Ahytherium
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''Ahytherium'' is an extinct genus of
megalonychid Megalonychidae is an extinct family of sloths including the extinct ''Megalonyx''. Megalonychids first appeared in the early Oligocene, about 35 million years (Ma) ago, in southern Argentina (Patagonia). There is actually one possible find dating ...
sloth Sloths are a group of Neotropical xenarthran mammals constituting the suborder Folivora, including the extant arboreal tree sloths and extinct terrestrial ground sloths. Noted for their slowness of movement, tree sloths spend most of their li ...
that lived during the Pleistocene of what is now
Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin America. At and with over 217 million people, Brazil is the world's fifth-largest country by area ...
. It contains a single species, ''A. aureum''.


Discovery and taxonomy

The almost-complete skeleton of ''Ahytherium'' alongside remains another extinct sloth species, ''
Australonyx ''Australonyx'' is an extinct genus of ground sloths, endemic to South America during the Late Pleistocene. It was found in Brazil. Discovery The holotype A holotype is a single physical example (or illustration) of an organism, known to ...
'', were discovered in Poço Azul, an underwater cave in
Chapada Diamantina National Park The Chapada Diamantina National Park (; pt, Parque Nacional da Chapada Diamantina) is a national park in the Chapada Diamantina region of the State of Bahia, Brazil. The terrain is rugged, and mainly covered by flora of the Caatinga biome. Loca ...
in 2005. It was described by Castor Cartelle of Pontifícia Universidade Católica de Minas Gerais. The bones, which had a length of about when put together, belong to an animal which presumably was still growing.


Description

This animal is known from well-preserved and nearly complete fossils, including a skeleton of an immature specimen, which in life must have been about three meters long, one meter tall and weighing perhaps half a ton. Like all ground sloths, ''Ahytherium'' was equipped with a robust body and legs equipped with powerful claws. ''Ahytherium'' was similar to other ground sloths such as ''
Megalonyx ''Megalonyx'' (Greek, "large claw") is an extinct genus of ground sloths of the family Megalonychidae, native to North America during the Pliocene and Pleistocene epochs. It became extinct during the Quaternary extinction event at the end of the ...
'' but possessed some features that clearly distinguished it: the head, for example, was extremely short and tall, and possessed dorsally swollen frontal bones. The zygomatic arches were wide, particularly the frontal processes, while the lacrimal bones were narrow, blade-shaped and directed anterolaterally. The dentition included
canine teeth In mammalian oral anatomy, the canine teeth, also called cuspids, dog teeth, or (in the context of the upper jaw) fangs, eye teeth, vampire teeth, or vampire fangs, are the relatively long, pointed teeth. They can appear more flattened howeve ...
that were thin, curved and oval in cross-section. The
humerus The humerus (; ) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius and ulna, and consists of three sections. The humeral upper extremity consists of a roun ...
was equipped with a poorly developed deltopectoral crest, while the
femur The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates with ...
possessed a large
trochanter A trochanter is a tubercle of the femur near its joint with the hip bone. In humans and most mammals, the trochanters serve as important muscle attachment sites. Humans are known to have three trochanters, though the anatomic "normal" includes ...
located distally. The shape of the caudal vertebrae indicates that the tail was dorsoventrally flattened.


