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.     

Repeatability of Male Mate Choice in the Mosquitofish, Gambusia holbrooki

Male mate choice has evolved in many species in which female fecundity increases with body size. In these species, males are thought to have been selected to favour mating with large females over smaller ones, thereby potentially increasing their reproductive success. While male mate choice is known to occur, it is less well studied than female mate choice and little is known about variation in mating preference among individual males. Here, we presented individual male eastern mosquitofish ( Gambusia holbrooki ) with paired females that differed in body size, and we quantified their mate preference on two consecutive days, allowing us to assess repeatability of preferences expressed. When males were allowed to view paired stimulus females, but not to acquire chemical or tactile cues from them, they exhibited a strong preference for large females over smaller ones. However, individual males were not consistent in the strength of their preference and repeatability was not significant. When individual males were allowed to fully interact with pairs of females, the males again exhibited a preference for large females over smaller ones, as revealed by a greater number of attempted copulations with large females than with smaller ones. In the latter social context, individual male preference was significantly repeatable. These results indicate that male eastern mosquitofish from our Florida study population possess, on average, a mating preference for larger females and that this preference is repeatable when males socially interact freely with females. The significant repeatability for mating preference, based on female body size, obtained for male mosquitofish in the current study is consistent with the presence of additive genetic variation for such preferences in our study population and thus with the opportunity for the further evolution of large body size in female mosquitofish through male mate choice.     

Mate Choice and Spawning Success in the Fighting Fish Betta splendens: the Importance of Body Size, Display Behavior and Nest Size

Courtship displays should be exaggerated enough to attract mates and yet tempered so as not to deter them. We tested this hypothesis in the fighting fish Betta splendens by studying courtship displays and body size and their relationships with male parental quality and female fecundity, as well as the effects of display behavior and body size on mate choice decisions and spawning success. Because of their high degree of parental investment, males are expected to be discriminating in their choice of mates. Males who displayed more frequently built larger nests, a measure of parental quality, but larger males did not. When females were paired with males with high display rates, however, the pair had fewer eggs in their nest, even when accounting for female body mass. In a mate choice test using computer‐generated male stimuli that differed only in display behavior, females showed no preferences for displaying males vs. non‐displaying males, or for males with higher display rates vs. lower display rates. In similar tests in which the computer‐generated males differed only in size, females preferred larger males, but also preferred males that differed with respect to body size (negative assortative mating). Males preferred computer‐generated females that performed courtship displays over non‐displaying females, but showed no preferences for female body size. Neither a female's body size nor her display behavior was a significant predictor of her fecundity as estimated by the number of eggs released during spawning. Thus, our results suggest that female B. splendens must balance male parental quality (nest size) with the risk of potentially disruptive or dangerous behavior during spawning, and that females may minimize these risks through negative size‐assortative mating. Female display behavior, while unrelated to fecundity in our study, may attract males because it indicates reproductive readiness or serves a species‐recognition function.     

The relative importance of orange spot coloration and total length of males in female guppy mate preference

Female mate choice by multiple male traits is an important current topic in animal behavior. However, the relative importance among the multiple cues in female choice is not explored in most cases. Female guppies Poecilia reticulata use both the color saturation of orange spots and the total length of males as mate choice criteria. In the present study, we used digitally modified video playbacks to examine the relative importance of these two male traits to female mate preferences. We initially examined the effective difference in the color saturation of orange spots as well as that in total length between two stimulus male images. Females only showed a strong preference for a bright male image (compared to the dull image) when the difference in color saturation was large (91% versus 25%). Conversely, females only exhibited a preference for larger size when they were presented a choice between two relatively small male images (total length 26.0 mm versus 23.0 mm). When two male images in which both the two traits were modified were presented to females, they prioritized male images possessing higher color saturation of orange spots, indicating the color saturation of male orange spots to be a more important factor than the total length in their mate choice. The color saturation of orange spots may convey more reliable information about the males to the females than their total lengths. These findings imply that females may rank multiple male criteria depending on relative benefits or costs derived from their mate choice based on each criterion.     

Mating preference of female zebrafish, Danio rerio, in relation to male dominance

Mating success tends to be skewed toward dominant males, thoughfemale mate preferences may not always correlate with male dominance.In this study, we investigated the mating preferences of femalezebrafish, Danio rerio, in the absence of male–male competition.We paired females sequentially with males of known dominancerank, using a nested, repeated measures design, with egg productionas a measure of female mate preference. We predicted that femaleswould spawn more frequently and produce larger clutches whenpaired with males of higher dominance rank. We found significantdifferences among females in the size of clutches produced andamong males in the size of clutches received, but these differenceswere independent of male dominance rank. Male body size wasnot related to either dominance rank or clutch size received.These results indicate that females vary clutch size in relationto the males with which they are paired but that they do notfavor dominant males. Thus, male competition may normally overridefemale mate preference in zebrafish.     

