unrelated clades assumed to be closely related (or ancestral) to Carnivora. With Hyaenodonta and Oxyaenodonta now being considered their own distinct orders. [15] [16] [17] [18] [19] [20] [21] [22] [23] [24] [25]

Taxonomy

Characteristics

Size

They generally ranged in size from 30–140 cm (1 ft 0 in – 4 ft 7 in) at the shoulder. [13] It was estimated that Simbakubwa kutokaafrika may have weighed up to 1,500 kg (3,300 lb), with Megistotherium estimated to have reached an even larger size of 1,794–3,002 kg (3,955–6,618 lb). [10] However estimate is suspect due to being based on skull-body size ratios derived from felids, which have much smaller skulls for their body size. Additionally, it was estimated that the maximum size a terrestrial mammalian carnivores could reach was 1,100 kg (2,400 lb), further suggesting the higher body mass estimates for Megistotherium was unlikely. [26]

Other large hyaenodonts include the hyainailourine Hyainailouros and the much earlier-living Hyaenodon gigas and Hyaenodon mongoliensis (The largest species of the genus, Hyaenodon), both species had a skull length of 60 cm (2.0 ft) and are estimated to be similar in size to Hyainailouros. [27] Most hyaenodonts, however, were in the 5–15 kg (11–33 lb) range, equivalent to a mid-sized dog. [28]

Skull and dentition

Skull of Hyaenodon horridus Hyaenodon horridus skull.jpg
Skull of Hyaenodon horridus
Comparison of carnassial teeth of a wolf, Hyaenodon, and Oxyaena Comparison of Carnivoran and Creodont Carnassials.png
Comparison of carnassial teeth of a wolf, Hyaenodon, and Oxyaena

Compared to carnivorans, who have one pair of carnassial teeth, hyaenodonts possessed as many as three sequential pairs of carnassials or carnassial-like molar teeth in their jaws. Their skulls are known to high have sagittal crests which supported attachment for powerful temporalis muscles. [29] Hyaenodonts, like all "creodonts", lacked post-carnassial crushing molar teeth, such as those found in many carnivoran families, especially the Canidae and Ursidae. [29]

The anatomy of their skulls show that they had a particularly acute sense of smell, while their teeth were adapted for shearing, rather than crushing. [13] Hyaenodonts differed from Carnivora in that they replaced their deciduous dentition slower in development than carnivorans. [16] [17] Studies on Hyaenodon show that juveniles took 3 to 4 years in the last stage of tooth eruption, implying a very long adolescent phase. In North American forms, the first upper premolar erupts before the first upper molar, while European forms show an earlier eruption of the first upper molar. [17]

Postcranial anatomy

Hyaenodonts were ancestrally plantigrade, [30] but the later, larger forms were generally digitigrade or semidigitigrade. [10] [30] At least one hyaenodont lineage, subfamily Apterodontinae, was specialized for aquatic, otter-like habits. [18] Hyainailouros had a humerus as robust as an average felid. Compared to the humerus of a tiger, the deltoid scar of Hyainailouros was more distally than tigers. [31]

Diet

While many hyaenodonts were hypercarnivorous predators, [10] [32] [33] some hyaenodonts, such as Anasinopa , Limnocyon , and Pakakali , were found to have been mesocarnivores or omnivorous. [32] [34] Lesmesodon was found to have been an insectivorous based on chewing cycle analysis. [35] Despite this, hyaenodonts weren't as versatile in their diet as carnivorans. [36] [29]

Evolution

Hyaenodonts are believed to have evolved in Europe either during the Late Cretaceous or Early Paleocene. [8] [1] [10] Hyaenodonts soon dispersed into Africa and India, implying close biogeographical connections between these areas. [18] [19] This dispersal later resulted in a between the evolution of the Afro-Arabia clade. [8] Afterwards, hyaenodonts dispersed into Asia from either Europe or India, and finally, North America from either Europe or Asia. [37] [38] During the middle to late Eocene, hyaenodonts, along with carnivorans, would replace oxyaenids, mesonychid, and miacoids. [39]

