The effects of temperature and body size on the mating pattern of a gregariously nesting bee, Colletes cunicularius (Hymenoptera: Colletidae)

Abstract. 1. In a 3-year study of the solitary bee Colletes cunicularius L. in Sweden, average body size and population density fluctuated greatly between years.
2. In this protandrous population, females mated just once and the sex ratio was slightly male biased. Males were smaller than females.
3. Size assortative mating (homogamy), associated with an increase in population density during the central days of female emergence and mating, was observed in two out of three years. Homogamy was also observed in pairs with remating males.
4. Most of the mating males had emerged the day they mated, but 42% were older. We found no support for a general large-male mating advantage.
5. Weight of emerging females and mating males were negatively correlated with ground temperature, indicating thermoregulatory influence on the process of sexual selection in this species.     

Body size and mating success inDrosophila willistoni are uncorrelated under laboratory conditions

Mating activity and wing length were investigated in the F₁ progeny ofDrosophila willistoni females collected in the field to examine any possible relationship between body size and mating success. The flies were observed in a mating chamber under laboratory conditions. No significant differences in wing length were observed between copulating and noncopulating flies, and there was no significant correlation between wing length and copulation latency for both males and females. These results therefore suggest that the commonly accepted view that large body size is positively correlated with mating success inDrosophila does not always hold true. The results support the view that the extent of environmentally induced variation in body size may be an important factor in determining whether an association between body size and mating success is observed inDrosophila species.     

DOES LARGE BODY SIZE IN MALES EVOLVE TO FACILITATE FORCIBLE INSEMINATION? A STUDY ON GARTER SNAKES

A trend for larger males to obtain a disproportionately high number of matings, as occurs in many animal populations, typically is attributed either to female choice or success in male-male rivalry; an alternative mechanism, that larger males are better able to coercively inseminate females, has received much less attention. For example, previous studies on garter snakes (Thamnophis sirtalis parietalis) at communal dens in Manitoba have shown that the mating benefit to larger body size in males is due to size-dependent advantages in male-male rivalry. However, this previous work ignored the possibility that larger males may obtain more matings because of male-female interactions. In staged trials within outdoor arenas, larger body size enhanced male mating success regardless of whether a rival male was present. The mechanism involved was coercion rather than female choice, because mating occurred most often (and soonest) in females that were least able to resist courtship-induced hypoxic stress. Males do physically displace rivals from optimal positions in the mating ball, and larger males are better able to resist such displacement. Nonetheless, larger body size enhances male mating success even in the absence of such male-male interactions. Thus, even in mating systems where males compete physically and where larger body size confers a significant advantage in male-male competition, the actual selective force for larger body size in males may relate to forcible insemination of unreceptive females. Experimental studies are needed to determine whether the same situation occurs in other organisms in which body-size advantages have been attributed to male-male rather than male-female interactions.     

Factors influencing male mating behaviour in Gambusia affinis (Baird & Girard) with a coercive mating system

The effects of male and female body size, and correlated characteristics, on male mating behaviour were investigated in the western mosquitofish Gambusia affinis . Because larger females typically have larger broods in Gambusia sp., it was predicted that males would attempt more copulations with larger females. Two-way ANOVA showed that female body size was a significant predictor of male mating behaviour but male size was not. The effects of a suite of additional traits (both male and female) on male mating attempts were also tested. In a stepwise multiple regression, female standard length ( L _S), size of the female gravid spot and male testes mass were significant predictors of male mating attempts, accounting for c. 27% of variation in male mating. Path analysis showed that differences between male and female L _S, male body condition and male testes mass were significant predictors of male mating attempts, and also accounted for 27% of the variation in male mating attempts. The two statistical models were very similar in their predictive power, but differed slightly in significant predictor variables. Results confirm that factors other than female size are important predictors of male mating behaviour in the western mosquitofish.     