Male Mate Choice in the Guppy (Poecilia reticulata): Do Males Prefer Larger Females as Mates?

Although females are the choosier sex in most species, male mate choice is expected to occur under certain conditions. Theoretically, males should prefer larger females as mates in species where female fecundity increases with body size. However, any fecundity‐related benefits accruing to a male that has mated with a large female may be offset by an associated fitness cost of shared paternity if large females are more likely to be multiply mated than smaller females in nature. We tested the above hypothesis and assumption using the Trinidadian guppy (Poecilia reticulata) by behaviourally testing for male mate choice in the laboratory and by ascertaining (with the use of microsatellite DNA genotyping) patterns of male paternity in wild‐caught females. We observed significant positive relationships between female body length and fecundity (brood size) and between body length and level of multiple paternity in the broods of females collected in the Quaré River, Trinidad. In laboratory tests, a preference for the larger of two simultaneously‐presented virgin females was clearly expressed only when males were exposed to the full range of natural stimuli from the females, but not when they were limited to visual stimuli alone. However, as suggested by our multiple paternity data, males that choose to mate with large females may incur a larger potential cost of sperm competition and shared paternity compared with males that mate with smaller females on average. Our results thus suggest that male guppies originating from the Quaré River possess mating preferences for relatively large females, but that such preferences are expressed only when males can accurately assess the mating status of encountered females that differ in body size.     

Mate discrimination by females in the burying beetle Nicrophorus orbicollis: the influence of male size on attractiveness to females

1. Female burying beetles behave differently towards males of different sizes, avoiding mating with large males that are not defending resources but mating with small males regardless of the presence of resources. Females of the burying beetle Nicrophorus orbicollis were therefore examined to determine whether they discriminate among males using only pheromonal signals. The influence of female size on its own mate choice was also examined. 2. Females do use male pheromonal signals to discriminate among males and these signals do appear to convey information about male body size to females. Overall, females were more likely to be attracted to larger males than to smaller males. 3. Female choice of a male was influenced by both the female's own body size and the size of the female relative to the size of the two males available to it. 4. While there is an overall mating advantage for larger males, resulting from female preferences based on odour cues, smaller males are also attractive to some females under some circumstances. 5. It is argued that there are different costs and benefits of mating with different sized males, leading to the evolution of context‐dependent mate choice for females and the need to be able to discriminate males of different sizes from a distance.     

Dynamics of House Sparrow Biparental Care: What Contexts Trigger Partial Compensation?

Nest attendance during incubation is characterized by an inequitable division of labor in house sparrows, Passer domesticus, with females spending more time at the nest than males. Previous research has shown that if male contributions are reduced experimentally via testosterone (T) implants, females compensate partially for those reductions, consistent with predictions from most models of negotiated biparental care. In this study, we attempted to identify the cues and contexts generating partial compensation, using data from both unmanipulated parents and pairs with T‐males. Both males and females of this species sometimes leave the nest before their mate returns to relieve them, and we found that these unrelieved departures by unmanipulated individuals occur when partners are on lengthy recesses. Females compensated partially for long male recesses by marginally extending their bouts; most females also slightly reduced their next recess. By contrast, when males left before their mate returned, they left earlier than when they waited for the female. Neither males nor females adjusted their recess lengths after returning to the nest and discovering that their partner was absent. More pronounced changes in nest attendance of unmanipulated parents occurred in the context of ‘visits’, when individuals returned to the nest but then left without relieving their mate. Such visits effectively prolonged the bout of the on‐duty partner and extended the visitor’s recess. Analyses of behavior of T‐males and their mates revealed that T‐males had significantly longer recesses than control males, and that their mates, in turn, had elevated rates of unrelieved departures. T‐males also visited their on‐duty mates more often than control males, whereas female visits to T‐males were rare. Collectively, the predicted changes in female nest attendance associated with lengthy male recesses and male and female visits account reasonably well for the compensatory response of females paired to T‐males. The majority of female compensation was attributable to changes in visit behavior, however, suggesting that much of the negotiation over nest attendance in this species occurs during direct interactions between mates.     

