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New material attributed to Agerinia smithorum from Casa Retjo-1 (early Eocene, NE Iberian Peninsula), consisting of 13 isolated teeth and a fragment of calcaneus, is studied in this work. These fossils allow the first description of the calcaneus and the upper premolars for the genus Agerinia, as well as the first description of the P2 and M2 for A. smithorum. The newly recovered lower teeth are virtually identical to the holotype of A. smithorum and are clearly distinguishable from the other species of Agerinia. The upper teeth also show clear differences with Agerinia marandati. The morphology of the calcaneal remains reveals that A. smithorum practiced a moderately active arboreal quadrupedal mode of locomotion, showing less leaping proclivity than notharctines but more than asiadapids. All the morphological features observed in the described material reinforce the hypothesis of a single lineage consisting of the species A. smithorum, A. marandati, and Agerinia roselli. Furthermore, the phylogenetic analysis developed in this work, which incorporates the newly described remains of A. smithorum, maintains the position of Agerinia as closely related to sivaladapids and asiadapids.

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BACKGROUND: The Eocene was the warmest epoch of the Cenozoic and recorded the appearance of several orders of modern mammals, including the first occurrence of Euprimates. During the Eocene, Euprimates were mainly represented by two groups, adapiforms and omomyiforms, which reached great abundance and diversity in the Northern Hemisphere. Despite this relative abundance, the record of early Eocene primates from the European continent is still scarce and poorly known, preventing the observation of clear morphological trends in the evolution of the group and the establishment of phylogenetic relationships among different lineages. However, knowledge about the early Eocene primates from the Iberian Peninsula has been recently increased through the description of new material of the genus Agerinia from several fossil sites from Northeastern Spain. METHODS: Here we present the first detailed study of the euprimate material from the locality of Masia de l'Hereuet (early Eocene, NE Spain). The described remains consist of one fragment of mandible and 15 isolated teeth. This work provides detailed descriptions, accurate measurements, high-resolution figures and thorough comparisons with other species of Agerinia as well with other Eurasian notharctids. Furthermore, the position of the different species of Agerinia has been tested with two phylogenetic analyses. RESULTS: The new material from Masia de l'Hereuet shows several traits that were previously unknown for the genus Agerinia, such as the morphology of the upper and lower fourth deciduous premolars and the P2, and the unfused mandible. Moreover, this material clearly differs from the other described species of Agerinia, A. roselli and A. smithorum, thus allowing the erection of the new species Agerinia marandati. The phylogenetic analyses place the three species of Agerinia in a single clade, in which A. smithorum is the most primitive species of this genus. DISCUSSION: The morphology of the upper molars reinforces the distinction of Agerinia from other notharctids like Periconodon. The analysis of the three described species of the genus, A. smithorum, A. marandati and A. roselli, reveals a progressive change in several morphological traits such as the number of roots and the position of the P1 and P2, the molarization of the P4, the reduction of the paraconid on the lower molars and the displacement of the mental foramina. These gradual modifications allow for the interpretation that these three species, described from the early Eocene of the Iberian Peninsula, are part of a single evolutionary lineage. The stratigraphical position of Masia de l'Hereuet and Casa Retjo-1 (type locality of A. smithorum) and the phylogenetic analyses developed in this work support this hypothesis.

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The study of Eocene primates is crucial for understanding the evolutionary steps undergone by the earliest members of our lineage and the relationships between extinct and extant taxa. Recently, the description of new material from Spain has improved knowledge of European Paleogene primates considerably, particularly regarding microchoerines. Here we describe the remains of Microchoerus from Sossís (late Eocene, Northern Spain), consisting of more than 120 specimens and representing the richest sample of Microchoerus from Spain. This primate was first documented in Sossís during the 1960s, on the basis of scarce specimens that were ascribed to Microchoerus erinaceus. However, the studied material clearly differs from M. erinaceus at its type locality, Hordle Cliff, and shows some characters that allow the erection of a new species, Microchoerus hookeri. This new species is characterized by its medium size, moderate enamel wrinkling, generally absent mesoconid and small hypoconulid in the M1 and M2, single paracone in the upper molars and premolars and, particularly, by the lack of mesostyle in most M1 and M2, a trait not observed in any other species of Microchoerus. Some specimens from Eclépens B (late Eocene, Switzerland), determined previously to be Microcherus aff. erinaceus, are also ascribed to M. hookeri. M. hookeri represents the first step of a lineage that differentiated from Necrolemur antiquus and, later, gave rise to several unnamed forms of Microchoerus, such as those from Euzet and Perrière, finally leading to M. erinaceus. This discovery sheds new light on the complex evolutionary scheme of Microchoerus, indicating that it is most probably a paraphyletic group. A detailed revision of the age of the localities containing remains of Microchoerus and the description of the still unpublished material from some European localities, are necessary to clarify the phylogenetic relationships among the members of this microchoerine group.

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The new species Agerinia smithorum (Adapiformes, Primates) from the early Eocene of the Iberian Peninsula is erected in this work. An emended diagnosis of the genus is provided, together with a broad description of the new species and comparisons with other samples assigned to Agerinia and other similar medium-sized cercamoniines. The new species is based on the most complete specimen of this genus published to date, a mandible preserving the alveoli of the canine and P1 , the roots of the P2 and all teeth from P3 to M3 . It was found in Casa Retjo-1, a new early Eocene locality from Northeastern Spain. The studied specimen is clearly distinguishable from other cercamoniines such as Periconodon, Darwinius, and Donrussellia, but very similar to Agerinia roselli, especially in the similar height of P3 and P4 and the general morphology of the molars, therefore allowing the allocation to the same genus. However, it is undoubtedly distinct from A. roselli, having a less molarized P4 and showing a larger paraconid in the M1 and a tiny one in the M2 , among other differences. The body mass of A. smithorum has also been estimated, ranging from 652 to 724 g, similar to that of A. roselli. The primitive traits shown by A. smithorum (moderately molarized P4 , large paraconid in the M1 and small but distinct in the M2 ) suggest that it could be the ancestor of A. roselli.

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The species Pseudoloris parvulus, identified in several Middle and Late Eocene European sites, was previously known in the Iberian Peninsula by a single mandible preserving P4-M3 from Sossís (Southern Pyrenean Basins, northeastern Spain), described in the 1960s. Further field work at this Late Eocene site has led to the recovery of a large number of mammal remains, including the additional material of P. parvulus described in this paper. Some specimens of P. parvulus from this locality have also been recently found in the collections of the Naturhistorisches Museum Basel, Switzerland. The whole sample consists of 11 mandible fragments including several teeth, three upper dental series and nearly 80 isolated teeth including all of the dental elements, and represents the most complete sample of the genus described from the Iberian Peninsula. This abundant material allows us to provide an emended diagnosis for the species and to observe several directional changes in the dental morphology of the lineage including the species Pseudoloris saalae, Pseudoloris isabenae, Pseudoloris pyrenaicus and P. parvulus. These directional changes include the progressive reduction of the paraconid in the lower molars and the increase in size of the hypocone, metaconule and paraconule in the upper molars. Moreover, despite the overall resemblance among all of the samples ascribed to P. parvulus, we also recognize some differences, particularly an increase in size and better development of the hypocone from the oldest populations of the species, such as Le Bretou, to the most recent ones, like Sossís and Perrière. Therefore, this study sheds new light on the evolution of this genus, which inhabited Europe from the Middle Eocene to the Early Oligocene.

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