Hyaenodonta

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Hyaenodonta
Hyaenodon, drawing by Heinrich Harder

Hyaenodon , drawing by Heinrich Harder

Temporal occurrence
Middle Paleocene ( Zealand ) to Middle Miocene ( Serravallian )
61.1 to 11.1 million years
Locations
Systematics
Synapsids (Synapsida)
Mammals (mammalia)
Higher mammals (Eutheria)
Laurasiatheria
Ferae
Hyaenodonta
Scientific name
Hyaenodonta
Van Valen , 1967

The Hyaenodonta (rarely also Hyaenodontida) are an extinct group of carnivorous mammals that were found in North America , Europe , Asia and Africa . They appeared with Lahimia in the mid Paleocene and died out in North America and Europe with the end of the Oligocene , in Africa and Asia during the Miocene .

features

The representatives of the Hyaenodonta had an external resemblance to today's dogs and hyenas . They had a plantigrade or completely digitigrade (v. Latin digitus "toe") foot ( toe walker ) and, like today's dogs and hyenas, were partially adapted to running fast. The main axis of the foot was on the middle toe. The fibula was steered with the calcaneus , an articulation of the talus with the cuboid bone was missing. The phalanges of the toes were notched. Early forms were still small, Lesmesodon , whose fossils were found in the Messel Pit , was only 20 cm long, Arfia was already as big as a fox . The largest forms of the Hyaenodonta, Hyaenodon , Hyainailouros, and Megistotherium had skulls twice the size of today's lions. In general, the skull was characterized by a long, narrow structure, a narrow base and a high occiput .

Their jaws housed a scissor bite , which was formed by the entire row of molars, with an emphasis on the second molar in the upper jaw and the third molar in the lower jaw. In the possibly related Oxyaenida , the first molar in the upper jaw and the second in the lower jaw were of greater importance, in recent predators the fourth premolar in the upper jaw and the first molar in the lower jaw.

External system

The Hyaenodonta were originally (as Hyaenodontidae) together with the cat- or marten- like Oxyaenodonta in the order of the Creodonta , also called original predators in German. The Oxyaenodonta appeared in the fossil record in the middle Paleocene and died out earlier than the Hyaenodonta. Both groups hardly share synapomorphies , so that the validity of the Creodonta taxon is questioned. Along with the recent predators (Carnivora) and the shed animals (Manidae) form the Hyaenodonta and Oxyaenodonta the taxon ferae .

Internal system

General

Internal systematics of the Hyaenodonta according to Solé & Mennecart 2019
  Hyaenodonta  


 " Sinopa Clade"


   

 Limnocyoninae


   

 Arfiinae




   
  Hyaenodontoidea  
  Hyaenodontidae  


 Prionogalidae


   

 Hyaenodontinae



   

 " Cynohyaenodon clade"



   

 Proviverrinae



   

 " Galecyon Clade"


  Hyainailouroidea  

 Teratodontinae


  Hyainailouridae  

 Apterodontinae


   

 Hyainailourinae







Template: Klade / Maintenance / Style

The structure of the hyaenodonta is currently not fully clarified and is under discussion. In general, two larger taxonomic units can be distinguished: the Hyaenodontidae and the Hyainailouridae . Both groups can be relatively easily separated from one another on the basis of skeletal anatomical features. The Hyaenodontidae have a postorbital process , which the Hyainailouridae usually lack. On the occiput, the prominent bony bulge as the attachment point of the neck muscles extends laterally down to the mastoid process in the former , but not in the latter. In addition, the joint surfaces there for connection with the cervical spine are small in the Hyaenodontidae and large in the Hyainailouridae. In the Hyaenodontidae , the mental foramen on the lower jaw usually appears doubled and the angular process is extremely strong. The Hyainailouridae often have more than two foramina, and their angular process is rather weak. There are also significant differences in the teeth. The typical sectorial molars have several pointed cusps, the three largest include the para-, meta- and protoconus in the upper jaw and the para-, meta- and protoconid in the lower jaw. In both the Hyaenodontidae and the Hyainailouridae, the para and metaconus of the upper molars are fused to form the amphiconus. However, in the Hyaenodontidae the Metaconus towers over the Paraconus, in the Hyainailouridae this is exactly the opposite.

