FIGHTING BEHAVIOR OF MUNTJAC AND THE EVOLUTION OF ANTLERS.

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Phylogenetically and morphologically, the Miocene Muntiacinae, represented today by the Asian Muntiacus and Elaphodus fit between the Oligocene tusk bearer antlerless cervids and tragulids, and the Pliocene and living tuskless antlered deer (Romer, 1966). In addition to being intermediate in size and phylogenetic age, the muntjacs are among the first Cervidae to have acquired antlers and still have preserved upper canine tusks (Fig. 1). Antlers and tusks are two different kinds of weapon. Tusks are a strictly offensive weapon (Geist, 1966a). Antlers however serve two kinds of functions. First, they serve as visual social rank symbols (Geist, 1966a; Henshaw, 1971; Lincoln, 1972) and second, in combat, they have the dual function of a weapon and a shield (Geist, 1966a). Muntjacs' antlers have an additional role: they are a necessary instrument permitting the tusks to be put to use in intra-specific combats. The available literature on the fighting behavior of muntjacs is rare and imprecise (Aitchison, 1946; Antonius, 1939; Dubost, 1971; Scherpe, 1971). Until recently, the same was true for other tusk bearers, but antlerless cervids and tragulids (Aitchison, 1946; Dubost, 1965; Eibl-Eibesfeldt, 1957; Fradrich, 1966; Kirchshofer, 1961; Scherpe, 1971). Recently, Dubost (1975) and Ralls et al. (1975) have described the fighting behavior of male Hyemoschus aquaticus and Tragulus napu respectively. From the shape of tusks, the location of wounds on adult males, and direct observa-