Biology:Ortotherium

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Short description: Extinct genus of ground sloth

Ortotherium
Temporal range: Late Miocene Huayquerian
~9–6.8 Ma
↓
Ortotherium laticurvatum type MLP M 62.jpg
Illustration of the cast of the holotype dentary (MLP M 62).
Scientific classification edit
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Pilosa
Family: †Megalonychidae
Genus: †Ortotherium
Ameghino, 1885
Species:
†O. laticurvatum
Binomial name
†Ortotherium laticurvatum
Ameghino, 1885
Synonyms
  • Menilaus affinis Ameghino, 1891
  • Ortotherium schlosseri Ameghino, 1891
  • Ortotherium seneum Ameghino, 1891
  • Amphiocnus seneum (Ameghino, 1891)
  • Ortotherium scrofum Bordas, 1942

Ortotherium ("upright beast") is a genus of megalonychid ground sloth from the Late Miocene (Huayquerian SALMA, around 9 to 6.8 million years ago) Ituzaingó Formation of Entre Rios Province, Argentina . Although many species were described, the only valid species of the genus is Ortotherium laticurvatum, with many species being junior synonyms. Ortotherium is known from very fragmentary material, all of which is material from the mandible (lower jaw) and teeth. The holotype (specimen a scientific name is applied to) of O. laticurvatum consists of an incomplete left dentary that had been unearthed from a series of sediments known as ‘Conglomerado osifero’ in Paraná, Argentina. Argentina paleontologist Florentino Ameghino named the species in 1885, though he would go on to name four more, invalid, species of the genus. One species however, O. brevirostrum, has been reclassified as Mesopotamocnus.

Due to Ortotherium being known from very scant fossils, little is definitively known about the animal. However, much can be inferred based on related taxa. Ortotherium was average-sized for a Huayquerian megalonychid, being around 65 kg (143.3 lbs) using the similarly proportioned genus Eucholoeops. Ortotherium has a long mandible, with large, rectangular molars preceded by a giant caniniform. It had long, robust forelimbs terminating in a series of enlarged claws, using a mix of quadrapedral and bipedal movement, possibly permitting tree climbing. The hindlimbs were large and pillar-like supported by a lengthy tail.

Being a megalonychid, Ortotherium was a herbivorous mammal that likely consumed a large amount of leafy plant material. Their jaws and teeth were built for a shearing and cutting method of mastication, bearing sharp teeth and a wide array of jaw movement. The Ituzaingó Formation that Ortotherium inhabited was subtropical, with a mix of swampy woodlands and more open grasslands present. Fossils of mangroves indicate parts of the formation bore a shoreline marine ecosystem along the warm, salty waters of the Paranaense Sea. This allowed for a wide array of fauna to inhabit the area, which included many different genera of ground sloths, "native ungulates", and rodents, in addition to a menagerie of birds, including the large carnivorous "terror birds", as well as fish and reptiles.

History

Portrait of Florentino Ameghino, describer of Ortotherium.

The earliest published record of Ortotherium was in 1885, when a partial left dentary (lower jawbone) lacking preserved teeth that had been unearthed from the Upper Miocene strata of "Conglomerado osifero" (ossiferous conglomerate) near Paraná in Entre Rios Province, Argentina. It was described by prolific Argentine paleontologist Florentino Ameghino in his monograph on fossils from the area.[1] Ameghino named the type species Ortotherium laticurvatum, the specific name coming from the laterally curved lower jaw.[1] The holotype (name-bearing) specimen of O. laticurvatum has been lost, though a plaster cast exists in the Museo de La Plata.[2] Ameghino, mistakenly believed that the rock layers of Parana dated to the Oligocene, a much older epoch.[3][1] Ameghino noted the large size of the molariform and caniniform cavities, believing that Ortotherium may be synonymous with the taxon Olygodon, which he had named on the basis of large molariforms.[4][1] Ortotherium, however, is a distinct taxon that comes from different aged sediments.[4] O. laticurvatum was one of three species Ameghino named, fostering the novel species of O.schlosseri and O. seneum in an 1891 paper, both of which were also based on fragmented mandibular material.[5] Ameghino dubbed a new genus and species of ground sloth, Menilaus affinis, in the same 1891 work. M. affinis was named on the basis of another imperfect left mandible from "Conglomerado osifero".[6][5] Menilaus has since been synonymized with Ortotherium laticurvatum, lacking any distinct traits that differ itself from the species.[7][2]

Map of Argentina showing where all fossils of Ortotherium have been unearthed (marked by a star)

