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

Morgan L Guignard,Marina B Soares,Agustin G Martinelli,Leon Claessens

doi:10.1371/journal.pone.0216672

Abstract

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.

Highlights

Much attention has been paid to the craniomandibular changes in the non-mammaliaform cynodont to mammaliaform transition (e.g., [1,2,3,4,5,6,7,8])
Posterolaterally facing humeral head, lesser tubercle larger than the greater one, Postcranial anatomy of Brasilodon quadrangularis long and anteriorly expanded deltopectoral crest, wide bicipital groove, teres major tuberosity, wide distal humeral extremity, shallow olecranon fossa and bulbous ulnar condyle are features seen in other non-mammaliaform cynodonts
The postcranium of B. quadrangularis differs from other non-mammaliaform cynodonts and is similar to early mammaliaforms and extant therians

Summary

Introduction

Much attention has been paid to the craniomandibular changes in the non-mammaliaform cynodont to mammaliaform transition (e.g., [1,2,3,4,5,6,7,8]). A general view is that the postcranial skeleton of non-mammaliaform cynodonts is quite conservative compared to craniodental adaptations, having an “intermediate bauplan” between basal therapsids and early mammaliaforms [11]. The postcranial anatomy in Triassic cynodonts permits to evaluate the main changes and locomotor advances in mammalian evolution and provide information about the transition from the sprawling posture of basal synapsids to a more erect posture. We present a detailed description of the postcranial elements of Brasilodon quadrangularis, including previously mentioned specimens and new ones. It adds more information about its postcranial morphology and provides new insights into the anatomical transition between the advanced non-mammaliaform cynodonts and early mammaliaforms. We investigate functional implications of its postcranium to improve our knowledge on posture and locomotion of mammalian precursors

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