Within the Hyainailouridae, the Apterodontinae and the Hyainailourinae are distinguished from one another. Both groups differ from one another in specific tooth characteristics. The Hyainailourinae form a hypercarnivore dentition, that is, there is a decrease in the complexity of the tooth structure. As a result, the bit largely loses its perforating and breaking function and develops into a powerful cutting tool. This happens in the lower jaw, for example, through the reduction of the metaconid (perforating) and the talonid (breaking; this is a deeper tooth surface of the lower molars, into which the protoconus of the upper molars engages when the bite is closed). As a result, numerous representatives of the Hyainailouridae can be regarded as scavengers. Today's hypercarnivorous predators obtain over 70% of their food from vertebrates . The Apterodontinae also lose individual cusps (metaconid and paraconid) on the lower molars, but the talonid remains. This rather unusual tooth design is often interpreted as specializing in fish or invertebrates .

The exact position of the teratodontinae is problematic . Some researchers see them as more closely related to the Hyaenodontidae and justify this with the developed postorbital process or the strong angular process. Other scientists combine the Teratodontinae with the Hyainailouridae in a common higher group, the Hyainailouroidea. This happens, among other things, because of the less strong occipital ridge or the multiple foramina on the lower jaw. Like the Hyaenodontidae and the Hyainailouridae, the Teratodontinae also have a fused para- and metaconus in the upper jaw, whereby both are only united at the base and the tips are free. In accordance with the Hyaenodontidae, the metaconus is higher than the paraconus.

Apart from these larger groups, the Limnocyoninae are differentiated. These are relatively small representatives of the hyaenodonta, the main characteristic of which is the loss or reduction of the upper and lower third molars. The upper molars have a fused para- and metaconus, which, however, differs gradually within the individual genera. The Proviverrinae, on the other hand, combine a rather primitive trunk group of the Hyaenodonta with clearly distinguishable humps on the molars. Due to their very original characteristics, they are sometimes in their own higher group, the Proviverroidea, but a study from 2019 refers them to the close relationship of the Hyaenodontidae. The Prionogalidae were originally classified as higher mammals with imprecise relationships due to their unusual molar design. They represent dwarfish predators in which the rearmost molar has receded, while the remaining molars show extremely hypercarnivorous properties. Recent phylogenetic studies refer them to the Hyaenodonta, depending on the weighting of the characteristics, they either form a clade with the Hyaenodontidae or with the Hyainailouridae.

The systematic assignment of the saber-toothed Machaeroidinae is unclear. On the one hand they are regarded as more closely related to the Limnocyoninae, on the other hand they are also included in the Oxyaenodonta . A study published in 2018 of an almost complete skeleton from Utah suggests a close relationship with the Oxyaenodonta due to the structure of the musculoskeletal system.

Overview of the families and genera of the Hyaenodonta

The structure shown here is largely based on the studies by Borths et al. 2016 and 2017. Accordingly, the hyaenodonta are composed as follows:

  • Order Hyaenodonta Van Valen , 1967
  • Subfamily Arfiinae Solé , 2013 (also Arfianinae)
  • Family incertae sedis
  • Family incertae sedis
  • Sinopa clade
  • Cynohyaenodon clade
  • Family incertae sedis
  • Galecyon clade
  • Family incertae sedis

