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 artiodactyl that lived during the Middle Eocene and is only known from the Iberian Peninsula. The genus is a member of the family Anoplotheriidae and the subfamily Anoplotheriinae, and contains one species, D. sudrei. Like other anoplotheriids, it was endemic to Western Europe. The genus was described based on a left fragment of a maxilla (upper jaw) from the Mazaterón Formation of the Duero Basin, from which its name derives, in 2009. Its dentition is mostly typical of the Anoplotheriinae but differs from related genera in the elongated and triangular third upper premolar and traits of the molars. It is thought to have been part of an endemic fauna that evolved in the Iberian Peninsula during the Middle Eocene, when 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 origin of anoplotheriids remain uncertain, but they were exclusively distributed in Europe 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 artiodactyl groups are not well-resolved. [2] [3]

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

In 2022, Weppe published a phylogenetic analysis in his PhD thesis regarding Palaeogene artiodactyl lineages, focusing on the endemic European families. This phylogenetic tree is the first to include all anoplotheriid genera, although not all individual species were included. In this tree, the Anoplotheriidae, Mixtotheriidae, and Cainotherioidea form a clade based on synapomorphic dental traits (newly evolved tooth features they have in common). Ephelcomenus , Duerotherium, and Robiatherium form a clade within Anoplotheriidae. [6]

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. [7] [8] Anoplotheriids have selenodont (crescent-shaped ridge form) or bunoselenodont (bunodont and selenodont) premolars (P/p) and molars (M/m) made for leaf-browsing diets. The canines (C/c) of the Anoplotheriidae are overall undifferentiated from the incisors (I/i). 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 (or rounded) lingual cuspids. The subfamily Anoplotheriinae differs from the Dacrytheriinae by the molariform premolars with crescent-shaped paraconules and the lower molars that lack a third cusp between the metaconid and entoconid. [3]

Duerotherium is diagnosed based on its dental traits based on the maxilla fragment. The P3 (third upper premolar) is elongated mesiodistally (front-to-back) and triangular in shape and has a protocone cusp that is positioned back in relation to the mouth and noticeable talon positioned slightly back. 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 in being mesiodistally elongated compared to the former. The morphology of P4 is typical of the Anoplotheriinae and has only has one 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 and increase in size 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 a small-sized anoplotheriine that is slightly larger than Robiatherium but smaller 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 (lophs) for folivorous diets. [9] [10]

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 (56–33.9 Ma), the western Eurasian continent was separated into three landmasses: western Europe (an archipelago), Balkanatolia, and eastern Eurasia (Balkanatolia was in between the Paratethys Sea to the north and the Neotethys Ocean to the south). [2] The Holarctic mammalian faunas of western Europe were therefore mostly isolated from other landmasses including Greenland, Africa, and eastern Eurasia, allowing for endemism to occur within western Europe. [10] The European mammals of the late Eocene (MP17–MP20) were therefore mostly descendants of endemic Middle Eocene groups. [11]

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 mammalian 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). [12]

Related Research Articles

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

Palaeotherium is an extinct genus of equoid that lived in Europe and possibly the Middle East from the Middle Eocene to the Early Oligocene. It is the type genus of the Palaeotheriidae, a group exclusive to the Palaeogene that was closest in relation to the Equidae, which contains horses plus their closest relatives and ancestors. Fossils of Palaeotherium were first described in 1782 by the French naturalist Robert de Lamanon and then closely studied by another French naturalist, Georges Cuvier, after 1798. Cuvier erected the genus in 1804 and recognized multiple species based on overall fossil sizes and forms. As one of the first fossil genera to be recognized with official taxonomic authority, it is recognized as an important milestone within the field of palaeontology. The research by early naturalists on Palaeotherium contributed to the developing ideas of evolution, extinction, and succession and demonstrating the morphological diversity of different species within one genus.

<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.

Brachyhyops is an extinct genus of entelodont artiodactyl mammal that lived during the Eocene Epoch of western North America and southeastern Asia. The first fossil remains of Brachyhyops are recorded from the late Eocene deposits of Beaver Divide in central Wyoming and discovered by paleontology crews from the Carnegie Museum of Natural History during the early 20th century. The type species, Brachyhyops wyomingensis, is based on a single skull and was named by E.H. Colbert in 1937, but was not officially described until 1938. During the latter half of the 20th century, additional specimens from North America have been recorded from Saskatchewan and as far south as Texas, indicating that Brachyhyops had a broad distribution and was well-dispersed throughout western North America.

