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. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Solitary nesting as an alternative breeding tactic in colonial nesting bluegill sunfish (Lepomis macrochirus)

Colonial breeding can evolve in response to benefits afforded by clumped individuals, such as reduced predation and increased ease of assessing potential mates. However, colonial breeding can also impose costs such as increased disease transmission or increased cuckoldry. Here, we investigate solitary nesting as a potential alternative breeding tactic in colonial breeding bluegill sunfish (Lepomis macrochirus). Most male bluegill, termed parentals, compete for nesting sites in colonies and then court and spawn with females and provide sole care of the eggs. Although nesting in a colony results in reduced predation and fungal infection of broods, it comes at a cost of increased parasitism by specialized cuckolder males that do not nest. We found that 4.5% of parentals forgo spawning in a colony and instead construct nests solitarily. Solitary males were of similar size and age to colonial males, but were in significantly better condition. Solitary males obtained as many eggs as males nesting in the center of colonies, and significantly more than males nesting on the periphery of colonies. Thus, females do not appear to discriminate against solitary males. Solitary males had smaller ear tabs, a presumed sexually selected character used by parental males in intrasexual competition, than colonial males. Tracking data revealed consistency in nesting tactic (but not nest position within the colony) between spawning attempts. We suggest that solitary nesting represents either a facultative decision made by parental males in top condition at the onset of breeding, or a life history decision to forgo spawning in colonies.Communicated by K. Lindström     

Protein content of spermatophores in relation to monandry/polyandry in butterflies

The evolution of a mating system, and specifically mating frequency, is dependent on the costs and benefits to both sexes of mating once or several times. In butterflies, males transfer a spermatophore that contains both sperm and accessory gland products. Accessory gland substances contain nutrients which, in some species, females use to increase their reproductive output and longevity. Nutrients contained in these packaged ejaculates represent investment by males in reproduction. Consequently, the nutritional composition of spermatophores may vary depending on the mating system. There are two lines of arguments concerning the evolution of the nutrient content of ejaculates. One hypothesis argues that male nuptial gifts evolved in the context of certainty of paternity and ease of finding mates; thus spermatophores of polyandrous males (with lower certainty of paternity and greater ease of finding mates) should contain less protein than those of monandrous males, since more spermatophores are produced on average. The other hypothesis argues that polyandry evolved in the context of maximization of male transfer of nutrients to females, and hence spermatophores of polyandrous males should contain more protein than those of monandrous males. In an attempt to distinguish between these two hypotheses, we determined how protein content of ejaculates varied with the degree of polyandry in nine species of pierid and two species of satyrid butterflies. We found that both relative ejaculate mass and protein content increased with the degree of polyandry. Hence our results are consistent with the view that polyandry has evolved in the context of male transfer of nutrients to females, and provides another example of a male adaptation to multiple mating in butterflies.     

Extra-pair paternity and the opportunity for sexual selection in a socially monogamous bird (Dendroica petechia)

We used multi-locus DNA fingerprinting to characterise the genetic mating system of the socially monogamous yellow warbler (Dendroica petechia). Over 2 years there were no instances of brood parasitism, but 59% of families (n = 90) contained extrapair sired young and 37% of offspring (n = 355) were of extra-pair paternity. Most hypotheses for extra-pair mating in monogamous species assume a paternity benefit to extra-pair sires, and focus on the benefit(s) to females. However, the assumption of male benefit has been little tested. Among yellow warblers, known extra-pair sires were just as likely to be cuckolded as any male in the population, and there was at least one reciprocal exchange of extra-pair paternity. Nevertheless, among known extra-pair sires, the paternity gains from extra-pair paternity were, on average, greater than the losses in their own families. These results show there is a paternity benefit to certain males. However, the benefit is not absolute but relative and therefore more difficult to measure. The results also suggest that patterns of extra-pair fertilisation are not determined by female choice alone. Most confirmed extra-pair mates were territorial neighbours, but some resided as far as three territories apart, and greater spatial separation was implied in other cases. Thus, the opportunity for extra-pair mating is great. We estimate that as a result of extra-pair fertilisations, variance in male mating success is increased somewhere between 3-fold and 15-fold over that which would result from within-pair reproduction alone. These findings affirm the potential importance of extra-pair reproduction for sexual selection in monogamous species and they support earlier suggestions that extra-territorial forays by male yellow warblers are for the purpose of extra-pair mating.     

The cause of size-assortative mating in the leaf beetle Trirhabda canadensis (Coleoptera : Chrysomelidae)

Summary Examples of positive assortative mating by body size are abundant but its causes remain controversial. I show that size-assortative mating occurs in the chrysomelid beetle Trirhabda canadensis and I test a series of alternative hypotheses to explain how this mating pattern comes about. Results suggest that assortative mating in this beetle is due to the greater ease with which size-matched pairs can achieve intromission, and not due to size-biased skews in the availability of mates or mate choice favoring large individuals. There was no correlation between male and female elytron length (a measure of body size) at the initiation of courtship, but pairs assorted positively by size at the onset of intromission. Moreover, in the laboratory, there was a negative correlation between male and female size for pairs engaged in courtship that terminated without mating. Assortative mating was not associated with a large-male mating advantage and there was no evidence of female choice of large males. Nor was there unequivocal evidence for male choice of large females; although mating females were slightly larger and considerably heavier than solitary females, males did not differ in the frequency with which they rejected large and small females. Assortative mating in T. canadensis appeared to be caused by the lower ability of mismatched pairs to achieve intromission after an encounter, both when males were larger and when they were smaller than the female.     

