Duerotherium

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Duerotherium
Temporal range: Middle Eocene 41–37  Ma
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Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Artiodactyla
Family: Anoplotheriidae
Subfamily: Anoplotheriinae
Genus: Duerotherium
Cuesta & Badiola, 2009
Species:
D. sudrei
Binomial name
Duerotherium sudrei
Cuesta & Badiola, 2009

Duerotherium is an extinct genus of Palaeogene artiodactyls known only from the Iberian Peninsula during the Middle Eocene, which contains one species D. sudrei. It, like other members of the Anoplotheriidae, was endemic to western Europe. The anoplotheriine was described from a left fragment of a maxilla from the Mazaterón Formation of the Duero Basin (where its name derives from) in 2009. Its dentition is mostly typical of the Anoplotheriinae but differs by an elongated plus triangular 3rd upper premolar and very specific traits of the molars. It is thought to have been part of an endemic faunal assemblage that evolved within the Iberian Peninsula by the Middle Eocene, where climates were subtropical.

Contents

Taxonomy

In 2009, Spanish palaeontologists Miguel-Ángel Cuesta and Ainara Badiola described a newly erected anoplotheriine genus from the Mazaterón Formation near the village of Mazaterón, which are located within the Duero Basin. The genus and type species Duerotherium sudrei was created based on a fragment of a left fragment of a maxilla with a dental series of P3-M3 (specimen STUS 11562), which was deposited in the "Sala de las Tortugas" of the University of Salamanca. The genus etymology derives from the Duero Basin for where the fossil was described plus the Greek θήρ/therium meaning "beast" or "wild animal." The etymology of the species name was dedicated in honour of Jean Sudre for his studies on endemic European Palaeogene artiodactyls. [1]

Classification

Skeleton of Anoplotherium commune, National Museum of Natural History, France Anoplotherium commune.jpg
Skeleton of Anoplotherium commune, National Museum of Natural History, France

Duerotherium belongs to the Anoplotheriidae, a Palaeogene artiodactyl family endemic to western Europe that lived from the middle Eocene to the early Oligocene (~44 to 30 Ma, possible earliest record at ~48 Ma). The exact evolutionary origins and dispersals of the anoplotheriids are uncertain, but they exclusively resided within the continent when it was an archipelago that was isolated by seaway barriers from other regions such as Balkanatolia and the rest of eastern Eurasia. The Anoplotheriidae's relations with other members of the Artiodactyla are not well-resolved, with some determining it to be either a tylopod (which includes camelids and merycoidodonts of the Palaeogene) or a close relative to the infraorder and some others believing that it may have been closer to the Ruminantia (which includes tragulids and other close Palaeogene relatives). [2] [3]

The Anoplotheriidae consists of two subfamilies, the Dacrytheriinae and Anoplotheriinae, the latter of which is the subfamily that Duerotherium belongs to. The Dacrytheriinae is the older subfamily of the two that first appeared in the middle Eocene (since the Mammal Palaeogene zones unit MP13, possibly up to MP10), although some authors consider them to be a separate family in the form of the Dacrytheriidae. [1] [4] [5] Anoplotheriines made their first appearances by the late Eocene (MP15-MP16), or ~41-40 Ma, within western Europe with Duerotherium and Robiatherium . After a significant gap of anoplotheriines in MP17a-MP17b, the derived anoplotheriids Anoplotherium and Diplobune made their first appearances in western Europe by MP18, although their exact origins are unknown. [1]

Conducting studies focused on the phylogenetic relations within the Anoplotheriidae has proven difficult due to the general scarcity of fossil specimens of most genera. [1] The phylogenetic relations of the Anoplotheriidae as well as the Xiphodontidae, Mixtotheriidae, and Cainotheriidae have also been elusive due to the selenodont morphologies of the molars, which were convergent with tylopods or ruminants. [6] Some researchers considered the selenodont families Anoplotheriidae, Xiphodontidae, and Cainotheriidae to be within Tylopoda due to postcranial features that were similar to the tylopods from North America in the Palaeogene. [7] Other researchers tie them as being more closely related to ruminants than tylopods based on dental morphology. Different phylogenetic analyses have produced different results for the "derived" selenodont Eocene European artiodactyl families, making it uncertain whether they were closer to the Tylopoda or Ruminantia. [8] [9]

