Eric De Bast

Euarchontan affinity of Paleocene Afro-European adapisoriculid mammals and their origin in the late Cretaceous Deccan Traps of India.

The controversial family Adapisoriculidae, a group of shrew-sized Paleocene mammals, had proposed relationships with insectivores, marsupials and more recently to plesiadapiforms. Adapisoriculid remains are numerous in the early Paleocene locality of Hainin, Belgium, and allow us a test of these different phylogenetic hypotheses. Here, we identify the first tarsal bones of adapisoriculid mammals. The highly specialised bones indicate an arboreal mode of life with euarchontan affinity. Moreover, the tarsal bones are morphologically very close to those of the late Cretaceous Deccanolestes from the Deccan intertrappean beds of India, and also share several characters with the Paleocene plesiadapiforms and the extant cynocephalid dermopterans. The adapisoriculid affinities of Deccanolestes are also confirmed by tooth morphology, indicating that Deccanolestes is a primitive member of this family. These phylogenetic affinities suggest a paleobiogeographic scenario for the family with dispersal either via East Africa or across the Tethys area.

First Clarkforkian equivalent Land Mammal Age in the latest Paleocene basal Sparnacian facies of Europe: fauna, flora, paleoenvironment and (bio)stratigraphy.

The Paleocene-Eocene Thermal Maximum (PETM) is correlated with the first occurrences of earliest modern mammals in the Northern Hemisphere. The latest Paleocene Clarkforkian North American Land Mammal Age, that has yielded rodents and carnivorans, is the only exception to this rule. However, until now no pre-PETM localities have yielded modern mammals in Europe or Asia. We report the first Clarkforkian equivalent Land Mammal Age in the latest Paleocene deposits of the basal Sparnacian facies at Rivecourt, in the north-central part of the Paris Basin. The new terrestrial vertebrate and macroflora assemblages are analyzed through a multidisciplinary study including sedimentologic, stratigraphic, isotopic, and palynological aspects in order to reconstruct the paleoenvironment and to evaluate biochronologic and paleogeographic implications. The mammals are moderately diverse and not abundant, contrary to turtles and champsosaurs. The macroflora is exceptional in preservation and diversity with numerous angiosperms represented by flowers, fruits, seeds and wood preserved as lignite material, revealing an abundance of Arecaceae, Betulaceae, Icacinaceae, Menispermaceae, Vitaceae and probably Cornaceae. Results indicate a Late Paleocene age based on carbon isotope data, palynology and vertebrate occurrences such as the choristoderan Champsosaurus, the arctocyonid Arctocyon, and the plesiadapid Plesiadapis tricuspidens. However, several mammal species compare better with the earliest Eocene. Among these, the particular louisinid Teilhardimys musculus, also recorded from the latest Paleocene of the Spanish Pyrenees, suggests a younger age than the typical MP6 reference level. Nevertheless, the most important aspect of the Rivecourt fauna is the presence of dental remains of a rodent and a “miacid” carnivoran, attesting to the presence of two modern mammalian orders in the latest Paleocene of Europe. Interestingly, these two groups are also the only modern groups recorded from the latest Paleocene of North America, making Rivecourt the first direct equivalent to the Clarkforkian Land Mammal Age outside of North America.

Dental and tarsal anatomy of ‘Miacis’ latouri and a phylogenetic analysis of the earliest carnivoraforms (Mammalia, Carnivoramorpha)

ABSTRACT One of the earliest basal carnivoraforms, Miacis latouri, previously known by only two teeth from the earliest Eocene of Dormaal, Belgium, is here described based on about 280 new specimens from Dormaal, allowing illustration of almost the entire deciduous and permanent dentition and thus giving information on the dentition of an early basal carnivoraform species and its variability. Based on the dental features, we refer the species to a new genus, Dormaalocyon. We identify possible sexual dimorphism in D. latouri that is less pronounced than in Uintacyon rudis. We also describe for the first time the tarsal bones (calcaneum and astragalus) of D. latouri; these indicate arboreal capabilities for this species. In order to ascertain the position of Dormaalocyon among basal carnivoraforms, we performed a phylogenetic analysis of the carnivoramorphans. Among basal carnivoraforms, three groups are recovered: the Uintacyon group, Oodectes group, and the Vulpavus group. Dormalocyon is one of the most primitive carnivoraforms and is closely related to North American Vulpavus and Miacis species. We propose that the two latter genera are North American with an ancestry that involves the European Dormaalocyon; this implies a dispersal of carnivoraforms from Europe to North America near the Paleocene-Eocene boundary. Finally, the topology of the phylogenetic tree supports a Paleocene radiation of the carnivoraforms, which is presently unknown.

