The postcranial anatomy of Brasilodon quadrangularis and the acquisition of mammaliaform traits among non-mammaliaform cynodonts

Brasilodon quadrangularis (Cynodontia, Probainognathia) is an iconic non-mammaliaform cynodont from the Late Triassic of Brazil (Riograndia Assemblage Zone, Candelária Sequence), being considered as the sister taxon of Mammaliaformes. Although its phylogenetic position is very important, several aspects of its postcranial anatomy remain unclear or unstudied. Here, we present a detailed description of the postcranial elements referred to Brasilodon, including previously mentioned specimens and new ones, which add relevant information about its postcranial morphology and provide a new insight into the anatomical transition between advanced non-mammaliaform cynodonts and early mammaliaforms. Functional and ecological implications are also investigated, based on the postcranial morphology and muscular reconstructions. The postcranium of Brasilodon differs from most non-mammaliaform cynodonts and presents similarities with tritylodontids, early mammaliaforms and extant therians, such as a ventrally oriented scapular glenoid facet, a distinct and ossified greater humeral tubercle, lack of ectepicondylar foramen, olecranon process, hemispherical humeral and femoral heads and a prominent intertrochanteric crest. The humeral torsion, the length of the deltopectoral crest, the large bicipital groove and the well-developed lesser tubercle, indicate that the forelimb of Brasilodon was hold in a semi-sprawling position, with well-developed adductor muscles to maintain the body off the ground. The short femoral neck and the strong medial projection of the femoral head indicate the femur was held in a more erect posture than in basal non-mammaliaform cynodonts. The anterodorsally projected iliac blade with reduced postacetabular process, reduction of the anterior part of the pubis, medially located lesser trochanter indicate a basically mammalian pattern of pelvic musculature, able to swing the femur in a nearly parasagittal plane.