Classification

''Ahytherium'' is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of the extinct
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
Megalonychidae Megalonychidae is an extinct family of sloths including the extinct ''Megalonyx''. Megalonychids first appeared in the early Oligocene, about 35 million years (Ma) ago, in southern Argentina (Patagonia). There is actually one possible find dating ...
within the
suborder Order ( la, ordo) is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between family and class. In biological classification, the order is a taxonomic rank used in the classification of organisms and ...
Folivora Sloths are a group of Neotropical xenarthran mammals constituting the suborder Folivora, including the extant arboreal tree sloths and extinct terrestrial ground sloths. Noted for their slowness of movement, tree sloths spend most of their li ...
and the superorder
Xenarthra Xenarthra (; from Ancient Greek ξένος, xénos, "foreign, alien" + ἄρθρον, árthron, "joint") is a major clade of placental mammals native to the Americas. There are 31 living species: the anteaters, tree sloths, and armadillos. Ex ...
. The Megalonychidae represent a very diverse group. The closest relatives of the Megalonychidae are the
Megatheriidae Megatheriidae is a family of extinct ground sloths that lived from approximately 23 mya—11,000 years ago. Megatheriids appeared during the Late Oligocene (Deseadan in the SALMA classification), some 29 million years ago, in South America ...
and the
Nothrotheriidae Nothrotheriidae is a family of extinct ground sloths that lived from approximately 17.5 mya—10,000 years ago, existing for approximately . Previously placed within the tribe Nothrotheriini or subfamily Nothrotheriinae within Megatheriidae, they ...
. The former include the largest known representatives of the sloths, the latter consist of rather smaller members of the sloths. All three families together are relegated to the superfamily of
Megatherioidea Sloths are a group of Neotropical xenarthran mammals constituting the suborder Folivora, including the extant arboreal tree sloths and extinct terrestrial ground sloths. Noted for their slowness of movement, tree sloths spend most of their li ...
. Within the sloths, the Megalonychidae form a very old lineage, with the oldest known fossils dating to the
Oligocene The Oligocene ( ) is a geologic epoch of the Paleogene Period and extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that define the epoch are well identified but the ...
of
Patagonia Patagonia () refers to a geographical region that encompasses the southern end of South America, governed by Argentina and Chile. The region comprises the southern section of the Andes Mountains with lakes, fjords, temperate rainforests, and gl ...
. Characteristic features are found in the
caniniform In mammalian oral anatomy, the canine teeth, also called cuspids, dog teeth, or (in the context of the upper jaw) fangs, eye teeth, vampire teeth, or vampire fangs, are the relatively long, pointed teeth. They can appear more flattened howeve ...
and incisiform design of the anterior tooth in each case, as well as in the molar-like (molariform) designed posterior teeth. The latter are characterized by two transverse ridges (
bilophodont The molars or molar teeth are large, flat teeth at the back of the mouth. They are more developed in mammals. They are used primarily to grind food during chewing. The name ''molar'' derives from Latin, ''molaris dens'', meaning "millstone to ...
) on the occlusal surface, which points to a rather leaf-eating diet in the Megalonychidae. Unlike the Megatheriidae and the Nothrotheriidae, the hind foot is
plantigrade 151px, Portion of a human skeleton, showing plantigrade habit In terrestrial animals, plantigrade locomotion means walking with the toes and metatarsals flat on the ground. It is one of three forms of locomotion adopted by terrestrial mammals. T ...
in shape and not twisted, thus retaining its original shape.H. Gregory McDonald und Gerardo de Iuliis: ''Fossil history of sloths.'' In: Sergio F. Vizcaíno und W. J. Loughry (eds.): ''The Biology of the Xenarthra.'' University Press of Florida, 2008, S. 39–55. The Megalonychidae were once widespread, with fossil remains found in both
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the southe ...
and
Central America Central America ( es, América Central or ) is a subregion of the Americas. Its boundaries are defined as bordering the United States to the north, Colombia to the south, the Caribbean Sea to the east, and the Pacific Ocean to the west. ...
, as well as in
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...
as far north as the
Arctic The Arctic ( or ) is a polar regions of Earth, polar region located at the northernmost part of Earth. The Arctic consists of the Arctic Ocean, adjacent seas, and parts of Canada (Yukon, Northwest Territories, Nunavut), Danish Realm (Greenla ...
. In a classic view based on skeletal anatomy comparisons-based view, the Megalonychidae originally included the sloths of the
West Indies The West Indies is a subregion of North America, surrounded by the North Atlantic Ocean and the Caribbean Sea that includes 13 independent island countries and 18 dependencies and other territories in three major archipelagos: the Greater A ...
as well as the extant
