Clint D. Kelly

Sexual selection and sperm quantity: meta-analyses of strategic ejaculation

Multiple mating or group spawning leads to post‐copulatory sexual selection, which generally favours ejaculates that are more competitive under sperm competition. In four meta‐analyses we quantify the evidence that sperm competition (SC) favours greater sperm number using data from studies of strategic ejaculation. Differential investment into each ejaculate emerges at the individual level if males exhibit phenotypic plasticity in ejaculate properties in response to the likely risk and/or intensity of sperm competition after a given mating. Over the last twenty years, a series of theoretical models have been developed that predict how ejaculate size will be strategically adjusted in relation to: (a) the number of immediate rival males, with a distinction made between 0 versus 1 rival (‘risk’ of SC) and 1 versus several rivals (‘intensity’ of SC); (b) female mating status (virgin or previously mated); and (c) female phenotypic quality (e.g. female size or condition). Some well‐known studies have reported large adjustments in ejaculate size depending on the relevant social context and this has led to widespread acceptance of the claim that strategic sperm allocation occurs in response to each of these factors. It is necessary, however, to test each claim separately because it is easy to overlook studies with weak or negative findings. Compiling information on the variation in outcomes among species is potentially informative about the relevance of these assumptions in different taxa or mating systems.

Geographic variation in multiple paternity within natural populations of the guppy (Poecilia reticulata).

Mating can increase an individuals risk of mortality by predation. In response to predation hazards, males in some species court females less often, but alternatively engage in coerced copulations more frequently and females become less selective. Such predator‐mediated shifts in mating tactics may result in higher levels of multiple inseminations in females and, thus, in greater frequencies of females with broods of mixed paternity. We tested this hypothesis using two polymorphic microsatellite loci to estimate conservatively multiple paternity in broods of female guppies (Poecilia reticulata) originating from ten natural populations that have evolved under different fish predation regimes in Trinidad. The frequency of broods that were multiply sired was significantly greater on average in populations experiencing high predation pressure compared to populations experiencing a relatively low predation risk. These results suggest that the intensity of male sperm competition covaries geographically with predation pressure in this species and that the local risk of predation mediates the opportunity for sexual selection within populations.

Replicating Empirical Research In Behavioral Ecology: How And Why It Should Be Done But Rarely Ever Is

That empirical evidence is replicable is the foundation of science. Ronald Fisher, a founding father of biostatistics, recommended that a null hypothesis be rejected more than once because “no isolated experiment, however significant in itself, can suffice for the experimental demonstration of any natural phenomenon” (Fisher 1974:14). Despite this demand, animal behaviorists and behavioral ecologists seldom replicate studies. This practice is not part of our scientific culture, as it is in chemistry or physics, due to a number of factors, including a general disdain by journal editors and thesis committees for unoriginal work. I outline why and how we should replicate empirical studies, which studies should be given priority, and then elaborate on why we do not engage in this necessary endeavor. I also explain how to employ various statistics to test the replicability of a series of studies and illustrate these using published studies from the literature.

Fighting for harems : assessment strategies during male-male contests in the sexually dimorphic Wellington tree weta

Females often aggregate at particular sites for feeding or shelter, thus giving adult males the opportunity to defend harems and increase male reproductive success. Rival males compete for control of harems via ritualized displays or direct combat using weaponry. Contests for harems or the resources required by females can be settled based on asymmetries in fighting ability or resource ownership. Males that accurately assess a rivals fighting ability prior to engaging in potentially costly combat should be favoured by selection. Game theory and optimality models provide three models to explain how individuals decide to persist in or flee from a fight. These models are the energetic war of attrition, the sequential assessment model and the cumulative assessment model. Using staged contests in the laboratory, I tested predictions of these models using the Wellington tree weta, Hemideina crassidens, a sexually dimorphic insect native to New Zealand. Male H. crassidens use their enlarged mandibles as weapons in fights for access to adult females that reside in cavities in trees. My results supported a prediction common to each assessment model: contest duration was negatively correlated with the asymmetry in opponents weapon size. The sequential assessment model of contest settlement was partially supported but the strongest support was for the cumulative assessment model. Predictions of the latter model were supported because: (1) fights are apparently settled based on own-size assessment; (2) fights occur in a single phase and escalate; and (3) contests involve physical combat and injury. I suggest that, in nocturnal species, cumulative assessment will generally be most applicable.

The interrelationships between resource-holding potential, resource-value and reproductive success in territorial males: How much variation can we explain?

A long-standing hypothesis in behavioural ecology posits that males with greater resource-holding potential (RHP) control resource sites deemed more valuable by sexually-receptive females and, thereby, males controlling such sites accrue greater reproductive success (RS). This hypothesis has historically been investigated using three separate but non-mutually exclusive relationships (male RHP vs. resource value, resource value vs. male RS and male RHP vs. RS). The relationships between these three variables are predicted to be strongly positive, however, due to measurement error and biological noise, perfect correlations (r = 1.0) are rare in biology even for well-established relationships. Moreover, the inaccurate identification of either the male trait(s) important to RHP or the resource characteristic sought by females will weaken the observed strength of the relationships. Here, I use meta-analysis to quantitatively describe the general pattern of these relationships in animals. I predict that the relationships between male RHP, resource-value and RS should be significantly positive (male RHP and resource-value should explain a large amount of the variation in male RS). My meta-analysis supports this hypothesis; however, in the best case scenario only ca. 20% of the variation in the response variable was explained. I conclude by identifying areas in which we need to improve our investigations of resource-defence animals and recommending approaches to meet these needs.

