When should a trophically transmitted parasite manipulate its host?

We investigate evolution of two categories of adaptive host manipulation by trophically transmitted helminths: (1) predation suppression decreases the host's mortality before the helminth is capable of establishing in its next host; (2) predation enhancement increases the existing host's mortality after it can establish in its next host. If all parasite mortality is purely random (time-independent), enhancement must increase predation by the next host sufficiently more (depending on manipulative costs) than it increases the average for all forms of host mortality; thus if host and parasite die only through random predation, manipulation must increase the "right" predation more than the "wrong" predation. But if almost all parasites die in their intermediate host through reaching the end of a fixed life span, enhancement can evolve if it increases the right predation, regardless of how much it attracts wrong predators. Although enhancement is always most favorable when it targets the right host, suppression aids survival to the time when establishment in the next host is possible: it is most favorable if it reduces all aspects of host (and hence parasite) mortality. If constrained to have selective effects, suppression should reduce the commonest form of mortality.

Sperm competition and the evolution of male reproductive anatomy in rodents.

Sperm competition is a pervasive selective force in evolution, shaping reproductive anatomy, physiology and behaviour. Here, we present comparative evidence that varying sperm competition levels account for variation in the male reproductive anatomy of rodents, the largest and most diverse mammalian order. We focus on the sperm-producing testes and the accessory reproductive glands, which produce the seminal fluid fraction of the ejaculate. We demonstrate a positive association between relative testis size and the prevalence of within-litter multiple paternity, consistent with previous analyses in which relative testis size has been found to correlate with sperm competition levels inferred from social organization and mating systems. We further demonstrate an association between sperm competition level and the relative size of at least two accessory reproductive glands: the seminal vesicles and anterior prostate. The size of the major product of these glands-the copulatory plug-is also found to vary with sperm competition level. Our findings thus suggest that selection for larger plugs under sperm competition may explain variation in accessory gland size, and highlight the need to consider both sperm and non-sperm components of the male ejaculate in the context of post-copulatory sexual selection.

Spermatozoal traits and sperm competition in Atlantic salmon: relative sperm velocity is the primary determinant of fertilization success.

Sperm competition occurs when sperm from more than one male compete for fertilizations. This form of post-copulatory sexual selection is recognized as a significant and widespread force in the evolution of male reproductive biology and as a key determinant of differential male reproductive success. Despite its importance, however, detailed mechanisms of sperm competition at the gamete level remain poorly understood. Here, we use natural variation in spermatozoal traits among wild Atlantic salmon (Salmo salar), a species naturally adapted to sperm competition, to examine how the relative influences of sperm (i) number, (ii) velocity, (iii) longevity, and (iv) total length determine sperm competition success. Atlantic salmon fertilize externally, and we were therefore able to conduct controlled in vitro fertilization competitions while concurrently measuring spermatozoal traits within the aqueous micro-environment to which salmon gametes are naturally adapted. Microsatellite DNA fingerprinting revealed that a male's relative sperm velocity was the primary determinant of sperm competition success. There was no significant relationship between fertilization success and either relative sperm number or total length; sperm longevity showed an inverse relationship with competition success. These relationships were consistent for two experimental repeats of the in vitro fertilization competitions. Our results therefore show, under the natural microenvironment for salmon gametes, that relative sperm velocity is a key spermatozoal component for sperm competition success. Atlantic salmon sperm can be considered to enter a competition analogous to a race in which the fastest sperm have the highest probability of success.

Sexual selection and speciation in mammals, butterflies and spiders.

Recently refined evolutionary theories propose that sexual selection and reproductive conflict could be drivers of speciation. Male and female reproductive optima invariably differ because the potential reproductive rate of males almost always exceeds that of females: females are selected to maximize mate 'quality', while males can increase fitness through mate 'quantity'. A dynamic, sexually selected conflict therefore exists in which 'competitive' males are selected to override the preference tactics evolved by 'choosy' females. The wide variation across taxa in mating systems therefore generates variance in the outcome of intrasexual conflict and the strength of sexual selection: monandry constrains reproductive heterozygosity and allows female choice to select and maintain particular (preferred) genes; polyandry promotes reproductive heterozygosity and will more likely override female choice. Two different theories predict how sexual selection might influence speciation. Traditional ideas indicate that increased sexual selection (and hence conflict) generates a greater diversity of male reproductive strategies to be counteracted by female mate preferences, thus providing elevated potentials for speciation as more evolutionary avenues of male-female interaction are created. A less intuitively obvious theory proposes that increased sexual selection and conflict constrains speciation by reducing the opportunities for female mate choice under polyandry. We use a comparative approach to test these theories by investigating whether two general measures of sexual selection and the potential for sexual conflict have influenced speciation. Sexual size dimorphism (across 480 mammalian genera, 105 butterfly genera and 148 spider genera) and degree of polyandry (measured as relative testes size in mammals (72 genera) and mating frequency in female butterflies (54 genera)) showed no associations with the variance in speciosity. Our results therefore show that speciation occurs independently of sexual selection.

ADVANTAGES AND DISADVANTAGES OF EGRET AND HERON BROOD REDUCTION.

Data from great egrets and great blue herons were used to test a fundamental assumption of Lack's brood-reduction hypothesis, that mortality is brood-size dependent. This was confirmed for the largest brood sizes (4 and 3), which, in egrets, also have the highest sib-fighting rates. Broods of one, however, experienced paradoxically high mortality, especially early in the season. The hypothesis is advanced that parents desert unprofitably small broods when sufficient time remains for production of a larger brood. A simple game-theory model shows that this parental desertion may hinge primarily on the overall costs of renesting. Egret brood reduction caused by sibling aggression (siblicide) occurred later than less aggressive forms of brood reduction. The inclusive fitness of senior broodmates is maximized by the successful fledging of all sibs, and the physical superiority of seniors (in food-handling for herons; food-handling and aggression for egrets) usually suffices to guarantee their own welfare in brood competitions. Finally, it is shown that the last chick in asynchronously hatching broods represents two kinds of reproductive value (RV) to the parents-"extra RV" (obtained despite the survival of elder sibs) and "insurance RV" (obtained only when at least one elder sib dies first)-which can be distinguished from field data. This approach can be used in comparisons with other asynchronous species for partitioning the fitness contributions of marginal offspring.