In an article published in 2019, Romain Weppe et al. conducted a phylogenetic analysis on the Cainotherioidea within the Artiodactyla based on mandibular and dental characteristics, specifically in terms of relationships with artiodactyls of the Palaeogene. The results retrieved that the superfamily was closely related to the Mixtotheriidae and Anoplotheriidae. They determined that the Cainotheriidae, Robiacinidae, Anoplotheriidae, and Mixtotheriidae formed a clade that was the sister group to the Ruminantia while Tylopoda, along with the Amphimerycidae and Xiphodontidae split earlier in the tree. [9] The phylogenetic tree used for the journal and another published work about the cainotherioids is outlined below: [10]

In 2022, Weppe created a phylogenetic analysis in his academic thesis regarding Palaeogene artiodactyl lineages, focusing most specifically on the endemic European families. The phylogenetic tree, according to Weppe, is the first to conduct phylogenetic affinities of all anoplotheriid genera, although not all individual species were included. He found that the Anoplotheriidae, Mixtotheriidae, and Cainotherioidea form a clade based on synapomorphic dental traits (traits thought to have originated from their most recent common ancestor). The result, Weppe mentioned, matches up with previous phylogenetic analyses on the Cainotherioidea with other endemic European Palaeogene artiodactyls that support the families as a clade. As a result, he argued that the proposed superfamily Anoplotherioidea, composing of the Anoplotheriidae and Xiphodontidae as proposed by Alan W. Gentry and Hooker in 1988, is invalid due to the polyphyly of the lineages in the phylogenetic analysis. However, the Xiphodontidae was still found to compose part of a wider clade with the three other groups. He said that Ephelcomenus , Duerotherium, and Robiatherium compose a clade of the Anoplotheriidae. [6] [11]

Description

The dental formula of the Anoplotheriidae is 3.1.4.33.1.4.3 for a total of 44 teeth, consistent with the primitive dental formula for early-middle Palaeogene placental mammals. [12] [13] Anoplotheriids have selenodont or bunoselenodont premolars and molars made for folivorous/browsing diets, consistent with environment trends in the late Eocene of Europe. The canines of the Anoplotheriidae are premolariform in shape, meaning that the canines are overall undifferentiated from other teeth like incisors. The lower premolars of the family are piercing and elongated. The upper molars are bunoselenodont in form while the lower molars have selenodont labial cuspids and bunodont lingual cuspids. The subfamily Anoplotheriinae differs from the Dacrytheriinae by the lower molars lacking a third cusp between the metaconid and entoconid as well as molariform premolars with crescent-shaped paraconules. [3]

Duerotherium is diagnosed based on its small size and, most exclusively, its dental traits based on the maxilla fragment. The P3 tooth is in a mesiodistal position, is elongated plus triangular in shape, and has a distolingually positioned protocone cusp plus a noticeable posterolingual talon. The morphology of the tooth of Duerotherium is similar to the P3 tooth of Dacrytherium based on the positions of the cusps, although the latter differs by it being mesiodistally elongated compared to the former. The morphology of P4 is typical of the Anoplotheriinae and has only has externally-positioned cusp. [1]

The upper molars of Duerotherium also have similar morphologies to those of other anoplotheriines. They are bunoselenodont and have large and conical protocone cusps in the near-front of the paracone in the front areas of the teeth. The metaconule is slightly asymmetric, and the postmetaconule ridge is moderate in form. The parastyle and metastyle cusps are divergent, revealing a moderate W-shaped ectoloph ridge. The molars are heterodont in dentition form and increase in size sequencing from M1 to M3. In a top (or occlusal) outline view, the M1 is quadrate in shape while M2-M3 appears more trapezoidal. It differs from each anoplotheriine genus based on various specific morphologies of the molars. [1]

Duerotherium is described as being slightly larger than Robiatherium but having a smaller size than Ephelcomenus. D. sudrei is especially smaller than Anoplotherium and most species of Diplobune. It might be similar in size to Diplobune minor. [1]

Palaeoecology

Palaeogeography of Europe and Asia during the middle Eocene with possible artiodactyl and perissodactyl dispersal routes. Middle Eocene Paleogeography Tethys Dispersals.jpg
Palaeogeography of Europe and Asia during the middle Eocene with possible artiodactyl and perissodactyl dispersal routes.