Ameghino was a very active paleontologist, naming hundreds of new genera and species during his academic career in Argentina.[8][9] Following his death in 1911, analysis of his taxa and scrutiny of his work came about.[10][11] Lucas Kraglievich was one of these paleontologists, reclassifying O. seneum as a species of the other megalonychid Amphiocnus and considered Menilaus congeneric with Pliomorphus.[6] Later in 1942, paleontologist A. F. Bordas dubbed another two species of Ortotherium, the specimens composed of only mandible material.[12][2] The species, O. brevirostrum and O. scrofum, though the latter is currently considered a synonym of O. laticurvatum. The 21st century saw extensive review of the genus and its many species by Diego Brandoni first in 2010 in which he did a full overview of all species.[2] In his 2010 review, he synonymized all of Ortotherium's species with the exception of O. brevirostrum which he noted likely belonged to a separate genus.[13][7][14][2] Brandoni also described fossils of an indeterminate species, including a nearly complete left mandible that had a different placement of an opening in the mandibular canal (a ridge running along the sides of the mandible) compared to O. laticurvatum.[13][2] However, this was not seen as enough to justify naming a novel species according to Brandoni.[2] Four years later, Brandoni erected the generic name Mesopotamocnus for O. brevirostrum, the name deriving from its geographic provenance in Mespotamia, Argentina and ocnus meaning "slowness", a commonly used root for the generic names of ground sloths.[15][14]

Taxonomy

Life restoration of the related ground sloth Megalonyx.

Ortotherium was a megalonychid sloth, a family within the order Folivora which contains all of sloths. Megalonychids existed from the Deseadan SALMA (29–21 mya) to the Rancholabrean NALMA (240,000 BCE to 11,000 BCE), the last surviving genus being Megalonyx itself from North America.[16] Megalonychids were long-considered to be an extant group including the two-toed sloth genus Choeloepus, however analyses of the collagen and DNA of fossils of folivorans proved that Choloepus was instead related to Mylodontids, another family of ground sloths. Fossils of early megalonychids are rare, the oldest named being of the genus Deseadognathus from Argentina and Bolivia,[17] though even older fossils from the Early Oligocene have been tentatively reported from Puerto Rico.[18] The megalonychids later saw an explosion in diversity during the Middle-Upper Miocene in the Americas, primarily among the Santacrucian (17.5–16.3 mya) and Friasian (16.3–15.5) sites of Argentina, Uruguay, Venezuela, and Brazil. Extensive waterways formed in South America during this period, giving way to a more subtropical climate fostering a variety of flora and fauna to evolve in this environment. Megalonychids also expanded their range north, with genera like Zacatzontli from Mexico evolving on the North American continent prior to the development of the Isthmus of Panama.[19] Megalonychids had spread throughout the Caribbean and as far north as the Yukon Territory, Canada by their demise at the end of the Pleistocene,[20] though the closely related three-toed sloth Bradypus is extant. Research of evolutionary size trends suggests that in contrast to other ground sloth groups that grew over time, megalonychids did not exponentially increase in mass but instead varied greatly around the same amount until their extinction.[21][22]

The following sloth family phylogenetic tree is based on collagen and mitochondrial DNA sequence data (see Fig. 4 of Presslee et al., 2019).[23]

Xenarthra
 

Cingulata (armadillos and allies)

 
Pilosa
 

Vermilingua (anteaters)

 
Folivora
 

Megalocnidae (Caribbean sloths)

 
 
 
 
 
 

Nothrotheriidae (ground sloths)

 
 

Megatheriidae (giant ground sloths)

 
 
 
 

Megalonychidae

 
 

Bradypodidae (three-toed sloths)

 
 
 
 
 

Scelidotheriidae (two-toed sloths)

 
 
 
 
("Edentata")  

The internal phylogeny of Megalonychidae, specifically taxa like Ortotherium, is tenuous as many taxa bear only fragmentary or incomplete specimens. This has caused Ortotherium, a genus known purely from partial mandibles and teeth, to be excluded from the majority of Megalonychid cladistic analyses.[24][7] The only exception being by McDonald & Carranza-Castañeda (2017) in their description of Zacatzontli, a megalonychid from the Late Miocene of southern Mexico.[25] In their analysis, three separate clades related to biogeography were recovered, with a South American clade, including Ortotherium, Caribbean clade, and a North American clade. The Caribbean and North American taxa formed a separate subfamily from the South American, suggesting that the southern megalonychid branch broke off earlier in the Oligocene.[18][25] When originally named by Ameghino in 1885, he did not specify the phylogenetic position of the genus besides Edentata, though he did state similarities to Bradypus and the nomen dubium Olygodon. Family-level assessment in 1942, Bordas believed that Ortotherium was part of the clade Nothrotheriinae, then believed to be within Megalonychidae.[2] As part of his review of the genus, Brandoni moved Ortotherium to an indeterminate position within Megalonychidae, though noted similarities to other Miocene megalonychids.[14][26][2]

