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''Kelenkura'' is an
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
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 heavily armored mammals belonging to the subfamily
Glyptodont Glyptodonts are an extinct subfamily of large, heavily armoured armadillos. They arose in South America around 48 million years ago and spread to southern North America after the continents became connected several million years ago. The best-kn ...
inae, from the family
Chlamyphoridae Chlamyphoridae is a family of cingulate mammals. While glyptodonts have traditionally been considered stem-group cingulates outside the group that contains modern armadillos, there had been speculation that the extant family Dasypodidae could b ...
that contain most of the modern
armadillos Armadillos (meaning "little armored ones" in Spanish) are New World placental mammals in the order Cingulata. The Chlamyphoridae and Dasypodidae are the only surviving families in the order, which is part of the superorder Xenarthra, along w ...
. It was a medium-sized South American animal, distantly related to '' Doedicurus''. Fossils of this genus were recovered in the
Arroyo Chasicó Formation Arroyo often refers to: * Arroyo (creek), an intermittently dry creek Arroyo may also refer to: People * Arroyo (surname) Places United States ;California * Arroyo Burro Beach, a public beach park in Santa Barbara County, California * Arroyo ...
and in the Loma de Las Tapias Formation of
Argentina Argentina (), officially the Argentine Republic ( es, link=no, República Argentina), is a country in the southern half of South America. Argentina covers an area of , making it the second-largest country in South America after Brazil, th ...
in rocks dating back to the
Late Miocene The Late Miocene (also known as Upper Miocene) is a sub-epoch of the Miocene epoch (geology), Epoch made up of two faunal stage, stages. The Tortonian and Messinian stages comprise the Late Miocene sub-epoch, which lasted from 11.63 Ma (million ye ...
epoch.


Discovery and etymology

The presence of glyptodonts in the Arroyo Chasicó Formation was known from fragmentary remains since 1926. In 2005, a new, more complete specimen was unearthed from a river bed in the
formation Formation may refer to: Linguistics * Back-formation, the process of creating a new lexeme by removing or affixes * Word formation, the creation of a new word by adding affixes Mathematics and science * Cave formation or speleothem, a secondar ...
. Alfredo E. Zurita and Silvia A. Aramayo described it in 2007 as PV-UNS-260, a well preserved skeleton from the Arroyo Chasicó including a partially complete skull and
carapace A carapace is a Dorsum (biology), dorsal (upper) section of the exoskeleton or shell in a number of animal groups, including arthropods, such as crustaceans and arachnids, as well as vertebrates, such as turtles and tortoises. In turtles and tor ...
, a complete right
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 ...
, several caudal rings and a complete caudal tube, alongside several limb bones and isolated
osteoderms Osteoderms are bony deposits forming scales, plates, or other structures based in the dermis. Osteoderms are found in many groups of extant and extinct reptiles and amphibians, including lizards, crocodilians, frogs, temnospondyls (extinct ...
. They assigned the skeleton to the already existing species ''
Eosclerocalyptus ''Eosclerocalyptus'' is an extinct genus of glyptodont. It lived during the Late Miocene, and its fossilized remains were discovered in South America. Description This animal, like all glyptodonts, was protected by a carapace covering most of ...
'', and estimated it as
Huayquerian The Huayquerian ( es, Huayqueriense) age is a period of geologic time (9.0–6.8 Ma) within the Late Miocene epoch of the Neogene, used more specifically within the SALMA classification. It follows the Mayoan and precedes the Montehermosan age. ...
.> In 2011, new glyptodontid remains from the Loma de Las Tapias Formation, known as PVSJ-366, and including an almost complete crania and a fragment of the left femur, were tentatively assignated by Victor H. Contreras and Juan A. Baraldo to '' Palaehoplophorus'' and '' Hoplophractus'', the latter being now considered to be a
junior synonym The Botanical and Zoological Codes of nomenclature treat the concept of synonymy differently. * In botanical nomenclature, a synonym is a scientific name that applies to a taxon that (now) goes by a different scientific name. For example, Linna ...
of ''Eosclerocalyptus''. In 2016–2017, Cristian G. Oliva examined a number of
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
fragments from the Arroyo Chasicó Formation and suggested the existence of a still undescribed new
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
of glyptodont from the locality. Finally, in 2022, a new study conducted by Daniel Barasoain ''et al'' contested the referral of the Arroyo Chasicó material to ''Eosclerocalyptus'', and named the new genus and species ''Kelenkura castroi'', with PV-UNS-260 as
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of several ...
. Other material previously recovered from the Arroyo Chasicó Formation and the Loma de Las Tapias Formation, including PVSJ-366, were assigned to the genus. The holotype material was also reevaluated as belonging to the Chasicoan period. The genus name, ''Kelenkura'' was constructed on the words ''këlen'', which means "tail", and ''kura'', meaning "rock", in the local
Mapuche language Mapuche (, Mapuche & Spanish: , or Mapudungun; from ' 'land' and ' 'speak, speech') is an Araucanian language related to Huilliche spoken in south-central Chile and west-central Argentina by the Mapuche people (from ''mapu'' 'land' and ''che ...
, referencing the shape of its tail. The species epithet, ''castroi'', honors D. R. Castro, who participated in the discovery of PVSJ-366, one of the complete skulls assigned to the genus.


