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Tribosphenidans Temporal range: Early Cretaceous–Recent PreꞒ Ꞓ O S D C P T J K Pg N
Eomaia fossil
Kangaroo with her joey
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Clade:	Zatheria
Clade:	Tribosphenida McKenna, 1975
Clades
†Henosferidae? †Ambondro? Monotremaformes? †Ausktribosphenidae? †Bishopidae? Boreosphenida? Luo et al., 2001 †Aegialodontia †Eomaia †Hypomylos †Tribotherium Theria

Tribosphenida is a group (infralegion) of mammals that includes the ancestor of Hypomylos, Aegialodontia and Theria (the last common ancestor of marsupials and placentals plus all of its descendants). It belongs to the group Zatheria. The current definition of Tribosphenida is more or less synonymous with Boreosphenida.

Characteristics

Tribosphenid mammals were originally grouped on the basis of triangular or V-shaped (tribosphenic) molars. Since then, other unrelated mammal groups have been found to have tribosphenic molars, such as the australosphenidans (a group that includes the still extant monotremes), suggesting that as a synapomorphy this is fundamentally useless as it evolved multiple times among mammals.

However, a clade between the aforementioned groups, the "true Tribosphenida" or Boreosphenida, is still identifiable, united by characteristics such as the lack of a mesial cingulid and of a triangulated trigonid on the last premolar. They are also united by postcranial features such as the presence of a modern ear (though this too has evolved independently in many other groups, like monotremes), modern shoulder blades, and several features of the hindlimb.

Phylogeny

Below is a cladogram showing one hypothesis of mammal relationships based on Rowe (1988) and McKenna and Bell (1997):

Boreosphenida

Boreosphenida (from boreas, "northern wind" and sphen, "wedge") were early mammals that originated in the Northern Hemisphere and had tribosphenic molars (three-cusped cheek teeth). In boreosphenidans, the mandibular angle is placed posteriorly and the primitive postdentary trough (hole in the mandible) is absent (in contrast to Kuehneotheriidae, Eupantotheria, and Australosphenida.) They share the tribosphenic molars with the Australosphenida but differ from them by having cingulid cuspules but lacking a continuous mesial cingulid. Boreosphenidans also lack the triangulated trigonid on the last premolar found in Early Cretaceous mammals. They differ from Shuotherium (a monotreme-relative) in having the talonid placed posterior to the trigonid (like in modern tribosphenic mammals) in the lower molars, but upper molars similar to those of Shuotherium.

The oldest boreosphenidans are from the Berriasian (~145-140 mya). They were restricted to the Northern Hemisphere during the Early Cretaceous, but spread to South America and India during the end of the Cretaceous.

An alternative hypothesis is offered by Flannery and colleagues, who view part of australosphenidans as members of Tribosphenida (excluding monotremes and others australosphenidans with Steropodon, which they consider to be an unrelated group), with Barremian-Aptian Australian Bisphopidae as the sister taxon to all Northern Hemisphere Theria (Boreosphenida).

On the other hand, the study that cites Flannery's article, given in 2024, maintains Ambondro as a relative of monotremata. Implying support for the Australosphenida-Boreosphenida hypothesis. It may also be that both studies are right, consequently Tribosphenida would be a monophyletic group that contains the crown group Mammalia. Australosphenida possible paraphyly.

