Sexual dimorphism,body size,bite force and male mating success in tuatara

Sexual dimorphisms in body size and head size are common among lizards and are often related to sexual selection on male fighting capacity (organismal performance) and territory defence. However, whether this is generally true or restricted to lizards remains untested. Here we provide data on body and head size, bite performance and indicators of mating success in the tuatara (Sphenodon punctatus), the closest living relative to squamates, to explore the generality of these patterns. First, we test whether male and female tuatara are dimorphic in head dimensions and bite force, independent of body size. Next, we explore which traits best predict bite force capacity in males and females. Finally, we test whether male bite force is correlated with male mating success in a free‐ranging population of tuatara (Sphenodon punctatus). Our data confirm that tuatara are indeed dimorphic in head shape, with males having bigger heads and higher bite forces than females. Across all individuals, head length and the jaw closing in‐lever are the best predictors of bite force. In addition, our data show that males that are mated have higher absolute but not relative bite forces. Bite force was also significantly correlated to condition in males but not females. Whereas these data suggest that bite force may be under sexual selection in tuatara, they also indicate that body size may be the key trait under selection in contrast to what is observed in squamates that defend territories or resources by biting. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 287–292.     

The effects of male body size on territorial and mating success in the landmark-defending fly Hermetia comstocki (Stratiomyidae)

Abstract.

• 1 Males of Hermetia comstocki Williston compete for territorial control of certain agaves and yuccas. Winners copulate with females that visit these plants solely to acquire a mate.
• 2 Males vary in body weight by more than an order of magnitude and larger flies almost always defeat smaller ones in aerial contests for control of landmark territories.
• 3 The mean body size (as measured by wing-length) was significantly greater for males retaining residency at a site for at least one hour compared to males unable to do so. Likewise, males able to return to a perch site in the study area on more than one day were larger on average than males unable to do so.
• 4 Male preferences for landmark territories remained similar across years. Large males dominated the perch landmarks most likely to be occupied by males and most likely to be visited by females.
• 5 Despite the fighting and territorial advantages enjoyed by large males, the mean size of males found mating with females was not significantly larger than that of the general population.
• 6 The apparent failure of large males to secure a statistically significant mating advantage may be a statistical consequence of the small sample size of males observed mating. On the other hand, any mating advantage of large males may be reduced because (a) receptive females visit many different landmarks, (b) females mate with the first male they encounter at a landmark, regardless of his size, (c) there are usually many vacant landmarks available for smaller males, and (d) even popular territories are often open to small males, thanks to the low site-tenacity of territory owners.

     

The mating system of a bee fly (Diptera: Bombyliidae). II. Factors affecting male territorial and mating success

Males of an undescribed bombyliidfly (Comptosia sp.)occupy traditional territories on a Southeast Queensland hilltop, to which females come solely for the purpose of mating. Territorial fights between males involve aerial collisions during which modified spines on the wing margins produce scars on the bodies of opponents. Territory owners and mating males are not different in size or age from the remainder of the male population. Although residency is related to fighting success, the strength of the effect is ambiguous. Consequently, our data do not appear to fit predictions from game theoretical models for fighting protocol. Hilltop males lacked the extensive population variation typically found in territorial species, and thus, the presumed advantages of traits such as large size may be suppressed. Hilltop males were larger than males at a nonhilltop, resource-based mating site and the possibility of alternative mating tactics is discussed.     

Size and mating success in males of the western harvester ant,Pogonomyrmex occidentalis (Hymenoptera: Formicidae)

Mating success in males of the lek mating ant species,Pogonomyrmex occidentalis, increases with increased body size. We estimated the magnitude of the selection coefficients on components of size by collecting males in copula and comparing their morphology to that of males that were collected at the lek but that were not mating. Four characters, body mass, head width, wing length, and leg length, were measured for a sample of 225 mating and 324 nonmating males and 225 females. Significant direct selection favors increased wing length and leg length. Multiple regression of transformed variables (principal components) indicated that the increased mating success of larger males is a function of all four characters. We found no evidence of positive assortative mating on the basis of any individual character or on the multivariate general size variable (the first principal component).     