Several other subfamilies turned out to be problematic, such as the Koholiinae, introduced by Jean-Yves Crochet in 1988 , which bring together genera such as Koholia , Lahimia and Boualitomus . The same applies to the Sinopinae (or Sinopaninae) by Floréal Solé from 2013 around Sinopa , Prototomus , Tritemnodon and Pyrocyon as well as for the Indohyaenodontinae by Solé and colleagues in 2014 with Indohyaenodon , Paratritemnodon and Kyawdawia . Here, phylogenetic studies showed that the composition of the groups is usually not conclusive. In addition, there are numerous tribes within the Hyainailouridae , such as the Paroxyaenini, Hyainailourini, Dissopsalini and Metapterodontini. Here, too, the phylogenetic analyzes sometimes reveal different relationships.

literature

  • Thomas S. Kemp: The Origin and Evolution of Mammals. Oxford University Press, Oxford et al. 2005, ISBN 0-19-850760-7 .
  • Jordi Augusti, Mauricio Antón: Mammoths, Sabertooths, and Hominids. 65 Million Years of Mammalian Evolution in Europe. Columbia University Press, New York NY et al. 2002, ISBN 0-231-11640-3 .
  • Alan Turner, Mauricio Antón: Evolving Eden. An Illustrated Guide to the Evolution of the African Large-Mammal Fauna. Columbia University Press, New York NY 2004, ISBN 0-231-11944-5 .