Size Dependent Sexual Selection in Drosophila ananassae

Mate choice based on body size is widespread and can have numerous consequences. We present data, which show the effect of male and female body size on sexual selection in Drosophila ananassae. The relationships between wing size, locomotor activity, mating latency, courtship pattern, fertility and mating success were studied. Mating latency was negatively correlated with wing length and with locomotor activity, while wing length and locomotor activity was positively correlated in males as well as in females. In female- and male-choice, we found that mate choice influenced size-assortative mating by: (1) large and small males preferring to mate with large females, (2) large males successfully competing for large females, leaving small males to mate with small females. Males increased their reproductive success by mating with large and more fecund females. In addition, in pairs of long/short winged flies, long winged flies courted and mated more successfully than short winged flies and they also have longer duration of copulation and more progeny than short winged flies. We found sterile mating in pairs of small winged males and females.     

Female‐biased size dimorphism in a diapausing caddisfly,Mesophylax aspersus: effect of fecundity and natural and sexual selection

1. The effect of mating success, female fecundity and survival probability associated with intra‐sex variation in body size was studied in Mesophylax aspersus, a caddisfly species with female‐biased sexual size dimorphism, which inhabits temporary streams and aestivates in caves. Adults of this species do not feed and females have to mature eggs during aestivation. 2. Thus, females of larger size should have a fitness advantage because they can harbour more energy reserves that could influence fecundity and probability of survival until reproduction. In contrast, males of smaller size might have competitive advantages over others in mating success. 3. These hypotheses were tested by comparing the sex ratio and body size of individuals captured before and after the aestivation period. The associations between body size and female fecundity, and between mating success and body size of males, were explored under laboratory conditions. 4. During the aestivation period, the sex ratio changed from 1 : 1 to male biased (4 : 1), and a directional selection on body size was detected for females but not for males. Moreover, larger clutches were laid by females of larger size. Finally, differences in mating success between small and large males were not detected. These results suggest that natural selection (i.e. the differential mortality of females associated with body size) together with possible fecundity advantages, are important factors responsible of the sexual size dimorphism of M. aspersus. 5. These results highlight the importance of taking into account mechanisms other than those traditionally used to explain sexual dimorphism. Natural selection acting on sources of variation, such as survival, may be as important as fecundity and sexual selection in driving the evolution of sexual size dimorphism.     

Random size‐assortative mating despite size‐dependent fecundity in a Neotropical amphibian with explosive reproduction

Sexual selection theory predicts that, when body size is correlated with fecundity, there should be fitness advantages for mate choice of the largest females. Moreover, because larger males are expected to monopolise the largest females, this should result in an assortative mating based on body size. Although such patterns could be expected in both explosive and prolonged breeders, non‐assortative mating should be more widespread in species under time constraints. However, patterns of sexual selection are largely unexplored in explosive breeding species, and contrasting patterns have been found previously. We expect that the active choice of partners may be particularly risky when the time period during which sexual partners are available is severely limited. Therefore, to avoid missing an entire reproductive act, males and females should pair irrespective of traits, such as body size. We tested this hypothesis by investigating the mating patterns of the Pacific horned toad, Ceratophrys stolzmanni, a short‐lived fossorial species inhabiting Neotropical dry forests. This species is particularly adequate to test our prediction because it reproduces explosively over the course of a single night per year. Although the number of eggs laid was proportional to the size of females, and individuals of both sexes showed variation in body size, there was no assortative mating based either on size, body condition or age of mates. Egg size was not influenced by either female size or clutch size. The larger body size of females compared to males is likely due to fecundity selection, that is, the selective pressure that enhances reproductive output. Although we cannot dismiss the possibility that individuals could select their partners based on other criteria than those related to size or age, the results fit well our prediction, showing that the explosive breeding makes improbable an active choice of partners in both sexes and therefore favours a random mating pattern.     

Components of male fitness in relation to body size in Epirrita autumnata (Lepidoptera, Geometridae)

Abstract. 1. The effect of body size on different components of male fitness was studied for Epirrita autumnata , a geometrid known for its eruptive population dynamics. Body size is the main determinant of female fecundity in this species.
2. Longevity of males was found to have a weak negative correlation with body size at low temperatures. No significant correlation was found at higher temperatures.
3. We found no correlation between male size and female fecundity or egg size which is consistent with the small size of spermatophores in this species.
4. Small and large males were equally successful when allowed to compete for females in laboratory conditions.
5. In one or two field collections, males found mating were larger than males found singly. Large males also had an advantage in finding of virgin females, offered experimentally. No size-assortative mating was recorded.
6. We conclude that size-dependent mate location ability is the factor accounting for most of the variance in male fitness in E.autumnata. The dependence of fitness on body size may well be equally strong in males and females.     

