Catonyx

Catonyx; Temporal range: Early Pliocene-Early Holocene (Montehermosan-Lujanian); ~2.5–0.010 Ma PreꞒ Ꞓ O S D C P T J K Pg N ↓
	Replica of a Catonyx skull at the Museo delle Scienze
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	Pilosa
Family:	†Scelidotheriidae
Genus:	†Catonyx; Ameghino, 1891
Species
	C. cuvieri Lund, 1839; C. chiliensis Lydekker, 1886; C. tarijensis Gervais and Ameghino, 1880;

Catonyx is an extinct genus of ground sloth of the family Scelidotheriidae, endemic to South America during the Pliocene and Pleistocene epochs.^[1] It lived from 2.5 Ma to about 10,000 years ago, existing for approximately 2.49 million years.^[2] The most recent date obtained is about 9600 B.P.^[3]^[4]

Description

This animal, like many other terrestrial sloths, was of conspicuous size and mighty build. It had to reach and exceed 4 meters in length, and the skull alone was at least 50 centimeters long. Its weight has been estimated at over 1500 kg. ^[5] The snout of Catonyx was elongated, although not as in some similar forms (e.g., Scelidotherium). Unlike the latter, Catonyx possessed shorter premaxillae that formed a triangular (and not rectangular like Scelidotherium) snout tip, a pronounced rostrum bulge, a palate equipped with a median groove, and larger teeth. In addition, the mandibular symphysis was elongated and elevated, and the posterior lobe of the lower fourth molar was more curved than that of Scelidotherium. Like all terrestrial sloths, Scelidotherium possessed strong-boned limbs and large claws.

Taxonomy

At a cave in Lagoa Santa, Brazil, Peter Wilhelm Lund and his crew members collected the fragmentary fossils of a fossil sloth that Wilhelm named Megalonyx cuvieri in 1838. Catonyx, the genus name, was made by Ameghino in 1891. It was assigned to Scelidotheriinae by Gaudin in 1995.^[6] Scelidotheriinae was elevated back to family status by Presslee et al. in 2019.^[7]

The very first fossils of this animal were found in Upper Pleistocene strata of Brazil and were described by Lund in 1839, but for a long time there was considerable systematic confusion: the remains were gradually attributed to the genera Scelidotherium and Scelidodon. Only recently have revisions based on clear morphological anatomy been proposed, according to which the genera Scelidotherium and Catonyx are two valid taxa, while Scelidodon may or may not be a valid genus. Catonyx, in any case, is a member of the Scelidotheriidae, a group of terrestrial sloths known from the Miocene, Pliocene, and Pleistocene and characterized by an elongated snout; scelidotheres themselves part are usually placed as a subfamily of the Mylodontidae, although they are sometimes considered a separate family, Scelidotheriidae.^[7]

The type species of Catonyx is Catonyx cuvieri, named in 1839 by Peter Wilhelm Lund. C. cuvieri has been found in Brazil and Uruguay, and dates to the Late Pleistocene and likely the Early Holocene.^[8] Other species attributed to this genus but are sometimes considered to belong to a separate genus, Scelidodon, are C. tarijensis and C. chiliense, found in Chile, Argentina, Uruguay, Bolivia, and Ecuador.

Below is a phylogenetic tree of the Scelidotheriinae, based on the work of Nieto and colleagues (2020), showing the position of Catonyx.^[9]

Scelidotheriinae

Sibyllotherium guenguelianum

Neonematherium flabellatum

Scelidotherium

Proscelidodon gracillimus

Proscelidodon patrius

	Proscelidodon rothi

	Valgipes bucklandi

	Catonyx cuvieri

	Catonyx tarijensis

	Catonyx chiliense

Palaeobiology

Hypothetical life reconstruction of *C. tarijensis* showing its inferred feeding behavior. The reconstruction is based on the skull MNHN-Bol V 13364 from the Pleistocene of Oruro (Bolivian Altiplano).