Covariation and repeatability of male mating effort and mating preferences in a promiscuous fish

Although mate choice by males does occur in nature, our understanding of its importance in driving evolutionary change remains limited compared with that for female mate choice. Recent theoretical models have shown that the evolution of male mate choice is more likely when individual variation in male mating effort and mating preferences exist and positively covary within populations. However, relatively little is known about the nature of such variation and its maintenance within natural populations. Here, using the Trinidadian guppy (Poecilia reticulata) as a model study system, we report that mating effort and mating preferences in males, based on female body length (a strong correlate of fecundity), positively covary and are significantly variable among subjects. Individual males are thus consistent, but not unanimous, in their mate choice. Both individual mating effort (including courtship effort) and mating preference were significantly repeatable. These novel findings support the assumptions and predictions of recent evolutionary models of male mate choice, and are consistent with the presence of additive genetic variation for male mate choice based on female size in our study population and thus with the opportunity for selection and further evolution of large female body size through male mate choice.     

Sexual selection, sexual size dimorphism and Rensch's rule in Odonata

Odonata (dragonflies and damselflies) exhibit a range of sexual size dimorphism (SSD) that includes species with male-biased (males > females) or female-biased SSD (males < females) and species exhibiting nonterritorial or territorial mating strategies. Here, we use phylogenetic comparative analyses to investigate the influence of sexual selection on SSD in both suborders: dragonflies (Anisoptera) and damselflies (Zygoptera). First, we show that damselflies have male-biased SSD, and exhibit an allometric relationship between body size and SSD, that is consistent with Rensch's rule. Second, SSD of dragonflies is not different from unit, and this suborder does not exhibit Rensch's rule. Third, we test the influence of sexual selection on SSD using proxy variables of territorial mating strategy and male agility. Using generalized least squares to account for phylogenetic relationships between species, we show that male-biased SSD increases with territoriality in damselflies, but not in dragonflies. Finally, we show that nonagile territorial odonates exhibit male-biased SSD, whereas male agility is not related to SSD in nonterritorial odonates. These results suggest that sexual selection acting on male sizes influences SSD in Odonata. Taken together, our results, along with avian studies (bustards and shorebirds), suggest that male agility influences SSD, although this influence is modulated by territorial mating strategy and thus the likely advantage of being large. Other evolutionary processes, such as fecundity selection and viability selection, however, need further investigation.     

Male mating success in the Azorean rock-pool blenny: the effects of body size, male behaviour and nest characteristics

Larger male Azorean rock-pool blennies Parablennius sanguinolentus parvicornis received more female visits, were courted more often by females and received more spawnings. Larger males also received a higher number of male intrusions, attacked more conspecifics and defended larger territories. Larger males showed more nest cleaning behaviour and a marginally non-significant trend for higher egg fanning rate. Male courtship, male attack rate against conspecifics and parental behaviour were all correlated with the frequency of female spawnings received by each male even when controlling for male size, suggesting that these behaviour patterns influence male mating success. On the other hand, a positive partial correlation was found between female courtship and the frequency of female spawnings, controlling for the number of female visits, which suggests a role for female mate choice on male mating success. Finally, males nesting in chambers in the bottom of pools received more spawnings than males nesting either in crevices or under boulders. However, nest opening area was associated significantly negatively with male mating success, when controlling for male size. Thus, the present data suggest strongly that male characteristics overrule nest characteristics in determining male mating success in the Azorean rock-pool blenny.     

Fat and sassy: factors underlying male mating success in a butterfly

Theory predicts that male mating success depends on resource holding potential (RHP), which is reflected by proxies of condition, such as body mass, fat content, strength, or weaponry. In species lacking any physical means to inflict injuries upon combatants, such as butterflies, the factors determining mating success are less clear. Against this background, we explored the determinants of male mating success in the tropical butterfly Bicyclus anynana Butler (Lepidoptera: Nymphalidae: Satyrinae), by comparing physiological, immunological, and morphological traits between winning and losing males. Our results showed that successful males are characterized by a better flight performance, evidenced by having longer wings, a heavier thorax, a lighter abdomen, a higher fat content, and higher phenoloxidase expression levels than their unsuccessful counterparts, when being compared after their first mating. Males that won three consecutive trials against the same combatant were also characterized by a better flight performance, having larger forewings, a higher body mass, and a higher fat content. Thus, successful males were larger and in better condition than unsuccessful ones. Strikingly, many differences found indicated an enhanced flight performance for the former, which we suggest ultimately plays the key role for male mating success in B. anynana. As fat is the main energy source for flying insects, being crucial to flight endurance and in turn presumably to male mating success, it may represent a key determinant at the proximate level.     