Diversity of the adapisoriculid mammals from the early Palaeocene of Hainin, Belgium

Adapisoriculidae are an enigmatic group of small mammals known from the late Cretaceous of India, and from the early Palaeocene to early Eocene of Europe and Africa. Based on their primitive dental morphology, they have been classified as didelphids, nyctitheriids, leptictids, mixodectids, tupaiids, and palaeoryctids. While the latest hypothesis based on dental morphology suggests an affinity with Lipotyphla, postcranial remains indicate a close relationship with Euarchonta. Here, we present new adapisoriculid dental remains from the early Palaeocene locality of Hainin (Belgium). Adapisoriculidae are particularly abundant in Hainin, where they represent about one third of the mammalian fauna, offering new insights into both their specific and generic phylogenetic interrelationships. We describe three new species (Afrodon gheerbranti sp. nov., Bustylus folieae sp. nov. and Proremiculus lagnauxi gen. et sp. nov.) and document the previously unknown lower dentition of Bustylus marandati. The diversity of dental morphologies observed in the Hainin fauna suggests different interrelationships than previously suggested. In particular, the genus Proremiculus is considered morphologically intermediate between Afrodon and Remiculus, and the latter is no longer recognised as the sister group of Adapisoriculus. Although the highest diversity of adapisoriculids occurs in Europe, the oldest and most primitive members of the family were found in India and Africa, respectively. The geographic origin of the family could thus be located in any of these three continents, depending on the importance attributed to each of these factors. The coexistence of primitive and derived adapisoriculids at Hainin might indicate a very quick diversification in Europe, probably starting around the K-T boundary.

Reassessment of the small ‘arctocyonid’ Prolatidens waudruae from the early Paleocene of Belgium, and its phylogenetic relationships with ungulate-like mammals

ABSTRACT ‘Arctocyonids’ are generally considered as including some of the most primitive ‘ungulates’ from the Paleocene. Although more than 15 genera are known from North America, European members of this order are less common and mainly belong to derived genera such as Arctocyon. However, one species of primitive arctocyonid, Prolatidens waudruae, was described from the early Paleocene of Hainin, Mons Basin, Belgium. Here we describe new dental positions of this small taxon, including for the first time upper molars and upper fourth premolar. Morphological comparisons confirm the position of P. waudruae among primitive ‘ungulates,’ with the closest North American arctocyonids being Prothryptacodon furens and Oxyprimus galadrielae. Oxyprimus galadrielae features slightly more primitive morphological traits than both other species. Apheliscids share several characters with Prolatidens, but the latter lacks the apomorphies defining the family. Among ‘arctocyonids,’ Prolatidens shares with only Protungulatum and Oxyprimus the incomplete lingual cingulum at the base of the protocone of M1. Prolatidens waudruae is unique among Procreodi in its combination of primitive and derived characters. The cladistic analysis places P. waudruae close to the base of the ingroup, indicating that this species is among the most primitive members of the Paleocene ‘ungulates.’ However, the lower nodes of the trees are not well supported and definitive conclusions should await more complete specimens and analysis. Apheliscids are situated relatively far from Prolatidens, suggesting that the resemblances between them are better considered as convergences. Based on the morphological comparisons, the arctocyonid from Hainin correlates best with North American Torrejonian taxa.

The oldest Cenozoic mammal fauna of Europe: implication of the Hainin reference fauna for mammalian evolution and dispersals during the Paleocene

The mammal fauna of Hainin is particularly interesting as the oldest in the Cenozoic of Europe, and the earliest reference level (MP1–5) of the mammalian biochronological scale for the European Palaeogene. This paper summarizes the mammal taxa discovered in the fauna, presents an analysis of the fauna as a whole (relative abundance and species richness), and describes four new eutherians: Belgoryctes thaleri gen. et sp. nov., Eurolestes dupuisi gen. et sp. nov., Quadratodon sigei gen. et sp. nov. and Cingulodon magioncaldai gen. et sp. nov. The assemblage is relatively small (about 400 dental specimens), characterized by a high diversity and abundance of small insectivorous species and very low abundance of ‘plesiadapiforms’ and ‘condylarths’. By comparison with younger European Paleocene faunas, ‘condylarths’ and ‘plesiadapiforms’ became more and more abundant and diverse through the Paleocene but collapsed at the Paleocene–Eocene Boundary. ‘Proteutherians’ declined steadily, while multituberculates remained diverse, although the early Paleocene was populated mainly by Kogaionidae whereas the late Paleocene was dominated by Neoplagiaulacidae. The palaeoecology of Hainin is deduced from the mammal assemblage: the local environment was likely a forested area. Stratigraphically, the Hainin deposits are most likely of late Danian age, and biochronologically its fauna represents a partial equivalent of the North American Torrejonian Land Mammal Age. When compared to younger Paleocene faunas of Europe, the composition of the Hainin fauna reveals that a relatively important intercontinental dispersal of mammals occurred around the Danian–Selandian boundary, roughly corresponding to the Torrejonian–Tiffanian boundary. This dispersal is marked by the arrival in Europe of typically North American taxa such as arctocyonids, plesiadapids and neoplagiaulacid multituberculates. Additional exchanges of lesser magnitude probably also occurred around the Selandian–Thanetian boundary (i.e. during the Tiffanian), although the evidence is less compelling and mainly concerns the plesiadapids Chiromyoides and Plesiadapis. http://zoobank.org/urn:lsid:zoobank.org:pub:A240BAE2-7446-4DBD-A597-7BEFE58447DF