Kin recognition and incest avoidance in a group-living insect

Mate choice theories predict that animals evolved strategiesto mate with optimally genetically dissimilar partners, providingfitness benefits. In group-living species, when adults do notdisperse, assessment of relatedness between conspecifics canbe a key factor for choosing mates. Here, we report for thefirst time, kin recognition abilities and their implicationin mate choice in the gregarious cockroach, Blattella germanica(L.). Binary choice tests showed that females mated preferentiallywith nonsibling rather than with sibling males, thus avoidingincest. In addition, inbreeding induced an important decreaseof their reproductive success. Contrary to what could be expectedwhen females had the choice between a nonsibling strain memberand a nonstrain member, they did not avoid mating with distantlyrelated nonstrain members, and extreme outbreeding induced anincrease of their reproductive success. Furthermore, our matechoice experiments disentangled the influences of familiarityfrom those of relatedness and evidenced that kin discriminationwas based on genetic cues independently of familiarity. Phenotypematching was a plausible mechanism for kin recognition. Contraryto many insect species, body size was not a salient criterionfor mate choice and had no consequences on reproductive success.     

Body Size and the Maintenance of Reproductive Isolation in Stickleback,Genus Gasterosteus

The blackspotted stickleback Gasterosteus wheatlandi and the widely studied threespine stickleback G. aculeatus are sympatric throughout the former’s range and share many aspects of life history and reproductive behaviour. These two species differ significantly in size, with G. wheatlandi of both sexes measured at approximately 60% of the standard length of their G. aculeatus counterparts. This study concentrated on G. wheatlandi courtship behaviour and investigated its role in the maintenance of reproductive isolation with G. aculeatus. Specifically, the roles that (1) female body size plays in influencing male courtship preferences and (2) male body size and behaviour play in female courtship preferences were investigated through dummy and live conspecific and heterospecific stimulus presentations. Male G. wheatlandi courtship preferences are consistent with previously described patterns for G. aculeatus. Males of both species preferentially approach and court the larger of two simultaneously presented live or dummy females. Thus, the smaller G. wheatlandi males are indiscriminate with respect to assortative mate choice; not only preferring to approach and court more fecund conspecific females but, more significantly, G. aculeatus‐sized females. In contrast, females of both species demonstrate strong assortative courtship preferences. When presented with pairs of flask‐enclosed males, females of both species preferentially orient and court the conspecific male over the heterospecific. Similarly, when presented with a conspecific male and a heterospecific male presented singly, females prefer to enter the nest of the conspecific. Systematic analysis of the interactions between these pairs of fish (one male, one female) demonstrates that the breakdown of courtship in heterospecific courtship occurs late in the courtship sequence when the widely differing forms of male leading behaviour results in drastically differing female responses. I suggest that, as previously described in G. aculeatus, the supernormality effect plays a significant role in mediating adaptive mate choice behaviour in G. wheatlandi. However, the added element of a larger sympatric species introduces a possible cost in time and energy devoted to courting heterospecific, and sympatric, females that the larger G. aculeatus do not likely incur. There is substantial evidence from many sympatric G. aculeatus species pairs that there is assortative mate choice based on size and/or courtship behaviour. Courtship trials suggest a more pervasive role for females in assortative mate choice. Whether it is male body size per se, or in combination with behaviour, morphology or other cues, is unresolved in the present study.     

Male mate choice in the Andrew’s toad <Emphasis Type="Italic">Bufo andrewsi</Emphasis>: a preference for larger females

In amphibians, theory predicts that male mate choice with respect to female body size can be expected to occur when female fecundity is related to body size and when the time and energy invested into one mating are relatively large. Based on experimental observations, we tested whether male mate choice occurs in a population of the Andrew’s toad (Bufo andrewsi), a species in which both assumptions are likely to be met. When a male B. andrewsi was placed with a gravid female and a non-gravid similar-sized female, the male did not discriminate between them. When two gravid females with distinct size differences were provided to a male, the male preferred the larger one. In an experiment in which two different-sized gravid females were put in two separate transparent cylinders to exclude potential chemical cues, males spent more time in proximity to the larger gravid females and jumped more frequently towards the larger gravid females than the smaller ones. These findings suggest that male B. andrewsi recognizes female body size, exhibits mate choice, and prefers to mate with larger females that provide greater reproductive potential.     