The following cladistic diagram is adapted from McDonald & Caranza-Castañeda (2017), their description of Zacatzontli:[25]

Megalonychidae
 

Hapalops

 
 
 
 

Megistonyx

 
 
 

Zacatzontli

 
 
 

Ortotherium

 
 
 

Ahytherium

 
 
 

Deseadognathus

 
 
 

Proschismotherium

 
 

Mesopotamocnus

 
 

Australonyx

 
 
 
 
 
 
 
 
 

Eucholoeops

 
 
 

Megalonyx

 
 
 

Aratocnus

 
 

Neocnus

 
 
 

Megalocnus

 
 
 

Parocnus

 
 

Pliometanastes

 
 
 
 
 
 
 
 

Description

Although scant material is known, estimations of the mass and length of related genera of similar size and proportions have been made. Ortotherium was slightly larger than Eucholaeops and around the same size as Paranabradys.[14][2] Eucholaeops was between 60 and 65 kg in weight, though some estimates reach 80 kg, giving an idea for the mass of Ortotherium.[27][22] Megalonychids like Ortotherium were heavily built for their size, with large, plantigrade (flat-footed) hindlimbs that supported its large mass. It has been suggested that, with their long, stiff tails as support, megalonychids could reach a semi-erect position that could enable eating at greater heights.[28] The hands were tridactyl terminating in large claws, the namesake for the family, that had defensive as well as browsing capabilities.[28]

Mandibles and teeth

Reconstruction of the mandible based on the type cast (white) with teeth labeled.

O. laticurvatum preserves a wide array of individual variation, with the size of teeth, mandible length, and height varying greatly. The proportions also fluctuate, causing many specimens to be misinterpreted previously as their own species.[7][2] The mandible is exceptionally deep and robust compared to related taxa, with the largest specimen preserving a ramus that reaches 6 centimeters thick, though the size varies widely among individuals.[2] The ventral margin is convex, transversely and anterioposteriorily, creating the thickness and robusticity seen. An opening on the posterolateral face of the mandibular canal is on the coronoid process in O. laticurvatum, however specimen MACN Pv-8916 of Ortotherium sp. bears this opening anterior to the process. The anatomy of O. laticurvatum is more similar to genera like Megalonyx and Paranabradys in this aspect, suggesting that MACN Pv-8916 may be from a distinct species.[29][30][2] All megalonychids had hypsodont (high-crowned) teeth, as in the teeth of Ortotherium. The third molariform (cheek tooth) lines up with the anterior margin of the coronoid process, making the entire tooth visible in lateral (side) view. All megalonychids bear only four teeth on their lower jaws that is divided by a large diastema (toothless gap) between the caniniform and the first molar.[2] Caniniforms of Ortotherium are ovate with a posteromedial bulge, their position strongly protruding from the anterior end of the mandible. The tip of the mandible was U-shaped, with long and boxy sides composing the ramus.[2] Along the tooth row, alveoli are present along the mandibular near the molars, the first being triangular, second oval, and third suboval.[2]

The diagnostic traits (characteristics that distinguish a taxon from others) of the genus include: an anterior margin of coronoid process lateral to midpoint of the molariform 3 (molariform abbreviated as m), more anterior than in the fellow megalonychids Eucholaeops and Paranabradys, and more posterior than in Megalocnus, Megalonyx, and Pliometanastes.[31][30] The posterior edge of mandibular symphysis inferomedial to the caniniform–m1 diastema; inferomedial to m1/m2 interval; inferomedial to the caniniforms. This is a combination of traits not present in other megalonychids, with taxa like Megalocnus preserving the former trait but lacking the latter two, giving credence to Ortotherium's independence as a genus.[2] Alveolus of the caniniforms are elliptical in cross-section and small relative to molariforms, in contrast to the larger molariforms of Megalonyx and Megalocnus.[30] The first molariform is subtrapezoidal in cross-section, while it is subrectangular in Megalonyx and Megalocnus. The m2 and m3 are oval in cross-section, a distinct trait compared to Megalonyx and Megalocnus.[32][30] As in most genera in the megalonychid grade, the mandible has a diastema between the caniniforms and molar row, differing from Diabolotherium nordenskioldi in which it is absent.[33][2] This unique combination of characteristics proves Ortotherium's validity.[34][14][26][2]

Paleoenvironment

Life restoration of the “terror bird” Andalgalornis, which coexisted with Ortotherium.