The cranial anatomy of the species C. tarijensis indicates it may have been a browser which used its strong lips to grab vegetation,^[10] and Santos Pereira et al. (2013) tentatively suggested browsing habits for C. cuvieri.^[11] Isotopic analysis of Smilodon populator remains from Brazil indicates that Catonyx was one of its main prey items.^[12]

Fossil distribution

Fossils of Catonyx have been uncovered in Brazil, the San José Formation of Uruguay, the Tezanos Pinto Formation of Argentina,^[13] and the Tarija Valley of Bolivia.^[10]

Paleoecology

Life restoration of *C. cuvieri* (upper left) and contemporary animals

In the Brazilian Intertropical Region in eastern Brazil, Catonyx was a browser in arboreal savannahs and forested grasslands. Large, mesoherbivorous mammals in the BIR were widespread and diverse, including the cow-like toxodontids Toxodon platensis and Piauhytherium, the macraucheniid litoptern Xenorhinotherium and equids such as Hippidion principale and Equus neogaeus. Toxodontids were large mixed feeders as well and lived in forested areas, while the equids were nearly entirely grazers. Other xenarthran fossils are present in the area as well from several different families, like the giant megatheriid ground sloth Eremotherium, the fellow scelidotheriid Valgipes, the mylodontids Glossotherium, Ocnotherium, and Mylodonopsis. Smaller ground sloths such as the megalonychids Ahytherium and Australonyx and the nothrotheriid Nothrotherium have also been found in the area. Eremotherium was a generalist, while Nothrotherium was a specialist for trees in low density forests, and Valgipes was an intermediate of the two that lived in arboreal savannahs. Other glyptodonts and cingulates like the grazing glyptodonts Glyptotherium and Panochthus and the omnivorous pampatheres Pampatherium and Holmesina were present in the open grasslands. A proboscidean species has also been found in the BIR, Notiomastodon platensis, which was also present and was a mixed grazer on the open grasslands. Carnivores included some of the largest known mammalian land carnivores, like the giant felid Smilodon populator and the bear Arctotherium wingei.^[14]^[15] Several extant taxa are also known from the BIR, like guanacos, giant anteaters, collared peccaries, and striped hog-nosed skunks.^[16] Two crab-eating types of extant mammals are also known from the BIR, the crab-eating raccoon and the crab-eating fox, indicating that crabs were also present in the region.^[16] The environment of the BIR is unclear, as there were both several species that were grazers, but the precede of the arboreal fossil monkeys Protopithecus and Caipora in the area causes confusion over the area's paleoenvironment. Most of Brazil was thought to have been covered in open tropical cerrado vegetation during the Late Pleistocene, but if Protopithecus and Caipora were arboreal, their presence suggests that the region may have supported a dense closed forest during the Late Pleistocene.^[16]^[17] It is possible that the region alternated between dry open savannah and closed wet forest throughout the climate change of the Late Pleistocene.^[18]

