Bats are among the most numerous and widespread mammals today, but their fossil record is comparatively meagre and their early evolution poorly understood. Here we describe a new fossil bat from dental remains recovered from late Early Eocene sediments at Prémontré, northern France. This 50 million-year-old bat exhibits a mosaic of plesiomorphic and apomorphic dental features, including the presence of three lower premolars, a single-rooted p3, short p4 with metaconid, myotodont lower molars and a tall coronoid process of the dentary. This combination of features suggests it is an early member of Vespertilionidae, today’s most speciose and geographically widespread bat family. The Prémontré bat has bearing on hypotheses about the origins of vesper or evening bats (Family Vespertilionidae), as well as crown-group chiropterans.


  

The study of endocasts, or casts of the endocranial space, have played an important role in shaping understanding of mammalian, and particularly primate, brain evolution. Recently, the reconstructions of three-dimensional virtual endocasts from high-resolution computed tomography images have allowed for the visualization and quantification of endocasts in several Paleocene and Eocene primate species. Here we present the virtual endocast of MaPhQ 289 (informally known as the Montauban 9 skull), a specimen of Necrolemur antiquus Filhol 1873, a middle to late Eocene European primate of the family Microchoeridae. The virtual endocast of MaPhQ 289 reveals a lissencephalic surface morphology with expanded temporal poles and minimal overlap of the cerebellum or olfactory bulb by the cerebrum, which closely resembles the morphology of the endocast of its contemporary relative, Microchoerus erinaceus (Primates, Microchoeridae). MaPhQ 289 yields an endocranial volume (ECV) of 2.36 cm3, about 60% smaller than the volume of the most commonly cited ECV of N. antiquus. Thus, the size of the brain of N. antiquus relative to its body size is likely to be smaller than has been reported in previous literature, highlighting the importance of corroborating older ECV estimates with new evidence using 3-D imaging techniques. 

The stapes is the last of the middle ear ossicle chain and the smallest bone of the mammalian skeleton. Because it rests on the membrane of the fenestra vestibuli during life, the stapes may often fall within the bony labyrinth cavity when soft structures decay after death. In this work, we highlight the unexpected role that the bony labyrinth plays in the preservation of the stapes. Systematic investigation of the bony labyrinth of 50 petrosal bones of extinct and extant artiodactyls led to the discovery of eight cases of “intralabyrinthine” stapes. Three dimensional reconstructions of these stapes allow documenting stapes morphology of four extinct artiodactyl taxa: Microstonyx erymanthius (Suidae), Elomeryx borbonicus (Hippopotamoidea), ?Helohyus plicodon (Helohyidae), and an undetermined Cainotheriidae; and four extant ones Choeropsis and Hippopotamus (Hippopotamidae), and Tayassu and Phacochoerus (Suoidea). ?Helohyus plicodon from the Middle Eocene documents the oldest stapes known for the order Artiodactyla. Morphological study and metric analyses of our sample of artiodactylan stapes show that stapes are likely to carry relevant phylogenetic characters/signal within artiodactyls, and a potential Euungulata signature. 

