Hilarcotherium Skull
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''Hilarcotherium'' is an extinct genus of astrapotheriid
mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...
s that lived in South America during the
Middle Miocene The Middle Miocene is a sub-epoch of the Miocene Epoch made up of two stages: the Langhian and Serravallian stages. The Middle Miocene is preceded by the Early Miocene. The sub-epoch lasted from 15.97 ± 0.05 Ma to 11.608 ± 0.005 Ma (million y ...
( Laventan). The type species is ''H. castanedaii'', found in sediments of the
La Victoria Formation The Honda Group ( es, Grupo Honda, Tsh, Ngh) is a geological group of the Upper and Middle Magdalena Basins and the adjacent Central and Eastern Ranges of the Colombian Andes. The group, in older literature also defined as formation, is in its ...
, part of the
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in the department of Tolima in
Colombia Colombia (, ; ), officially the Republic of Colombia, is a country in South America with insular regions in North America—near Nicaragua's Caribbean coast—as well as in the Pacific Ocean. The Colombian mainland is bordered by the Car ...
. In 2018, Carrillo ''et al.'' described a partial skull and mandible of a second species ''H. miyou'' from the
Castilletes Formation The Castilletes Formation ( es, Formación Castilletes, N1c) is a fossiliferous geological formation of the Cocinetas Basin in the northernmost department of La Guajira, Colombia. The formation consists of fossiliferous mudstones, siltstones an ...
in the Cocinetas Basin of northern Colombia, and estimated the body weight of the animal at .Carillo, 2018, p.142


Discovery

The remains of ''Hilarcotherium castanedaii'' were discovered by José Alfredo Castañeda, who found them in the Malnombre Creek in the Hilarco village located near the town of Purificación, in Tolima. This area, in the valley of the Upper Magdalena River, correspond to the La Victoria Formation, that along with the Villavieja Formation makes the
Honda Group is a Japanese public multinational conglomerate manufacturer of automobiles, motorcycles, and power equipment, headquartered in Minato, Tokyo, Japan. Honda has been the world's largest motorcycle manufacturer since 1959, reaching a product ...
, dating from the Middle Miocene, made between 13-11.8 million years ago and it has provided a remarkable fossil site known as La Venta, although the lack of key fossils does not allow accurate establishment of the membership of the Hilarco site to the formation. The remains of ''Hilarcotherium'' were collected by dr. María Páramo and Gerardo Vargas, who took them to the Geological Museum José Royo y Gómez of
Ingeominas The Colombian Geological Survey (CGS) ( es, Servicio Geológico Colombiano; formerly known as INGEOMINAS) is a scientific agency of the Colombian government in charge of contributing to the socioeconomic development of the nation through research ...
(former name of Colombian Geological Service) in
Bogotá Bogotá (, also , , ), officially Bogotá, Distrito Capital, abbreviated Bogotá, D.C., and formerly known as Santa Fe de Bogotá (; ) during the Spanish period and between 1991 and 2000, is the capital city of Colombia, and one of the larges ...
for study and preservation. In February 2015 these remains were officially named, although the name ''Hilarcotherium castanedaii'' was released earlier and informally in conference abstracts. The genus name is derived from the Hilarco site and the term in Latinized Greek ''therium'', meaning "beast", while the species name ''castanedaii'' is in recognition to Castañeda, by finding and reporting the find. A second species, ''H. miyou'' was described by Carrillo ''et al''. in 2018. The remains of ''H. miyou'' were discovered in Patajau, in the Castilletes Formation of
