Triisodon

Triisodon
Temporal range: Early Paleocene (Torrejonian), 63.5–62 Ma
T. quivirensis skull, National Museum of Natural History
Restoration of T. quivirensis
Scientific classification Edit this classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Family: †Triisodontidae
Genus: †Triisodon
Cope, 1881
Type species
Triisodon quivirensis
Cope, 1881
Species
  • T. quivirensis (Cope, 1881)
  • T. crassicuspis (Cope, 1882)
Synonyms
  • T. antiquus (Cope, 1882)
  • T. rusticus (Cope, 1883)

Triisodon (ancient Greek: "Tri" (three),"isos" (equal), and modern Greek: "donti" (tooth/teeth)[citation needed], supposedly describing tritubercular lower cheek teeth) is a genus of an extinct family of hoofed mammals known as Triisodontidae. Triisodon existed during the Early Paleocene of New Mexico, North America, from about 63.5-62.0 Ma. The genus was named by Edward Drinker Cope in 1881 as a member of the Creodonta, a now invalid group that encompassed hyaenodonts, oxyaenids, mesonychians and certain arctocyonians. Cope described the type specimen of T. quivirensis as "about the size of a wolf." A smaller species, T. crassicuspis, has also been identified from the same region. Since material from this genus is incomplete, the exact size of adults and whether they showed sexual dimorphism or regional variations in size is unknown.

Triisodon would go extinct during a faunal turnover between the Tj4-5 and Tj6 biozones, caused by the U27r excursion, a warming period that saw an increase in aridity.

Taxonomy

Triisodon is the type genus of the family Triisodontidae, a group of hoofed mammals. Other triisodontid genera, including Goniacodon, Eoconodon, Stelocyon, and Oxyclaenus. Like many very early mammals, the relationship of triisonodontids to other living and fossil mammals has been uncertain. Originally, Triisodontidae were considered to be a family within Creodonta. Matthew (1909) would reclassify triisodontids as triisodontines within Mesonychidae within the suborder Acreodi. In a later paper, Matthew recovered triisodontines within Arctocyonidae.

Arctocyonids, as well as mesonychids, would be moved from Creodonta and into Condylarthra, also known as archaic ungulates by Van Valen (1966). Triisodontines would later be reclassified as Triisodontidae by McKenna and Bell (1997), as a family within Mesonychia. However most experts recover the family outside of the order. Tabuce et al. (2011) and Shelley et al. (2015) recovered triisodontids as a paraphyletic group stem to mesonychians. On the other hand, more recent analysis suggesting a closer relation to living ungulates.

Species

Description

Triisodon was a rather large and powerful terrestrial mammal, as evident from the robust forelimb morphology. The body mass of T. crassicuspis is estimated at 32–44 kg (71–97 lb) making it similar in size to a striped hyena and gray wolf. T. quivirensis is considered to be even larger than T. crassicuspis.

Skull and dentition

Skull reconstruction of T. crassicuspis, with known material in grey, with areas in white restored after other triisodontids, including T. quivirensis.

The skull of T. crassicuspis exhibits rostral constriction, particularly above the upper second premolar, similar to Arctocyon primaevus. The maxilla of T. crassicuspis barely extends onto the zygomatic arch laterally. The posterior zygomatic arch was particularly robust. T. crassicuspis had a large and particularly deep mandibular fossa that is more elongated mediolaterally than anteroposteriorly. The depth of the fossa suggests that it formed a relatively tight hinge-like articulation with the condyle of the mandible. As in Arctocyon, the articular surface of the mandibular fossa in T. crassicuspis is smooth and continuous.

The paired hemi-mandibles of T. crassicuspis were not co-ossified towards the front, as indicated by the rugose texture of the mandibular symphysis. The mandibular body of T. crassicuspis is proportionately deeper dorsoventrally and significantly more robust compared to that of Eoconodon coryphaeus, or mesonychids like Sinonyx jiashanensis and Dissacus praenuntius. In T. crassicuspis, a massive and mediolaterally broad coronoid crest forms the anterior surface of the coronoid process. The coronoid crest ends below the lower third molar, as in T. quivirensis, Eoconodon, and Arctocyon.

The teeth of Triisodon exhibit several morphological similarities to those of other triisodontids including the diagnostic characteristics of the group: basin-shaped talonids and tribosphenic upper molars.