Persistent size variation in the anthophorine bee Centris pallida (Apidae) despite a large male mating advantage

Abstract.

• 1 Despite apparent directional sexual selection in favour of large body size, males of the anthophorine bee Centris pallida remain highly variable in body size.
• 2 One possible cause of persistent size variation among males is geographic variation in the extent of the large male mating advantage. However, a study of a population in an area not previously investigated revealed that the large male mating advantage was as strong here as it has been elsewhere in other years.
• 3 Although the reproductive benefits of being large were consistent in populations separated spatially and temporally, the intensity of bird predation on mate-searching males varied greatly between locations.
• 4 The bee-killing birds focused exclusively on bees which were digging down to meet emerging females or fighting on the ground, never on flying males. Males which were collected on the ground by hand (to simulate avian predation) were significantly larger on average than flying males collected by sweep netting.
• 5 Therefore, in some location in some years, sexual selection in favour of large body size may be opposed by natural selection exerted by predators, perhaps contributing to the maintenance of size variation in this bee.

     

The effect of male mating history on paternal investment, fecundity and female remating in the seed beetle Callosobruchus maculatus

1. In many organisms, males provide nutrients to females via ejaculates that can influence female fecundity, longevity and mating behaviour. The effect of male mating history on male ejaculate size, female fecundity, female longevity and female remating behaviour in the seed beetle Callosobruchus maculatus was determined.
2. The quantity of ejaculate passed to females declined dramatically with successive matings. Despite the decline, a male's ability to fertilize a female fully did not appear to decline substantially until his fourth mating.
3. When females multiply mated with males of a particular mated status, the pattern of egg production was cyclic, with egg production increasing after mating. Females multiply mated to virgins had higher fecundity than females mated to non-virgins, and females mated to twice-mated males had disproportionately increased egg production late in their life.
4. Females that mated to multiple virgins, and consequently laid more eggs, experienced greater mortality than females mated only once or mated to non-virgins, suggesting that egg production is costly, and rather than ameliorating these costs, male ejaculates may increase them by allowing or stimulating females to lay more eggs.
5. Females mating with non-virgin males remated more readily than did females mated to virgins. Females given food supplements were less likely to remate than females that were nutritionally stressed, suggesting that females remate in part to obtain additional nutrients.     

Social and extra-pair mating in relation to major histocompatibility complex variation in common yellowthroats

Females are thought to gain better-quality genes for their offspring by mating with particular males. Genes of the major histocompatibility complex (MHC) play a critical role in adaptive immunity, and several studies have examined female mate choice in relation to MHC variation. In common yellowthroats, females prefer males that have larger black facial masks, an ornament associated with MHC variation, immune function and condition. Here we also tested whether mating patterns are directly correlated with MHC diversity or similarity. Using pyrosequencing, we found that the presence of extra-pair young in the brood was not related to male MHC diversity or similarity between the female and her within-pair mate. Furthermore, extra-pair sires did not differ in overall diversity from males they cuckolded, or in their similarity to the female. MHC diversity is extremely high in this species, and it may limit the ability of females to assess MHC variation in males. Thus, mating may be based on ornaments, such as mask size, which are better indicators of overall male health and genetic quality.     

Colour-associated mating success in a polymorphic Ladybird Beetle, Harmonia axyridis

1. Body size is often an important character in mating success, but has been only infrequently mentioned in regard to colour polymorphism. In this study, mating success was investigated in a colour polymorphic Ladybird Beetle, Harmonia axyridis , with reference both to colour morph and to body size.
2. In the non-melanic males the mating individuals were significantly larger than solitary individuals, while in melanic males there was no significant difference.
3. The mating pattern was close to random mating with respect to colour morph and there was no significant deviation.
4. The results suggest both body size and colour morph affect the male mating success and males of different body size obtain mating advantage according to the colour morph. Colour polymorphism in this species is controlled by alleles on a single locus. Thus, the alleles on that locus significantly influence the effect of selection on the quantitative character.     