For much of the Eocene, a hothouse climate with humid, tropical environments with consistently high precipitations prevailed. Modern mammalian orders including the Perissodactyla, Artiodactyla, and Primates (or the suborder Euprimates) appeared already by the early Eocene, diversifying rapidly and developing dentitions specialized for folivory. The omnivorous forms mostly either switched to folivorous diets or went extinct by the middle Eocene (47 - 37 Ma) along with the archaic "condylarths". By the late Eocene (approx. 37 - 33 Ma), most of the ungulate form dentitions shifted from bunodont cusps to cutting ridges (i.e. lophs) for folivorous diets. [14] [15]

Land-based connections to the north of the developing Atlantic Ocean were interrupted around 53 Ma, meaning that North America and Greenland were no longer well-connected to western Europe. From the early Eocene up until the Grande Coupure extinction event (56 Ma - 33.9 Ma), the western Eurasian continent was separated into three landmasses, the former two of which were isolated by seaways: western Europe (an archipelago), Balkanatolia, and eastern Eurasia (Balkanatolia was in between the Paratethys Sea of the north and the Neotethys Ocean of the south). [2] The Holarctic mammalian faunas of western Europe were therefore mostly isolated from other continents including Greenland, Africa, and eastern Eurasia, allowing for endemism to occur within western Europe. [15] The European mammals of the late Eocene (MP17 - MP20) were mostly descendants of endemic middle Eocene groups as a result. [16]

The Mazaterón Formation of the Duero Basin dates to the middle Late Eocene (Robiacian of Europe) and ranges in faunal level from MP15-MP16. Fossils of testudines, crocodilians, rodents, primates, hyaenodonts, artiodactyls, and perissodactyls were collected from the site. The mammal taxa collected from the Eocene of the Iberian region differ from contemporary faunas in other areas of Europe, supporting the hypothesis of a division of the Iberian Peninsula as a semi-separate bioregion. [1] The taxa known from the Mazaterón fossil site with Duerotherium include the testudines Hadrianus and Neochelys , alligatoroid Diplocynodon , baurusuchid Iberosuchus , adapoid Mazateronodon , omomyid Pseudoloris , pseudosciurid rodent Sciuroides , theridomyid rodents Pseudoltinomys and Remys , hyaenodont Proviverra , palaeotheres ( Paranchilophus , Plagiolophus , Leptolophus , Palaeotherium , Cantabrotherium , Franzenium , and Iberolophus ), dacrytheriines (cf. Dacrytherium), and xiphodonts (cf. Dichodon). [17]

Related Research Articles

<span class="mw-page-title-main">Artiodactyl</span> Order of mammals

Artiodactyls are placental mammals belonging to the order Artiodactyla. Typically, they are ungulates which bear weight equally on two of their five toes: the third and fourth, often in the form of a hoof. The other three toes are either present, absent, vestigial, or pointing posteriorly. By contrast, most perissodactyls bear weight on an odd number of the five toes. Another difference between the two is that many artiodactyls digest plant cellulose in one or more stomach chambers rather than in their intestine as perissodactyls do. The advent of molecular biology, along with new fossil discoveries, found that cetaceans fall within this taxonomic branch, being most closely related to hippopotamuses. Some modern taxonomists thus apply the name Cetartiodactyla to this group, while others opt to include cetaceans within the existing name of Artiodactyla. Some researchers use "even-toed ungulates" to exclude cetaceans and only include terrestrial artiodactyls, making the term paraphyletic in nature.

<span class="mw-page-title-main">Tylopoda</span> Suborder of mammals

Tylopoda is a suborder of terrestrial herbivorous even-toed ungulates belonging to the order Artiodactyla. They are found in the wild in their native ranges of South America and Asia, while Australian feral camels are introduced. The group has a long fossil history in North America and Eurasia. Tylopoda appeared during the Eocene around 50 million years ago.

<i>Andrewsarchus</i> Extinct genus of carnivorous ungulate from Eocene epoch

Andrewsarchus is an extinct genus of ungulate that lived during the Middle Eocene in China. It contains two species, A. mongoliensis and A. crassum. It was formerly placed in the families Mesonychidae or Arctocyonidae, but is now the sole member of a distinct family, Andrewsarchidae. Only known from a largely complete skull as well as isolated teeth, It is notable for being estimated as the largest terrestrial, carnivorous mammal, but that status has been disputed.

<i>Anoplotherium</i> Extinct genus of endemic Paleogene European artiodactyls

Anoplotherium is the type genus of the extinct Palaeogene artiodactyl family Anoplotheriidae, which was endemic to western Europe. It lived from the late Eocene to the earliest Oligocene. It was the fifth fossil mammal genus to be described with official taxonomic authority, with a history extending back to 1804 when its fossils from Montmartre in Paris, France were first described by the French naturalist Georges Cuvier. Discoveries of incomplete skeletons of A. commune in 1807 led Cuvier to thoroughly describe unusual features for which there are no modern analogues. His drawn skeletal and muscle reconstructions of A. commune in 1812 were amongst the first instances of anatomical reconstructions based on fossil evidence. Cuvier's contributions to palaeontology based on his works on the genus were revolutionary for the field, not only proving the developing ideas of extinction and ecological succession but also paving the way for subfields such as palaeoneurology. Today, there are four known species.

