Sperm wars and the evolution of male fertility

Females frequently mate with several males, whose sperm then compete to fertilize available ova. Sperm competition represents a potent selective force that is expected to shape male expenditure on the ejaculate. Here, we review empirical data that illustrate the evolutionary consequences of sperm competition. Sperm competition favors the evolution of increased testes size and sperm production. In some species, males appear capable of adjusting the number of sperm ejaculated, depending on the perceived levels of sperm competition. Selection is also expected to act on sperm form and function, although the evidence for this remains equivocal. Comparative studies suggest that sperm length and swimming speed may increase in response to selection from sperm competition. However, the mechanisms driving this pattern remain unclear. Evidence that sperm length influences sperm swimming speed is mixed and fertilization trials performed across a broad range of species demonstrate inconsistent relationships between sperm form and function. This ambiguity may in part reflect the important role that seminal fluid proteins (sfps) play in affecting sperm function. There is good evidence that sfps are subject to selection from sperm competition, and recent work is pointing to an ability of males to adjust their seminal fluid chemistry in response to sperm competition from rival males. We argue that future research must consider sperm and seminal fluid components of the ejaculate as a functional unity. Research at the genomic level will identify the genes that ultimately control male fertility.</jats:p

Rival Male Relatedness Does Not Affect Ejaculate Allocation as Predicted by Sperm Competition Theory

When females are sexually promiscuous, the intensity of sperm competition for males depends on how many partners females mate with. To maximize fitness, males should adjust their copulatory investment in relation to this intensity. However, fitness costs associated with sperm competition may not only depend on how many males a female has mated with, but also how related rival males are. According to theoretical predictions, males should adjust their copulatory investment in response to the relatedness of their male rival, and transfer more sperm to females that have first mated with a non-sibling male than females that have mated to a related male. Here, for the first time, we empirically test this theory using the Australian field cricket Teleogryllus oceanicus. We expose male crickets to sperm competition from either a full sibling or non-sibling male, by using both the presence of a rival male and the rival male's actual competing ejaculate as cues. Contrary to predictions, we find that males do not adjust ejaculates in response to the relatedness of their male rival. Instead, males with both full-sibling and non-sibling rivals allocate sperm of similar quality to females. This lack of kin biased behaviour is independent of any potentially confounding effect of strong competition between close relatives; kin biased behaviour was absent irrespective of whether males were raised in full sibling or mixed relatedness groups

Nuptial gifts fail to resolve a sexual conflict in an insect

Background Because of the potential benefits to individuals of saving investment for future mating opportunities, there is conflict between mates over most aspects of reproduction. Males of many species transfer compounds in the ejaculate that manipulate female reproductive physiology to increase male reproductive success. These seminal compounds are often associated with direct and/or indirect costs to females. In contrast, in some species ejaculates also contain nutrients used by females for somatic maintenance and increased reproductive output. In general, the extent to which male seminal components are detrimental or beneficial to females is poorly understood, and interactions between seminal compounds with different effects have been almost completely neglected. Here we examine the impact of male receptivity-suppressing factors and nutrient donations on female longevity and lifetime reproductive output in the bushcricket Requena verticalis. Results We show that receiving multiple ejaculates reduces longevity in female R. verticalis, indicating a cost of male derived receptivity-suppressing compounds. Consumption of male nutrient donations does not appear to ameliorate this longevity cost, and there was no effect of nutrient provisioning on female lifetime fecundity. Conclusion These results indicate that nutrient provisioning does not provide a resolution to sexual conflict over female receptivity in this bushcricket species

Evolution of Sexual Dimorphism and Male Dimorphism in the Expression of Beetle Horns: Phylogenetic Evidence for Modularity, Evolutionary Lability, and Constraint

Beetle horns are enlarged outgrowths of the head or thorax that are used as weapons in contests over access to mates. Horn development is typically confined to males (sexual dimorphism) and often only to the largest males (male dimorphism). Both types of dimorphism result from endocrine threshold mechanisms that coordinate cell proliferation near the end of the larval period. Here, we map the presence/absence of each type of dimorphism onto a recent phylogeny for the genus Onthophagits (Coleoptera: Scarabaeidae) to explore how horn development has changed over time. Our results provide empirical support for several recent predictions regarding the evolutionary lability of developmental thresholds, including uncoupled evolution of alternative phenotypes and repeated fixation of phenotypes. We also report striking evidence of a possible developmental constraint. We show that male dimorphism and sexual dimorphism map together on the phylogeny; whenever small males have horns, females also have horns (and vice versa). We raise the possibility that correlated evolution of these two phenomena results from a shared element in their endocrine regulatory mechanisms rather than a history of common selection pressures. These results illustrate the type of insight that can be gained only from the integration of developmental and evolutionary perspectives