Ortotherium is known from the Ituzaingó Formation, the fossils being deposited in a series of sediments made up of fine quartz gravel, clay, and chalcedony clasts called the "Conglomerado osifero".[35] The strata of the Ituzaingó Formation date to the Late Miocene, a period in which animal biodiversity boomed in South America, with the advent of the faunas of formations such as the Santa Cruz, Pebas, Cerro Azul, and Urumaco strata.[36] During the Late Miocene, water levels in South America rose and a selection of tropical habitats formed in large river basins such as the Pebas System and Paranese Sea,[37] the latter bordering the Ituzaingó Formation.[38] The Ituzaingó Formation's deposits are from the beaches of the Paranese Sea,[39][40] transforming the environment into a subtropical one, as demonstrated by the strata's paleoflora and paleofauna.[41][42] The paleoflora is diverse, but includes the aquatic mangroves and terrestrial legumes,[38][43] bamboos,[44] flowering shrubs,[45] and more. Bamboos, coconut palms, and other palms were prevalent, being very common.[46] The Ituzaingó Formation also had savannahs in addition to the coastal, riverine, and tropical segments.[47][43]

Image of Amazonian tidal mudflats, a similar environment to Ituzaingó.

The Ituzaingó Formation preserves vast tidal flats similar to those in the modern day Amazon and a warm climate.[48] Other ground sloths are endemic to the site such as the megalonychids Pliomorphus, Amphiocnus, and Mesopotamocnus, megatheriids Promegatherium[49] and Pyramiodontherium,[50] and mylodontids Octomylodon, Megabradys, and Promylodon.[51][35] Notoungulate mammals in the Ituzaingó Formation were widespread, including the toxodontids Xotodon and Adinotherium,[52] and litopterns such as Brachytherium, Cullinia, Diadiaphorus, Neobrachytherium, Oxyodontherium, Paranauchenia, Promacrauchenia, Proterotherium and Scalabrinitherium.[53] Large, armored glyptodonts like Palaehoplophorus, Eleutherocercus, and Plohophorus[54] lived in the area as well as other cingulates like the pampatheres Kraglievichia[54] and Scirrotherium.[55] Carnivores included the phorusrhacids Devincenzia and Andalgalornis[56] and sparassodonts,[57] with giant crocodilians like Gryposuchus and Mourasuchus in the freshwater.[58] Ortotherium was likely a leaf-eater based on studies of megalonychid masticatory and dental morphology and capabilities, of which there was an abundance of greenery that could be comfortably chewed and digested.[27]

