Diplobune

It was well-adapted for purely folivorous diets, with dentition capable of chewing through hard leaf material and an implied presence of tapered tongues for reaching branches similar to modern-day giraffids.

It, like Anoplotherium, arose long after a shift towards drier but still subhumid conditions that led to abrasive plants and the extinctions of the large-sized Lophiodontidae, becoming a regular component of late Eocene faunal communities.

In 1870, German palaeontologist Oscar Fraas wrote about a mammal with numerous remains from the locality of Munich, its molars being similar but not identical to A. commune in terms of typical species diagnoses.

In 1876, Paul Gervais named a newer genus Thylacomorphus based on skull that he thought to belong to an animal closely related to thylacines, the type species being T.

[13][14][15] 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.

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 Palaeogene zones unit MP13, possibly up to MP10), although some authors consider them to be a separate family in the form of the Dacrytheriidae.

[28] Some researchers considered the selenodont families Anoplotheriidae (represented below by Anoplotherium and Dacrytherium), Xiphodontidae, and Cainotheriidae to be within Tylopoda due to postcranial features that were similar to the tylopods from North America in the Palaeogene.

[30][31] 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.

[31] The phylogenetic tree published in the article and another work about the cainotherioids is outlined below:[32] Eurodexis russelli Dichobune leporina Amphimeryx murinus Xiphodon castrense Paratoceras coatesi Eotylopus reedi Parvitragulus priscus Lophiomeryx chalaniati Archaeomeryx optatus Mixtotherium cuspidatum Anoplotherium latipes Dacrytherium ovinum Robiacina lavergnesis Robiacina minuta Robiacina quercyi Palembertina deplasi Paroxacron bergeri Paroxacron valdense Oxacron courtoisii Cainotherium laticurvatum Caenomeryx filholi Caenomeryx procommunis Plesiomeryx cadurcensis Plesiomeryx huerzeleri In 2020, Vincent Luccisano et al. created a phylogenetic tree of the basal artiodactyls, a majority endemic to western Europe, from the Palaeogene.

The phylogenetic tree as produced by the authors is shown below:[30] 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.

The prominent part of the petrosal of Diplobune shows complicated positions and barely overlap with the skull's underside whereas that of Anoplotherium protrudes strongly around the internal auditory meatus and straightens towards the back area.

The internal acoustic meatus canal of the ear has a deep, oval shape with fixed boundaries from clear edges, containing two roughly equal in size foramina.

The upper surface of the cerebral hemispheres of Diplobune is flatter, and the neocortex lowering forward from the approximate back third of its length so that the latter can connect with the base of the olfactory peduncle.

The femur of D. minor is characterized by its lesser trochanter being close to the spheroidal femoral head, the distance separating them being equivalent to 1/4 of the bone's length as opposed to 1/3 for A. commune and D. secundaria.

[27] 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.

Although it is not possible to eliminate it from the Category 1 score (extremely slow like sloths), the lack of variability of inner ear morphologies compared to tragulids suggest unlikeliness of such low speed.

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.

[51] 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, which weighed over 2,000 kg (4,400 lb).

The extinction of the Lophiodontidae was part of a faunal turnover, which likely was 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.

[22][29] Diplobune 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 (recently determined to be polyphyletic, however), Cebochoeridae, Amphimerycidae, and Cainotheriidae.

[51] 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).

[54] Other mammal groups present in the late Eocene of western Europe represented the leptictidans (Pseudorhyncocyonidae),[62] primates (Adapoidea and Omomyoidea),[63] eulipotyphlans (Nyctitheriidae),[64] chiropterans,[50] herpetotheriids,[65] apatotherians,[66] and endemic rodents (Pseudosciuridae, Theridomyidae, and Gliridae).

[67] The alligatoroid Diplocynodon, present only in Europe since the upper Paleocene, coexisted with pre-Grande Coupure faunas as well, likely consuming insects, fish, frogs, and eggs due to prey partitioning previously with other crocodylomorphs that had since died out by the late Eocene.

[70] The MP18 locality of La Débruge of France indicates that D. secundaria coexisted with a wide variety of mammals, 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 Anoplotherium, tapirulid Tapirulus, xiphodonts Xiphodon and Dichodon, cainothere Oxacron, amphimerycid Amphimeryx, and the anthracothere Elomeryx.

The MP19 locality of Escamps has similar faunas but also includes the herpetotheriid Amphiperatherium, pseudorhyncocyonid Pseudorhyncocyon, bats (Hipposideros, Vaylatsia, Vespertiliavus, Stehlinia), primates (Microchoerus, Palaeolemur), cainothere Paroxacron, and xiphodont Haplomeryx.

The Turgai Strait 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.

[79][80][72][81] The Grande Coupure saw the extinctions of many artiodactyl genera previously endemic of Europe, including Anoplotherium and all representatives of "choeropotamids" (Amphirhagatherium, Choeropotamus), xiphodontids (Xiphodon, Dichodon) and amphimerycids (Amphimeryx).

[15] The deposit of Itardies, in addition to D. minor, yielded remains of the herpetotheriid Amphiperatherium, nyctithere Darbonetus, erinaceid Tetracus, various bats, large assemblages of rodents, hyaenodonts Hyaenodon and Thereutherium, amphicynodont Amphicynodon, enigmatic feliforms (Stenogale, Stenoplesictis, and Palaeogale), nimravid Nimravus, palaeothere Plagiolophus, rhinocerotid Ronzotherium, cainotheres Plesiomeryx and Caenomeryx, tragulid Iberomeryx, and the bachitheriid Bachitherium.

1877 illustrations of Diplobune bavarica fossil remains
Illustrations of Diplobune quercyi cranial and dental remains, 1891-1893
Portrait of Ludwig Rütimeyer , who originally erected the name Diplobune as a subgenus, which then was determined to be a distinct genus
D. secundaria jaw remains, National Museum of Natural History, France
Endocranial cast of Diplobune , Stuttgart State Museum of Natural History
Illustrations of the front foot (A) and hind foot (B) of Diplobune quercyi , 1891–1893. The fingers on each foot are labeled as "II," "III," and "IV."
Estimated size comparisons of D. minor and D. secundaria based on known fossil remains
Estimated body masses (kg) of Palaeogene artiodactyls based on recalculated trochlear widths (Li1) in comparison to estimates from Martinez and Sudre (1995)
A southern tree hyrax ( Dendrohyrax arboreus ), an arboreal folivore
Palaeogeography of Europe and Asia during the middle Eocene with possible artiodactyl and perissodactyl dispersal routes.
Reconstruction of A. commune . Anoplotherium and Diplobune commonly coexisted with each other based on being often found in many of the same Late Eocene localities.
Restoration of Anthracotherium magnum , an anthracothere genus that arrived in western by the Grande Coupure