Dj. Emlen, ALTERNATIVE REPRODUCTIVE TACTICS AND MALE-DIMORPHISM IN THE HORNED BEETLE ONTHOPHAGUS-ACUMINATUS (COLEOPTERA, SCARABAEIDAE), Behavioral ecology and sociobiology, 41(5), 1997, pp. 335-341
Adult dung beetles (Onthophagus acuminatus) exhibit continuous variati
on in body size resulting from differential nutritional conditions exp
erienced during larval development. Males of this species have a pair
of horns that protrude from the base of the head, and the lengths of t
hese horns are bimodally distributed in natural populations. Males gro
wing larger than a threshold body size develop long horns, and males t
hat do not achieve this size grow only rudimentary horns or no horns a
t all. Previous studies of other horned beetle species have shown that
horned and hornless males often have different types of reproductive
behavior. Here I describe the mating behaviors of the two male morphs
of O. acuminatus during encounters with females, Females excavate tunn
els beneath dung, where they feed, mate and provision eggs. Large, hor
ned males were found to guard entrances to tunnels containing females.
These males fought with all other males that attempted to enter these
tunnels. In contrast, small, hornless males encountered females by sn
eaking into tunnels guarded by other males. In many instances, this wa
s accomplished by digging new tunnels that intercepted the guarded tun
nels below ground. Side-tunneling behavior allowed sneaking males to e
nter tunnels beneath the guarding male, and mate with females undetect
ed. Both overall body size and relative horn length significantly affe
cted the outcome of fights over tunnel ownership. These results sugges
t that alternative reproductive tactics may favor divergence in male h
orn morphology, with long horns favored in males large enough to guard
tunnels, and hornlessness favored in smaller males that adopt the ''s
neaking'' behavioral alternative.