However, the order itself, saw a massive decline globally at the end of the Eocene to early Oligocene. [33] [40] [39] [41] However, in Afro-Arabia, the hyainailouroids experienced a rebound in diversity from the late Oligocene to the early Miocene. [41]

Several Miocene representatives of this order, include hyainailourids Megistotherium , Simbakubwa , Hyainailouros , Sectisodon , Exiguodon , Sivapterodon , Metapterodon , and Isohyaenodon , the prionogalid Prionogale , the teratodontid Dissopsalis and the youngest species of the genus Hyaenodon , Hyaenodon weilini. [11] [12]

Extinction

In North America, hyaenodonts experienced a terminal decline from the Eocene to Oligocene, with Hyaenodon being the only genus present in North America before going extinct in the Oligocene. [40] [33] In Afro-Arabia, hyainailouroids experienced a massive decline with only 5 lineages persisting into the early Oligocene. [41] By the late Oligocene, hyaenodontoids went extinct in Europe, with their last occurrence being dating to MP30. [42] After a period of diversification, hyainailouroids experienced a permanent decline during the early Miocene onwards. [43] [41] By the late Miocene, the order was represented by Dissopsalis , Hyainailouros , and Metapterodon , with Dissopsalis, the last hyaenodont, going extinct around 10 Ma. [12]

Many experts argued that the decline and extinction of the hyaenodonts was due to competition with the carnivorans. [10] [39] [40] Lang and colleagues found that the evolutionary success of carnivorans compared to hyaenodonts may have been largely influenced by the retention of a basal morphotype throughout their evolutionary history. The authors also suggested that carnivorans likely contributed in some way to the extinction of hyaenodonts, with the difference in functional morphology and adaptive potential of their carnassials possibly being a factor. [36]

In North America, Friscia and Valkenburgh argued that carnivorans may have limited the evolution of “creodonts”, resulting with only the large, hypercarnivorous ecomorphs remaining. [40] In addition, Serio and colleagues found that North American “creodonts” had a significant degree of morphological differentiation until the middle Eocene, around this time disparity among carnivorans increasing around the same time. The authors argue that carnivoran disparity negatively impacted the disparity of creodonts, suggesting carnivorans may have outcompeted hyaenodonts. [44]

Within Africa, experts had hypothesized that hyaenodonts suffered from competitive displacement, as the invading carnivorans forced hyainailourids to tend towards carnivory. In support of this hypothesis, as even some of the smallest Miocene hyainailourids, Isohyaenodon and Mlanyama , showed adaptations towards hypercarnivory compared to older, mesocarnivorous hyaenodonts such as Boualitomus and Pakakali . [32] [45] Borths and Stevens, in their 2019 paper, argued that cooperative carnivorans had larger and more complex brains, suggesting this enabled them to steal carcasses from large, solitary hyainailourines. [10] In Europe, the decline of the proviverrines coincided with the arrival of carnivorans, hyaenodontines, and hyainailourines. [46] [30]

However, this hypothesis has been contested by experts. [33] [43] [47] Among large hyainailourids, the discovery of Simbakubwa suggests the evolution of large hyainailourines was thought to have been changes in the herbivore fauna instead of competition with carnivorans. [10] Additionally, studies have found brain size have little to no correlation with sociality in carnivorans, [48] [49] [50] with the relative size of the anterior brain having a larger correlation to sociality than overall brain size. [51] [52] [53] Morales and colleagues argued the decline of hyainailourids was due to the increase in aridity, as they were more adapted for forested environments as opposed to savannas, steppes, or deserts. [43] The decline of hyainailourids in Afro-Arabia from the late Eocene to early Oligocene was due to the decline of their preferred prey such as anthropoids and hystricognaths. It was also linked to the uplifting of the Ethiopian plateau, volcanic super eruptions in Ethiopia, the outpouring of flood basalts of Arabian Large Igneous Province, as well as the opening of the Red Sea and Gulf of Aden. [41]

