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Theory predicts the evolution of alternative male social tactics when intense competition coupled with the superior competitive ability of some individuals limits access to reproductive opportunities by others. How selection has shaped alternative social tactics may be especially interesting in long-lived species where size among sexually mature males varies markedly. We conducted experimental studies on long-lived eastern Australian water dragons living where competition was intense to test the hypotheses that mature males adopt alternative social tactics that are plastic, and that large size and body condition determine resource-holding potential. Approximately one-half of mature males (N = 14) defended territories using high rates of patrol and advertisement display, whereas 16 smaller mature males having lower body condition indices utilized non-territorial social tactics. Although territorial males were larger in absolute size and head dimensions, their heads were not allometrically larger. Territorial males advertised very frequently using displays involving stereotypical movements of the head and dewlap. More aggressive displays were given infrequently during baseline social conditions, but increased during periods of social instability. Female home ranges overlapped those of several territorial and non-territorial males, but females interacted more frequently with territorial males. The extreme plasticity of social tactics in this species that are dependent on body size was confirmed by two instances when relatively large non-territorial males spontaneously evicted territory owners, and by marked shifts in tactics by non-territorial males in response to temporary experimental removals of territory owners, followed (usually) by their expulsion when original owners were reinstated. The high level of social plasticity in this population where same-sex competitors are densely concentrated in preferred habitat suggests that chronic high energetic costs of defense may select for males to cycle between territorial and non-territorial social tactics depending upon their changing energetic status and their current capacity for competition with rivals.

2001: Intra-and inter-seasonal variation in the socio-spatial behavior of adult male collared lizards, Crotaphytus collaris (Reptilia, Crotaphytidae). Ethology 107, 15Ð32. AbstractWhen individuals maintain strong inter-seasonal philopatry to the same territories, males may be able to re-establish territory occupancy without intense intrasexual aggression, and instead spend more time courting females early in the reproductive season. Furthermore, when some males have prior experience defending the same territories, it may be necessary for young males to exhibit higher levels of aggression because they are establishing a territory for the ®rst time. We tested these hypotheses by examining within-season (1992 and 1997) temporal variation in the social behavior of adult male collared lizards of known age and prior territorial experience in a population where inter-season philopatry to territories is high. Contrary to expectations, the frequency of aggression exhibited by males with and without prior territorial experience did not dier. The frequency of intra-sexual aggression was higher in 1992 than in 1997, perhaps because male competitors were more abundant in 1992. Although there was an interactive eect of year, male display and patrol were low at the beginning of the reproductive season in Apr. and May, reached peaks during midseason in June, and then decreased as reproduction ended in July. The size of territories showed a similar pattern, with males defending larger areas in June. Our data support the philopatry hypothesis in that the establishment of territories occurred without high levels of aggression early in the season, perhaps because territory boundaries have been well de®ned by high rates of patrol and advertisement during the middle of the previous season. Inter-sexual interactions were most frequent in June rather than at the beginning of the reproductive season. Adult females are producing their second clutches and yearling females are producing their ®rst clutches in June. The high rate of inter-sexual encounters in June supports the hypothesis that males allocate more time to courtship when females are receptive because there are more reproductively active females at this time. The temporal pattern of activities in adult Crotaphytus collaris appears to function as a compromise between U. S. Copyright Clearance Center Code Statement: 0179-1613/2001/1071±0015$15.00/0 competing intra-and inter-sexual social demands on males, allowing males to maximize mating opportunities as well as maintain future access to productive territories.

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