Individual evidence

  1. ↑ a b c d e f Floréal Solé, Jocelyn Falconnet and Dominique Vidalenc: New fossil Hyaenodonta (Mammalia, Placentalia) from the Ypresian and Lutetian of France and the evolution of the Proviverrinae in Southern Europe. Palaeontology 58 (6), 2015, pp. 1049-1072
  2. ↑ a b Michael Morlo, Gregg Gunnell, and P. David Polly: What, if not nothing, is a creodont? Phylogeny and classification of Hyaenodontida and other former creodonts. Journal of Vertebrate Paleontology 29 (3 suppl), 2009, p. 152A
  3. ↑ a b c d e f Floréal Solé: New proviverrine genus from the Early Eocene of Europe and the first phylogeny of Late Paleocene-Middle Eocene hyaenodontidans (Mammalia). Journal of Systematic Paleontology 11, 2013, pp. 375-398
  4. ↑ a b c d Floréal Solé and Bastien Mennecart: 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, 2019 doi: 10.4202 / app.00581.2018
  5. ↑ a b c d e f g Matthew R. Borths, Patricia A. Holroyd and Erik R. Seiffert: 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, 2016, p. E2639 doi: 10.7717 / peerj.2639
  6. ↑ Floréal Solé and Sandrine Ladevèze: Evolution of the hypercarnivorous dentition in mammals (Metatheria, Eutheria) and its bearing on the development of tribosphenic molars. Evolution & Development 19 (2), pp. 56-68
  7. ^ A b c d e Matthew R. Borths and Nancy J. Stevens: The first hyaenodont from the late Oligocene Nsungwe Formation of Tanzania: Paleoecological insightsinto the Paleogene-Neogene carnivore transition. PLoSONE 12 (10), 2017, p. E0185301 doi: 10.1371 / journal.pone.0185301
  8. ^ A b Michael Morlo and Gregg F. Gunnell: Small Limnocyonines (Hyaenodontidae, Mammalia) From the Bridgerian Middle Eocene of Wyoming: Thinocyon, Prolimnocyon, And Iridodon, New Genus. Contributions from the Museum of Paleontology, University of Michigan 31 (2), 2003, pp. 43-78
  9. ^ A b Floréal Solé, Thierry Smith, Rodolphe Tabuce and Bernard Marandat: New Dental Elements of the Oldest Proviverrine Mammal from the Early Eocene of Southern France Support Possible African Origin of the Subfamily. Acta Palaeontologica Polonica 60 (3), 2015, pp. 527-538
  10. ↑ Matthew R. Borths and Nancy J. Stevens: Taxonomic affinities of the enigmatic Prionogale breviceps, early Miocene, Kenya. Historical Biology, 2017 doi: 10.1080 / 08912963.2017.1393075
  11. ↑ Jorge Morales and Martin Pickford: A reassessment of Prionogale and Namasector (Prionogalidae, Hyaenodonta, Mammalia) with descriptions of new fossils from Napak, Uganda and Koru, Kenya. Communications of the Geological Survey of Namibia 20, 2018, pp. 114-139
  12. ↑ a b c Floréal Solé, Eli Amson, Matthew Borths, Dominique Vidalenc, Michael Morlo and Katharina Bastl: 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), 2015, p. E0135698 doi: 10.1371 / journal.pone.0135698
  13. ^ Shawn P. Zack, A skeleton of a Uintan machaeroidine 'creodont' and the phylogeny of carnivorous eutherian mammals. Journal of Systematic Palaeontology, 2018 doi: 10.1080 / 14772019.2018.1466374
  14. ↑ Leigh Van Valen: New Paleocene Insectivores and Insectivore classification. Bulletin of the American Museum of Natural History 135, 1967, pp. 217-284
  15. ↑ a b c d Floréal Solé, Jocelyn Falconnet and Laurent Yves: New proviverrines (Hyaenodontida) from the early Eocene of Europe; phylogeny and ecological evolution of the Proviverrinae. Zoological Journal of the Linnean Society 171, 2014, pp. 878-917
  16. ↑ a b c d Floréal Solé, Julie Lhuillier, Mohammed Adaci, Mustapha Bensalah, Mohammed Mahboubi and Rodolphe Tabuce: The hyaenodontidans from the Gour Lazib area (? Early Eocene, Algeria): implications concerning the systematics and the origin of the Hyainailourinae and Teratodontinae . Journal of Vertebrate Paleontology 12 (3), 2014, pp. 303-322
  17. ↑ Rajendra S. Rana, Kishor Kumar, Shawn P. Zack, Floreal Solé, Kenneth D. Rose, Pieter Missiaen, Lachham Singh, Ashok Sahni and Thierry Smith: Craniodental and Postcranial Morphology of Indohyaenodon raoi from the Early Eocene of India, and Its Implications for Ecology, Phylogeny, and Biogeography of Hyaenodontid Mammals. Journal of Vertebrate Paleontology 35 (5), 2015, p. E965308 doi: 10.1080 / 02724634.2015.965308
  18. ↑ a b c d e Jorge Morales and Martin Pickford: New hyaenodonts (Ferae, Mammalia) from the Early Miocene of Napak (Uganda), Koru (Kenya) and Grillental (Namibia). Fossil Imprint 73 (3/4), 2017, pp. 332–359