<span class="mw-page-title-main">Xiphodontidae</span> Extinct family of mammals

Xiphodontidae is an extinct family of herbivorous even-toed ungulates, endemic to Europe during the Eocene 40.4—33.9 million years ago, existing for about 7.5 million years. Paraxiphodon suggests that they survived into the Lower Oligocene, at least.

<span class="mw-page-title-main">Anoplotheriidae</span> Extinct family of mammals

Anoplotheriidae is an extinct family of artiodactyl ungulates. They were endemic to Europe during the Eocene and Oligocene epochs about 44—30 million years ago. Its name is derived from the Ancient Greek: ἂνοπλος ("unarmed") and θήριον ("beast"), translating as "unarmed beast".

<i>Pterodon</i> (mammal) Extinct genus of mammals

Pterodon is an extinct genus of hyaenodont in the family Hyainailouridae, containing five species. The type species Pterodon dasyuroides is known exclusively from the late Eocene to the earliest Oligocene of western Europe. The genus was first erected by the French zoologist Henri Marie Ducrotay de Blainville in 1839, who said that Georges Cuvier presented one of its fossils to a conference in 1828 but died before he could make a formal description of it. It was the second hyaenodont genus with taxonomic validity after Hyaenodon, but this resulted in taxonomic confusion over the validities of the two genera by other taxonomists. Although the taxonomic status of Pterodon was revised during the late 19th and early 20th centuries, it became a wastebasket taxon for other hyaenodont species found in Africa and Asia. Today, only the type species is recognized as belonging to the genus while four others are pending reassessment to other genera.

<i>Agriochoerus</i> Extinct genus of mammals

Agriochoerus is an extinct genus of scansorial herbivore of the tylopod family Agriochoeridae, endemic to North America. Agriochoerus and other agriochoerids possessed claws, which is rare within Artiodactyla, as well as likely being scansorial. Agriochoerus was first described in 1869.

<i>Xiphodon</i> Extinct genus of endemic Palaeogene European artiodactyls

Xiphodon is the type genus of the extinct Palaeogene artiodactyl family Xiphodontidae. It, like other xiphodonts, was endemic to western Europe and lived from the middle Eocene up to the earliest Oligocene. Fossils from Montmartre in Paris, France that belonged to X. gracilis were first described by the French naturalist Georges Cuvier in 1804. Although he designated the species to Anoplotherium, he recognized that it differed from A. commune by its dentition and limb bones, later designating it to its own subgenus in 1822. Xiphodon was promoted to genus rank by other naturalists in later decades. It is today defined by the type species X. gracilis and two other species X. castrensis and X. intermedium.

<span class="mw-page-title-main">Cainotheriidae</span> Extinct family of mammals

Cainotheriidae is an extinct family of artiodactyls known from the Late Eocene to Middle Miocene of Europe. They are mostly found preserved in karstic deposits.

<span class="mw-page-title-main">Helohyidae</span> Family of extinct artiodactyl mammals

Helohyidae were a group of artiodactyl mammals. They were most prominent in the mid-to-upper Eocene.

<i>Dichodon</i> (mammal) Extinct genus of endemic Palaeogene European artiodactyls

Dichodon is an extinct genus of Palaeogene artiodactyls belonging to the family Xiphodontidae. It was endemic to western Europe and lived from the middle Eocene up to the earliest Oligocene. The genus was first erected by the British naturalist Richard Owen in 1848 based on dental remains from the fossil beds in Hordle, England. He noticed similar dentitions to contemporary artiodactyls like those of the Anoplotheriidae and Dichobunidae and based the genus name off of Dichobune. Eventually, it was found to be more closely related to Xiphodon and now composes of 11 species, although one of them may be synonymous.

Research history of <i>Anoplotherium</i> Studies of a genus of endemic Paleogene European artiodactyls

The research history of Anoplotherium spans back to 1804 when Georges Cuvier first described the fossils of this extinct artiodactyl and named the genus after describing Palaeotherium, making it one of the first fossil mammal genera to be described as well as having one of the earliest official taxonomic authorities. It was also amongst the first fossil genera to be reconstructed by drawings and biomechanics. Subsequent descriptions of fossil evidence by Cuvier are also said to have been some of the earliest instances of palaeoneurology and palaeopathology. Anoplotherium was a significant find in palaeontological history and was once an iconic element of text and classroom sources of palaeontology, geology, and natural history. Today, it has a lessened cultural status compared to the 19th century as a result of public interest in Mesozoic dinosaurs or Neogene-Quaternary mammals, but it is still regularly acknowledged in sources of the history of palaeontology.