Description

The skull of ''Kelenkura'' was elongated, with a length of 211 mm for the holotype, with an underdevelloped
sagittal crest A sagittal crest is a ridge of bone running lengthwise along the midline of the top of the skull (at the sagittal suture) of many mammalian and reptilian skulls, among others. The presence of this ridge of bone indicates that there are exceptiona ...
and a narrow occipital area. The few preserved molars were trilobed. The skull was protected by a head shield made of relatively large osteoderms similar to its carapace, which were poorly preserved. The well preserved, 295 mm long
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 ...
of ''Kelenkura'' was intermediate in shape between ancient and more modern genus of austral glyptodonts. ''Kelenkuras total weight in its lifetime was estimated at 160 kg. The carapace, mainly known from the holotype, was made of 35 rows of osteoderms forming a repeated rosette pattern, and was 910 mm high and 1050 mm long. The tail was protected by a caudal armor, composed of caudal rings made of two rows of osteoderm and finished by a completely fused and ornamented caudal tube, known from five complete specimen from the Arroyo Chasicó Formation, and described by its namers as the earliest fully modern caudal tube known for a glyptodont.


Phylogeny

While being originally recovered as a specimen of ''Eosclerocalyptus tapinocephalus'', Kelenkura was erected as an entirely new genus and species on the basis of morphological differences and an earlier age. As a new genus, it stands as the sister group of all the other late
neogene The Neogene ( ), informally Upper Tertiary or Late Tertiary, is a geologic period and system that spans 20.45 million years from the end of the Paleogene Period million years ago ( Mya) to the beginning of the present Quaternary Period Mya. ...
and
quaternary The Quaternary ( ) is the current and most recent of the three periods of the Cenozoic Era in the geologic time scale of the International Commission on Stratigraphy (ICS). It follows the Neogene Period and spans from 2.58 million years ...
glyptodonts from the so-called "Austral lineage", whose late members are distinguished from every other mammals by a characteristic caudal tube. Depicted below is a reproduction of the
phylogenetic tree A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spec ...
presented by Barasoain ''et al'' (2022) for glyptodonts, including the newly described ''Kelenkura''.


