New data on Propyrotherium (Mammalia, Pyrotheria) from the middle Eocene age (Chubut, Argentina): anatomy, age constraints, and phylogeny.

Pyrotheria is one of the most peculiar orders of South American native ungulates, whose members evolved from the early? Eocene to the late Oligocene period when they became extinct. Here, we described the most complete specimen of Propyrotherium saxeum ever found, one of the lesser-known representatives of pyrotheres that characterized the middle-late Eocene period of Patagonia (Argentina). It includes a nearly complete mandible and a tusk-like tooth of the same individual, as well as other isolated upper and lower teeth. Propyrotherium saxeum has a dental formula that includes at least P2-M3 and i2?-p3-m3 (lack of p2). It is characterized by some peculiar features of the mandible (e.g., ascending ramus longer than high, hiding the m3 and straight incisura mandibular) and dentition (e.g., cristid obliqua in p3-m3, cristid between posterior lophid and distal cingulid in m1-m3, P3-M3 and p3-m3 bilophodont, P2 and p3-m3 bi-rooted, P3-M3 three-rooted, paraconid in p3). The phylogenetic analysis reveals that Propyrotherium is more closely related to Pyrotherium and Baguatherium, differing from the previous hypothesis, and supports the monophyly of Pyrotheriidae including Carolozittelia, Griphodon, Pyrotherium, Baguatherium, and Propyrotherium. The absolute age obtained through U-Pb zircon dating of the Sarmiento Formation at Cañadón Pelado, the fossil's original locality, indicates that the fossil-bearing tuff would have been deposited between 39.65 and 40.41 Ma, with a weighted mean age of 40.03 ± 0.38 Ma (Bartonian). This implies a biochron much longer than previously thought for Propyrotherium and provides a chronological framework for the fauna of Cañadón Pelado.