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''Hoffstetterius'' is an extinct
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
toxodontid Toxodontidae is an extinct family of notoungulate mammals, known from the Oligocene to the Holocene (11,000 BP) of South America, with one genus, ''Mixotoxodon'', also known from the Pleistocene of Central America and southwestern North Americ ...
notoungulate 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 ...
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 ...
, belonging to the subfamily
Toxodontinae Toxodontidae is an extinct family of notoungulate mammals, known from the Oligocene to the Holocene (11,000 BP) of South America, with one genus, ''Mixotoxodon'', also known from the Pleistocene of Central America and southwestern North Americ ...
whose remains were discovered in the Middle to Late Miocene (
Mayoan The Mayoan ( es, Mayoense) age is a period of geologic time from 11.8 to 10 Ma, within the Middle to Late Miocene epoch of the Neogene, used more specifically within the SALMA classification in South America. It follows the Laventan and precedes ...
to
Montehermosan The Montehermosan age is a period of geologic time (6.8–4.0 Ma) within the Miocene and Pliocene epochs of the Neogene used more specifically with South American Land Mammal Ages. It follows the Huayquerian and precedes the Chapadmalalan The C ...
)
Mauri Formation Mauri (from which derives the English term "Moors") was the Latin designation for the Berber population of Mauretania, located in the part of North Africa west of Numidia, in present-day northern Morocco and northwestern Algeria. Name ''Maur ...
in the La Paz Department in
Bolivia , image_flag = Bandera de Bolivia (Estado).svg , flag_alt = Horizontal tricolor (red, yellow, and green from top to bottom) with the coat of arms of Bolivia in the center , flag_alt2 = 7 × 7 square p ...
.''Hoffstetterius''
at
Fossilworks Fossilworks is a portal which provides query, download, and analysis tools to facilitate access to the Paleobiology Database The Paleobiology Database is an online resource for information on the distribution and classification of fossil animals ...
.org
The only described species is the type ''Hoffstetterius imperator''.Saint-Andre, P. A. (1993). ''Hoffstetterius imperator'' ng, n. sp. du Miocène supérieur de l'Altiplano bolivien et le statut des Dinotoxodontinés (Mammalia, Notoungulata). Comptes rendus de l'Académie des sciences. Série 2, Mécanique, Physique, Chimie, Sciences de l'univers, Sciences de la Terre, 316(4), 539-545.


Etymology

The genus is named after paleontologist
Robert Hoffstetter Robert Julien Hoffstetter (11 June 1908 in Fargniers – 29 December 1999 in Gennevilliers) was a French taxonomist and herpetologist who was influential in categorizing reptiles. He described the snake families Bolyeriidae and Madtsoiidae Mad ...
, who discovered the fossil in 1969.


Discovery

The remains of ''Hoffstetterius'' were discovered in a volcanic tuff in the 6th member of the
Mauri Formation Mauri (from which derives the English term "Moors") was the Latin designation for the Berber population of Mauretania, located in the part of North Africa west of Numidia, in present-day northern Morocco and northwestern Algeria. Name ''Maur ...
, located in the northwestern part of the Bolivian highlands, to . near to the village of Achiri in the
Pacajes Province Pacajes is a province in the Bolivian department of La Paz. Its capital is Coro Coro. Geography Some of the highest mountains of the province are listed below:Boliviam IGM map Rosario Hoja 5841-IV The most important river of the province ...
, Bolivia. These volcanic sediments have been dated in 10.35 million years, which indicates that they are of the Late Miocene and would correspond to the
Chasicoan The Chasicoan ( es, Chasiquense or es, Chasicoense) age is a period of geologic time from 10–9 Ma within the Late Miocene epoch of the Neogene, used more specifically within the SALMA classification in South America. It follows the Mayoan and ...
mammal-age. The uncovered remains include the
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 ...
specimen MNHN ACH1, a partial skull and mandible of an adult individual, plus
postcrania Postcrania (postcranium, adjective: postcranial) in zoology and vertebrate paleontology is all or part of the skeleton apart from the skull. Frequently, fossil remains, e.g. of dinosaurs or other extinct tetrapods, consist of partial or isolated sk ...
l remains including an
axis An axis (plural ''axes'') is an imaginary line around which an object rotates or is symmetrical. Axis may also refer to: Mathematics * Axis of rotation: see rotation around a fixed axis *Axis (mathematics), a designator for a Cartesian-coordinate ...
, right
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 ...
, both
radius In classical geometry, a radius ( : radii) of a circle or sphere is any of the line segments from its center to its perimeter, and in more modern usage, it is also their length. The name comes from the latin ''radius'', meaning ray but also the ...
, part of the left
ulna The ulna (''pl''. ulnae or ulnas) is a long bone found in the forearm that stretches from the elbow to the smallest finger, and when in anatomical position, is found on the medial side of the forearm. That is, the ulna is on the same side of t ...
and left
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 ...
, retrieved by the expedition of the paleontologist
Robert Hoffstetter Robert Julien Hoffstetter (11 June 1908 in Fargniers – 29 December 1999 in Gennevilliers) was a French taxonomist and herpetologist who was influential in categorizing reptiles. He described the snake families Bolyeriidae and Madtsoiidae Mad ...
in 1969. A second specimen, the
paratype In zoology and botany, a paratype is a specimen of an organism that helps define what the scientific name of a species and other taxon actually represents, but it is not the holotype (and in botany is also neither an isotype nor a syntype). Of ...
MNHN BOL V 3291 corresponds to the partial skull of a juvenile individual, which comes from Cerro Pisakeri, also near to Achiri, recovered by the joint expedition of the French Institute of Andean Studies (IFEA) and The Geological Service of Bolivia (GEOBOL) in 1989. These remains were the basis for the designation of the new genus and species ''Hoffstetterius imperator'' in 1993 by French paleontologist Pierre-Antoine Saint-André. The name of the genus commemorates the scientific work of Professor Hoffstetter, while the name of the species, ''imperator'' ("emperor" in
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
) refers to the great
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 ...
of the animal, reminiscent of the crown of the emperors of the
Holy Roman Empire The Holy Roman Empire was a Polity, political entity in Western Europe, Western, Central Europe, Central, and Southern Europe that developed during the Early Middle Ages and continued until its Dissolution of the Holy Roman Empire, dissolution i ...
.


