Anoplotherium

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.

It was an evolutionarily advanced and unusual artiodactyl, sporting three-toed feet in certain species like A. latipes, a long and robust tail, and a highly-developed brain with strong support for both sense of smell and sensory perception.

Its overall robust build may have allowed it to stand bipedally to browse on plants at greater heights, reaching approximately 3 m (9.8 ft) tall, effectively competing with the few other medium to large herbivores it lived with.

Its exact origins are unknown, but it arose long after a shift towards drier but still subhumid conditions that led to abrasive plants and the extinctions of the large-sized Lophiodontidae, achieving gigantism and establishing itself as a dominant herbivore throughout the entirety of the western European region given its abundant fossil evidence.

Its success was abruptly halted by the Grande Coupure extinction and faunal turnover event in the earliest Oligocene of western Europe, which was caused by shifts towards further glaciation and seasonality.

Tropical and subtropical forests were rapidly replaced by more temperate environments, and most ocean barriers previously separating western Europe from eastern Eurasia closed, allowing for large faunal dispersals from Asia.

[1][2] The genus name Anoplotherium means "unarmed beast" and is a compound of the Greek words αν- (an, 'not'), ὅπλον (hóplon, 'armor, large shield'), and θήρ (thēr, 'beast, wild animal').

[42] In 1883, Max Schlosser made Eurytherium a synonym of Anoplotherium because he argued that the limb anatomies and dentitions were specific differences in characteristics rather than major ones that defined an entire genus.

[43] In 1922, Wilhelm Otto Dietrich erected the fourth species A. pompeckji from the locality of Mähringen in Germany, named in honor of German palaeontologist Josef Felix Pompeckj.

[23][45] In 1964, palaeontologist Louis de Bonis reviewed briefly the taxonomic synonyms of Anoplotherium, considering that A. duvernoyi was based on a young individual with incisor characteristics that Pomel did not specify and that A. cuvieri does not differ in metacarpal dimensions from A. laurillardi.

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 Dacrytheriinae is the older subfamily of the two that first appeared in the Middle Eocene (since the Mammal Palaeozone Zones unit MP13, possibly up to MP10), although some authors consider them to be a separate family in the form of the Dacrytheriidae.

[54][55] 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 phylogenetic tree as produced by the authors is shown below:[54] Bunophorus Gunophorus Diacodexis Protodichobune Eurodexis Buxobune Mouillacitherium Meniscodon Hyperdichobune Dichobune robertiana Dichobune leporina Homacodon Gobiohyus Khirtharia Entelodon Palaeocheorus Perchoerus Haplobunodon Cuisitherium Lophiobunodon Mixtotherium Robiacina Dacrytherium Diplobune Xiphodon Paraxiphodon Cainotherium Paroxacron Archaeomeryx Amphimeryx Pseudamphimeryx Aumelasia Hallebune Amphirhagatherium Cebochoerus Gervachoerus Choeropotamus Siamotherium In 2022, Weppe created a phylogenetic analysis in his academic thesis regarding Palaeogene artiodactyl lineages, focusing most specifically on the endemic European families.

Cyril Gagnaison and Jean-Jacques Leroux proposed in the case of D. secundaria that the elongated nasal region supports the presence of a very tapered tongue, which similar to giraffes may have allowed it to pull plant branches.

[75] The type species is Anoplotheriipus lavocati, which Ellenberger named in honor of palaeontologist René Lavocat and considered the "most majestic" of the three ichnospecies due to the displayed specific mobility of the metatarsals.

[52] In 2014, Takehisa Tsubamoto reexamined the relationship between astragalus size and estimated body mass based on extensive studies of extant terrestrial mammals, reapplying the methods to Palaeogene artiodactyls previously tested by Sudre and Martinez.

[45] In 2007, Hooker made size estimates of A. latipes based on an incomplete skeleton of an immature individual from the Hamstead Member of the Bouldnor Formation in the Isle of Wight, United Kingdom.

[23] Its large size and ability to bipedally browse may have given Anoplotherium few sources of terrestrial competition other than from Palaeotherium magnum, a large-sized palaeothere with a long neck that may have reached 240.3 kg (530 lb) in body mass.

[80] Ciaran Clark et al. (including J.J. Hooker) found from micro-CT scans that Anoplotherium being a facultative bipedal browser was not supported by the trabecular architecture of the proximal area of the femur.

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.

[85] The appearances of derived anoplotheriines by MP18 occurred long after the extinction of the endemic European perissodactyl family Lophiodontidae in MP16, including the largest lophiodont Lophiodon lautricense, likely the result of a shift from humid and highly tropical environments to drier and more temperate forests with open areas and more abrasive vegetation.