two-toed sloth ''Choloepus'' is a genus of xenarthran mammals of Central and South America within the monotypic family Choloepodidae, consisting of two-toed sloths, sometimes also called two-fingered sloths. The two species of ''Choloepus'' (which means "lame ...
s of the genus ''Choloepus'',Timothy J. Gaudrin: ''Phylogenetic relationships among sloths (Mammalia, Xenarthra, Tardigrada): the craniodental evidence.'' Zoological Journal of the Linnean Society 140, 2004, S. 255–305. which are now considered to belong to the superfamily
Mylodontoidea Sloths are a group of Neotropical xenarthran mammals constituting the suborder Folivora, including the extant arboreal tree sloths and extinct terrestrial ground sloths. Noted for their slowness of movement, tree sloths spend most of their li ...
.Frédéric Delsuc, Melanie Kuch, Gillian C. Gibb, Emil Karpinski, Dirk Hackenberger, Paul Szpak, Jorge G. Martínez, Jim I. Mead, H. Gregory McDonald, Ross D.E. MacPhee, Guillaume Billet, Lionel Hautier und Hendrik N. Poinar: ''Ancient mitogenomes reveal the evolutionary history and biogeography of sloths.'' Current Biology 29 (12), 2019, S. 2031–2042, doi:10.1016/j.cub.2019.05.043Samantha Presslee, Graham J. Slater, François Pujos, Analía M. Forasiepi, Roman Fischer, Kelly Molloy, Meaghan Mackie, Jesper V. Olsen, Alejandro Kramarz, Matías Taglioretti, Fernando Scaglia, Maximiliano Lezcano, José Luis Lanata, John Southon, Robert Feranec, Jonathan Bloch, Adam Hajduk, Fabiana M. Martin, Rodolfo Salas Gismondi, Marcelo Reguero, Christian de Muizon, Alex Greenwood, Brian T. Chait, Kirsty Penkman, Matthew Collins und Ross D. E. MacPhee: ''Palaeoproteomics resolves sloth relationships.'' Nature Ecology & Evolution 3, 2019, S. 1121–1130, doi:10.1038/s41559-019-0909-z
Molecular genetic Molecular genetics is a sub-field of biology that addresses how differences in the structures or expression of DNA molecules manifests as variation among organisms. Molecular genetics often applies an "investigative approach" to determine the ...
However, studies together with
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, respo ...
analyses failed to establish closer relationships between these individual groups. Much of the fossil material of Megalonychidae is fragmentary and largely incomplete. The systematic Relationships of the individual representatives of the group could be worked out therefore so far only insufficiently. However, due to the richness of forms, different lines of development can be traced. One includes largely South American representatives such as ''
Megistonyx ''Megistonyx'' is an extinct genus of ground sloth endemic to South America during the Late Pleistocene (Lujanian). It is known from one skeleton collected in the Andes of Venezuela, and is closely related to '' Ahytherium''. History of disco ...
'' or ''Ahytherium'' and '' Ortotherium'', respectively, another consists of the North American forms ''
Megalonyx ''Megalonyx'' (Greek, "large claw") is an extinct genus of ground sloths of the family Megalonychidae, native to North America during the Pliocene and Pleistocene epochs. It became extinct during the Quaternary extinction event at the end of the ...
'' and ''
Pliometanastes ''Pliometanastes'' is an extinct genus of ground sloths of the family Megalonychidae endemic to North America during the Late Miocene epoch through very early Pliocene epoch (Hemphillian in the NALMA classification). Its fossils have been found ...
'' (here, since based on skeletal features, also Caribbean sloths such as ''
Megalocnus ''Megalocnus'' ("great sloth" in Greek) is a genus of extinct large ground sloths that were native to Cuba during the Pleistocene and Holocene epochs. They were among the largest of the Caribbean sloths (Megalocnidae), with individuals estimate ...
'' or ''
Neocnus ''Neocnus'' is an extinct genus of ground sloth, whose species ranged across Cuba and Hispaniola (today split between Haiti and the Dominican Republic). ''Neocnus'' would have resembled a typical ground sloth, though much smaller, with a lon ...
''). Currently, however, it is not possible to determine direct ancestors for the North American representatives of Megalonychidae. Consequently, their relationship to the South American forms is rather unknown. Below is a phylogenetic tree of the Megalonychidae, based on the work of Stinnesbeck and colleagues (2021).


Paleobiology

''Ahytherium'' must have been a large, rather slow-moving terrestrial sloth that could defend itself from carnivores with its powerful claws. The flattened shape of the tail suggests that ''Ahytherium'' might have been a good swimmer. The related ''Megistonyx,'' on the other hand, would seem to have adapted to high-altitude environments. Another swimming sloth, more specialized for marine life is ''
Thalassocnus ''Thalassocnus'' is an extinct genus of semiaquatic ground sloths from the Miocene and Pliocene of the Pacific South American coast. It is monotypic within the subfamily Thalassocninae. The five species—''T. antiquus'', ''T. natans'', ''T. lit ...
''.


See also

* ''
Thalassocnus ''Thalassocnus'' is an extinct genus of semiaquatic ground sloths from the Miocene and Pliocene of the Pacific South American coast. It is monotypic within the subfamily Thalassocninae. The five species—''T. antiquus'', ''T. natans'', ''T. lit ...
''


References

{{Taxonbar, from=Q4696453 Prehistoric sloths Pleistocene xenarthrans Prehistoric placental genera Holocene extinctions Pleistocene mammals of South America Lujanian Pleistocene Brazil Fossils of Brazil Fossil taxa described in 2008 Monotypic mammal genera