Sexual size dimorphism and selection in the wild in the waterstrider Aquarius remigis: Body size,components of body size and male mating success

Sexual size dimorphism is assumed to be adaptive and is expected to evolve in response to a difference in the net selection pressures on the sexes. Although a demonstration of sexual selection is neither necessary nor sufficient to explain the evolution of sexual size dimorphism, sexual selection is generally assumed to be a major evolutionary force. If contemporary sexual selection is important in the evolution and maintenance of sexual size dimorphism then we expect to see concordance between patterns of sexual selection and patterns of sexual dimorphism. We examined sexual selection in the wild, acting on male body size, and components of body size, in the waterstrider Aquarius remigis, as part of a long term study examining net selection pressures on the two sexes in this species. Selection was estimated on both a daily and annual basis. Since our measure of fitness (mating success) was behavioral, we estimated reliabilities to determine if males perform consistently. Reliabilities were measured as ? statistics and range from fair to perfect agreement with substantial agreement overall. We found significant univariate sexual selection favoring larger total length in the first year of our study but not in the second. Multivariate analysis of components of body size revealed that sexual selection for larger males was not acting directly on total length but on genital length. Sexual selection for larger male body size was opposed by direct selection favoring smaller midfemoral lengths. While males of this species are smaller than females, they have longer genital segments and wider forefemora. Patterns of contemporary sexual selection and sexual size dimorphism agree only for genital length. For total length, and all other components of body size examined, contemporary sexual selection was either nonsignificant or opposed the pattern of size dimporhism. Thus, while the net pressures of contemporary selection for the species may still act to maintain sexual size dimorphism, sexual selection alone does not.     

Couch potatoes do better: Delayed dispersal and territory size affect the duration of territory occupancy in a monogamous mammal

In territorial, socially monogamous species, the establishment and defense of a territory are an important strategy to maximize individual fitness, but the factors responsible for the duration of territory occupancy are rarely studied, especially in long‐lived mammals. A long‐term monitoring program in southeast Norway spanning over 18 years allowed us to follow the individual life histories of Eurasian beavers (Castor fiber) from adolescence in their natal family group to dispersal and territory establishment until the end of territory occupancy. We investigated whether territory size, resource availability, population density, and dispersal age could explain the duration of territory occupancy, which ranged from 1 to 11 years. The duration of territory occupancy was positively related to dispersal age, suggesting that individuals that delayed dispersal had a competitive advantage due to a larger body mass. This is in support with the maturation hypothesis, which states that an animal should await its physical and behavioral maturation before the acquisition of a territory. Further, we found that individuals that established in medium‐sized territories occupied them longer as compared to individuals in small or large territories. This suggests that large territories are more costly to defend due to an increased patrolling effort, and small territories might not have sufficient resources. The lifetime reproductive success ranged from zero to six kits and generally increased with an increasing duration of territory occupancy. Our findings show the importance of holding a territory and demonstrate that dispersal decisions and territory selection have important consequences for the fitness of an individual.     

Dynamics of mating success in experimental groups ofDrosophila melanogaster (Diptera: Drosophilidae)

Determinants of male courtship success in Drosophila melanogasterwere examined in groups of five males sequentially presented with five individual females. Thirty-three percent of males never mated, while approximately half of the males mated two or three times. Rapid courtship initiation was associated with male success in early matings only. Male size was important for courtship outcome, but the size distributions of mating and nonmating males and their progeny numbers indicate balancing rather than directional selection on size- dependent courtship success.     