Palaeoecology

The Arroyo Chasicó formation was, in the Miocene, on the tip of a peninsula bordered by the Paranaense Sea. ''Kelenkura'' was the only glyptodont from the "Austral lineage" extant in the Chasicoan period, but it lived alongside various genera of Cingulata, including the last horned armadillo ''
Epipeltephilus ''Epipeltephilus'' is an extinct genus of armadillo, belonging to the family Peltephilidae, the "horned armadillos", whose most famous relative was Peltephilus. ''Epipeltephilus'' is the last known member of its family, becoming extinct during th ...
'', the Dasypodidae '' Vetelia'' and '' Chasicotatus'', the
Euphractinae Euphractinae is an armadillo subfamily in the family Chlamyphoridae. Euphractinae are known for having a well developed osteoderm that has large cavities filled with adipose tissue, and more hair follicles with well developed sebaceous glands i ...
''
Proeuphractus ''Proeuphractus'' is an extinct genus of xenarthran, related to the modern armadillos. It lived from the Early to the Late Miocene, and its fossilized remains were discovered in South America. Description This animal was quite similar to the ...
'', and the
pampathere Pampatheriidae ("Pampas beasts") is an extinct family of large plantigrade armored armadillos related to extant armadillos in the order Cingulata. However, pampatheriids have existed as a separate lineage since at least the middle Eocene Mus ...
''
Kraglievichia ''Kraglievichia'' is an extinct genus of cingulate belonging to the family Pampatheriidae. It lived from the Late Miocene to the Early Pliocene, and its fossilized remains were discovered in South America. Description This animal looked like ...
''. Other xenarthrans includes several genera of
ground sloths Ground sloths are a diverse group of extinct sloths in the mammalian superorder Xenarthra. The term is used to refer to all extinct sloths because of the large size of the earliest forms discovered, compared to existing tree sloths. The Carib ...
, such as the
Mylodontidae Mylodontidae is a family of extinct South American and North American ground sloths within the suborder Folivora of order Pilosa The order Pilosa is a clade of xenarthran placental mammals, native to the Americas. It includes the anteaters a ...
''
Octomylodon ''Octomylodon'' is an extinct genus of ground sloth of the family Mylodontidae, living during the Late Miocene (Huayquerian). Fossil remains of ''Octomylodon'' have been found uniquely in the Ituzaingó Formation, Argentina, South America ...
'', the
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 ...
'' Protomegalonyx'', 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 ...
'' Xyophorus'' and ''
Chasicobradys ''Chasicobradys'' is an extinct genus of ground sloths of the family Nothrotheriidae that lived in what is now Argentina. ''Chasicobradys'' was discovered in the Arroyo Chasicó Formation, in Buenos Aires Province. It is only known from jaw fra ...
'', and the Megatheriidae '' Anisodontherium''. Several genera of
rodents Rodents (from Latin , 'to gnaw') are mammals of the order Rodentia (), which are characterized by a single pair of continuously growing incisors in each of the upper and lower jaws. About 40% of all mammal species are rodents. They are nat ...
were recovered from the formation, such as the earliest genus of
tuco-tucos A tuco-tuco is a neotropical rodent in the family Ctenomyidae.Parada, A., G. D’Elia, C.J. Bidau, and E.P. Lessa. 2011. Species Groups and the Evolutionary Diversification of Tuco-Tucos, genus ''Ctenomys'' (Rodentia: Ctenomyidae). ''Journal of M ...
, maras and capybaras such as '' Cardiomys'', '' Procardiomys'' and '' Cardiatherium'', Octodontidae such as '' Chasicomys'' and '' Chasichimys'', the Echimyidae '' Pattersomys'', the plain viscacha ''Lagostomus telenkechanum'' and its relative '' Prolagostomus'', and large-sized Dinomyidae like '' Carlesia''.
Meridiungulates South American native ungulates, commonly abbreviated as SANUs, are extinct ungulate-like mammals of controversial affinities that were indigenous to South America prior to the Great American Biotic Interchange. They comprise five major groups c ...
were also present in the formation, with
Litopterna Litopterna (from grc, λῑτή πτέρνα "smooth heel") is an extinct order of fossil hoofed mammals from the Cenozoic era. The order is one of the five great orders of South American ungulates that were endemic to the continent, until the G ...
genera such as the Macraucheniidae ''
Cullinia ''Cullinia'' is an extinct genus of litoptern, an order of South American native ungulates that included horse-like and camel-like animals such as ''Macrauchenia''. It is only known from fragmentary remains. ''Cullinia levis'' is known from Cha ...