By then, Anoplotherium and Diplobune lived in Central Europe (then an island) and the Iberian Peninsula, only the former genus of which later dispersed into southern England by MP19 due to the apparent lack of ocean barriers.

[45][23] Anoplotherium coexisted with a wide diversity of artiodactyls in western Europe by MP18, ranging from the more widespread Dichobunidae, Tapirulidae, and Anthracotheriidae to many other endemic families consisting of the Xiphodontidae, Choeropotamidae, Cebochoeridae, Amphimerycidae, and Cainotheriidae.

[85] Late Eocene European groups of the clade Ferae represented predominantly the Hyaenodonta (Hyaenodontinae, Hyainailourinae, and Proviverrinae) but also contained Carnivoramorpha (Miacidae) and Carnivora (small-sized Amphicyonidae).

[88] Other mammal groups present in the Late Eocene of western Europe represented the leptictidans (Pseudorhyncocyonidae),[95] primates (Adapoidea and Omomyoidea),[96] eulipotyphlans (Nyctitheriidae),[97] chiropterans,[84] herpetotheriids,[98] apatotherians,[99] and endemic rodents (Pseudosciuridae, Theridomyidae, and Gliridae).

[101] In addition to snakes, frogs, and salamandrids, rich assemblage of lizards are known in western Europe as well from MP16-MP20, representing the Iguanidae, Lacertidae, Gekkonidae, Agamidae, Scincidae, Helodermatidae, and Varanoidea.

remains were found with undetermined frog and squamate groups, alligatoroid Diplocynodon, the herpetotheriid Peratherium, rodents (Theridomys, Elfomys, Pseudoltinomys, Remys), omomyid Microchoerus, carnivoraformes Quercygale and Paramiacis, dichobunid Dichobune, xiphodonts Xiphodon and Haplomeryx, and palaeotheres (Palaeotherium, Leptolophus, Iberolophus, Pachynolophus, Paranchilophus).

[103] As part of a separate landmass at the time, La Débruge of France, dating to MP18, yielded slightly different faunas that coexisted with A. commune, A. latipes, and A. laurillardi, namely the herpetotheriid Peratherium, rodents (Blainvillimys, Theridomys, Plesiarctomys, Glamys), hyaenodonts (Hyaenodon and Pterodon), amphicyonid Cynodictis, palaeotheres (Plagiolophus, Anchilophus, Palaeotherium), dichobunid Dichobune, choeropotamid Choeropotamus, cebochoerids Cebochoerus and Acotherulum, anoplotheriids Dacrytherium and Diplobune, tapirulid Tapirulus, xiphodonts Xiphodon and Dichodon, cainothere Oxacron, amphimerycid Amphimeryx, and anthracothere Elomeryx.

The Turgai Strait, which once separated much of Europe from Asia, is often proposed as the main European seaway barrier prior to the Grande Coupure, but some researchers challenged this perception recently, arguing that it completely receded already 37 Ma, long before the Eocene-Oligocene transition.

Sculptures of an Anoplotherium commune herd as part of the Crystal Palace Dinosaurs sculptures on the Tertiary Island of the Crystal Palace Park , United Kingdom
Illustrations of Anoplotherium (foreground left), Xiphodon (background left), and Palaeotherium (right)
A. commune jaw fragment, Teylers Museum
Portrait of Georges Cuvier , the French naturalist who described Palaeotherium and Anoplotherium in 1804
Reconstructed skeleton of the related Cainotherium
Upper part of the skull of Anoplotherium commune , National Museum of Natural History, France
Anoplotherium laurillardi lower dental remains, Natural History Museum of Basel
A. commune tail, NMHN, France
Foot of A. commune at the National Museum of Natural History, France
A. commune foot, Teylers Museum
Estimated Anoplotherium size comparisons based on known fossil material
Estimated body masses (kg) of Palaeogene artiodactyls based on recalculated trochlear widths (Li1) in comparison to estimates from Martinez and Sudre (1995)
Reconstruction of A. commune in a quadrupedal position
Reconstruction of A. commune in a bipedal position, most likely to browse on taller vegetation
Palaeogeography of Europe and Asia during the Middle Eocene with possible artiodactyl and perissodactyl dispersal routes.
Restoration of Palaeotherium magnum , which coexisted with Anoplotherium
A panorama of the Headon Hill Formation in the Isle of Wight, from which Anoplotherium material has been collected. The stratigraphy of it and the Bouldnor Formation led to better understandings of faunal chronologies from the Late Eocene up to the Grande Coupure.