Abstract
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.
Highlights
Male vertebrates under strong sexual selection sometimes evolve alternative reproductive tactics (ART) characterized by distinctive behavior patterns, and/or morphological differences that are associated with resource holding potential ( = RHP)
Male Social Behavior and Alternative Tactics During baseline social conditions, 98% of displays given by males were stereotypical movements of the head alone
Our observations of spontaneous evictions of territory owners, rapid territory takeovers in response to experimental removals, high baseline rates of advertisement display by territory owners, as well as prominent scars on large males all indicate that intrasexual male competition is intense in this high density population of water dragons, which likely makes the costs of territory maintenance very high
Summary
Male vertebrates under strong sexual selection sometimes evolve alternative reproductive tactics (ART) characterized by distinctive behavior patterns, and/or morphological differences that are associated with resource holding potential ( = RHP). Alternative male tactics may be either genetically fixed, or ontogenetically plastic. Males switch between/ among two or more alternative tactics [4,5]. The usual case is for males to adopt socially dominant behavioral tactics, often defense of reproductive territories, when they obtain high RHP. Males that are not able to acquire and defend territories may adopt subordinate tactics characterized by sneaking copulations when they can do so without being detected by socially-dominant males [16,17], or by mimicking females [18]. Especially in long-lived philopatric species, subordinate males may acquire territories by aggressive eviction of current territory owners [20,21]
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