Résumé :
L’étude des rongeurs Theridomorpha permet de suivre le déroulement d’une radiation adaptative pendant toute sa durée (Eocène moyen-Oligocène terminal), sur un territoire restreint à l’extrémité ouest de l’Europe Occidentale. Dans ce papier, la phylogénie de ce groupe est établie à partir d’une analyse cladistique reposant sur l’examen de 315 caractères (310 dentaires). Le groupe d’intérêt comprend 110 des 132 espèces (24 genres) de Theridomyoidea et deux genres encore inclus jusqu’ici dans les Reithroparamyinae qui rejoignent les Theridomorpha. Les groupes externes comprennent des Glires basaux, Cocomys, Tanquammys et 16 Ischyromyiformes. Un cadre phylogénétique robuste est produit, qui permet de clarifier la systématique des Theridomorpha. La position des Remyoidea (nov. sup.fam.) au sein des Ischyromyiformes, extérieure aux Theridomorpha, est confortée. Les Protadelomys et Tardenomys sont à la base des Theridomyoidea, avant la séparation en deux clades correspondant aux familles Pseudosciuridae et Theridomyidae. Les sous-familles sont consolidées : Pseudosciurinae et Sciuroidinae pour les Pseudosciuridae ; Issiodoromyinae, Oltinomyinae, Columbomyinae, Theridomyinae, auxquelles s’ajoute au moins une nouvelle sous-famille (Patriotheridomyinae), pour les Theridomyidae. La topologie des chrono-espèces (sensu Simpson), traitées antérieurement comme lignées évolutives, apparaît dans la plupart des cas sous forme de clades successifs dans lesquels les espèces sont le plus souvent arrangées de manière pectinée, émergeant dans l’ordre stratigraphique. L’analyse des caractères aux principaux nœuds permet de consolider les caractères diagnostiques des taxons et les tendances évolutives, ainsi que de discuter des divers parallélismes et convergences dans l’évolution des structures et patrons dentaires (e.g., émail des incisives unisérié chez les Issiodoromyinae et les Patriotheridomyinae, ou pseudo-multisérié chez les Blainvillimys les plus hypsodontes, les Protechimys et Archaeomys ; patrons dentaires téniodontes ; allongement des dents déciduales chez les Patriotheridomyinae, Issiodoromyinae et Theridomyidae ; sélénodontie ou lophodontie). Les dynamiques évolutives traduites par les changements morphologiques sont mises en relation avec les variations environnementales. Enfin, les implications biochronologiques de l’évolution des Theridomyoidea sont discutées.
Abstract:
The adaptive radiation of the rodents Theridomorpha occurred during a limited time window (middle Eocene to late Oligocene), on an area restricted to Western Europe. In this paper, the phylogeny of this group is established via a cladistic assessment of 315 morphological characters (310 dental). The group of interest encompasses 110 upon 132 species (24 genera) of Theridomyoidea, and two genera formerly included within the Reithroparamyinae, and which are included here within the Theridomorpha. The outgroups include basal Glires, Cocomys, Tanquammys and 16 Ischyromyiformes. A robust phylogenetic frame is produced, which allows clarifying the systematics of the Theridomorpha. Within the Ischyromyiformes, the Remyoidea (nov. supfam.) are set apart from the Theridomorpha. Protadelomys and Tardenomys represent the earliest offshoots of the Theridomyoidea, before the dichotomy between Pseudosciuridae and Theridomyidae. It supports the former subfamilies Pseudosciurinae and Sciuroidinae within the Pseudosciuridae; and for the Theridomyidae: the Issiodoromyinae, Oltinomyinae, Columbomyinae, Theridomyinae, with at least one new subfamily (Patriotheridomyinae). The topologies of the chronospecies (sensu Simpson), formerly considered as evolutionary lineages, appear in most cases as successive clades, in which the species are generally pectinately arranged and emerging in the stratigraphic order. The analysis of characters supporting the main nodes allow consolidating the diagnosic characters of the taxa and their evolutionary trends, as well as discussing the various cases of parallelism and convergence in the evolution of structures and dental patterns (e.g., uniserial incisor enamel for Issiodoromyinae and Patriotheridomyinae, or pseudo-multiserial for the most hypsodont Blainvillimys, Protechimys and Archaeomys; taeniodont dental patterns; lengthening of deciduous premolars for Patriotheridomyinae, Issiodoromyinae and Theridomyidae; selenodonty or lophodonty).
Evolutionary dynamics are analysed with respect to environmental changes. Finally, biochronological implications of the evolution of Theridomyoidea are discussed.
  

HGL50 is a latest Early-early Middle Eocene vertebrate-bearing locality located in Western Algeria. It has produced the richest and most diverse fauna of amphibians and squamate reptiles reported from the Palaeogene of Africa. Moreover, it is one of the rare faunas including amphibians and squamates known from the period of isolation of Africa. The assemblage comprises 17 to 20 taxa (one gymnophionan, one probable caudate, three to six anurans, seven ‘lizards’, and five snakes). Two new taxa were recovered: the anuran Rocekophryne ornata gen. et sp. nov. and the snake Afrotortrix draaensis gen. et sp. nov. The locality has also yielded the first confirmed anilioid snake, the first Palaeogene gymnophionan, and probably the first caudate from the Palaeogene (and possibly from the Tertiary) of Africa. The presence of a caudate at that time in Africa would be of particular interest; unfortunately, the available material does not permit a definitive identification. The fauna comprises Gondwanan and more specifically West Gondwanan vicariants, probably autochthonous groups and a Eurasian immigrant (assuming that the identification of the caudate is accurate). The fauna from HGL50 is clearly distinguished from the few other Eocene assemblages of Africa. However, if this results largely from differences in geological ages, geographic positions of the localities and mainly differences in environments took a part in the composition of the faunas. 