<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>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 assigned the species to Anoplotherium, he recognized that it differed from A. commune by its dentition and limb bones, later moving 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.

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

Plagiolophus is an extinct genus of equoids belonging to the family Palaeotheriidae. It lived in Europe from the middle Oligocene to the early Oligocene. The type species P. minor was initially described by the French naturalist Georges Cuvier in 1804 based on postcranial material including a now-lost skeleton originally from the Paris Basin. It was classified to Palaeotherium the same year but was reclassified to the subgenus Plagiolophus, named by Auguste Pomel in 1847. Plagiolophus was promoted to genus rank by subsequent palaeontologists and today includes as many as seventeen species. As proposed by the French palaeontologist Jean A. Remy in 2004, it is defined by three subgenera: Plagiolophus, Paloplotherium, and Fraasiolophus.

<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 references the name of the genus Dichobune. Eventually, it was found to be more closely related to Xiphodon and now includes 11 species, although one of them may be synonymous.

<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 recognised in his studies that it had dentition similar to the anoplotheriids Anoplotherium and Diplobune but differed from them by a deep preorbital fossa and a lacrimal fossa, the latter of which is where the genus name derives from. D. ovinum, originally classified in Dichobune, is the type species of Dacrytherium. Henri Filhol named D. elegans in 1884, and Hans Georg Stehlin named the species D. priscum and D. saturnini in 1910.

<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.

Anatomy of <i>Palaeotherium</i> Studies of a genus of palaeothere

The anatomy of Palaeotherium has been historically well-studied due to at least several of its species being known from common and good fossil material. As the type genus of the Palaeotheriidae, one of two families within the Equoidea, it shares common traits such as orbits that are wide in its back plus located in the skull's midlength, long nasal bones, selenodont form molars, and the presence of diastemata between the canine and other teeth between it. Palaeotherium itself differs from other palaeotheres primarily based on various cranial and dental traits; the subgenus Palaeotherium is likewise distinguished from the other subgenus Franzenitherium based on specialized and specific cranial traits. While not as often studied, Palaeotherium is also known by viable limb bone material, leading to the locomotion of different species being hypothesized. P. magnum, unlike other species, is known by complete skeletal material such as that from Mormoiron in France that is informative about its overall anatomy, sharing similar and different traits from equines and other perissodactyls.

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

Amphimeryx is an extinct genus of Palaeogene artiodactyls belonging to the Amphimerycidae that was endemic to the central region of western Europe and lived from the Late Eocene to the Early Oligocene. It was erected in 1848 by the French palaeontologist Auguste Pomel, who argued that its dentition was roughly similar to those of ruminants. Hence, the etymology of the genus name means "near ruminant," of which it derives from the ancient Greek words ἀμφί (near) and μήρυξ (ruminant). The type species A. murinus was previously recognized as a species of Dichobune by the French palaeontologist Georges Cuvier in 1822 before its eventual reclassification to its own genus. Two other species A. collotarsus and A. riparius are recognized also today although the former may be synonymous with A. murinus while the latter is known solely by a now-lost fossil specimen.

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

Pseudamphimeryx is an extinct genus of Palaeogene artiodactyls belonging to the Amphimerycidae that was endemic to the central region of western Europe and lived from the Middle to Late Eocene. It was first erected in 1910 by the Swiss palaeontologist Hans Georg Stehlin, who assigned to it multiple species and noted specific differences from another amphimerycid Amphimeryx. As of present, it is known by six species, although the validity of P. valdensis has been questioned while the earliest-appearing species P. schosseri has been suggested to not be an amphimerycid.