Sexual Selection for Male Mobility in a Giant Insect with Female‐Biased Size Dimorphism

Female‐biased size dimorphism, in which females are larger than males, is prevalent in many animals. Several hypotheses have been developed to explain this pattern of dimorphism. One of these hypotheses, the mobility hypothesis, suggests that female‐biased size dimorphism arises because smaller males are favored in scramble competition for mates. Using radiotelemetry, we assessed the mobility hypothesis in the Cook Strait giant weta (Deinacrida rugosa), a species with strong female‐biased size dimorphism, and tested the prediction that male traits promoting mobility (i.e., longer legs, smaller bodies) are useful in scramble competition for mates and thus promote reproductive success. Our predictions were supported: males with longer legs and smaller bodies exhibited greater mobility (daily linear displacement when not mating), and more mobile males had greater insemination success. No phenotypic traits predicted female mobility or insemination success. In species with female‐biased size dimorphism, sexual selection on males is often considered to be weak compared to species in which males are large or possess weaponry. We found that male giant weta experience sexual selection intensities on par with males of a closely related harem‐defending polygynous species, likely because of strong scramble competition with other males.

Predation risk reduces male-male sexual competition in the Trinidadian guppy (Poecilia reticulata)

Abstract. Because agonistic behaviour can increase an individuals risk of predation, natural selection should favour individuals that modify their behaviour in the face of predation hazard in ways that maximise fitness. We investigated experimentally the effects of an increase in the apparent risk of predation on male-male competition within mixed-sex shoals of the Trinidadian guppy (Poecilia reticulata). We show that males engaged in significantly fewer aggressive interactions with conspecific males in the presence of a fish predator. However, they continued to court and attempt copulations with females at the same rate irrespective of the level of predation risk. In comparison, under predation hazard, female guppies were less responsive sexually and avoided fewer male copulation attempts. Such predator-mediated changes in male-male agonistic interactions and female sexual responsiveness towards males have important implications for the mode and intensity of sexual selection within populations.

Transparency in ecology and evolution: Real problems, real solutions

To make progress scientists need to know what other researchers have found and how they found it. However, transparency is often insufficient across much of ecology and evolution. Researchers often fail to report results and methods in detail sufficient to permit interpretation and meta-analysis, and many results go entirely unreported. Further, these unreported results are often a biased subset. Thus the conclusions we can draw from the published literature are themselves often biased and sometimes might be entirely incorrect. Fortunately there is a movement across empirical disciplines, and now within ecology and evolution, to shape editorial policies to better promote transparency. This can be done by either requiring more disclosure by scientists or by developing incentives to encourage disclosure.

Resource quality or harem size: what influences male tenure at refuge sites in tree weta (Orthoptera: Anostostomatidae)?

Rather than seeking females directly, males in many animal species locate and defend sites that contain spatially limited resources essential for female survival and reproduction. In these cases, resident males successfully repelling conspecific rivals will mate with sexually receptive females that seek to use the resident’s resources. Theory predicts that if resources are limiting in nature, are costly to procure, and if residency at the resource site increases male reproductive success, each site should be monopolised by a single adult male. Moreover, if females aggregate at these sites, males should be sedentary and monopolise harems. This predicts that males should reside at resource sites longer than females and male tenure should be positively correlated with harem size. I address these hypotheses using a wild population of the Wellington tree weta, Hemideina crassidens, a sexually dimorphic insect in which males use mandibular weapons in fights to control resource sites (galleries in trees) required by females for diurnal refuge. In a longitudinal study using artificial galleries, I show that male tenure in a gallery is positively related to harem size and, contrary to prediction, not gallery size per se. Moreover, contrary to prediction, females are more sedentary than males: males appear to move frequently between galleries to search for novel mates. I show experimentally that galleries represent mating sites to males: males prefer galleries housing adult females whereas females prefer unoccupied galleries. Females likely avoid male-occupied galleries because they incur injuries when interacting with males.

Sexual Selection, Ontogenetic Acceleration, and Hypermorphosis Generates Male Trimorphism in Wellington Tree Weta

Strong sex-specific selection on traits common to both sexes typically results in sexual dimorphism. Here we find that Wellington tree weta (Hemideina crassidens) are sexually dimorphic in both head shape and size due to differential selection pressures on the sexes: males use their heads in male-male combat and feeding whereas females use theirs for feeding only. Remarkably, the sexes share a common ontogenetic trajectory with respect to head growth. Male head shape allometry is an extension of the female’s trajectory despite maturing two instars earlier, a feat achieved through ontogenetic acceleration and hypermorphosis. Strong sexual selection also favours the evolution of alternative reproductive strategies in which some males produce morphologically different weapons. Wild-caught male H. crassidens are trimorphic with regard to weapon size, a rare phenomenon in nature, and weapon shape is related to each morph’s putative mating strategy.