References

^ McDonald, H. G.; Perea, D. (2002). "The large scelidothere Catonyx tarijensis (Xenarthra, Mylodontidae) from the Pleistocene of Uruguay". Journal of Vertebrate Paleontology. 22 (3): 677. doi:10.1671/0272-4634(2002)022[0677:TLSCTX]2.0.CO;2. JSTOR 4524258. S2CID 86021240.
^ PaleoBiology Database: Catonyx, basic info
^ Turvey, Sam (2009). Holocene extinctions. Oxford University Press. pp. 20–33, 42–50, 352. ISBN 978-0-19-953509-5.
^ Fiedal, Stuart (2009). "Sudden Deaths: The Chronology of Terminal Pleistocene Megafaunal Extinction". In Haynes, Gary (ed.). American Megafaunal Extinctions at the End of the Pleistocene. Vertebrate Paleobiology and Paleoanthropology. Springer. pp. 21–37. doi:10.1007/978-1-4020-8793-6_2. ISBN 978-1-4020-8792-9.
^ Toledo, N.; Bargo, M. S.; Vizcaíno, S. F.; De Iuliis, G.; Pujos, F. (2017). "Evolution of body size in anteaters and sloths (Xenarthra, Pilosa): phylogeny, metabolism, diet and substrate preferences". Earth and Environmental Science Transactions of the Royal Society of Edinburgh. 106 (4): 289–301. Bibcode:2015EESTR.106..289T. doi:10.1017/S1755691016000177. hdl:11336/56403. S2CID 90400372.
^ Gaudin, T. J. (1995). "The ear region of edentates and the phylogeny of the Tardigrada (Mammalia, Xenarthra)". Journal of Vertebrate Paleontology. 15 (3): 672–705. Bibcode:1995JVPal..15..672G. doi:10.1080/02724634.1995.10011255. JSTOR 4523658.
^ ^a ^b Presslee, S.; Slater, G. J.; Pujos, F.; Forasiepi, A. M.; Fischer, R.; Molloy, K.; Mackie, M.; Olsen, J. V.; Kramarz, A.; Taglioretti, M.; Scaglia, F.; Lezcano, M.; Lanata, J. L.; Southon, J.; Feranec, R.; Bloch, J.; Hajduk, A.; Martin, F. M.; Gismondi, R. S.; Reguero, M.; de Muizon, C.; Greenwood, A.; Chait, B. T.; Penkman, K.; Collins, M.; MacPhee, R.D.E. (2019). "Palaeoproteomics resolves sloth relationships" (PDF). Nature Ecology & Evolution. 3 (7): 1121–1130. Bibcode:2019NatEE...3.1121P. doi:10.1038/s41559-019-0909-z. PMID 31171860. S2CID 174813630.
^ Corona, Andrea; Perea, Daniel; McDonald, H. Gregory (2013). "Catonyx cuvieri (Xenarthra, Mylodontidae, Scelidotheriinae) from the late Pleistocene of Uruguay, with comments regarding the systematics of the subfamily". Journal of Vertebrate Paleontology. 33 (5): 1214–1225. Bibcode:2013JVPal..33.1214C. doi:10.1080/02724634.2013.764311. ISSN 0272-4634. S2CID 84078331.
^ Nieto, Gastón L.; Haro, J. Augusto; McDonald, H. Gregory; Miño-Boilini, Ángel R.; Tauber, Adan A.; Krapovickas, Jerónimo M.; Fabianelli, Maximiliano N.; Rosas, Federico M. (2021-06-01). "The Skeleton of the Manus of Scelidotherium (Xenarthra, Mylodontidae) Specimens from the Pleistocene of the Province of Córdoba, Argentina, and its Systematic Implications". Journal of Mammalian Evolution. 28 (2): 221–243. doi:10.1007/s10914-020-09520-x. ISSN 1573-7055. S2CID 226319627.
^ ^a ^b Boscaini, Alberto; Iurino, Dawid A.; Mamani Quispe, Bernardino; Andrade Flores, Rubén; Sardella, Raffaele; Pujos, François; Gaudin, Timothy J. (2020). "Cranial Anatomy and Paleoneurology of the Extinct Sloth Catonyx tarijensis (Xenarthra, Mylodontidae) From the Late Pleistocene of Oruro, Southwestern Bolivia". Frontiers in Ecology and Evolution. 8. doi:10.3389/fevo.2020.00069. hdl:2434/959871. ISSN 2296-701X.
^ dos Santos Pereira, I. C.; et al. (2013). "Record of the giant sloth Valgipes bucklandi (Lund, 1839) (Tardigrada, Scelidotheriinae) in Rio Grande do Norte state, Brazil, with notes on taphonomy and paleoecology". Journal of South American Earth Sciences. 43: 42–45. Bibcode:2013JSAES..43...42P. doi:10.1016/j.jsames.2012.11.004.
^ Dantas, Mário André Trindade; Cherkinsky, Alexander; Lessa, Carlos Micael Bonfim; Santos, Luciano Vilaboim; Cozzuol, Mario Alberto; Omena, Érica Cavalcante; Silva, Jorge Luiz Lopes; Sial, Alcides Nóbrega; Bocherens, Hervé (2020-07-14). "Isotopic paleoecology (δ13C, δ18O) of a late Pleistocene vertebrate community from the Brazilian Intertropical Region". Revista Brasileira de Paleontologia. 23 (2): 138–152. doi:10.4072/rbp.2020.2.05. ISSN 2236-1715.
^ "Fossilworks: Catonyx".
^ Keeley, J. E., & Rundel, P. W. (2003). Evolution of CAM and C4 carbon-concentrating mechanisms. International journal of plant sciences, 164(S3), S55-S77.
^ Omena, Érica Cavalcante; Silva, Jorge Luiz Lopes da; Sial, Alcides Nóbrega; Cherkinsky, Alexander; Dantas, Mário André Trindade (2021-10-03). "Late Pleistocene meso-megaherbivores from Brazilian Intertropical Region: isotopic diet (δ13C), niche differentiation, guilds and paleoenvironmental reconstruction (δ13C, δ18O)". Historical Biology. 33 (10): 2299–2304. Bibcode:2021HBio...33.2299O. doi:10.1080/08912963.2020.1789977. ISSN 0891-2963. S2CID 225543776.
^ ^a ^b ^c Cartelle, Castor; Hartwig, W. C. (1996). "A new extinct primate among the Pleistocene megafauna of Bahia, Brazil". Proceedings of the National Academy of Sciences. 93 (13): 6405–6409.
^ Eisenberg, John F.; Redford, Kent H. (1989). Mammals of the Neotropics, Volume 3: Ecuador, Bolivia, Brazil. University of Chicago Press. p. 247. ISBN 978-0-226-19542-1.
^ Halenar, Lauren B. (December 2011). "Reconstructing the Locomotor Repertoire of Protopithecus brasiliensis". The Anatomical Record. 294 (12): 2048–2063.