A large avian femur recently discovered at the Late Cretaceous Montplo-Nord locality at Cruzy (Hérault, southern France) is referred to the giant bird Gargantuavis philoinos. The estimated mass of the bird is 57 kg, within the range of living cassowaries. The specimen provides new evidence about the anatomy of G. philoinos, notably showing that the distal end of the femur was similar to that of modern birds in having a condylus lateralis subdivided into two semicondyles. A new diagnosis of Gargantuavis philoinos is provided and the taxon is placed in a new family of basal ornithurines.
  

The discovery of new hippopotamid material from the late Miocene Baynunah Formation (Abu Dhabi, United Arab Emirates) has prompted the revision of the existing material of this as yet unnamed fossil taxon. The Baynunah hippopotamid appears to be distinct from all other contemporary and later species in having a relatively more elongate symphysis, a feature similar to the earlier (and more primitive) Kenyapotamus. Yet, the Baynunah hippopotamid presents a dentition typical of the Hippopotaminae. It is therefore a distinct species attributed to the later subfamily, described and named in this contribution. This species provides further evidence for a ca. 8 Ma evolutionary event (termed “Hippopotamine Event”) that initiated the spread and ecological significance of the Hippopotaminae into wet habitats across Africa and Eurasia. The morphological affinities of the new species from Abu Dhabi suggest that the Arabian Peninsula was not a dispersal route from Africa toward southern Asia for the Hippopotamidae at ca. 7.5 Ma to 6.5 Ma. 

Abstract: A very tiny cuckoo, Chambicuculus pusillus, was recently described after a few fragments of tarsometatarsi from the late Early/early Middle Eocene of Djebel Chambi, Tunisia. New remains, notably a coracoid, confirm the attribution of this genus to the recent family Cuculidae. This coracoid shows a very elongate and strap-like processus procoracoideus. This morphological feature is otherwise only known in the family Cuculidae. The characteristics of the coracoid and tarsometatarsi show that Chambicuculus is morphologically more advanced over the other stem cuculids described in Europe and North America. Chambicuculus is the oldest Cuculidae known so far. 

This paper provides a revision of the early and middle Eocene European rodents previously referred to as Ischyromyoidea, including taxa considered to be at the origin of the Theridomorpha. The use of an accurate dental terminology and a better understanding of the size and shape of their infra-orbital foramen (i.o.f.) led us to a substantial revision of this group, which allowed to better characterize them and to appreciate their variability. On these bases, phylogenetic analyses (cladistic and standard Bayesian
approaches) of early Ypresian to late Priabonian European rodent species were undertaken in order to highlight the root of the early Theridomorpha and its content. In this paper, the phylogeny was established based on 343 characters (338 dental) through 45 early Paleogene taxa using both cladistic and bayesian analyses. The ingroup included on one hand a few North American genera (Reithroparamys, Microparamys, and Acritoparamys) and European ones (Eogliravus, Ailuravus, Corbarimys, Meldimys, Euromys, Plesiarctomys, and Pseudoparamys) considered until now as being related with the North American superfamily Ischyromyoidea. On the other hand, it included genera close to the root of the Theridomorpha (Sparnacomys, Pantrogna, and Hartenbergeromys) and early Theridomyoidea (Masillamys, Protadelomys, and some Pseudosciuridae). The phylogenetic results obtained via the two
distinct reconstruction approaches are consistent in virtually all relationships. The proposed systematics here derives from these phylogenetic results. This phylogenetic context led us to change the suprafamilial, familial, subfamilial or generic attribution of several species. Characters of Theridomorpha, like the obliquely developed postprotocristid allied with the occurrence of a metalophulid I, have been found in genera previously considered as Ischyromyidae (Pseudoparamys, Euromys, Sparnacomys, Meldimys, Pantrogna, and Hartenbergeromys) as well as the large i.o.f., when preserved (Pseudoparamys, Hartenbergeromys, and Masillamys). Based on these morphological observations and new phylogenetic considerations, the content of the Theridomorpha clade is here enlarged, thereby extending back the first theridomorph radiations to the early Eocene. Aside, a new taxon (Reinomys rhomboides gen and sp. nov.) is described from Avenay. In addition, a new genus, Auroremys, is created for the species subita (Comte et al., 2012) from Chery-Chartreuve. 