Christison and colleagues analyzed hyaenodonts and carnivorans from Calf Creek locality of Cypress Hills Formation and found that only the smallest hyaenodont, Hyaenodonmicrodon, and carnivorans, had significant niche overlap. On the other hand, larger hyaenodonts, Hemipsalodon grandis and Hyaenodon horridus, and carnivorans, Hoplophoenus , had very distinct niches. They argue this would suggest that competitive replacement wasn't the driving factor the decline and extinction of North American hyaenodonts. Instead, global climatic cooling during the earliest Oligocene resulted in drier, more open landscapes and resulted in the extinction of large browsing herbivores, including brontotheres, who were replaced by grazing mammals such as equids and rhinoceroses, which were better suited for the open environments. Because rhinoceroses didn't reach their large sizes until the Miocene, hyaenodonts were at a disadvantage due to the lack of large prey available. In addition, because of their shorter legs, they were likely at a disadvantage in the increasingly open environments. [33] Despite recovering North American hyaenodonts as a relatively cursorial clade, Castellanos also hypothesized due their short distal limbs, hyaenodonts had less cursorial adaptations than contemporary amphicyonids and couldn't exploit open environments as well which likely led to their extinction in addition to the extinction of large browsing prey. [47]

In Europe, the late Eocene to early Oligocene extinction of hyaenodonts, such as Pterodon , was thought to have been the result of Grande Coupure, a faunal turnover that was contemporary with the Eocene-Oligocene extinction. [54] [55] Grande Coupure saw an extinction rate of 60% of western European mammalian lineages. [56] [57] [58] The event correlates with abrupt shift towards a icehouse world, which stems from the expansion of the Antarctic ice sheets and saw sea level decline by 70 m (230 ft). [59]

Additionally, Hyaenodon, who persisted the Grande Coupure, and amphicyonids were able to coexist by preferring different habitats, with Hyaenodon preferring more open environments compared to amphicyonids, resulting in niche partitioning between the two predators. [42] Instead, the extinction of European hyaenodonts in the Oligocene is believed to have been caused by climatic changes rather than competition with carnivorans. [45] [42]