New carnivoraforms from the latest Paleocene of Europe and their bearing on the origin and radiation of Carnivoraformes (Carnivoramorpha, Mammalia)

ABSTRACT We report the discovery of the earliest European carnivoraforms, based on two new taxa from the latest Paleocene of France and Romania. A new species of Vassacyon, V. prieuri, from the locality of Rivecourt (MP6b; Oise, France) (MP = Mammalian Paleogene reference levels) is described based on a dentary fragment and isolated teeth. This species displays several primitive features compared with species of Vassacyon known from the early Eocene of North America and Europe. A second Paleocene carnivoraform, cf. Gracilocyon sp., is described based on fragmentary specimens from the locality of Jibou (MP6b; Transylvania, Romania). Carnivoraformes were previously unknown in Europe before the Eocene, and Vassacyon prieuri and cf. Gracilocyon sp. are their oldest records in Europe. These discoveries favor the hypothesis of a dispersal of these two genera from Europe to North America during the Paleocene-Eocene Thermal Maximum. Vassacyon prieuri and cf. Gracilocyon sp. strengthen support for the level MP6b and its correlation with the Clarkforkian. The geographic origin of the Carnivoraformes remains unknown. However, based on the observed biochronological and geographic distributions of the carnivoraforms, we infer a possible Asian origin for this group. In this scenario, Gracilocyon and Vassacyon dispersed to Europe during the Clarkforkian MP6b time, probably together with rodent taxa, whereas Uintacyon dispersed from Asia to North America shortly after rodents and tillodonts.

A Strigogyps-like bird from the middle Paleocene of China with an unusual grasping foot

ABSTRACT We describe a new avian species, Qianshanornis rapax, gen. et sp. nov., from the middle Paleocene Wanghudun Formation of the Qianshan Basin in Anhui Province, China. The holotype consists of an incomplete articulated foot and a few associated bones, mainly of the leg, wing, and pectoral girdle. Qianshanornis rapax is characterized by a derived foot morphology and may have had a hyperextendible second toe, which has so far only been reported for some Mesozoic taxa. The new species is markedly different from all other known Cenozoic birds and is here classified in the new taxon Qianshanornithidae. The leg bones, especially the distal end of the tibiotarsus, most closely resemble those of Strigogyps (Ameghinornithidae) from the Eocene, and possibly Oligocene, of Europe, but unlike the latter, the much smaller Q. rapax appears to have had well-developed flight capabilities.

Plesiadapid mammals from the latest Paleocene of France offer new insights on the evolution of Plesiadapis during the Paleocene-Eocene transition

ABSTRACT Plesiadapidae are among the most successful mammal families of the Paleocene, but in North America they disappear abruptly around the Paleocene-Eocene boundary. In contrast, in Europe, they survive a few million years into the Eocene, although only as the genus Platychoerops. The latest Paleocene deposits of Petit-Pâtis (Paris Basin, France) have produced three new plesiadapid species, one of each genus known in Europe: Plesiadapis ploegi, sp. nov., Platychoerops boyeri, sp. nov., and Chiromyoides mauberti, sp. nov. Each of these new species is represented by the very characteristic upper incisor, thus ascertaining their concomitant presence and in particular the spatial and temporal coexistence of Plesiadapis and Platychoerops. Plesiadapis ploegi, sp. nov., is morphologically intermediate between Plesiadapis tricuspidens and Platychoerops russelli, with a tricuspid I1 typical of Plesiadapis and a semimolariform p4 closer to Platychoerops. Its relatively high morphological variability is illustrated. Platychoerops boyeri, sp. nov., has the simple derived I1 of all Platychoerops and a p4 slightly more molariform than that of Ples. ploegi. Chiromyoides mauberti, sp. nov., is closest to Chiromyoides campanicus, but it is smaller and has a particular I1 with multiple posterocones. The systematic position of ‘Platychoerops’ georgei is discussed; this taxon is considered a chimera, and its type I1 belongs to either Chiromyoides or Plesiadapis. Cladistic analysis highlights the paraphyly or polyphyly of all genera of Plesiadapidae. Finally, there is some indication of morphological convergences between European and North American plesiadapids, which may be the result of similar environmental changes on both continents just before the Paleocene-Eocene boundary.