  19. ^ Morgane Dubied, Floréal Solé and Bastein Mennecart: The cranium of Provierra typica (Mammalia, Hyaenodonta) and its impact on hyaenodont phylogeny and endocranial evolution. Palaeontology 2019, pp. 1–19 doi: 10.1111 / pala.12437
  20. ^ Emmanuel Gheerbrant: Les mammifères paléocènes du bassin d'Ouarzazate (Maroc). III. Adapisoriculidae et autres mammifères (Carnivora,? Creodonta, Condylarthra,? Ungulata et incertae sedis). Palaeontographica, Division A 237, 1995, pp. 39-132
  21. ↑ Michael Morlo and Jörg Habersetzer: The Hyaenodontidae (Crerodonta, Mammalia) from the lower Eocene (MP 11) of Messel (Germany) with special remarks on new x.ray methods. Courier Forschungsinstitut Senckenberg 216, 1999, pp. 31–73
  22. ↑ a b Jorge Morales, Martin Pickford and MJ Salesa: Creodonta and Carnivora from the Early Miocene of the Northern Sperrgebiet, Namibia. Memoirs of the Geological Survey of Namibia 20, 2008, pp. 291-310
  23. ^ Paul David Polly, Brigitte Lange-Badré: A new genus Eurotherium (Mammalia, Creodonta) in reference to taxonomic problems with some Eocene hyaenodontids of Eurasia. Comptes Rendus de l'Académie des Sciences Paris. Série 2 317, 1993, pp. 991-996
  24. ↑ AV Lavrov and AO Averianov: The Oldest Asiatic Hyaenodontidae (Mammalia, Creodonta) from the Early Eocene of the Southern Fergana Basin (Andarak-2 Locality). Paleontological Journal 32 (2), 1998, pp. 200-205
  25. ↑ a b Floréal Solé, Emmanuel Gheerbrant, Mbarek Amaghzaz and Baâdi Bouya: Further evidence of the African antiquity of hyaenodontid ('Creodonta', Mammalia) evolution. Zoological Journal of the Linnean Society 156 (4), 2009, pp. 827-846
  26. ↑ Emmanuel Gheerbrant, Mohamed Iarochene, Mbarek Amaghzaz and Baâdi. Bouya: Early African hyaenodontid mammals and their bearing on the origin of the Creodonta. Geological Magazine 143 (4), 2006, pp. 75-489
  27. ↑ Jean-Yves Crochet, Stéphane Peigné and Mohamed Mahboubi: Ancienneté des Carnivora (Mammalia) en Afrique. In: C. Denys, L. Granjon and A. Poulet (Eds.): African Small Mammals. Petits mammifères africains. Paris, 2001, pp. 91-100
  28. ↑ Sunil Bajpai, Vivesh V. Kapur and Hans. Thewissen: Creodont and condylarth from the Cambay Shale (Early Eocene, 55-54MA), Vastan Lignite Mine, Gujarat, Western India. Journal of the Palaeontological Society of India 54 (1), 2009, pp. 103-109
  29. ↑ Naoko Egi, Patricia A. Holroyd, Takehisa Tsubamoto, Nobuo Shigehara, Masanaru Takai, Soe Thura Tun, Aye Kung Aung and Aung Naing Soe: A new genus and species of hyaenodontid creodont from the Pondaung Formation (Eocene, Myanmar). Journal of Vertebrate Paleontology 24 (2), 2004, pp. 502-506
  30. ↑ Naoko Egi, Patricia A. Holroyd, Takehisa Tsubamoto, Aung Naing Soe, Masanaru Takai and Russell L. Ciochon: Proviverrine hyaenodontids (Creodonta: Mammalia) from the Eocene of Myanmar and a phylogenetic analysis of the proviverrines from the para-Tethys area. Journal of Systematic Palaeontology 3 (4), 2005, pp. 337-358
  31. ↑ Elwyn L. Simons and Philip D. Gingerich: New carnivorous mammals from the Oligocene of Egypt. Annals of the Geological Survey of Egypt 4, 1974, pp. 157-166
  32. ↑ Naoko Egi, Takehisa Tsubamoto and Masanaru Takai: Systematic status of Asian "Pterodon" and early evolution of hyaenaelurine hyaenodontid creodonts. Journal of Paleontology 81 (4), 2007, pp. 770-778
  33. ↑ Patricia A. Holroyd: New Pterodontinae (Creodonta: Hyaenodontidae) from the late Eocene-early Oligocene Jebel Qatrani Formation, Fayum Province, Egypt. PaleoBios 19 (2), 1999, pp. 1-18
  34. ↑ Michael Morlo, Ellen R. Miller and Ahmed El-Barkkoky: Creodonta and Carnivora from Wadi Moghra, Egypt. Journal of Vertebrate Paleontology 27 (1), 2007, pp. 145-159
  35. Jump up ↑ D. Tab Rasmussen and Mercedes Gutierrez: A Mammalian Fauna from the Late Oligocene of Northwestern Kenya. Palaeontographica Department A 288 (1-3), 2009, pp. 1-52
  36. ↑ Matthew R. Borths and Nancy J. Stevens . Simbakubwa kutokaafrika, gen et sp. nov. (Hyainailourinae, Hyaenodonta, 'Creodonta,' Mammalia), A Gigantic Carnivore from the earliest Miocene of Kenya. Journal of Vertebrate Paleontology, 2019, p. E1570222 doi: 10.1080 / 02724634.2019.1570222

Web links

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