See also

References

  1. 1 2 Borths, Matthew R.; Holroyd, Patricia A.; Seiffert, Erik R. (2016). "Hyainailourine and teratodontine cranial material from the late Eocene of Egypt and the application of parsimony and Bayesian methods to the phylogeny and biogeography of Hyaenodonta (Placentalia, Mammalia)". PeerJ. 4 e2639. doi: 10.7717/peerj.2639 . PMC   5111901 . PMID   27867761.
  2. ↑ Borths, Matthew R.; Stevens, Nancy J. (2017). "The first hyaenodont from the late Oligocene Nsungwe Formation of Tanzania: Paleoecological insights into the Paleogene-Neogene carnivore transition". PLOS ONE. 12 (10) e0185301. Bibcode:2017PLoSO..1285301B. doi: 10.1371/journal.pone.0185301 . PMC   5636082 . PMID   29020030.
  3. ↑ Van Valen, L. (1967). "New Paleocene insectivores and insectivore classification." Bulletin of the American Museum of Natural History 135(5):217-284
  4. ↑ Solé F. (2010) "Les premiers placentaires carnassiers européens (Oxyaenodonta, Hyaenodontida et Carnivora): origine, évolution, paléoécologie et paléobiogéographie; apport des faunes de l'Eocène inférieur du Bassin de Paris." Paris: Muséum National d'Histoire Naturelle. 703 p.
  5. ↑ Morlo, M., Gunnell, G. F. and Polly, P. D. (2009). "What, if not nothing, is a creodont? Phylogeny and classification of Hyaenodontida and other former creodonts." Journal of Vertebrate Paleontology Program and Abstracts, 2009:152A.
  6. ↑ Solé, F.; Lhuillier, J.; Adaci, M.; Bensalah, M.; Mahboubi, M.; Tabuce, R. (16 July 2013). "The hyaenodontidans from the Gour Lazib area (?Early Eocene, Algeria): implications concerning the systematics and the origin of the Hyainailourinae and Teratodontinae". Journal of Systematic Palaeontology. 12 (3): 303–322. doi:10.1080/14772019.2013.795196. S2CID   84475034.
  7. ↑ Solé, F.; Amson, E.; Borths, M.; Vidalenc, D.; Morlo, M.; Bastl, K. (23 September 2015). "A New Large Hyainailourine from the Bartonian of Europe and Its Bearings on the Evolution and Ecology of Massive Hyaenodonts (Mammalia)". PLOS ONE. 10 (9) e0135698. Bibcode:2015PLoSO..1035698S. doi: 10.1371/journal.pone.0135698 . PMC   4580617 . PMID   26398622.
  8. 1 2 3 Borths, Matthew R.; Stevens, Nancy J. (2017). "The first hyaenodont from the late Oligocene Nsungwe Formation of Tanzania: Paleoecological insights into the Paleogene-Neogene carnivore transition". PLOS ONE. 12 (10) e0185301. Bibcode:2017PLoSO..1285301B. doi: 10.1371/journal.pone.0185301 . PMC   5636082 . PMID   29020030.
  9. ↑ Borths, Matthew R.; Holroyd, Patricia A.; Seiffert, Erik R. (2016). "Hyainailourine and teratodontine cranial material from the late Eocene of Egypt and the application of parsimony and Bayesian methods to the phylogeny and biogeography of Hyaenodonta (Placentalia, Mammalia)". PeerJ. 4 e2639. doi: 10.7717/peerj.2639 . PMC   5111901 . PMID   27867761.
  10. 1 2 3 4 5 6 7 8 Borths, M. R.; Stevens, N. J. (2019). "Simbakubwa kutokaafrika, gen. et sp. nov. (Hyainailourinae, Hyaenodonta, 'Creodonta,' Mammalia), a gigantic carnivore from the earliest Miocene of Kenya" . Journal of Vertebrate Paleontology. 39 (1) e1570222. Bibcode:2019JVPal..39E0222B. doi:10.1080/02724634.2019.1570222. S2CID   145972918.
  11. 1 2 Barry, J. C. (1988). "Dissopsalis, a middle and late Miocene proviverrine creodont (Mammalia) from Pakistan and Kenya". Journal of Vertebrate Paleontology. 48 (1): 25–45. Bibcode:1988JVPal...8...25B. doi:10.1080/02724634.1988.10011682.
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  16. 1 2 Borths, Matthew R.; Stevens, Nancy J. (2017). "Deciduous dentition and dental eruption of Hyainailouroidea (Hyaenodonta, "Creodonta," Placentalia, Mammalia)". Palaeontologia Electronica. 20 (3): 55A. Bibcode:2017PalEl..20..776B. doi: 10.26879/776 .
  17. 1 2 3 Bastl, Katharina Anna (2013). "First evidence of the tooth eruption sequence of the upper jaw in Hyaenodon (Hyaenodontidae, Mammalia) and new information on the ontogenetic development of its dentition". Paläontologische Zeitschrift. 88 (4): 481–494. doi:10.1007/s12542-013-0207-z. S2CID   85304920.