La Guajira department La Guajira () is a department of Colombia. It occupies most of the Guajira Peninsula in the northeast region of the country, on the Caribbean Sea and bordering Venezuela, at the northernmost tip of South America. The capital city of the departm ...
, on northern Colombia. The deposits of Castilletes corresponds to the older SALMAs
Santacrucian The Santacrucian age is a period of geologic time (17.5 – 16.3 Ma) within the Early Miocene epoch of the Neogene, used more specifically with SALMA classification in South America. It follows the Colhuehuapian and precedes the Friasian age. ...
-
Colloncuran The Colloncuran ( es, Colloncurense) age is a period of geologic time (15.5–13.8 Ma) within the Middle Miocene epoch of the Neogene, used more specifically within the SALMA classification in South America. It follows the Friasian and precedes ...
, 16.7–14.2 million years ago during the early Middle Miocene. The remains the holotype, IGMp 881327, consists on a partial mandible with the left ramus, some of the molars and the canines, the left condylar process, a second upper molar (M2) and the distal portion of the femur. Also were referred to ''H. miyou'' the specimens MUN-STRI 34216, a fragmentary skull with portion of the occipitals, palatines, and left upper canine, an upper fourth premolar P4 and the second upper M2, and a fragmentary mandibular symphysis with the base of the lower canines and alveoli for left three incisives (i3, i2, and i1) and the right first incisives (i1 and i2); and MUN-STRI 38073, which consists in a left P4 and upper molar fragments. Also were discovered a series of postcranial remains, including humerus, radii, vertebrae, patella, sacrum, scapulae, femora and rib fragments. Since that these fossils were not associated with dental remains is not possible refer them to ''H. miyou'', but their large size and the fact that ''Hilarcotherium'' is the only astrapotherid taxon recognized in Castilletes allow classify them tentatively as ''Hilarcotherium cf. miyou''. The name of this species derived from the word ''miyo’u'' that means "large" in the
Wayuunaiki Wayuu ( guc, Wayuunaiki ), or Guajiro, is a major Arawakan language spoken by 305,000 indigenous Wayuu people in northwestern Venezuela and northeastern Colombia on the Guajira Peninsula. There are 200,000 speakers of Wayuu in Venezuela and 1 ...
language of the Wayuu people of La Guajira. An additional discovery was made near to the wetlands of Zapatosa, in the municipality of Chimichagua in the
Cesar department Caesar Department ( es, Departamento del Cesar, links=no) or simply Caesar () is a department of Colombia located in the north of the country in the Caribbean region, bordering to the north with the Department of La Guajira, to the west with th ...
in northern Colombia. This specimen, a fragment of maxilar with a premolar (P4) and the first upper molar (M1) is catalogued as SGCMGJRG2018.V.7 and is housed in the Geological Museum Jose Royo y Gómez, which were donated by the discoverers of the specimen, Hasmet Florián and José Martínez. The site belongs to the Miocene but without a precise datation; the characteristics of the teeth support the assignation to ''Hilarcotherium'', but since that it shows some differences with the recognised species of the genus, is only classified as ''Hilarcotherium'' sp.