Impact of body melanization on mating success in Drosophila melanogaster

Mating speed and copulation duration respond rapidly to laboratory selection in Drosophila melanogaster Meigen (Diptera: Drosophilidae), but there is a lack of data on the evolutionary response to natural selection in the wild. Further, it is not clear whether body melanization and mating behavior are correlated traits. Accordingly, we tested whether variation in body color impacts on mating latency, copulation duration, and fecundity in latitudinal populations of D. melanogaster. We observed geographical variation (cline) for mating propensity, i.e., mating speed as well as copulation duration increased along latitude. Phenotypic plastic responses for body melanization at 17 and 25 °C also showed significant correlations with mating latency and copulation duration. Within‐population analysis based on assorted dark and light flies of five geographical populations showed significant positive correlations of copulation duration and fecundity with body melanization. To assess the role of males and/or females on mating speed and copulation duration, we used atypical body color strains (i.e., dark and light males of D. melanogaster) for no‐choice mating tests. Our data showed a major influence of males for copulation duration and of females for mating speed. Furthermore, a difference in impact of body melanization on mating speed and copulation duration was demonstrated between species, i.e., low melanization in Drosophila ananassae Doleschall is correlated with lower mating speed and shorter copulation duration than in D. melanogaster. Geographical changes in mating propensity were significantly correlated with body melanization at three levels, i.e., within and between populations and between species. Thus, we have shown that a relationship exists between body melanization and mating success. Further, we found seasonal changes in temperature and humidity to confer selection pressures on mating‐related traits.     

Mating ecology of beluga (Delphinapterus leucas) and narwhal (Monodon monoceros) as estimated by reproductive tract metrics

Narwhal and beluga whales are important species to Arctic ecosystems, including subsistence hunting by Inuit, and little is understood about their mating ecology. Reproductive tract metrics vary across species in relation to mating strategy, and have been used to infer mating ecology. Reproductive tracts from beluga and narwhal were collected between 1997 and 2008 from five beluga stocks and two narwhal stocks across the Canadian Arctic. Tract length for males and females, relative testes mass for males, and tusk length for male narwhal were measured. We assessed variation relative to species, body size, stock, maturity, and season. Significant variation was found in testes mass across month and stock for beluga, and no significant difference between stock or date of harvest for narwhal. Beluga had significantly larger testes relative to body size than narwhal, suggesting they were more promiscuous than narwhal. A significant relationship was found between narwhal tusk length and testes mass, indicating the tusk may be important in female mate choice. No significant differences were found between narwhal and beluga reproductive tract length for males or females. The mating systems suggested for narwhal and belugas by our results mean the two species may respond differently to climate change.     

Mating Group Size in Dusky Dolphins (Lagenorhynchus obscurus): Costs and Benefits of Scramble Competition

In fission–fusion social systems with scramble competition between males, multiple males join mating groups while surrounding an oestrous female. If male decisions to join a mating group have been shaped by natural selection, then there should be an optimal group size resulting from the trade‐offs between the benefits of monopolizing a female in small groups and the energy lost in defending her from rivals in large groups. Male dusky dolphins (Lagenorhynchus obscurus) off Kaikoura, New Zealand, provide a unique opportunity to assess the optimum mating group size because they join transient mating groups not confounded by foraging or predator evasion. Within aggregations of up to 1000 individuals, males search for oestrous females, encountering choices of staying with a large mating group or leaving to find a smaller group. Mating groups typically involve multiple males mating with a single female. We conducted focal follows of mating groups (N = 44) by vessel from November 2011 through January 2012. We used video and a GPS to record group size, behaviour and movement. For each group, we measured potential costs (Swim Speed, Loss of Monopolization Potential) and benefits (Copulation Rate and Duration, Energy Savings). Only Loss of Monopolization Potential was positively correlated with group size, while Energy Savings was negatively correlated. Using these two factors as utility functions, we constructed an optimality model and predicted the optimal mating group size to be seven individuals with a range of 4–11 individuals due to variance. The observed modal mating group size was five dolphins, with a range of 2–15. We compare variation in mating group currencies and sizes to past studies. We discuss potential limitations of applying optimality models to predict mating group size for socially complex and behaviourally plastic species such as dolphins.     