Postcranial skeleton

While the postcranial skeleton of Triisodon was less robust compared to other "condylarths" like Periptychus and Arctocyon, it was far from gracile. Much of what is known about the forelimb of Triisodon comes from the humerus, with both the greater and lesser tubercles placed low relative to the massive, hemispherical humeral head. In addition, the humerus also had a prominent, V-shaped deltopectoral region that extended anteriorly. The attachment site for the teres major and latissimus dorsi muscles is poorly defined. The distal humeral epiphysis is anteroposteriorly compressed. The humeral trochlea is asymmetrical, as it is in Arctocyon and other "triisodontids" including Goniacodon levisanus and Eoconodon. The humeral trochlea and rounded capitulum form a near continuous articular surface, isolated by a zona conoidea that is poorly-developed. The medial crest of the humeral trochlea is very prominent and is slightly flared, protruding mediodistally, while the lateral crest is smaller and protrudes posteriorly.

The ulna is conspicuously more gracile relative to the humerus, and has a gently convex ventral border. The ulna has a large coronoid process, while the trochlear notch has a saddle-shaped anterior surface. The ulnar diaphysis bears a deep fossa for the abductor pollicis longus muscle on the lateral surface, while the distal ulna was relatively thick, with a well-developed pronator quadratus crest on its medial surface. The radius is somewhat robust relative to the ulna, and exhibits an ovoid, shallowly concave radial fovea. The radial diaphysis is also rather straight in its proximal portion, with a bicipital tuberosity that is proximodistally long, but not particularly prominent compared to condylarths including Goniacodon, Arctocyon, and Ankalagon. Triisodon also had a massive, hemispherical femoral head with a smooth articular surface, and an ovoid, somewhat deep fovea capitis for attachment of the ligamentum teres.

Paleobiology

Diet

Skull of T. quivirensis

The canines of Triisodon are massive and single-rooted, with marked shearing edges. Other features of the teeth reveal that Triisodon and other triisodontids were showing evolutionary trends towards a carnivorous lifestyle – notably the mesiodistally oriented hypoconulid crest, tall trigonid and distinct protoconid of the lower molars, as well as the strongly-developed posterior premolars bearing marked shearing surfaces. Among triisodontids, Triisodon showed the most specialization towards carnivory. Despite showing signs of carnivory, Triisodon cheek dentition were poorly suited for slicing flesh compared to hypercarnivorous carnivorans such as felids. The broad, low-crowned, bunodont molars Triisodon were similar to that of bears, suggesting it had diet consisting of fruits, nuts, seeds, invertebrates, as well as medium to large sized vertebrates and carrion. The deepened glenoid fossa as well as inflated, conical molar cusps suggests it may have been engaging in durophagy.

Locomotion

Triisodon may have had strong shoulders, as indicated by the anteriorly prominent deltopectoral region. Triisodon may also have had moderate digging capabilities, as evident from the thick distal ulna and the robustness of the humerus. The fossa for the abductor pollicis longus muscle on the ulna suggests high-powered forelimb extension, which is movement possibly linked to brief bursts of quick terrestrial locomotion including running.

References

  1. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 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.
  2. 1 2 Cope, E. D. (1881). "On Some Mammalia of the Lowest Eocene Beds of New Mexico". Proceedings of the American Philosophical Society. 19 (109): 484–495. ISSN 0003-049X. JSTOR 982794.
  3. ↑ Secord, Ross; Peppe, Daniel J.; Williamson, Thomas E.; et al. (2026). "Recognition of two potential warming events in the Paleocene terrestrial record of North America and associated mammalian responses". PNAS Nexus. 5 (9). doi:10.1093/pnasnexus/pgag289.
  4. 1 2 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.
  5. ↑ "Mesonychians part III: Andrewsarchus and the triisodontines | ScienceBlogs". scienceblogs.com. Retrieved 2019-06-09.
  6. ↑ 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.
  7. ↑ 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.
  8. ↑ 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.
  9. ↑ 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.
  10. 1 2 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)
  11. 1 2 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.
  12. 1 2 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.
  13. 1 2 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.
  14. ↑ 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.
  15. ↑ Kondrashov, and Spencer Lucas, Peter (January 2006). "Early Paleocene (Puercan and Torrejonian) Archaic ungulates (Condylarthra, Procreodi and Acreodi) of the San Juan basin, New Mexico" – via ResearchGate.
  16. ↑ Journal of Mammalogy. American Society of Mammalogists. 1981. p. 683.
  17. ↑ Grossnickle, David M.; Newham, Elis (2016). "Therian mammals experience an ecomorphological radiation during the Late Cretaceous and selective extinction at the K–Pg boundary". The Royal Society Proceedings of the Royal Society B. 283 (1832). doi:10.1098/rspb.2016.0256. PMC 4920311.