The antler-like appendages of the primitive deer Dicrocerus elegans: morphology, growth cycle, ontogeny, and sexual dimorphism

Males and many females of the primitive deer Dicrocerus elegans from Sansan (Middle Miocene, France) bore antler-like appendages consisting of a simple-branched protoantler growing from a rather long pedicle and are decorated with ridges and furrows. The protoantler capacity to be rejected and subsequently re-grow is clearly evidenced by the presence of both pedicle and protoantler cast specimens. The youngest appendage is a long, laterally flattened shaft whose apex is usually forked with no appreciable limit between the pedicle and the protoantler. In females, the anterior and posterior appendage margins form a more acute angle than that of males, and are more parallel when viewed laterally. After the first casting, the protoantler base is larger than the pedicle top and a coronet-like structure appears developed only around the medial side. With successive castings, the pedicles become shorter and their section is more circular, while protoantlers become much bigger, and have much longer and more separated branches. Branches of females are shorter than those of males, especially the anterior one, and appear in a straight line, instead of being bent. In oldest appendages, the branches are shorter and more similar in size. Accessory branches and irregularities of this basic morphology are common. The separation between both sex morphotypes appears clearly evidenced by Discriminant and Principal Component Analyses. Histological features point to important differences with true antlers and suggest that casting could not occur annually. A core of spongy bone trabeculae is not developed. Once growth is completed, the mineralization progress from the core to the periphery and when the final ‘velvet’ protoantler becomes completely petrified, the tissues dies and the velvet-like skin is cleaned. A high degree of both wear and polish of the branch apices evidence the hard, bare, dead protoantler phase before casting. Due to the complete growth cycle and the presence of the coronet-like structure, Dicrocerus protoantlers and antlers seem to be homologous appendages. Histological differences could be related to differences in hormonal cycle regulation that can be caused by the fact that i) Dicrocerus inhabited a tropical environment, and ii) females also developed protoantlers. It should not be overlooked that true antlers appear several million years later in time than the development of protoantlers and other cranial appendages in ruminants, and coinciding with the Middle Miocene Climatic Transition. Tanto los machos como muchas de las hembras del ciervo primit