References

1. Panciroli E; Roger B.J. Benson; Richard J. Butler (2018). "New partial dentaries of amphitheriid mammalian Palaeoxonodon ooliticus from Scotland, and posterior dentary morphology in early cladotherians". Acta Palaeontologica Polonica. 63 (2). doi:10.4202/app.00434.2017.{{cite journal}}: CS1 maint: multiple names: authors list (link)
2. ^ Kielan-Jaworowska, Zofia; Cifelli, Richard L; Luo, Zhe-Xi (2004). Mammals from the Age of Dinosaurs: Origins, Evolution, and Structure. New York: Columbia University Press. ISBN 978-0-231-11918-4.
3. ^ Luo, Zhe-Xi; Cifelli, Richard L.; Kielan-Jaworowska, Zofia (2001). "Dual origin of tribosphenic mammals". Nature. 409 (6816): 53–57. Bibcode:2001Natur.409...53L. doi:10.1038/35051023. PMID 11343108. S2CID 4342585.
4. Rich, T. H.; Hopson, J. A.; Musser, A. M.; Flannery, T. F.; Vickers-Rich, P. (2005). "Independent origins of middle ear bones in monotremes and therians". Science. 307 (5711): 910–914. Bibcode:2005Sci...307..910R. doi:10.1126/science.1105717. PMID 15705848. S2CID 3048437.
5. Rowe, Timothy (1988). "Definition, diagnosis, and origin of Mammalia". Journal of Vertebrate Paleontology. 8 (3): 241–264. Bibcode:1988JVPal...8..241R. doi:10.1080/02724634.1988.10011708. ISSN 0272-4634.
6. McKenna, Malcolm C.; Bell, Susan K. (1997). Classification of Mammals Above the Species Level. Columbia University Press. pp. 9–10. ISBN 978-0-231-11012-9. Retrieved 16 March 2015.
7. Flannery, Timothy F.; Rich, Thomas H.; Vickers-Rich, Patricia; Ziegler, Tim; Veatch, E. Grace; Helgen, Kristofer M. (2022-01-02). "A review of monotreme (Monotremata) evolution". Alcheringa: An Australasian Journal of Palaeontology. 46 (1): 3–20. Bibcode:2022Alch...46....3F. doi:10.1080/03115518.2022.2025900. ISSN 0311-5518.
8. Flannery, Timothy F.; Rich, Thomas H.; Vickers-Rich, Patricia; Veatch, E. Grace; Helgen, Kristofer M. (2022-11-01). "The Gondwanan Origin of Tribosphenida (Mammalia)". Alcheringa: An Australasian Journal of Palaeontology. 46 (3–4): 277–290. Bibcode:2022Alch...46..277F. doi:10.1080/03115518.2022.2132288. ISSN 0311-5518. S2CID 253323862.
9. ^ Mao, Fangyuan; Li, Zhiyu; Wang, Zhili; Zhang, Chi; Rich, Thomas; Vickers-Rich, Patricia; Meng, Jin (2024-04-03). "Jurassic shuotheriids show earliest dental diversification of mammaliaforms". Nature. 628 (8008): 569–575. Bibcode:2024Natur.628..569M. doi:10.1038/s41586-024-07258-7. ISSN 0028-0836. PMID 38570681.
10. Flannery, Timothy F.; McCurry, Matthew R.; Rich, Thomas H.; Vickers-Rich, Patricia; Smith, Elizabeth T.; Helgen, Kristofer M. (2024-05-26). "A diverse assemblage of monotremes (Monotremata) from the Cenomanian Lightning Ridge fauna of New South Wales, Australia". Alcheringa: An Australasian Journal of Palaeontology. 48 (2): 319–337. Bibcode:2024Alch...48..319F. doi:10.1080/03115518.2024.2348753. ISSN 0311-5518.
11. Celik, Mélina A.; Phillips, Matthew J. (8 July 2020). "Conflict Resolution for Mesozoic Mammals: Reconciling Phylogenetic Incongruence Among Anatomical Regions". Frontiers in Genetics. 11: 0651. doi:10.3389/fgene.2020.00651. PMC 7381353. PMID 32774343.
12. Luo, Zhe-Xi (2011-12-01). "Developmental Patterns in Mesozoic Evolution of Mammal Ears". Annual Review of Ecology, Evolution, and Systematics. 42 (1): 355–380. doi:10.1146/annurev-ecolsys-032511-142302. ISSN 1543-592X.

External links

• "MESOZOIC MAMMALS; Stem zatherians, zatherians & Peramuridae, an internet directory". Home.arcor. Archived from the original on June 18, 2013.
• "THERIA И MARSUPIONTA". Elementy (in Russian). Archived from the original on October 31, 2013.
• "Tribosphenida (after McKenna & Bell, 1997)". Mikko's Phylogeny Archive. Archived from the original on August 22, 2023.
• "Infralegion Tribosphenida". Taxonomy. Archived from the original on March 3, 2016.