References

  1. ↑ 1.0 1.1 1.2 1.3 Ameghino, Florentino (1885). "Nuevos restos de mamíferos fósiles oligocenos, recogidos por el profesor Pedro Scalabrini y pertenecientes al Museo Provincial de la Ciudad del Paraná.". Boletin de la Academia Nacional de Ciencias en Córdoba, República Argentina 8. 
  2. ↑ 2.00 2.01 2.02 2.03 2.04 2.05 2.06 2.07 2.08 2.09 2.10 2.11 2.12 2.13 2.14 2.15 2.16 2.17 2.18 Brandoni, Diego (2010-05-18). "On the systematics of Ortotherium Ameghino (Xenarthra, Tardigrada, Megalonychidae) from the 'Conglomerado osífero' (late Miocene) of Argentina". Journal of Vertebrate Paleontology 30 (3): 975–980. doi:10.1080/02724631003763474. ISSN 0272-4634. Bibcode: 2010JVPal..30..975B. https://doi.org/10.1080/02724631003763474. 
  3. ↑ Torra, R. (1999). Ituzaingó Formation. A key for the interpretation of upper tertiary stratigraphy, Mesopotamia-Chaco Paraná Basin, Argentina. Ciência e Natura, 139–168.
  4. ↑ 4.0 4.1 Ameghino, F. (1883). Sobre una colección de mamíferos fósiles del Piso Mesopotámico de la formación patagónica: recogidos en las barrancas del Paraná por el Profesor Pedro Scalabrini/por Florentino Ameghino.
  5. ↑ 5.0 5.1 Ameghino, Florentino (1891). "Caracteres diagnósticos de cincuenta especies nuevas de mamíferos fósiles argentinos.". Revista Argentina de Historia Natural, 1 3: 129–167. 
  6. ↑ 6.0 6.1 Kraglievich, L. (1923). Un probable descendiente directo del género Hapalops del Oligoceno de Patagonia en la fauna miocena de Entre Ríos. Neohapalops rothi, 9–16.
  7. ↑ 7.0 7.1 7.2 7.3 Brandoni, Diego (2011-01-01). "The Megalonychidae (Xenarthra, Tardigrada) from the late Miocene of Entre Ríos Province, Argentina, with remarks on their systematics and biogeography" (in en). Geobios 44 (1): 33–44. doi:10.1016/j.geobios.2010.06.005. ISSN 0016-6995. Bibcode: 2011Geobi..44...33B. https://www.sciencedirect.com/science/article/pii/S0016699510001117. 
  8. ↑ Bargo, M. Susana; Iuliis, Gerardo De; Toledo, Néstor (2019-12-20). "EARLY MIOCENE SLOTHS (XENARTHRA, FOLIVORA) FROM THE RÍO SANTA CRUZ VALLEY (SOUTHERN PATAGONIA, ARGENTINA). AMEGHINO, 1887 REVISITED" (in en). Publicación Electrónica de la Asociación Paleontológica Argentina 19 (2). doi:10.5710/PEAPA.06.08.2019.297. ISSN 2469-0228. https://www.peapaleontologica.org.ar/index.php/peapa/article/view/297. 
  9. ↑ Fernicola, J. C. (2011). 1886-1888: Ascenso, auge y caída de la sociedad entre Florentino Ameghino y Francisco P. Moreno.
  10. ↑ Podgorny, Irina (2005). "Bones and Devices in the Constitution of Paleontology in Argentina at the End of the Nineteenth Century" (in en). Science in Context 18 (2): 249–283. doi:10.1017/S0269889705000475. ISSN 1474-0664. https://www.cambridge.org/core/journals/science-in-context/article/abs/bones-and-devices-in-the-constitution-of-paleontology-in-argentina-at-the-end-of-the-nineteenth-century/D1192B7FC32665A4F4879F9F33D89134. 
  11. ↑ Morrone, J. J. (2011). La teoría biogeográfica de Florentino Ameghino y el carácter episódico de la evolución geobiótica de los mamíferos terrestres de América del Sur. Vida y obra de Florentino Ameghino (Fernícola, JC; Prieto, AR; Lazo, DG, 81–89.
  12. ↑ Bordas, A. F. (1942). Observaciones sobre algunos Nothrotheriinae (Gravigrada). Physis, 19(52), 173–179.
  13. ↑ 13.0 13.1 Brandoni, D. (2008). Nuevos materiales de Ortotheriinae (Xenarthra, Tardigrada, Megalonychidae) procedentes del" Mesopotamiense"(Mioceno tardío) de Entre Ríos.
  14. ↑ 14.0 14.1 14.2 14.3 14.4 Brandoni, D. (2014). A new genus of Megalonychidae (Mammalia, Xenarthra) from the late Miocene of Argentina. Revista Brasileira de Paleontologia, 17, 33–42.
  15. ↑ McAfee, Robert; Beery, Sophia; Rimoli, Renato; Almonte, Juan; Lehman, Phillip; Cooke, Siobhan (2021-08-31). "New species of the ground sloth Parocnus from the late Pleistocene-early Holocene of Hispaniola" (in en). Vertebrate Anatomy Morphology Palaeontology 9 (1). doi:10.18435/vamp29369. ISSN 2292-1389. https://journals.library.ualberta.ca/vamp/index.php/VAMP/article/view/29369. 