References

  1. 1 2 3 4 5 6 7 Cuesta, Miguel-Ángel; Badiola, Ainara (2009). "Duerotherium sudrei gen. et sp. nov., a New Anoplotheriine Artiodactyl from the Middle Eocene of the Iberian Peninsula". Journal of Vertebrate Paleontology. 29 (1): 303–308. Bibcode:2009JVPal..29..303C. doi:10.1671/039.029.0110. JSTOR   20491092. S2CID   55546022.
  2. 1 2 Licht, Alexis; Métais, Grégoire; Coster, Pauline; İbilioğlu, Deniz; Ocakoğlu, Faruk; Westerweel, Jan; Mueller, Megan; Campbell, Clay; Mattingly, Spencer; Wood, Melissa C.; Beard, K. Christopher (2022). "Balkanatolia: The insular mammalian biogeographic province that partly paved the way to the Grande Coupure". Earth-Science Reviews. 226: 103929. Bibcode:2022ESRv..22603929L. doi: 10.1016/j.earscirev.2022.103929 .
  3. 1 2 Badiola, Ainara; De Vicuña, Nahia Jiménez; Perales-Gogenola, Leire; Gómez-Olivencia, Asier (2023). "First clear evidence of Anoplotherium (Mammalia, Artiodactyla) in the Iberian Peninsula: an update on the Iberian anoplotheriines". The Anatomical Record: Advances in Integrative Anatomy and Evolutionary Biology. doi: 10.1002/ar.25238 . PMID   37221992. S2CID   258864256.
  4. Erfurt, Jörg; Métais, Grégoire (2007). "Endemic European Paleogene Artiodactyls". In Prothero, Donald R.; Foss, Scott E. (eds.). The Evolution of Artiodactyls. Johns Hopkins University Press. pp. 59–84.
  5. Orliac, Maeva; Gilissen, Emmanuel (2012). "Virtual endocranial cast of earliest Eocene Diacodexis (Artiodactyla, Mammalia) and morphological diversity of early artiodactyl brains". Proceedings of the Royal Society B. 279 (1743): 3670–3677. doi:10.1098/rspb.2012.1156. PMC   3415922 . PMID   22764165.
  6. Weppe, Romain (2022). Déclin des artiodactyles endémiques européens, autopsie d'une extinction (Thesis) (in French). University of Montpellier.
  7. von Zittel, Karl Alfred (1925). Schlosser, Max (ed.). Text-Book of Paleontology. Volume III. Mammalia. Macmillan and Co. Limited. pp. 179–180.
  8. Lihoreau, Fabrice; Boisserie, Jean-Renaud; Viriot, Laurent; Brunet, Michel (2006). "Anthracothere dental anatomy reveals a late Miocene Chado-Libyan bioprovince". Proceedings of the National Academy of Sciences. 103 (23): 8763–8767. Bibcode:2006PNAS..103.8763L. doi: 10.1073/pnas.0603126103 . PMC   1482652 . PMID   16723392.
  9. Eronen, Jussi T.; Janis, Christine M.; Chamberlain, Charles Page; Mulch, Andreas (2015). "Mountain uplift explains differences in Palaeogene patterns of mammalian evolution and extinction between North America and Europe". Proceedings of the Royal Society B. 282 (1809). doi:10.1098/rspb.2015.0136. PMC   4590438 . PMID   26041349.
  10. 1 2 Maitre, Elodie (2014). "Western European middle Eocene to early Oligocene Chiroptera: systematics, phylogeny and palaeoecology based on new material from the Quercy (France)". Swiss Journal of Palaeontology . 133 (2): 141–242. doi: 10.1007/s13358-014-0069-3 . S2CID   84066785.
  11. Badiola, Ainara; Perales-Gogenola, Leire; Astibia, Humberto; Suberbiola, Xabier Pereda (2022). "A synthesis of Eocene equoids (Perissodactyla, Mammalia) from the Iberian Peninsula: new signs of endemism". Historical Biology. 34 (8): 1623–1631. Bibcode:2022HBio...34.1623B. doi:10.1080/08912963.2022.2060098. S2CID   248164842.
  12. Marigó, Judit; Minwer-Barakat, Raef; Moyà-Solà, Salvador (2010). "New Anchomomyini (Adapoidea, Primates) from the Mazaterón Middle Eocene locality (Almazán Basin, Soria, Spain)". Journal of Human Evolution. 58 (4): 353–361. doi:10.1016/j.jhevol.2010.01.011.