v t e Mammalia
Kingdom: Animalia Phylum: Chordata Clade: Synapsida Clade: Therapsida Clade: Cynodontia Clade: Mammaliaformes Synapsida see Synapsida Cynodontia see Cynodontia Mammalia see below↓
Mammalia Australosphenida? †Ambondro †Tendagurutherium? Monotremata? †Henosferidae Asfaltomylos Henosferus †Ausktribosphenidae Ausktribosphenos Bishops? Kryoparvus? Monotremata †Kollikodontidae? Kollikodon Kryoryctes? Sundrius †Steropodontidae Parvopalus? Steropodon †Teinolophidae Stirtodon? Teinolophos Ornithorhynchoidea †Opalios Ornithorhynchidae †Dharragarra? †Monotrematum? †Obdurodon Ornithorhynchus †Patagorhynchus? Tachyglossidae †Megalibgwilia †Murrayglossus Tachyglossus Zaglossus †Allotheria? see Allotheria includes multituberculates Theriimorpha see below↓
Theriimorpha Theriimorpha †Acinacodus †Amphidon †Amphilestes †Aploconodon †Chaoyangodens †Comodon †Condorodon †Dyskritodon? †Fruitafossor? †Hakusanodon †Indotriconodon †Juchilestes †Jueconodon †Kemchugia †Kryptotherium †Liaoconodon †Liaotherium †Manchurodon †Nakunodon †Phascolotherium †Sangarotherium †Tendagurodon †Allotheria? Monotremata? †Gobiconodontidae Fuxinoconodon Gobiconodon Hangjinia Huasteconodon? Meemannodon Repenomamus Spinolestes †Jeholodentidae Jeholodens Yanoconodon †Klameliidae? Ferganodon Klamelia †Triconodontidae Alticonodon Arundelconodon Astroconodon Corviconodon Eotriconodon Jugulator Meiconodon Priacodon Triconodon Trioracodon Victoriaconodon Volaticotheria? †Volaticotheria Argentoconodon Ichthyoconodon Triconolestes? Volaticotherium †Tinodontidae Gobiotheriodon? Tinodon Trishulotherium Yermakia Trechnotheria Monotremata? †Zhangheotheriidae Anebodon Kiyatherium Maotherium Ningchengodon Origolestes Zhangheotherium †Spalacotheriidae Akidolestes Aliaga Cifellitherium Heishanlestes Infernolestes Lactodens Shalbaatar Spalacolestes Spalacotheridium Spalacotherium Spalacotheroides Symmetrodontoides Symmetrolestes Yaverlestes Cladotheria see below↓
Cladotheria †Dryolestida Brancatherulum? Dorsetodon Drescheratherium Euthlastus Henkelotherium Paurodon Tathiodon Dryolestidae Achyrodon Amblotherium Anthracolestes Beckumia Crusafontia Dryolestes Groebertherium? Guimarotodus Hercynodon Krebsotherium Lakotalestes? Laolestes Minutolestes Phascolestes Portopinheirodon Treslagosodon †Meridiolestida Amarillodon Austrotriconodon Bondesius Casamiquelia Cronopio Groebertherium? Lakotalestes? Leonardus Necrolestes Paraungulatum Quirogatherium Solanutherium Mesungulatoidea Coloniatherium Mesungulatum Orretherium Peligrotherium Reigitherium †Donodontidae Amazighodon Anoualestes Donodon Stylodens Prototribosphenida †Abelodon †Amphibetulimus †Amphitherium †Arguimus †Brancatherulum? †Chunnelodon †Magnimus †Minimus †Mozomus †Nanolestes †Palaeoxonodon †Vincelestes Monotremata? †Peramuridae Afriquiamus? Kouriogenys Peramuroides Peramus Tendagurutherium? Tribosphenida / Boreosphenida †Aegialodon †Argaliatherium †Brasilestes †Carinalestes †Eomaia †Holoclemensia †Hypomylos †Juramaia †Kermackia †Kielantherium †Pappotherium †Picopsis †Tirotherium †Tribactonodon Theria †Patagomaia Eutheria see Eutheria includes placentals Metatheria see Metatheria includes marsupials
Other taxa Incertae sedis †Atlasodon †Chronoperates †Microderson †Thereuodon †Tingamarra Saint Bathans Mammal UA 8699 Other taxa Prototheria Yinotheria Eutriconodonta Amphilestidae Theriiformes Pantotheria Symmetrodonta Eupantotheria Paurodontidae Amphitheriidae Marsupionta

• Tribosphenida
• Mammal taxonomy