<i>Diplobune</i> Extinct genus of endemic Palaeogene European artiodactyls

Diplobune is an extinct genus of Palaeogene artiodactyls belonging to the family Anoplotheriidae. It was endemic to Europe and lived from the late Eocene to the early Oligocene. The genus was first erected as a subgenus of Dichobune by Ludwig Rütimeyer in 1862 based on his hypothesis of the taxon being a transitional form between "Anoplotherium" secundaria, previously erected by Georges Cuvier in 1822, and Dichobune. He based the genus etymology off of the two-pointed pillarlike shapes of the lower molars, which had since been a diagnosis of it. However, in 1870, Diplobune was elevated to genus rank by Oscar Fraas, who recognized that Diplobune was a distinct genus related to Anoplotherium and not Dichobune. After several revisions of the anoplotheriids, there are currently four known species of which D. minor is the type species.

<i>Dacrytherium</i> Extinct genus of endemic Palaeogene European artiodactyls

Dacrytherium is an extinct genus of Palaeogene artiodactyls belonging to the family Anoplotheriidae. It occurred from the middle to late Eocene of western Europe and is the type genus of the subfamily Dacrytheriinae, the older of the two anoplotheriid subfamilies. Dacrytherium was first erected in 1876 by the French palaeontologist Henri Filhol, who recognized in his studies that it had dentition similar to the anoplotheriids Anoplotherium and Diplobune but differed from them by a deep preorbital fossa, where the genus name derives from. Since then, there are currently four valid species within the genus, of which D. ovinum is the type species.

<i>Catodontherium</i> Extinct genus of Palaeogene artiodactyls

Catodontherium is an extinct genus of Palaeogene artiodactyls belonging to the family Anoplotheriidae. It was endemic to Western Europe and had a temporal range exclusive to the middle Eocene, although its earliest appearance depends on whether C. argentonicum is truly a species of Catodontherium. It was first named Catodus by the French palaeontologist Charles Depéret in 1906, who created two species for the genus and later changed the genus name to Catodontherium in 1908. The Swiss palaeontologist Hans Georg Stehlin renamed one species and classified two other newly erected species to Catodontherium in 1910. Today, there are four known species, although two remain questionable in genus placement.

Ephelcomenus is an extinct genus of Palaeogene artiodactyls endemic to western Europe. It contains one species E. filholi, which was first described by Richard Lydekker in 1889 but eventually classified to its own genus by the Swiss palaeontologist Johannes Hürzeler in 1938. It has an uncertain stratigraphic range, but some sources suggest that it was present in the Oligocene after the Grande Coupure turnover event of western Europe.

Robiatherium is an extinct genus of Palaeogene artiodactyls containing one species R. cournovense. The genus name derives from the locality of Robiac in France where some of its fossil were described plus the Greek θήρ/therium meaning "beast" or "wild animal." It was known only from the middle Eocene and, like other anoplotheriids, was endemic to western Europe. The genus was erected by Jean Sudre in 1988 for a species originally attributed to the xiphodont genus Paraxiphodon in 1978. Robiatherium had dentitions typical of the subfamily Anoplotheriinae, differing from other genera by specific differences in the molars. It is one of the earliest-appearing anoplotheriine species in the fossil record as well as the earliest to have appeared in central Europe.

<i>Mixtotherium</i> Extinct genus of endemic Palaeogene European artiodactyls

Mixtotherium is an extinct genus of Palaeogene artiodactyls belonging to the monotypic family Mixtotheriidae. Known informally as mixtotheriids or mixtotheres, these artiodactyls were endemic to western Europe and occurred from the middle to late Eocene. The genus and type species were both first established by the French naturalist Henri Filhol in 1880. Several species are well known by good skull fossils, which were informative enough to allow for classifications of the species to their own family. The Mixtotheriidae, first recognized by Helga Sharpe Pearson in 1927, is currently known by 7 valid species, although M. priscum is thought by several authors to be synonymous with M. gresslyi. The affinities of the Mixtotheriidae in relation to other artiodactyl families is uncertain, but it is currently thought to have been related to the Cainotherioidea and Anoplotheriidae.

Haplomeryx is an extinct genus of Palaeogene artiodactyls belonging to the family Xiphodontidae. It was endemic to western Europe and lived from the middle Eocene up to the earliest Oligocene. Haplomeryx was first established as a genus by the German naturalist Max Schlosser in 1886 based on a molar tooth set from Quercy Phosphorites deposits. Three additional species were erected and classified to the xiphodontid genus while one other species. first recognized in 1822, was tentatively classified to it and remains unresolved in affinity.

References

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