'' and the
Proterotheriidae Proterotheriidae is an extinction, extinct family of fossil ungulates from the Cenozoic era that displays toe reduction. Despite resembling primitive, small horses, they were only distantly related to them, and instead belonged to the native Sou ...
''
Neobrachytherium ''Neobrachytherium'' is an extinct genus of proterotheriid mammal from the Late Miocene of Argentina and Uruguay. It is represented by multiple species, including the type ''N. intermedium'', originally named in 1891 by Moreno and Mercerat as a ...
'', while
notoungulates Notoungulata is an extinct order of mammalian ungulates that inhabited South America from the early Paleocene to the Holocene, living from approximately 61 million to 11,000 years ago. Notoungulates were morphologically diverse, with forms resemb ...
were represented by genera such as the large-sized late surviving
Homalodotheriidae Homalodotheriidae is an extinct family comprising four genera of notoungulate mammals known from the Late Eocene (Tinguirirican) through Late Miocene (Chasicoan) of Argentina and Chile in South America South America is a continent enti ...
'' Chasicotherium'', the Toxodontidae '' Paratrigodon'', the
Interatheriidae Interatheriidae is an extinct family of notoungulate mammals from South America. Interatheriids are known from the Middle Eocene (Mustersan) to the Early Pliocene (Montehermosan).McKenna & Bell, 1997Linares, 2004 These animals were principally sm ...
''
Protypotherium ''Protypotherium'' is an extinct genus of notoungulate mammals native to South America during the Oligocene and Miocene epochs. A number of closely related animals date back further, to the Eocene. Fossils of ''Protypotherium'' have been found in ...
'', the
Mesotheriidae Mesotheriidae ("Middle Beasts") is an extinct family of notoungulate mammals known from the Oligocene through the Pleistocene of South America. Mesotheriids were small to medium-sized herbivorous mammals adapted for digging. Characteristics Meso ...
''
Typotheriopsis ''Typotheriopsis'' is an extinct genus of Notoungulate, belonging to the family Mesotheriidae, which included several small sized Meridiungulates specialized in digging. It is considered as the sister taxon of the clade including ''Mesotherium'' ...
'', and the
Hegetotheriidae Hegetotheriidae is an extinct family of notoungulate mammals known from the Oligocene through the Pliocene of South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a r ...
''
Paedotherium ''Paedotherium'' is an extinct, potentially paraphyletic genus of Notoungulate, belonging to the family Hegetotheriidae, composed of small-sized, rodent or lagomorph-like Meridiungulata, South American ungulates. Four species are unambiguously re ...
'', '' Pseudohegetotherium'' and ''
Hemihegetotherium ''Hemihegetotherium'' is an extinct genus of hegetotheriid notoungulate that lived from the Middle to the Late Miocene of what is now Argentina. Description It was a medium-sized animal, approximately the size of large lowland paca ; it may h ...
''. The largest
predators Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill the ...
were the
Sparassodonta Sparassodonta (from Ancient Greek, Greek to tear, rend; and , gen.
, ' The comma is a punctuation mark that appears in several variants in different languages. It has the same shape as an apostrophe or single closing quotation mark () in many typefaces, but it differs from them in being placed on the baseline ...
tooth) is an extinct order (biology), order of carnivore, carnivorous metatherian mammals native to South America, related to modern marsupials. They were once con ...
'' Pseudolycopsis cabrerai'' and ''
Lycopsis viverensis ''Lycopsis'' is an extinct genus of South American metatherian, that lived during the Miocene in Argentina and Colombia. History Although not named until 1927, Florentino Ameghino described a species now seen as synonymous with ''Lycopsis tor ...
'', and the small
terror bird Phorusrhacids, colloquially known as terror birds, are an extinct clade of large carnivorous flightless birds that were one of the largest species of apex predators in South America during the Cenozoic era; their conventionally accepted temporal ...
'' Psilopterus''.


References

{{Taxonbar, from=Q110588834 Armadillos Prehistoric placental genera Prehistoric cingulates Miocene xenarthrans Tortonian first appearances Miocene genus extinctions Miocene mammals of South America Chasicoan Neogene Argentina Fossils of Argentina Fossil taxa described in 2022