Description

The skull of ''Hoffstetterius'' is high and elongated; in the holotype specimen it measures in length, while in the juvenile specimen it reaches . The most notable characteristic of this is the elevation of the
frontal bone The frontal bone is a bone in the human skull. The bone consists of two portions.''Gray's Anatomy'' (1918) These are the vertically oriented squamous part, and the horizontally oriented orbital part, making up the bony part of the forehead, par ...
over the
nasals In phonetics, a nasal, also called a nasal occlusive or nasal stop in contrast with an oral stop or nasalized consonant, is an occlusive consonant produced with a lowered velum, allowing air to escape freely through the nose. The vast majorit ...
, forming a large sagittal crest, which is more developed in the adult individual, and also presents in the anterior frontal area a sort of triangular "shield", which may have served as a point of attachment for a
keratinous Keratin () is one of a family of structural fibrous proteins also known as ''scleroproteins''. Alpha-keratin (α-keratin) is a type of keratin found in vertebrates. It is the key structural material making up scales, hair, nails, feathers, horn ...
horn, as in modern
rhinoceros A rhinoceros (; ; ), commonly abbreviated to rhino, is a member of any of the five extant species (or numerous extinct species) of odd-toed ungulates in the family Rhinocerotidae. (It can also refer to a member of any of the extinct species o ...
es and some toxodontids such as ''
Trigodon ''Trigodon'' is an extinct genus of the family Toxodontidae, a large bodied notoungulate which inhabited South America during the Late Miocene to Early Pliocene (Mayoan to Montehermosan in the SALMA classification), living from 11.61 to 4.0 Ma ...
'' and '' Paratrigodon''. In dorsal view, the skull is triangular in shape. The mandible has a notorious projection pointing downwards into the mandibular ramus, at the level of the lower third molar (m3), a feature seen in other toxodontids such as ''
Mixotoxodon ''Mixotoxodon'' ("mixture ''Toxodon''") is an extinct genus of notoungulate of the family Toxodontidae inhabiting South America, Central America and parts of southern North America during the Pleistocene epoch, from 1,800,000—12,000 years ago ...
'' and '' Pericotoxodon''. The
dental formula Dentition pertains to the development of teeth and their arrangement in the mouth. In particular, it is the characteristic arrangement, kind, and number of teeth in a given species at a given age. That is, the number, type, and morpho-physiolo ...
is in the holotype; in the juvenile paratype it is . Molar teeth have lingual folds with less depths than in other toxodontids such as '' Ocnerotherium'', a well-developed middle lobe and a complex contour in the posterior lobe of the upper molar, apart from a sharp posterior edge in the upper third molar (M3), while in the second lower molar (m2) the lingual surface of the trigonid is much longer. The upper premolars lack of lingual folds and the upper molars lacking of the bifurcated lingual fold, a trait shared with advanced toxodontids such as ''Trigodon'', '' Stereotoxodon'' and ''Mixotoxodon''. The femur of ''Hoffstetterius'' has an elongated trochlear medial crest at its lower end, in the area of the
knee In humans and other primates, the knee joins the thigh with the leg and consists of two joints: one between the femur and tibia (tibiofemoral joint), and one between the femur and patella (patellofemoral joint). It is the largest joint in the hu ...
joint, as in ''
Toxodon ''Toxodon'' (meaning "bow tooth" in reference to the curvature of the teeth) is an extinct genus of South American mammals from the Late Miocene to early Holocene epochs (Mayoan to Lujanian in the SALMA classification) (about 11.6 million to 11 ...
''. This type of structure may have served these toxodontids to maintain static their legs while standing, facilitating them to maintain this posture for long periods throughout the day without fatigue, as in modern horses and rhinoceroses.