Estrus cycle asynchrony in wild female chimpanzees, Pan troglodytes schweinfurthii

Although estrous synchrony has been reported in a number of mammalian species, most often among primates, methodological and analytical problems make it difficult to interpret these results. We developed a novel estrous synchrony index and employed a randomization procedure to analyze long-term observations of female chimpanzee (Pan troglodytes schweinfurthii) estrous cycles at the Mahale Mountains National Park, Tanzania. Our results revealed that female chimpanzees at Mahale avoid synchronizing their estrous periods with each other. We also found that birthrates decreased as the breeding sex ratio increased. We suggest that estrous asynchrony decreases female–female competition for mates. Asynchrony may also reduce the potential for male sexual coercion by nonpreferred mating partners.     

Experimental and natural changes in the peacock's (Pavo cristatus) train can affect mating success

Petrie et al. (1991) demonstrated a correlation between the degree of elaboration of peacocks' trains and their mating success, and also showed that this correlation occurred because females preferred to mate with the male that had the most elaborate train of those sampled on the lek. Although these data suggest that female choice is responsible for non-random mating in this species, they do not conclusively show that train morphology is the cue that females respond to, because they do not rule out the possible influence of another unidentified variable which is correlated with train elaboration. This paper presents an experimental test of the importance of the peacock's train in determining male mating success. If the number or arrangement of eye-spots in the peacock's train influences mating success, then changing the number of eye-spots should change mating success. This prediction was tested in an experiment where the trains of male peafowl (Pavo cristatus) were manipulated by removing a number of eye-spots between mating seasons. Peacocks with eye-spots removed showed a significant decline in mating success between seasons compared with a control group. This result, together with the observational data, supports the hypothesis that the peacock's train has evolved, at least in part, as a result of female choice.     

Mating system and individual reproductive success of sympatric anadromous and resident brook charr, <Emphasis Type="Italic">Salvelinus fontinalis</Emphasis>, under natural conditions

Salmonids are known for the occurrence in sympatry of two life-history forms, one that undergoes migration to sea before returning to freshwater to reproduce (anadromous) and one that inhabits freshwater without a migration phase (resident). Whereas one breeding population is often suggested by population genetic studies, mating patterns have rarely been directly assessed, especially when both sexes are found within each life-history form. By using highly polymorphic microsatellite loci and parentage analysis in a natural population of sympatric anadromous and resident brook charr (Salvelinus fontinalis), we found that gene flow occurred between the two forms and was mediated by resident males mating with both resident and anadromous females. Determinants of reproductive success, estimated by the number of surviving juveniles (ages 1 and 2 years), differed between the sexes. No strong evidence of the influence of size on individual reproductive success was found for males, whereas larger females (and hence most likely to be anadromous) were more successful. The higher individual reproductive success of anadromous fish compared to residents was mainly explained by this higher reproductive success of anadromous females. We suggest that resident males adopt a “sneaking” reproductive tactic as a way of increasing their reproductive success by mating with females of all sizes in all habitats. The persistence of the resident tactic among females may be linked to their advantage in accessing spatially constrained spawning areas in small tributary streams unavailable to larger females.     

Environmental and genetic influences on mating strategies along a replicated food availability gradient in guppies (Poecilia reticulata)

Food availability is expected to influence the relative cost of different mating tactics, but little attention has been paid to this potential source of adaptive geographic variation in behavior. Associations between the frequency of different mating tactics and resource availability could arise because tactic use responds directly to food intake (phenotypic plasticity), because populations exposed to different average levels of food availability have diverged genetically in tactic use, or both. Different populations of guppies (Poecilia reticulata) in Trinidad experience different average levels of food availability. We combined field observations with laboratory “common garden” and diet experiments to examine how this environmental gradient has influenced the evolution of male mating tactics. Three independent components of variation in male behavior were found in the field: courtship versus foraging, dominance interactions, and interference competition versus searching for mates. Compared with low-food-availability sites, males at high-food-availability sites devoted more effort to interference competition. This difference disappeared in the common garden experiment, which suggests that it was caused by phenotypic plasticity and not genetic divergence. In the diet experiment, interference competition was more frequent and intense among males raised on the greater of two food levels, but this was only true for fish descended from sites with low food availability. Thus, the association between interference competition and food availability in the field can be attributed to a genetically variable norm of reaction. Genetically variable norms of reaction with respect to food intake were found for the other two behavioral components as well and are discussed in relation to the patterns observed in the field. Our results indicate that food availability gradients are an important, albeit complex, source of geographic variation in male mating strategies.     

Independence, not conflict, characterizes dart-shooting and sperm exchange in a hermaphroditic snail

Although the sexes are united in hermaphrodites, conflict can still occur because the male and female functions have separate interests. We examined the evidence for conflict in the mating system of the terrestrial snail Cantareus aspersus (formerly Helix aspersa) where sharp, calcareous darts are ‘shot’ during courtship. We predicted that the use of the dart would either reflect or create conflict and this would be evident in either the courtship behavior or the transference of sperm. Previous studies demonstrated that the dart functions after sperm transfer to increase sperm survival. Using detailed observations of mating snails, we examined the factors that determine dart shooting order, the behavioral responses after being hit by a dart, the accuracy of dart shooting, and the allocation of sperm resources. We found that each dart was shot independently, and each animal appeared to be interested only in getting off the best possible shot, probably one that penetrates deeply near the genital pore. There is no evidence of mating conflict. Every snail transfers sperm to its partner, and the size of the donation does not depend on the success or failure of either snail’s dart shot. Although the receipt of a dart does not appear to cause harm, it may produce indirect costs due to the partial loss of control over fertilization. We conclude that mating in C. aspersus is a partnership in which independent actors demonstrate unconditional reciprocity during courtship and sperm transfer.