  16. ↑ McDonald, H. Gregory; Stafford, Thomas W.; Gnidovec, Dale M. (2015). "Youngest Radiocarbon Age for Jefferson's Ground Sloth, Megalonyx Jeffersonii (Xenarthra, Megalonychidae)" (in en). Quaternary Research 83 (2): 355–359. doi:10.1016/j.yqres.2014.11.006. ISSN 0033-5894. Bibcode: 2015QuRes..83..355G. https://www.cambridge.org/core/journals/quaternary-research/article/abs/youngest-radiocarbon-age-for-jeffersons-ground-sloth-megalonyx-jeffersonii-xenarthra-megalonychidae/6C7A769A0C5CE333C5A9810DBCA5168F. 
  17. ↑ Carlini, A. A., & Scillato-Vane, G. J. (2004). The oldest Megalonychidae (Xenarthra: Tardigrada); phylogenetic relationships and an emended diagnosis of the family. Neues Jahrbuch für Geologie und Paläontologie-Abhandlungen, 423–443.
  18. ↑ 18.0 18.1 MacPhee, R. D. E., & Iturralde-Vinent, M. (1995). Origin of the Greater Antillean land mammal fauna. 1, New Tertiary fossils from Cuba and Puerto Rico. American Museum novitates; no. 3141.
  19. ↑ Stinnesbeck, Sarah R.; Frey, Eberhard; Olguín, Jerónimo Avíles; Stinnesbeck, Wolfgang; Zell, Patrick; Mallison, Heinrich; González González, Arturo; Aceves Núñez, Eugenio et al. (2017). "Xibalbaonyx oviceps, a new megalonychid ground sloth (Folivora, Xenarthra) from the Late Pleistocene of the Yucatán Peninsula, Mexico, and its paleobiogeographic significance" (in en). PalZ 91 (2): 245–271. doi:10.1007/s12542-017-0349-5. ISSN 0031-0220. http://link.springer.com/10.1007/s12542-017-0349-5. 
  20. ↑ McDonald, H. G., Harington, C. R., & De Iuliis, G. (2000). The ground sloth Megalonyx from Pleistocene deposits of the Old Crow Basin, Yukon, Canada. Arctic, 213–220.
  21. ↑ Toledo, N.; Bargo, M. S.; Vizcaíno, S. F.; Iuliis, G. De; Pujos, F. (2015). "Evolution of body size in anteaters and sloths (Xenarthra, Pilosa): phylogeny, metabolism, diet and substrate preferences" (in en). Earth and Environmental Science Transactions of the Royal Society of Edinburgh 106 (4): 289–301. doi:10.1017/S1755691016000177. ISSN 1755-6910. 
  22. ↑ 22.0 22.1 Toledo, Néstor (2012-09-07). "Mass estimation of Santacrucian sloths from the Early Miocene Santa Cruz Formation of Patagonia, Argentina" (in en). Acta Palaeontologica Polonica. doi:10.4202/app.2012.0009. ISSN 0567-7920. 
  23. ↑ Presslee, Samantha; Slater, Graham J.; Pujos, François; Forasiepi, Analía M.; Fischer, Roman; Molloy, Kelly; Mackie, Meaghan; Olsen, Jesper V. et al. (2019). "Palaeoproteomics resolves sloth relationships" (in en). Nature Ecology & Evolution 3 (7): 1121–1130. doi:10.1038/s41559-019-0909-z. ISSN 2397-334X. PMID 31171860. https://www.nature.com/articles/s41559-019-0909-z. 
  24. ↑ Varela, L., Tambusso, P. S., McDonald, H. G., & Fariña, R. A. (2019). Phylogeny, macroevolutionary trends and historical biogeography of sloths: insights from a Bayesian morphological clock analysis. Systematic Biology, 68(2), 204–218.
  25. ↑ 25.0 25.1 25.2 McDonald, H. Gregory; Carranza-Castañeda, Oscar (2017). "Increased xenarthran diversity of the Great American Biotic Interchange: a new genus and species of ground sloth (Mammalia, Xenarthra, Megalonychidae) from the Hemphillian (late Miocene) of Jalisco, Mexico" (in en). Journal of Paleontology 91 (5): 1069–1082. doi:10.1017/jpa.2017.45. ISSN 0022-3360. Bibcode: 2017JPal...91.1069M. 
  26. ↑ 26.0 26.1 Brandoni, D., Scillato Yané, G. J., Miño Boilini, Á. R., & Favotti, S. E. (2016). Los Tardigrada (Mammalia, Xenarthra) de Argentina: diversidad, evolución y biogeografía. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"
  27. ↑ 27.0 27.1 Bargo, M. Susana; Vizcaíno, Sergio F.; Kay, Richard F. (2009-09-12). "Predominance of orthal masticatory movements in the Early Miocene Eucholaeops (Mammalia, Xenarthra, Tardigrada, Megalonychidae) and other megatherioid sloths". Journal of Vertebrate Paleontology 29 (3): 870–880. doi:10.1671/039.029.0324. ISSN 0272-4634. Bibcode: 2009JVPal..29..870B. https://doi.org/10.1671/039.029.0324. 