Classification

''Hoffstetterius'' is clearly a toxodontid, identified by characteristics such as the
hypsodont Hypsodont is a pattern of dentition with high-crowned teeth and enamel extending past the gum line, providing extra material for wear and tear. Some examples of animals with hypsodont dentition are cows and horses; all animals that feed on gritt ...
molars and the development of four large frontal hypselodont incisors of continuous growth. However, it is more difficult to establish its position within this family, since it has features in common with ''Ocnerotherium'', which were considered as part of a subfamily called Dinotoxodontinae, while dental characteristics and the frontal prominence bring it closer to another subfamily, the Haplodontotheriinae; Saint-André considered it as part of the latter group in its initial description in 1992. However, it seems that the traits that distinguish these subfamilies are actually found in different genera and therefore is better only distinguish between the primitive forms grouped in Nesodontinae and the more advanced Toxodontinae, including ''Hoffstetterius''.
Cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...
based in the analysis of Analía Forasiepi ''et al''. (2014), showing the
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
position of ''Hoffstetterius'':


Paleoenviroment

''Hoffstetterius'' is one of the few toxodonts found in the Bolivian
Altiplano The Altiplano (Spanish for "high plain"), Collao (Quechua and Aymara: Qullaw, meaning "place of the Qulla") or Andean Plateau, in west-central South America, is the most extensive high plateau on Earth outside Tibet. The plateau is located at the ...
, apart from another Late Miocene genus, '' Andinotoxodon'', which has also been found in the Nabón Formation in
Ecuador Ecuador ( ; ; Quechua: ''Ikwayur''; Shuar: ''Ecuador'' or ''Ekuatur''), officially the Republic of Ecuador ( es, República del Ecuador, which literally translates as "Republic of the Equator"; Quechua: ''Ikwadur Ripuwlika''; Shuar: ''Eku ...
. The area is today at about above
sea level Mean sea level (MSL, often shortened to sea level) is an average surface level of one or more among Earth's coastal bodies of water from which heights such as elevation may be measured. The global MSL is a type of vertical datuma standardised g ...
, but during the Late Miocene it was at an altitude of , with an average temperature of .Gregory-Wodzicki, K. M. (2002). A late Miocene subtropical-dry flora from the northern Altiplano, Bolivia. Palaeogeography, Palaeoclimatology, Palaeoecology, 180(4), 331-348. In this warmer environment were found plants typical from a subtropical dry forest, with plants of the genus ''
Polylepis ''Polylepis'' is a genus comprising 28 recognised shrub and tree species, that are endemic to the mid- and high-elevation regions of the tropical Andes. This group is unique in the rose family in that it is predominantly wind-pollinated. They are ...
'', '' Zyzyphus'' and ''
Berberis ''Berberis'' (), commonly known as barberry, is a large genus of deciduous and evergreen shrubs from tall, found throughout Temperateness, temperate and subtropical regions of the world (apart from Australia). Species diversity is greatest in S ...
'', whereas the contemporary fauna includes the mesotheriid ''
Plesiotypotherium ''Plesiotypotherium'' is an extinct genus of Notoungulate, belonging to the suborder Typotheria. It lived from the Middle to the Late Miocene, and its fossilized remains were discovered in South America. Description This animal was somewhat si ...
'', a notoungulate that constitutes a fossil guide of the area; the typothere ''
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 ...
'', remains of macraucheniids, rodents like dynomyids, octodontoids, caviods and the chinchillid '' Prolagostomus''. Among the
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 ...
ns have been found remains of the
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 ...
'' Trachycalyptoides'', the dasypodid ''
Chorobates The chorobates, described by Vitruvius in Book VIII of the '' De architectura'', was used to measure horizontal planes and was especially important in the construction of aqueducts. Similar to modern spirit levels, the chorobates consisted of ...
'' and the pampatheriid ''
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 ...
'', along ground sloths like scelidotheriids, mylodontids, the
nothrotheriid 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 a
megatheriid 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 Amer ...
.Pujos, F., Antoine, P. O., Mamani Quispe, B., Abello, A., & Andrade Flores, R. (2012, September). The Miocene vertebrate faunas of Achiri, Bolivia. In Journal of Vertebrate Paleontology (Vol. 32, pp. 159-159).


References

{{Taxonbar, from=Q27794741 Toxodonts Miocene mammals of South America Montehermosan Huayquerian Chasicoan Mayoan Neogene Bolivia Fossils of Bolivia Fossil taxa described in 1993 Prehistoric placental genera