The Las Águilas site near Porvenir de Jalpa, Coahuila, Mexico, is extremely rich in tetrapod remains comprising both bones and trackways of several dinosaur taxa of late Campanian age. Within a 50 m thick section we identified at least nine layers with dinosaur bone assemblages. In one of these the dinosaur bones are associated with remnants of eusuchian crocodilians, turtles, plesiosaurs, pterosaurs, tyrannosaurids, dromaeosaurids, parksosaurid, hadrosaurids, ceratopsids, and ankylosaurs. This layer is also rich in coprolites of turtles, crocodilians and likely theropods, thus providing evidence for the wealth of Late Cretaceous vertebrate life in the area. 

Welcome to the 18th conference of the EAVP, the first online meeting of our association. The pandemic emergency made it impossible to organize the in-person meeting in Benevento as we all had hoped. However, we couldn’t miss another EAVP meeting. Therefore, this year we are meeting online, trying to make the experience the closest to the in-person meeting possible, in order to offer the delegates the opportunity to share knowledge, build new networks and reinforce the old ones. We have received 137 communications, with more than 150 delegates from 24 countries. All the abstracts have passed a peer review process and are part of this special volume of Palaeovertebrata, the official journal of the EAVP. This year we are also offering a variety of workshops, roundtables and symposia on different topics. These include the annual “Pride EAVP: An LGBTQ+ Roundtable” and “Women in Palaeontology Roundtable Discussion”, together with the workshops on “Gendered Perspective in Palaeontological Research: from Definition to Action”, “International Palaeontology Education: Virtual Teaching and Real-World Learning”, “Stepping out of Academia: Why, When and How?”, “Introduction to Hypothesis Testing in Statistics”, “The Early-Middle Pleistocene Transition: Marked Mammal Turnover and Ecosystem Dynamic” (included in the early event for the XXI INQUA Congress in Rome 2023, “A Mediterranean Perspective on Quaternary Sciences”). To conclude, we are hosting two symposia on “Palaeoart: Diversity on and behind the Canvas” and “3D fossils, Robotic and Experimental Palaeontology”. We wish you all a happy and productive meeting. And see you in Benevento next year! 

We report the first confirmed fossil bats from North Dakota, including a new species referable to the Vespertilionidae represented by a maxilla with P4-M3 from the Brule Formation, Fitterer Ranch local fauna, early Oligocene, Whitneyan North American Land Mammal Age. Unassociated postcranial fragments of the humerus and femur also represent a vespertilionoid, but appear to reflect a different, unidentified species. The new taxon, Quinetia frigidaria sp. nov., is referred to the genus Quinetia, previously known only from approximately contemporaneous deposits in Europe. The new species is larger than Quinetia misonnei from the early Oligocene of Belgium. It is similar in some morphological characters to Chadronycteris rabenae (Chiroptera incertae sedis) of the late Eocene (Chadronian) of northwestern Nebraska and to Stehlinia species (?Palaeochiropterygidae) from the Eocene and Oligocene of Europe, but differs from each in morphological details of the dentition and maxilla. An unassociated talonid of a lower molar from Fitterer Ranch shows myotodont morphology, unlike the nyctalodont lower molars in Q. misonnei, and thus represents a second chiropteran taxon in the fauna. Quinetia frigidaria is a member of a Paleogene radiation of bats near the low point of the Eocene-early Oligocene decline in global temperatures, increased seasonal aridity, and loss of tropical floras from mid-latitude North America. 