Description

The holotype specimen of ''Hilarcotherium castanedaii'' is named IGM p881231. This consists of fragments of skull, a partial jaw, the vertebral ramus of a dorsal
rib In vertebrate anatomy, ribs ( la, costae) are the long curved bones which form the rib cage, part of the axial skeleton. In most tetrapods, ribs surround the chest, enabling the lungs to expand and thus facilitate breathing by expanding the ches ...
, a complete left
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 ...
and an incisor tooth associated. The skull includes the rostrum area, the palate with the fourth
premolar The premolars, also called premolar teeth, or bicuspids, are transitional teeth located between the canine and molar teeth. In humans, there are two premolars per quadrant in the permanent set of teeth, making eight premolars total in the mouth ...
(P4) and the three upper molars (M1-M2-M3) plus part of the zygomatic arch and the
braincase In human anatomy, the neurocranium, also known as the braincase, brainpan, or brain-pan is the upper and back part of the skull, which forms a protective case around the brain. In the human skull, the neurocranium includes the calvaria or skul ...
, but lacks the upper canines and the top portion of the skull. The premaxilla shows no sign of having teeth, as in other astrapotheres. The jaw missing the incisors, the crown of the fourth
premolar The premolars, also called premolar teeth, or bicuspids, are transitional teeth located between the canine and molar teeth. In humans, there are two premolars per quadrant in the permanent set of teeth, making eight premolars total in the mouth ...
and the left mandibular ramus, but has the roots of the teeth of the right side and a broken canine with an oval cross section. It has however the sockets for three incisors on each side of the jaw, which is a
plesiomorphic In phylogenetics, a plesiomorphy ("near form") and symplesiomorphy are synonyms for an ancestral character shared by all members of a clade, which does not distinguish the clade from other clades. Plesiomorphy, symplesiomorphy, apomorphy, and ...
feature, as its other relatives as '' Xenastrapotherium'' had two. In 2018, Carillo ''et al.'' described a partial skull and mandibule of a second species ''H. miyou'' from the
Castilletes Formation The Castilletes Formation ( es, Formación Castilletes, N1c) is a fossiliferous geological formation of the Cocinetas Basin in the northernmost department of La Guajira, Colombia. The formation consists of fossiliferous mudstones, siltstones an ...
in the Cocinetas Basin of northern Colombia. The humerus of ''H. castanedaii'' measures , with a head of at its widest and oriented backwards with a convex form. Moreover, as in '' Astrapotherium'', the bone thickness decreases as it approaches its lower (distal) end. Analyses performed with his molars and humerus indicate that weight range of ''H. castanedaii'' would be between , which would be comparable to the modern black rhinoceros, close to in length. Between astrapotheres, this size is equivalent to an "intermediate" range, since it is greater than that of genus as ''
Comahuetherium ''Comahuetherium'' is an extinct genus of astrapotherian mammal from the Early Miocene (Colhuehuapian age). It is a basal astrapotheriid which lived in what is now Patagonia, Argentina. The holotype was found in the Cerro Bandera Formation in Ne ...
'' () and ''
Astrapothericulus ''Astrapothericulus'' is an extinct genus of mammals, belonging to the order Astrapotheria. It lived during the Lower Miocene in what is now South America. Description This animal was as large than a tapir, and may have looked like them, as it h ...
'' (), and similar to ''Xenastrapotherium'' and ''Astrapotherium? ruderarium'', but lower than in the larger genera as ''Astrapotherium'' (), '' Granastrapotherium'' () and ''
Parastrapotherium ''Parastrapotherium'' is an extinct genus of South American land mammal that existed from the Late Oligocene (Deseadan SALMA) to the Early Miocene (Colhuehuapian SALMA). The genus includes some of the largest and smallest known astrapotherian, bu ...
'' () This indicates that the large body size is a trait that developed early in the group, but the gigantic size, of more than , evolved only on three separate occasions. ''H. miyou'' had an estimated body weight of .


Phylogeny

''Hilarcotherium'' belongs to the family Astrapotheriidae, the family of advanced
astrapotheres Astrapotheria is an extinct order of South American and Antarctic hoofed mammals that existed from the late Paleocene to the Middle Miocene, ."The uruguaytheriine Astrapotheriidae from the rich middle Miocene Honda Group of the upper Magdalena R ...
characterized by its developed tusks, separated by a diastema of the other teeth, the retracted nasal bones that indicate the presence of a
trunk Trunk may refer to: Biology * Trunk (anatomy), synonym for torso * Trunk (botany), a tree's central superstructure * Trunk of corpus callosum, in neuroanatomy * Elephant trunk, the proboscis of an elephant Computing * Trunk (software), in rev ...
in his snout and the flattened astragalus. Within this group, ''Hilarcotherium'' shares with ''Xenastrapotherium'', ''
Uruguaytherium ''Uruguaytherium'' is an extinct genus of astrapotherid mammal from the Late Oligocene to Early Miocene of South America. It was named by the Argentinean paleontologist Lucas Kraglievich in 1928, from a fragmentary fossil found in the Fray B ...
'' and ''Granastrapotherium'' the lower canines that are horizontally inserted into the jaw, in addition to the characteristics of the molars (without labial cingulum, an inner protuberance of the molars), indicating that ''Hilarcotherium'' and the others genera already mentioned belongs to the subfamily
Uruguaytheriinae Astrapotheriidae is an extinct family of herbivorous South American land mammals that lived from the Late Eocene (Mustersan SALMA) to the Middle Miocene (Laventan SALMA) . The most derived of the astrapotherians, they were also the largest and mo ...
, a group that colonized the equatorial region of South America during the Miocene until their extinction in the middle of such period, in contrast with other astrapotheres only known in the south of the continent. Cladogram based in the phylogenetic analysis published in the original description article by Vallejo-Pareja ''et al''., 2015, showing the position of ''Hilarcotherium'':


References


Bibliography

* {{Taxonbar, from=Q19356902 Meridiungulata Miocene mammals of South America Laventan Friasian Neogene Colombia Fossils of Colombia Honda Group, Colombia Fossil taxa described in 2015 Prehistoric placental genera