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  22. ↑ Polly, P. D. (1996). "The skeleton of Gazinocyon vulpeculus gen. et comb. nov. and the cladisitic relationships of Hyaenodontidae (Eutheria, Mammalia)". Journal of Vertebrate Paleontology. 16 (2): 303–319. Bibcode:1996JVPal..16..303P. doi:10.1080/02724634.1996.10011318. S2CID   84853339.
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  25. ↑ Solé, Floréal; Marandat, Bernard; Lihoreau, Fabrice (2020). "The hyaenodonts (Mammalia) from the French locality of Aumelas (Hérault), with possible new representatives from the late Ypresian". Geodiversitas. 42 (13): 185–214. Bibcode:2020Geodv..42..185S. doi: 10.5252/geodiversitas2020v42a13 .
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  32. 1 2 3 Borths, Matthew R.; Stevens, Nancy J. (2017). "The first hyaenodont from the late Oligocene Nsungwe Formation of Tanzania: paleoecological insights into the Paleogene-Neogene carnivore transition". PLOS ONE. 12 (10) e0185301. Bibcode:2017PLoSO..1285301B. doi: 10.1371/journal.pone.0185301 . PMC   5636082 . PMID   29020030.
  33. 1 2 3 4 5 Christison, Brigid E; Gaidies, Fred; Pineda-Munoz, Silvia; Evans, Alistair R; Gilbert, Marisa A; Fraser, Danielle (25 January 2022). Powell, Roger (ed.). "Dietary niches of creodonts and carnivorans of the late Eocene Cypress Hills Formation". Journal of Mammalogy . 103 (1): 2–17. doi:10.1093/jmammal/gyab123. ISSN   0022-2372. PMC   8789764 . PMID   35087328 . Retrieved 17 October 2024– via Oxford Academic.
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  35. ↑ Weidtke, Jannik; Schultz, Julia A.; Morlo, Michael; et al. (October 2025). "Chewing cycle analysis of Lesmesodon edingeri (Mammalia; Hyaenodonta) and implications for carnivoran feeding adaptions". Palaeontologia Electronica. 28 (3): 1–35.
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  38. ↑ ""New Remains of Hyaenodontidae (Creodonta, Mammalia) From the Oligocene of Central Mongolia"" (PDF). Archived from the original (PDF) on 8 May 2010. Retrieved 4 May 2008.
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  40. 1 2 3 4 Friscia, Anthony; Van, Valkenburgh B. (2010). "Ecomorphology of North American Eocene carnivores: evidence for competition between Carnivorans and Creodonts". Carnivoran Evolution. Cambridge University Press. pp. 311–341. doi:10.1017/CBO9781139193436.012. ISBN   978-0-521-51529-0.
  41. 1 2 3 4 5 de Vries, Dorien; Heritage, Steven; Borths, Matthew R.; Sallam, Hesham M.; Seiffert, Erik R. "Widespread loss of mammalian lineage and dietary diversity in the early Oligocene of Afro-Arabia". Communications Biology. 4. doi: 10.1038/s42003-021-02707-9 .
  42. 1 2 3 Solé; Dubied; Le Verger; Mennecart (2018). "Niche partitioning of the European carnivorous mammals during the paleogene" . PALAIOS. 33 (11): 514–523. Bibcode:2018Palai..33..514S. doi:10.2110/palo.2018.022.
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  44. ↑ Serio, Carmela; Brown, Richard P.; Clauss, Marcus; Meloro, Carlo (2024). "Morphological disparity of mammalian limb bones throughout the Cenozoic: the role of biotic and abiotic factors". Palaeontology. 67 (4) e12720. Bibcode:2024Palgy..6712720S. doi: 10.1111/pala.12720 .
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  46. ↑ Lange-Badré, Brigitte (1979). "Les Creodontes (Mammalia) d'Europe occidentale de l'Eocene superieur a l'Oligocene superieur". Mémories du Muséum National d'Historie Naturelle, Série C, Sciences de la Terre. 42: 1–249.
  47. 1 2 Castellanos, Miguel (2025). "Hunting types in North American Eocene–Oligocene carnivores and implications for the 'cat-gap'" . Journal of Mammalian Evolution. 32 (2): 1–12. doi:10.1007/s10914-025-09767-2.
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  56. ↑ Hooker, Jerry J.; Collinson, Margaret E.; Sille, Nicholas P. (2004). "Eocene–Oligocene mammalian faunal turnover in the Hampshire Basin, UK: calibration to the global time scale and the major cooling event" (PDF). Journal of the Geological Society. 161 (2): 161–172. Bibcode:2004JGSoc.161..161H. doi:10.1144/0016-764903-091. S2CID   140576090. Archived (PDF) from the original on 8 August 2023. Retrieved 31 August 2023.
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