Body size influences mating success of the Eucalyptus longhorned borer (Coleoptera: Cerambycidae)

Both sexes of adultPhoracantha semipunctata F. (Coleoptera: Cerambycidae) congregate on stressedEucalyptus that are the larval hosts. In a field study, 721 adultP. semipunctata captured on host trees varied considerably in body size with the largest individuals being about twice the length of the smallest. Females that were paired with a mate were similar in size to solitary females, suggesting that the probability of a female being mated was not affected by her size. However, large males had greater success than smaller males in obtaining mates. MaleP. semipunctata rely on antennal contact to locate and identify females on the larval host. Therefore, the rate at which males search for mates is a function of the area swept by their antennae per unit time. Because of their greater antennal spread, large males were able to search for females at double the rate of the smallest males. Large males also dominated in aggressive contests for females. The superior abilities of large maleP. semipunctata in both locating and defending mates account for the influence of body size on mating success.     

Polymorphisms in a desaturase 2 ortholog associate with cuticular hydrocarbon and male mating success variation in a natural population of Drosophila serrata

Elucidating the nature of genetic variation underlying both sexually selected traits and the fitness components of sexual selection is essential to understanding the broader consequences of sexual selection as an evolutionary process. To date, there have been relatively few attempts to connect the genetic variance in sexually selected traits with segregating DNA sequence polymorphisms. We set out to address this in a well‐characterized sexual selection system – the cuticular hydrocarbons (CHCs) of Drosophila serrata – using an indirect association study design that allowed simultaneous estimation of the genetic variance in CHCs, sexual fitness and single nucleotide polymorphism (SNP) effects in an outbred population. We cloned and sequenced an ortholog of the D. melanogaster desaturase 2 gene, previously shown to affect CHC biosynthesis in D. melanogaster, and associated 36 SNPs with minor allele frequencies > 0.02 with variance in CHCs and sexual fitness. Three SNPs had significant multivariate associations with CHC phenotype (q‐value < 0.05). At these loci, minor alleles had multivariate effects on CHCs that were weakly associated with the multivariate direction of sexual selection operating on these traits. Two of these SNPs had pleiotropic associations with male mating success, suggesting these variants may underlie responses to sexual selection due to this locus. There were 15 significant male mating success associations (q‐value < 0.1), and interestingly, we detected a nonrandom pattern in the relationship between allele frequency and direction of effect on male mating success. The minor‐frequency allele usually reduced male mating success, suggesting a positive association between male mating success and total fitness at this locus.     

Brain size affects responsiveness in mating behaviour to variation in predation pressure and sex ratio

Despite ongoing advances in sexual selection theory, the evolution of mating decisions remains enigmatic. Cognitive processes often require simultaneous processing of multiple sources of information from environmental and social cues. However, little experimental data exist on how cognitive ability affects such fitness‐associated aspects of behaviour. Using advanced tracking techniques, we studied mating behaviours of guppies artificially selected for divergence in relative brain size, with known differences in cognitive ability, when predation threat and sex ratio was varied. In females, we found a general increase in copulation behaviour in when the sex ratio was female biased, but only large‐brained females responded with greater willingness to copulate under a low predation threat. In males, we found that small‐brained individuals courted more intensively and displayed more aggressive behaviours than large‐brained individuals. However, there were no differences in female response to males with different brain size. These results provide further evidence of a role for female brain size in optimal decision‐making in a mating context. In addition, our results indicate that brain size may affect mating display skill in male guppies. We suggest that it is important to consider the association between brain size, cognitive ability and sexual behaviour when studying how morphological and behavioural traits evolve in wild populations.     

Environmental and demographic drivers of male mating success vary across sequential reproductive episodes in a polygynous breeder

Ecological and social factors underpinning the inequality of male mating success in animal societies can be related to sex ratio, sexual conflict between breeders, effects of nonbreeders, resource dispersion, climatic conditions, and the various sequential stages of mating competition that constitute the sexual selection process. Here, we conducted an individual‐based study to investigate how local resource availability and demography interact with annual climate conditions to determine the degree of male mating inequality, and thus opportunity for sexual selection across two sequential reproductive episodes (harem and subsequent mate acquisition) in a naturally regulated (feral) horse population in Sable Island National Park Preserve, Canada. Using a 5‐year, spatially explicit, mark‐resight dataset and hierarchical mixed‐effects linear modeling, we evaluated the influence of adult sex ratio (ASR) on mating success and then tested for effects of freshwater availability, density, unpaired male abundance, and precipitation during each breeding season. Unpaired male abundance, freshwater availability, and ASR differed in their effects on male mating success according to year and selection episode. Opportunity for sexual selection in males associated with harem acquisition increased with ASR, and unpaired male abundance further explained weather‐related interannual variation after accounting for ASR. In contrast, once a harem was secured, ASR had little effect on male mating inequality in regard to acquiring additional females, while interannual variation in mating inequality increased with decreasing freshwater availability. Our findings show that local demography, resource availability, and weather effect opportunity for sexual selection in males differently depending on selection episode, and can attenuate or accentuate effects of ASR.     