Ornithocheirid pterosaurs were the largest of the toothed pterodactyloids and had a worldwide distribution, although their fossil record is fragmentary, with the exception of the north-eastern Brazilian Crato and Santana Formations (Aptian, ?Albian, Early Cretaceous). With Istiodactylidae, they were also the only toothed pterosaurs that survived into the Cenomanian (Late Cretaceous), becoming extinct at the end of this period. Here we report on an ornithocheirid metacapus from the Late Cenomanian laminated limestone of north-eastern Mexico discovered about 120 km north-west of Ciudad Acuña, northern Coahuila at the south banks of Rio Bravo. The specimen comprises a fragmentary distal syncarpal, a crushed but complete metacarpal IV, two fragmentary preaxial metacarpals and a possible fragmentary terminal left wing finger phalanx. It represents the geologically youngest known ornithocheirid worldwide. We suggest that ornithocheirid pterosaurs may have become extinct because of massive sea level fluctuations during the mid to late Cretaceous that may have obliterated their breeding sites on coastal plains and low lying islands. 

The Late Miocene site of Gritsev (MN 9, Ukraine) has yielded a very rich bat fauna, the remains of which are well preserved. Compared to other Neogene bat assemblages of Europe, the Gritsev bat community is unique in preserving plecotine bats, which are rare from Neogene sites. Some peculiar and new bat species, including a large plecotin Otonycteris, already were described from the Gritsev mammal site. Here we report new records of small plecotin bats from Gritsev, including a new taxon, Barbastella maxima nov. sp. This is the earliest reliable fossil record of this genus and it differs from more recent species of Barbastella in being considerably larger. The evolutionary patterns in the odontology within the tribe Plecotini, supported by biostratigraphical distribution of fossil records of Plecotus are discussed. The morphological peculiarities of the new fossils of plecotine bats from Gritsev are discussed in connection with its possible taxonomical affinity. 

This paper describes the first hexanchid tooth from the Tithonian (Late Jurassic) of New Zealand. For the moment, this tooth represents the earliest representative of the fossil genus Notidanodon in the world and one of the most ancient neoselachians in the Southern Hemisphere. Despite the perfect state of preservation of the unique tooth, the species is left in open nomenclature, pending the discovery of additional specimens. Few nominal species have been assigned to the genus Notidanodon. Four from Cretaceous deposits: N. antarcti Grande & Chatterjee, 1987, Notidanodon dentatus (Woodward, 1886), Notidanodon lanceolatus (Woodward, 1886), Notidanodon pectinatus (Agassiz, 1843) and only two from Paleocene: Notidanodon brotzeni Siverson, 1995, and Notidanodon loozi (Vincent, 1876). Considering the important morphological variations observed between some of these species, it seems obvious that the genus Notidanodon is not monophyletic and will need a revision in the future. 
  

Confident attribution of bipedal tridactyl dinosaur tracks to theropods or ornithopods can be challenging. Here we describe trackways produced by tetanuran dinosaurs, previously attributed to hadrosaurs, from Coahuila State, northeastern Mexico. Multiple trackways headed in the same direction suggest gregarious behaviour in these late Campanian theropods.  

A new Propalaeotherium species, clearly distinct from the genus Eurohippus, is described. It is characterized by having a similar size as P. voigti from the German Geiseltal localities (MP 11 to MP 13 reference-level), but differs in several features suggesting a slighty more derived morphology. It presents indeed less brachyodont crowns with less prominent and less elevated cingula, slightly larger relative surface of premolars, and a more marked metaconid splitting on cheek teeth. This new species is unknown from other European localities except the nearby Saint-Martin de Londres locality which has been considered older than the MP 13 level. 