The Resource-Based Mating System of the Mediterranean Pollen Wasp Ceramius fonscolombei Latreille 1810 (Hymenoptera, Vespidae, Masarinae)

Males of C. fonscolombei patrol and perch at water collection sites or at plants of Reseda, both of which are important resources for female brood care. The mating system can be classified as resource defense polygyny modified by the existence of alternative male mating tactics. Occupying temporary territories at watercollection sites constitutes the primary tactic which is more profitable for larger males. The secondary tactic of patrolling at flowers provides a nonaggressive alternative through which smaller males gain at least some mating success. Males at water collection sites occupy considerably smaller ranges but spend a higher proportion of time patrolling than males at flowers. They frequently grapple with other males, an activity that is absent at flowers. Males at water collection sites copulate about 2.5 times more frequently than males at flowers. The copulation frequency of the males at water collection sites is positively correlated with their body size, while copulation frequency is negatively correlated with body size at flowers. Males patrolling at water collection sites and males patrolling at flowers do not differ in body size, indicating that the decision between alternative mating tactics is not made relative to body size but is influenced by other factors.     

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).     

Mutual mate choice in crayfish: large body size is selected by both sexes, virginity by males only

Mate choice is often assumed to be a prerogative of females because of their putatively larger reproductive investment than males. However, recent evidence suggests that spermatogenesis is far from being limitless and that males show a high selectivity towards their mates, thus maximizing their reproductive success. We investigated mutual mate choice in the crayfish Procambarus clarkii through two experiments. The first experiment explored the effects of body size, chelar size and chelar symmetry and social status of a potential partner. In the second experiment, we asked whether this species can discriminate between partners of the same body size but with different mating status. We used a binary choice test paradigm, in which two 'targets' with opposing characteristics were simultaneously presented to a test animal, the 'chooser'. The results showed that P. clarkii males are more selective than expected. Similar to the other sex, they were significantly attracted by targets with large body sizes, but not by individuals with larger and symmetric chelae or with a dominant status. An inter-sexual difference was found in the second experiment, in which only males seemed to select virgin potential mates. The several adaptive explanations for these preferences, still under debate, are finally discussed.     

No evidence for size‐assortative mating in the wild despite mutual mate choice in sex‐role‐reversed pipefishes

Size‐assortative mating is a nonrandom association of body size between members of mating pairs and is expected to be common in species with mutual preferences for body size. In this study, we investigated whether there is direct evidence for size‐assortative mating in two species of pipefishes, Syngnathus floridae and S. typhle, that share the characteristics of male pregnancy, sex‐role reversal, and a polygynandrous mating system. We take advantage of microsatellite‐based “genetic‐capture” techniques to match wild‐caught females with female genotypes reconstructed from broods of pregnant males and use these data to explore patterns of size‐assortative mating in these species. We also develop a simulation model to explore how positive, negative, and antagonistic preferences of each sex for body size affect size‐assortative mating. Contrary to expectations, we were unable to find any evidence of size‐assortative mating in either species at different geographic locations or at different sampling times. Furthermore, two traits that potentially confer a fitness advantage in terms of reproductive success, female mating order and number of eggs transferred per female, do not affect pairing patterns in the wild. Results from model simulations demonstrate that strong mating preferences are unlikely to explain the observed patterns of mating in the studied populations. Our study shows that individual mating preferences, as ascertained by laboratory‐based mating trials, can be decoupled from realized patterns of mating in the wild, and therefore, field studies are also necessary to determine actual patterns of mate choice in nature. We conclude that this disconnect between preferences and assortative mating is likely due to ecological constraints and multiple mating that may limit mate choice in natural populations.     

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.