New Leontiniid Notoungulata (Mammalia) from Chile and Argentina: Comparative Anatomy, Character Analysis, and Phylogenetic Hypotheses

Abstract

Herein we describe and name two new species of leontiniid notoungulates, one being the first known from Chile, the other from the Deseadan South American Land Mammal Age (SALMA) of Patagonia, Argentina. The Chilean leontiniid is from the lower horizons of the Cura-Mallín Formation (Tcm1) at Laguna del Laja in the Andean Main Range of central Chile. This new species, Colpodon antucoensis, is distinguishable from Patagonian species of Colpodon by way of its smaller I2; larger I3 and P1; sharper, V-shaped snout; and squarer upper premolars. The holotype came from a horizon that is constrained below and above by 40Ar/39Ar ages of 19.53 ± 0.60 and 19.25 ± 1.22, respectively, suggesting an age of roughly 19.5 Ma, or a little older (∼19.8 Ma) when corrected for a revised age of the Fish Canyon Tuff standard. Either age is slightly younger than ages reported for the Colhuehuapian SALMA fauna at the Gran Barranca. Taxa from the locality of the holotype of C. antucoensis are few, but they (e.g., the mylodontid sloth, Nematherium, and a lagostomine chinchillid) also suggest a post-Colhuehuapian faunal age. The second leontiniid named in this paper has been known in the literature for over 75 years as Leontinia sp. Several specimens referable to this species were discovered at Pico Truncado (Deseadan SALMA) during the Field Museums first Marshall Field Expedition, led by Elmer Riggs in 1924. This “new” taxon, Elmerriggsia fieldia, is a small-bodied leontiniid, possessing grooved premolar protocones that lack intermediate lingual cingulae, but have well-developed labial cingulids on their lower molars. This new taxon is fairly common at Pico Truncado, in Santa Cruz, Argentina, but we have not encountered it at other localities.The character-taxon matrix that we constructed for this analysis differs from those previously developed for notoungulates by the substantially greater number of postcranial characters used (41). Colbertia magellanica was used as the outgroup in all analyses. Our initial phylogenetic analysis was limited to only taxa traditionally assigned to the Toxodontia. These included a dozen taxa traditionally considered to be leontiniids, two toxodontids, four notohippids, a homalodotheriid, and two isotemnids. The taxa traditionally classified as leontiniids formed a monophyletic group, in which V-shaped muzzle, caniniform i3, femur with medial suprapatellar ridge, and large wedge-shaped fibular facet of the calcaneum were unequivocal synapomorphies. Colpodon spp. nested within a clade that includes the “tropical” leontiniids, Taubatherium and Huilatherium. Toxodontids and notohippids formed a monophyletic group sister to the leontiniids, with these two clades forming a more inclusive clade that previously had been called the “advanced Toxodontia.” However, when five species of typotheres from three “families” were added to the analysis, the “notohippid” Eurygenium was identified as the nearest outgroup of leontiniids and an “advanced notohippid” plus toxodontid clade (nodes C F). Unequivocal synapomorphies uniting these two nodes were robust calcanonavicular articulation (“reverse alternating tarsus” as evidenced by a distinct navicular facet on the calcaneum) and a distal radius with a styloid process. The presence of an entolophid fossettid in the lower molars and the downturned olecranon process of the ulna were equivocal synapomorphies for this clade. Though lacking the character states that diagnose a more exclusive “notohippid-toxodontid-leontiniid” clade, Eurygenium shared several unequivocal synapomorphies that unite it with these taxa. These include a well-formed fossette of upper molars formed by the posterior cingulum, absence of an entepicondylar foramen of the humerus, lack of a neck on the astragalus, a transversely elongated astragalar head, and absence of the “astragalar buttress” of the navicular.Unconventionally, the interatheriids used in the analysis (Federicoanaya and Protypotherium, both interatheriin