Investigations of the Clinchfield Formation (middle Eocene, upper Bartonian) exposed at the Hardie Mine (Wilkinson County, Georgia, USA), produced 4,768 actinopterygian otoliths representing 14 taxa and increased the number of bony fishes threefold from the site. The somewhat limited richness was characterized by bonefishes, mud eels, conger eels, sea catfishes, cusk-eels, snooks, grunts, drums and croakers, and porgies. The assemblage had a relatively even distribution with Ophidiidae, Congridae, and Sciaenidae most common. Included in the otolith taxa was a new sciaenid genus and species, Eosciaena ebersolei, with unknown relationships to other Sciaenidae. The Clinchfield otoliths were compared to other middle and late Eocene in age otolith assemblages in Alabama, Mississippi, and Louisiana utilizing percentage similarity measurements. Analysis indicated that the Clinchfield otoliths were not greatly similar or greatly unlike the Moodys Branch and Yazoo Clay otolith assemblages. However, the Clinchfield showed little relationship to the slightly older Lisbon Formation in adjacent Alabama and is postulated to be related to global climatic and plate tectonic events. Biostratigraphically, the Clinchfield otolith taxa are essentially the same as the other formations except for the Lisbon, which has at least ten unique species. Abundances of Clinchfield otolith taxa indicate a possible sub-bioprovince in the eastern Gulf Coastal Plain. The Clinchfield otoliths indicate a tropical to perhaps subtropical, soft substrate, mainly normal marine to slightly reduced salinities, inner shelf (0–20 m) paleoenvironment with indications of proximal continental coastlines. This investigation represents an initial step in addressing the immensely understudied Paleogene otolith assemblages in Georgia.
  

Micro-CT imaging is an increasingly popular method in paleontology giving access to internal structures with a high resolution and without destroying precious specimens. However, its potential for the study of hypoplasia defects has only recently been investigated. Here, we propose a preliminary study to test whether hypoplastic defects can be detected with micro-CT (μCT) scan and we assess the costs and benefits of using this method instead of naked eye. To do so, we studied 13 fossil rhinocerotid teeth bearing hypoplasia from Béon 1 (late early Miocene, Southwestern France) as positive control and 11 teeth of the amynodontid Cadurcotherium (Oligocene, Phosphorites du Quercy, Southwestern France), for which enamel was partly or totally obscured by cement. We showed that all macroscopically-spotted defects were retrieved on 3D reconstructions and selected virtual slices. We also detected additional defects using μCT scan compared to naked eye identification. The number of defects detected using μCT was greater in the Cadurcotherium dataset (paired-sample Wilcoxon test, p-value = 0.02724) but not for our control sample (paired-sample Wilcoxon test, p-value = 0.1171). Moreover, it allowed for measuring width and depth of the defects on virtual slices (sometimes linked to stress duration and severity, respectively), which we could not do macroscopically. As μCT imaging is both expensive and time consuming while not drastically improving the results, we recommend a moderate and thoughtful use of this method for hypoplasia investigations, restricted for instance to teeth for which enamel surface is obscured (presence of cement, uncomplete preparation, or unerupted germs). 

Isolated elasmobranch teeth (sharks and rays) from the middle Eocene (Bartonian) Bandah Formation in the Jaisalmer District of Rajasthan, India are described. The remains improve our knowledge of the environment represented by this lithostratigraphic unit and the ecology preserved therein. Seventeen unequivocal taxa were identified, including Nebrius sp., Striatolamia aff. S. macrota, Brachycarcharias atlasi, B. lerichei, cf. Jaekelotodus sp., Carcharhinus mancinae, Rhizoprionodon sp., Physogaleus sp., Galeocerdo clarkensis, G. eaglesomei, Odontorhytis aff. O. pappenheimi, “Rhinobatos” sp., “Dasyatis” sp., Coupatezia sp., “Aetomylaeus” sp., “Rhinoptera” sp., and Ouledia aff. O. lacuna. Of these, “Aetomylaeus” sp., B. atlasi, C. mancinae, G. clarkensis, G. eaglesomei, cf. Jaekelotodus sp., Nebrius sp., Odontorhytis aff. O. pappenheimi, Ouledia aff. O. lacuna, and “Rhinoptera” sp. are reported from the middle Eocene of India for the first time. The Bandah Formation elasmobranch palaeofauna has close affinities to the Palaeocene-Eocene Tethyan/Paratethyan faunas of Africa, Madagascar, Asia, and Europe, and some taxa indicate a western hemisphere influence from North America. The Bandah Formation palaeofauna indicates that deposition occurred in a moderately shallow marine environment. The Bartonian age is primarily based on foraminifera but is corroborated by the presence of elasmobranch taxa that also occur in contemporaneous deposits elsewhere. The marine regression started during the early Palaeogene, and our study indicates that the sea completely withdrew from the Jaisalmer Basin after the deposition of the Bandah Formation. This event may have been synchronous with the middle Eocene uplift of the Himalayan-Tibetan Plateau.