In polyandrous species, a male’s reproductive success depends on his ability to fertilize females, which, in turn, depends on his mating ability and his ability to produce competitive ejaculates. In many species, sperm traits differ between old and young males in ways that are likely to decrease the sperm competitiveness and fertility of older males. This age–ejaculate quality relationship is attributed to male ageing (i.e., senescence). In a natural setting, male age and mating history are usually confounded: older males have usually mated and replenished their sperm supplies more often, so they have made a greater lifetime reproductive effort. In principle, the costs of reproduction, independent of any causal effect of male age, could generate an age‐related decline in ejaculate quality. To date, only a handful of studies have determined how male age, reproductive effort or their interaction affect ejaculate quality. Here, we experimentally manipulated the long‐term mating history of 209 adult male mosquitofish (Gambusia holbrooki) over 14 weeks (N = 1,118 sperm samples). Males either had direct access to females and could mate freely, or had only visual and olfactory access to females. We documented the effect of mating history, adult male age (3, 9 and 14 weeks post‐maturation) and their interaction on sperm velocity, sperm reserves and the rate of sperm replenishment. For sperm velocity, we additionally examined the effects of sperm age, because when older males mate less (or more) often than younger males there will be a correlation between mean sperm age and male age. Sexually active males produced fewer sperm and replenished their sperm at a lower rate, and their sperm had lower velocity than males prevented from mating. Though older males produced more sperm, the rate of replenishment and velocity of their sperm was lower than the sperm of younger males. We also tested for a difference in the velocity of recently replenished (<24 hr) and older sperm (i.e., post‐meiotic sperm senescence). There was no evidence that male age or mating history affects the extent of sperm senescence, but older sperm swam faster than recently produced sperm. Crucially, complex interactions are evident between male age and male mating history with respect to sperm number and the proportion of sperm that are replenished. These results suggest that male age and mating history will interact to determine the reproductive success of a male under sperm competition. They reveal a complex relationship between a male’s age and his ejaculate quality. We suggest that both mating history and sperm age should be controlled for when measuring the intrinsic rate of senescence for male reproductive traits if the goal is to isolate effects that are solely attributable to a male’s chronological age. A plain language summary is available for this article.
As cities continue to grow it is increasingly important to understand the long-term responses of wildlife to urban environments. There have been increased efforts to determine whether urbanization imposes chronic stress on wild animals, but empirical evidence is mixed. Here, we conduct a meta-analysis to test whether there is, on average, a detrimental effect of urbanization based on baseline and stress-induced glucocorticoid levels of wild vertebrates. We found no effect of urbanization on glucocorticoid levels, and none of sex, season, life stage, taxon, size of the city nor methodology accounted for variation in the observed effect sizes. At face value, our results suggest that urban areas are no more stressful for wildlife than rural or non-urban areas, but we offer a few reasons why this conclusion could be premature. We propose that refining methods of data collection will improve our understanding of how urbanization affects the health and survival of wildlife.
BackgroundThe optimal allocation of resources to sexual signals and other life history traits is usually dependent on an individual’s condition, while variation in the expression of sexual traits across environments depends on the combined effects of local adaptation, mean condition, and phenotypic responses to environment-specific cues that affect resource allocation. A clear contrast can often be drawn between natural habitats and novel habitats, such as forest plantations and urban areas. In some species, males seem to change their sexual signals in these novel environments, but why this occurs and how it affects signal reliability is still poorly understood.ResultsThe relative size of sexual traits and level of immune responses were significantly lower for male palmate newts Lissotriton helveticus caught in pine and eucalyptus plantations compared to those caught in native forests, but there was no habitat-dependent difference in body condition (n = 18 sites, 382 males). The reliability with which sexual traits signalled body condition and immune responses was the same in all three habitats. Finally, we conducted a mesocosm experiment in which males were maintained in pine, eucalypt or oak infused water for 21 days. Males in plantation-like water (pine or eucalypt) showed significantly lower immune responses but no change in body condition. This matches the pattern seen for field-caught males. Unlike field-caught males, however, there was no relationship between water type and relative sexual trait size.ConclusionsPine and eucalyptus plantations are likely to be detrimental to male palmate newt because they are associated with reduced immune function and smaller sexual traits. This could be because ecological aspects of these novel habitats, such as high water turbidity or changes in male-male competition, drive selection for reduced investment into sexual traits. However, it is more probable that there are differences in the ease of acquisition, hence optimal allocation, of resources among habitats. Our mesocosm experiment also provides some evidence that water toxicity is a causal factor. Our findings offer insights into how plantations affect amphibian life histories, and how novel habitats might generate long-term selection for new resource allocation strategies in native species.Electronic supplementary materialThe online version of this article (doi:10.1186/s12862-016-0706-0) contains supplementary material, which is available to authorized users.
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