Evolution Of Alternative Reproductive Tactics Research Articles

An eco-evolutionary feedback loop between population dynamics and fighter expression affects the evolution of alternative reproductive tactics.

Surprisingly, little is known about how eco‐evolutionary feedback loops affect trait dynamics within a single population. Polymorphisms of discrete alternative phenotypes present ideal test beds to investigate this, as the alternative phenotypes typically exhibit contrasting demographic rates mediated through frequency or density dependence, and are thus differentially affected by selection.Alternative reproductive tactics (ARTs), like male fighters and sneakers, are an extreme form of discrete phenotype expression and occur across many taxa. Fighters possess weapons for male–male competition over access to mates, whereas sneakers are defenceless but adopt tactics like female‐mimicking. Because fighters in some species mortally injure conspecifics, this raises the question whether fighter expression can feed back to affect population size and structure, thereby altering the selection gradient and evolutionary dynamics of ART expression in an eco‐evolutionary feedback loop.Here, we investigated how the eco‐evolutionary feedback loop between fighter expression and population size and structure affects the evolution and maintenance of ARTs. We introduced intraspecific killing by fighters in a two‐sex, two‐ART population model parameterized for the male dimorphic bulb mite (Rhizoglyphus robini) that includes life‐history differences between the ARTs and a mating‐probability matrix analogous to the classic hawk–dove game.Using adaptive dynamics, we found that the intraspecific killing by fighters can extend the range of life‐history parameter values under which ARTs evolve, because fighters that kill other fighters decrease fighter fitness. This effect can be nullified when benefits from killing are incorporated, like increased reproduction through increased energy uptake.The eco‐evolutionary feedback effects found here for a dimorphic trait likely also occur in other fitness‐related traits, such as behavioural syndromes, parental care and niche construction traits. Current theoretical advances to model eco‐evolutionary processes, and empirical steps towards unravelling the underlying drivers, pave the way for understanding how selection affects trait evolution in an eco‐evolutionary feedback loop.

“Failure to launch”: is there a reproductive cost to males living at home?

Differential reproductive success commonly reflects variations in reproductive physiology, behavior, and morphology. In some species, competition among males results in the evolution of alternative reproductive tactics that confer a fitness advantage relating to social status, density, or myriad other factors. In the Cape ground squirrel (Xerus inauris), a species that is highly competitive but lacks typical mammalian aspects of intrasexual competition (e.g., territoriality and aggression), 2 alternative reproductive tactics occur relating to dispersal. While physiological and behavioral differences have been demonstrated between dispersed males and males that delay dispersal, we used microsatellite markers to quantify variations in reproductive success between tactics. We found that dispersed males are in better body condition with larger home ranges likely allowing greater encounter frequencies with estrous females. However, we found no difference in copulation frequency between tactics, and the decisio...

Behavioral and Morphological Traits Interact to Promote the Evolution of Alternative Reproductive Tactics in a Lizard

Alternative reproductive tactics (ARTs) are predicted to be the result of disruptive correlational selection on suites of morphological, physiological, and behavioral traits. ARTs are most obvious when they occur in discrete morphs with concomitant behavioral tactics. However, ARTs driven by behavior in species lacking obvious phenotypic differences are rarely documented and poorly understood. We quantified selection acting on phenotypic traits predicted to characterize ARTs by observing marked lizards in six seminatural populations. We quantified reproductive fitness for each male using six microsatellite DNA loci from 226 offspring born to 56 females. Candidate models containing directional and correlational selection gradients were equally supported. As predicted, large males with large home ranges and large males that were observed frequently had the highest reproductive success. We also found evidence that large males that moved little but that were observed frequently and large males that moved frequently but that were observed little were predicted to have high fitness. Model predictions support our verbal hypothesis regarding the phenotypes characterizing ARTs and suggest that large males may be adopting subtly different tactics to acquire paternity. Our results suggest that disruptive correlational selection between behavioral traits may drive the evolution of ARTs in "cryptic" systems that lack overt polymorphisms.