Threshold Evolution in Exotic Populations of a Polyphenic Beetle

Polyphenic development is thought to play an important role in the evolution of phenotypic diversity and morphological novelties, yet the evolution of polyphenisms has rarely been documented in natural populations. Here we compare the morphologies of male dung beetles (Onthophagus taurus; Coleoptera: Scarabaeidae) from populations introduced to Australia and the eastern United States. Males in this species express two alternative morphologies in response to larval feeding conditions. Males encountering favourable conditions grow larger than a threshold body size and develop a pair of horns on their heads, whereas males that encounter poor conditions do not reach this threshold size and remain hornless. Australian and US populations did not differ in overall body size ranges, but exhibited significant differences in the location of the critical body size threshold that separates alternative male morphs. Australian males remained hornless at much larger body sizes than males in US populations, resulting in substantial and significant differences in the average body size-horn length allometry between exotic populations, as well as significant differences in morph ratios. The phenotypic divergence observed between field populations was maintained in laboratory populations after two generations under identical environmental conditions, suggesting a genetic basis to allometric divergence in these populations. Divergence between exotic O. taurus populations was of a magnitude and kind typically observed between species. We use our results to examine potential causes of allometric divergence in onthophagine beetles, and discuss the evolutionary potential of threshold traits and polyphenic development in the origin of morphological and behavioural diversity

X-ray micro-CT scanning reveals temporal separation of male harm and female kicking during traumatic mating in seed beetles

In the seed beetle Callosobruchus maculatus, the male intromittent organ is covered in sharp spines that pierce the female copulatory tract wall during mating. Although the fitness consequences of traumatic mating are well studied in this species, we know much less about how the male and female genitalia interact during mating. This is partly due to the fact that genital interactions occur primarily inside the female, and so are difficult to observe. In this study, we use X-ray micro-CT scanning to examine the proximate mechanisms of traumatic mating in C. maculatus in unprecedented detail. We show that this technique can be used to identify female tissue damage before the melanization of wound sites. We visualize the positioning of the male intromittent organ inside the female copulatory tract during mating, and show how this relates to tract wounding in three dimensions. By scanning pairs flash-frozen at different times during mating, we show that significant tract wounding occurs before the onset of female kicking. There is thus some degree of temporal separation between the onset of wounding and the onset of kicking, which supports recent suggestions that kicking is not an effective female counter-adaptation to reduce copulatory wounding in this species. We also present evidence that the sharp teeth protruding from the female tract wall are able to pierce the spermatophore as it is deposited, and may thus function to aid sperm release

Future Distribution of Suitable Habitat for Pelagic Sharks in Australia Under Climate Change Models.

Global oceans are absorbing over 90% of the heat trapped in our atmosphere due to accumulated anthropogenic greenhouse gases, resulting in increasing ocean temperatures. Such changes may influence marine ectotherms, such as sharks, as their body temperature concurrently increases toward their upper thermal limits. Sharks are high trophic level predators that play a key role in the regulation of ecosystem structure and health. Because many sharks are already threatened, it is especially important to understand the impact of climate change on these species. We used shark occurrence records collected by commercial fisheries within the Australian continental Exclusive Economic Zone (EEZ) to predict changes in future (2050-2099) relative to current (1956-2005) habitat suitability for pelagic sharks based on an ensemble of climate models and emission scenarios. Our predictive models indicate that future sea temperatures are likely to shift the location of suitable shark habitat within the Australian EEZ. On average, suitable habitat is predicted to decrease within the EEZ for requiem and increase for mackerel sharks, however, the direction and severity of change was highly influenced by the choice of climate model. Our results indicate the need to consider climate change scenarios as part of future shark management and suggest that more broad -scale studies are needed for these pelagic species

Shark conservation hindered by lack of habitat protection

Many of the world's shark populations are in decline, indicating the need for improved conservation and management. Well managed and appropriately located marine parks and marine protected areas (MPAs) have potential to enhance shark conservation by restricting fisheries and protecting suitable habitat for threatened shark populations. Here, we used shark occurrence records collected by commercial fisheries to determine suitable habitat for pelagic sharks within the Australian continental Exclusive Economic Zone (EEZ), and to quantify the amount of suitable habitat contained within existing MPAs. We developed generalised linear models using proportional occurrences of pelagic sharks for three families: Alopiidae (thresher), Carcharhinidae (requiem), and Lamnidae (mackerel) sharks. We also considered aggregated species from the Lamnidae and Carcharhinidae families ('combined sharks' in the models). Using a set of environmental predictors known to affect shark occurrence, including chlorophyll-a concentration, salinity, sea surface temperature, and turbidity, as well as geomorphological, geophysical, and sedimentary parameters, we found that models including sea surface temperature and turbidity were ranked highest in their ability to predict shark distributions. We used these results to predict geographic regions where habitat was most suitable for pelagic sharks within the Australian EEZ, and our results revealed that suitable habitat was limited in no-take zones within MPAs. For all shark groupings, suitable habitats were found mostly at locations exposed to fishing pressure, potentially increasing the vulnerability of the pelagic shark species considered. Our predictive models provide a foundation for future spatial planning and shark management, suggesting that strong fisheries management in addition to MPAs is necessary for pelagic shark conservation