  28. ↑ 28.0 28.1 Semken, Holmes A.; Gregory McDonald, H.; Graham, Russell W.; Adrain, Tiffany; Artz, Joe Alan; Baker, Richard G.; Bryk, Alexander B.; Brenzel, David J. et al. (2022-06-30). "Paleobiology of Jefferson's Ground Sloth (Megalonyx jeffersonii) derived from three contemporaneous, ontogenetically distinct individuals recovered from Southwestern Iowa, U.S.A.". Journal of Vertebrate Paleontology 42 (1): e2124115. doi:10.1080/02724634.2022.2124115. ISSN 0272-4634. Bibcode: 2022JVPal..42E4115S. 
  29. ↑ Scillato-Yane, G. (2013). Nuevo Megalonychidae (Edentata, Tardigrada) del "Mesopotamiense" (Mioceno, Tardioplioceo) de la Provincia de Entre Rios. Ameghiniana, 17(3), 193–199.
  30. ↑ 30.0 30.1 30.2 30.3 Stock, C. (1925). Cenozoic gravigrade edentates of Western North America: with special reference to the Pleistocene Megalonychinae and Mylodontidae of Rancho La Brea (No. 331). Carnegie Institution of Washington.
  31. ↑ Hirschfeld, S. E., Webb, S. D., & Webb, S. D. (1968). Plio-Pleistocene megalonychid sloths of North America: Bulletin of the Florida State Museum.
  32. ↑ Matthew, W. D., Couto, C. D. P., & Simpson, G. G. (1959). The Cuban edentates. Bulletin of the AMNH; v. 117, article 1.
  33. ↑ Pujos, F., De Iuliis, G., Argot, C., & Werdelin, L. (2007). A peculiar climbing Megalonychidae from the Pleistocene of Peru and its implication for sloth history. Zoological Journal of the Linnean Society, 149(2), 179–235.
  34. ↑ Brandoni, D. (2013). Los mamíferos continentales del “Mesopotamiense”(Mioceno tardío) de Entre Ríos, Argentina. Diversidad, edad y paleobiogeografía. Publicación Electrónica de la Asociación Paleontológica Argentina, 14(1).
  35. ↑ 35.0 35.1 Cione, A. L., Azpelicueta, M. D. L. M., Bond, M., Carlini, A. A., Casciotta, J. R., Cozzuol, M. A., ... & Vucetich, M. G. (2000). Miocene vertebrates from Entre Ríos province, eastern Argentina. El Neógeno de Argentina. Serie Correlación Geológica, 14, 191–237.
  36. ↑ Wesselingh, F., & Salo, J. A. (2006). A Miocene perspective on the evolution of the Amazonian biota. Scripta Geologica, 133, 439–458.
  37. ↑ Hoorn, C., Boschman, L. M., Kukla, T., Sciumbata, M., & Val, P. (2022). The Miocene wetland of western Amazonia and its role in Neotropical biogeography. Botanical Journal of the Linnean Society, 199(1), 25–35.
  38. ↑ 38.0 38.1 Brea, Mariana; Zucol, Alejandro F.; Franco, María Jimena (2013-05-15). "A new Bambusoideae (Poaceae: Bambusoideae: Bambuseae: Guaduinae) from the Ituzaingó Formation (Pliocene–Pleistocene), Entre Ríos, Argentina" (in en). Review of Palaeobotany and Palynology 192: 1–9. doi:10.1016/j.revpalbo.2012.12.006. ISSN 0034-6667. Bibcode: 2013RPaPa.192....1B. https://www.sciencedirect.com/science/article/pii/S0034666713000080. 
  39. ↑ Aceñolaza, F. G. (2000). La Formación Paraná (Mioceno medio): estratigrafía, distribución regional y unidades equivalentes. El Neógeno de Argentina. Serie Correlación Geológica, 14, 9–28.
  40. ↑ Candela, Adriana M.; Bonini, Ricardo A.; Noriega, Jorge I. (2012-11-01). "First continental vertebrates from the marine Paraná Formation (Late Miocene, Mesopotamia, Argentina): Chronology, biogeography, and paleoenvironments" (in en). Geobios 45 (6): 515–526. doi:10.1016/j.geobios.2012.05.003. ISSN 0016-6995. Bibcode: 2012Geobi..45..515C. https://www.sciencedirect.com/science/article/pii/S0016699512000824. 
  41. ↑ Franco, M. J., & Brea, M. (2010). Microlobiusxylon paranaensis gen. et sp. nov.(Fabaceae, Mimosoideae) from the Pliocene-Pleistocene of Ituzaingó formation, Paraná basin, Argentina.
  42. ↑ Gutstein, C. S., Cozzuol, M. A., & Pyenson, N. D. (2014). The antiquity of riverine adaptations in Iniidae (Cetacea, Odontoceti) documented by a humerus from the late Miocene of the Ituzaingó Formation, Argentina. The Anatomical Record, 297(6), 1096–1102.