Alternative reproductive tactics in male Cape ground squirrels Xerus inauris

In some animal societies, males vary in the strategies and tactics that they use for reproduction. Explanations for the evolution of alternative tactics have usually focussed on extrinsic factors such as social status, the environment or population density and have rarely examined proximate differences between individuals. Anecdotal evidence suggests that two alternative reproductive tactics occur in cooperatively breeding male Cape ground squirrels. Here we show that there is strong empirical support for physiological and behavioural differences to uphold this claim. ‘Dispersed’ males have higher resting metabolic rates and a heightened pituitary activity, compared with philopatric ‘natal’ males that have higher circulating cortisol levels. Dispersed males also spend more time moving and less time feeding than natal males. Additionally, lone males spend a greater proportion of their time vigilant and less of their time foraging than those that were in groups. The choice of whether to stay natal or become a disperser may depend on a number of factors such as age, natal group kin structure and reproductive suppression, and the likelihood of successful reproduction whilst remaining natal. Measuring proximate factors, such as behavioural and endocrine function, may provide valuable insights into mechanisms that underlie the evolution of alternative reproductive tactics.

BREEDING COMPETITION IN A PACIFIC SALMON (COHO: ONCORHYNCHUS KISUTCH): MEASURES OF NATURAL AND SEXUAL SELECTION.

In the breeding system of Pacific salmon, females compete for oviposition territories, and males compete to fertilize eggs. The natural selection in females and sexual selection in males likely has been responsible for their elaborate breeding morphologies and the dimorphism between the sexes. We quantified direct-selection intensities during breeding on mature coho salmon (Oncorhynchus kisutch), measured for seven phenotypic characters, including three secondary sexual characters. Wild and sea-ranched hatchery coho were used to enhance the range of phenotypes over which selection could be examined. The fish were allowed to breed in experimental arenas where we could quantify components of breeding success as well as estimate overall breeding success. We found that without competition, natural selection acts only on female body size for increased egg production; there is no detectable selection on males for the phenotypic distribution we used. Under competition, the opportunity for selection increased sixfold among females. Natural selection favored female body size and caudal-peduncle (tail) depth. Increased body size meant increased egg production and access to nesting territories. The caudal peduncle, used in burst swimming and nest digging, influenced both successful egg deposition and nest survival. Increasing density increased competition among females, though it did not significantly intensify natural selection on their characters. In males, competition increased the opportunity for selection 52-fold, which was nine times greater than for females. Sexual selection favored male body size and hooked snout length, both characters directly influencing male access to spawning opportunities. Selection on male body size was also affected significantly by breeding density. The ability of large males to control access to spawning females decreased at higher densities reflecting an increase in the operational sex ratio. Further, the relative success of small males, which could sneak access to spawning females, appeared to increase as that of intermediate-sized males decreased. Such disruptive selection may be responsible for the evolution of alternative reproductive tactics in salmon.

Inter- and Intrasexual Dimorphisms in the Vocal Control System of a Teleost Fish: Motor Axon Number and Size

In one species of vocalizing fish, the plainfin midshipman (Porichthys notatus), large, nest-guarding males ('type I') use striated muscles to produce acoustic communication signals that include short duration (less than 1 s) 'burps' important in agonistic encounters and long duration (in the order of minutes) 'hums' which function in attracting females to nest sites during the breeding season. Females, and a second group of smaller reproductively active males ('type II') that 'sneak' spawn, do not generate hums, although they produce burps. Differences in vocal behavior are paralled by a relative increase of 6-fold in the sonic muscle of type I males. Inter- and intrasexual dimorphisms in sonic muscle mass were matched by those in the cross-sectional area of sonic motor axons, but not by those in number of axons. Thus, axon size was 2- to 3-fold larger in type I males than in females, type II males, or juveniles, none of which differed significantly from each other. Axon number was similar between type I males and females of a similar body size, despite the extreme dimorphism in sonic muscle mass. Axon number, however, was slightly greater (0.1-fold) in type I males and females compared to the smaller-sized juveniles and type II males. Type II males, in comparison to the non-reproductive juvenile males, have gonads that are about 20-fold larger and produce mature sperm. Nevertheless, the two groups resemble each other in body and swimbladder size, as well as sonic motor axon size and number. This suggests that type II males represent a subset of juvenile males that undergo precocious gonadal hypertrophy and spermiogenesis, but retain juvenile-like nongonadal traits. The results are discussed within the context of the development of vertebrate motor systems as well as the evolution of alternative reproductive tactics among teleost fishes.