Male size does not affect the strength of male mate choice for high-quality females in Drosophila melanogaster

Theory predicts that the strength of male mate choice should vary depending on male quality when higher-quality males receive greater fitness benefits from being choosy. This pattern extends to differences in male body size, with larger males often having stronger pre- and post-copulatory preferences than smaller males. We sought to determine whether large males and small males differ in the strength (or direction) of their preference for large, high-fecundity females using the fruit fly, Drosophila melanogaster. We measured male courtship preferences and mating duration to show that male body size had no impact on the strength of male mate choice; all males, regardless of their size, had equally strong preferences for large females. To understand the selective pressures shaping male mate choice in males of different sizes, we also measured the fitness benefits associated with preferring large females for both large and small males. Male body size did not affect the benefits that males received: large and small males were equally successful at mating with large females, received the same direct fitness benefits from mating with large females, and showed similar competitive fertilization success with large females. These findings provide insight into why the strength of male mate choice was not affected by male body size in this system. Our study highlights the importance of evaluating the benefits and costs of male mate choice across multiple males to predict when differences in male mate choice should occur.     

Male mating success in a North American pitviper: influence of body size,testosterone, and spatial metrics

Males with enhanced traits relative to conspecifics often show increased mating and reproductive success and thus have a fitness advantage. The opportunity or potential for sexual selection is predicted to occur under these conditions. Here, we investigated proximate determinants of mating success in male copperhead snakes (Agkistrodon contortrix), a medium‐sized pitviper of North America. Specifically, we investigated the relationships of body size (snout‐vent length, body mass), body condition index, spatial metrics (total distance moved, home range size), and plasma testosterone concentration on mating success in males. The single mating season lasts from August through September. We compared a set of candidate linear mixed models and selected the best‐fitting one using the adjusted Akaike Information Criterion (AICc). The AICc‐selected model (model 2), with testosterone, body condition index, and home range size as predictor variables, showed that male mating success was positively correlated with testosterone. To our knowledge, this is the first report to show the relationship of testosterone and individual mating success in any snake species. A parallel study conducted on male fitness in A. contortrix of the same population used microsatellite markers to assign parentage of fathers (known mothers). Unlike our study, they found that snout‐vent length was positively correlated with reproductive success and that males were experiencing greater sexual selection. This relationship has been detected under natural conditions in other species of snakes. Although behavioural data are important in any mating system analysis, they should not stand alone to infer parentage, relationships or selection metrics (e.g. Bateman gradients). Long‐term sperm storage by females, female cryptic choice, and other factors contribute to the complexity of mating success of males. Accordingly, we thus conclude that estimates of reproductive success and fitness in cryptic species, such as copperheads and other snakes, require robust molecular methods to draw accurate conclusions regarding proximate and evolutionary responses. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 115 , 185–194.     

Body size and alternative mating tactics in the beewolf Philanthus zebratus (Hymenoptera; Sphecidae)

A study of the mating behaviour of males of the beewolf Philanthus zebratus revealed that in one population males display variability in mating tactics and that this variability is related to male body size. There was a tendency for large males to patrol the airspace above the nesting area while smaller males were territorial adjacent to it. The mean sizes of the two groups of males were significantly different, although the size ranges of the two groups overlapped. Only 2.5% of the males were observed to undertake both mating tactics, at different times. Observations are presented on daily and seasonal activity patterns and on the relative location of nests, territories, and patrolling males. A second population, with lower nest density, was observed for several days, revealing only territorial males. It is suggested that the presence of patrolling males is related to the higher nest density of the one population. The fact that patrolling males tend to be relatively large is possibly related to flight energetics or simply to the ability of large males to seize females, which are usually larger than males, in mid-air.