  43. ↑ 43.0 43.1 Moya, Eliana; Brea, Mariana (2020-10-01). "Combretaceous fossil wood from Ituzaingó Formation (Late Miocene?), Argentina, indicate a coastal marine environment" (in en). Review of Palaeobotany and Palynology 281: 104270. doi:10.1016/j.revpalbo.2020.104270. ISSN 0034-6667. Bibcode: 2020RPaPa.28104270M. https://www.sciencedirect.com/science/article/pii/S0034666719303586. 
  44. ↑ Brea, M., & Zucol, A. F. (2007). Guadua zuloagae sp. nov., the first petrified bamboo culm record from the Ituzaingó Formation (Pliocene), Paraná Basin, Argentina. Annals of Botany, 100(4), 711–723.
  45. ↑ Franco, María Jimena (2018-07-04). "Small Celastraceae and Polygonaceae twigs from the Upper Cenozoic (Ituzaingó Formation) of the La Plata Basin, Argentina". Historical Biology 30 (5): 646–660. doi:10.1080/08912963.2017.1313840. ISSN 0891-2963. https://doi.org/10.1080/08912963.2017.1313840. 
  46. ↑ Franco, M. J., & Brea, M. (2015). First extra-Patagonian record of Podocarpaceae fossil wood in the Upper Cenozoic (Ituzaingó Formation) of Argentina. New Zealand Journal of Botany, 53(2), 103–116.
  47. ↑ Tambussi, C. P., Degrange, F., Tambussi, C. P., & Degrange, F. J. (2013). Neogene Birds of South America. South American and Antarctic Continental Cenozoic Birds: Paleobiogeographic Affinities and Disparities, 59–86.
  48. ↑ Cione, A. L., Dahdul, W. M., Lundberg, J. G., & Machado-Allison, A. (2009). Megapiranha paranensis, a new genus and species of Serrasalmidae (Characiformes, Teleostei) from the upper Miocene of Argentina. Journal of Vertebrate Paleontology, 29(2), 350–358.
  49. ↑ Brandoni, D. (2005). Los Megatheriinae (Xenarthra, Tardigrada) de la Formación Ituzaingó (Mioceno Superior-Plioceno) de la provincia de Entre Ríos.
  50. ↑ Brandoni, D., & Carlini, A. A. (2009). On the presence of Pyramiodontherium (Mammalia, Xenarthra, Megatheriidae) in the late Miocene of northeastern Argentina and its biogeographical implications. Revista Italiana di Paleontologia e Stratigrafia, 115.
  51. ↑ Scillato-Yané, G. J. (1981). Nuevo Mylodontinae (Edentata, Tardigrada) del" Mesopotamiense" de la Provincia de Entre Ríos. Ameghiniana, 18(1–2), 29–34.
  52. ↑ Schmidt, G. I. (2013). Los ungulados nativos (Litopterna y Notoungulata: Mammalia) del “Mesopotamiense”(Mioceno Tardío) de Entre Ríos, Argentina. Publicación Electrónica de la Asociación Paleontológica Argentina, 14(1).
  53. ↑ Schmidt, G. I. (2013). Litopterna y Notoungulata (Mammalia) de la Formación Ituzaingó (Mioceno tardío-Plioceno) de la Provincia de Entre Ríos: sistemática, bioestratigrafía y paleobiogeografía (Doctoral dissertation, Universidad Nacional de La Plata).
  54. ↑ 54.0 54.1 Scillato, G. J. (2013). Los Cingulata (Mammalia, Xenarthra) del “Conglomerado Osífero”(Mioceno tardío) de la Formación Ituzaingó de Entre Ríos, Argentina. Publicación Electrónica de la Asociación Paleontológica Argentina, 14(1).
  55. ↑ Góis, Flávio; Scillato-Yané, Gustavo Juan; Carlini, Alfredo Armando; Guilherme, Edson (2013-06-01). "A new species of Scirrotherium Edmund & Theodor, 1997 (Xenarthra, Cingulata, Pampatheriidae) from the late Miocene of South America". Alcheringa: An Australasian Journal of Palaeontology 37 (2): 177–188. doi:10.1080/03115518.2013.733510. ISSN 0311-5518. Bibcode: 2013Alch...37..177G. https://doi.org/10.1080/03115518.2013.733510. 
  56. ↑ Alvarenga, Herculano M.F.; Höfling, Elizabeth (2003). "Systematic revision of the Phorusrhacidae (Aves: Ralliformes)" (in en). Papéis Avulsos de Zoologia 43 (4). doi:10.1590/S0031-10492003000400001. ISSN 0031-1049. 
  57. ↑ Babot, J. M., & Ortiz, P. E. (2009). Primer registro de Borhyaenoidea (Mammalia, Metatheria, Sparassodonta) en la provincia de Tucumán (Formación India Muerta, Grupo Choromoro; Mioceno tardío). Acta Geológica Lilloana, 34–48.
  58. ↑ Riff, D., Romano, P. S. R., Oliveira, G. R., & Aguilera, O. A. (2010). Neogene crocodile and turtle fauna in northern South America. Amazonia. Landscapes and Species Evolution: A Look Into the Past, 259–280.

Wikidata ☰ Q118145239 entry



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