Is egg carrying attractive? Mate choice in the golden egg bug (Coreidae,Heteroptera)

In several species of fish, females select males that are already guarding eggs in their nests. It is a matter of debate as to whether a female selects a good nest site for her offspring (natural selection) or a male for his attractiveness (sexual selection). The golden egg bug, Phyllomorpha laciniata Vill, resembles fish in the sense that mating males carry more eggs than single males, but in the bugs, female mate choice is decoupled from egg site choice. The sexual selection hypothesis predicts that if females select males using male egg load as a cue for male quality, they should not mate with a male when eggs are removed, regardless of his mating attempts. When individual females were enclosed with an egg-loaded male and an unloaded male, they mated equally often with both males, although the loaded males courted more. In addition, when only successful males were used, females mated equally often with the loaded male and the unloaded male irrespective of sex ratio. Male choice rather than female choice affected mating frequency when sex ratio was equal. Therefore, females do not select the male by the eggs he carries, but successful males may receive many eggs due to egg dumping by alien females while they mate or as a consequence of mate guarding.     

Male body length influences mate-choice copying in the sailfin molly Poecilia latipinna

Females can choose a male independently of other females' matepreferences, or they can copy the mate choice of other females.Alternatively, mate-choice copying and independent mate choicecan interact if females assess male traits when deciding whetheror not to copy. We investigated how mate-choice copying interactswith a preference for large males in the sailfin molly (Potecilialatipinna). Sailfin molly females exhibited a preference forlarger males. They also copied the mate choice of other femaleswhen males were of similar body length. Females did not copy,however, when males differed substantially in body length. Ourresults show that conspecific mate copying occurs in the sailfinmolly but does not override a preference for larger males.     

Potential contribution of prenatal estrogens to the sexual differentiation of mate preferences in mice

The neural mechanisms controlling sexual behavior are sexually differentiated by perinatal actions of gonadal hormones. We recently observed using female mice deficient in alpha-fetoprotein (AFP-KO) and which lack the protective actions of AFP against maternal estrogens, that exposure to prenatal estrogens completely defeminized their potential to show lordosis behavior in adulthood. Therefore, we determined here whether mate preferences were also affected in female AFP-KO mice. We observed a robust preference for an estrous female over an intact male in female AFP-KO mice, which were ovariectomized in adulthood and subsequently treated with estradiol and progesterone, whereas similarly treated WT females preferred the intact male over the estrous female. Gonadally intact WT males preferred the estrous female over the male, but only when visual cues were blocked by placing stimulus animals behind opaque partitions. Furthermore, when given the choice between an intact male and a castrated male, WT females preferred the intact male, whereas AFP-KO females showed no preference. Finally when given the choice between an estrous female and an ovariectomized female, WT males preferred the estrous female whereas AFP-KO females preferred the ovariectomized female or showed no preference depending on whether they could see the stimulus animals or not. Taken together, when AFP-KO females are tested under estrous conditions, they do not show any male-directed preferences, indicating a reduced sexual motivation to seek out the male in these females. However, they do not completely resemble males in their mate preferences suggesting that the male-typical pattern of mate preferences is not solely organized by prenatal estrogens.     

Female Size Influences Mate Preferences of Male Guppies

Guppies (Poecilia reticulata) have a promiscuous mating system in which female choice for brightly coloured males plays an important role. Consequently, much research on guppies has examined how mate choice by females has lead to the evolution of male colour patterns. Much less attention has been devoted to mate choice by males in this species. In this study, we show that male guppies are choosy when selecting a female to associate with, significantly preferring the larger female when presented with two females that differed by ≥2 mm in standard length (SL). The strength of their preference for each female increased with absolute female size. The relative sizes of the females, however, also influenced male mating preferences: males showed stronger preferences for the larger female as the difference in SL between the two females increased. Such a preference for larger females is not unexpected as fecundity generally increases with body size in female fish. Thus, males choosing to mate with the larger female should have higher reproductive success. An apparent, but non‐significant anomaly, whereby males appear to prefer the smaller of the two females when the difference between female SL was <4 mm, deserves further investigation.     

Environmental change mediates mate choice for an extended phenotype,but not for mate quality

Sexual cues, including extended phenotypes, are expected to be reliable indicators of male genetic quality and/or provide information on parental quality. However, the reliability of these cues may be dependent on stability of the environment, with heterogeneity affecting how selection acts on such traits. Here, we test how environmental change mediates mate choice for multiple sexual traits, including an extended phenotype–‐the structure of male‐built nests – in stickleback fish. First, we manipulated the dissolved oxygen (DO) content of water to create high or low DO environments in which male fish built nests. Then we recorded the mate choice of females encountering these males (and their nests), under either the same or reversed DO conditions. Males in high DO environments built more compact nests than those in low DO conditions and males adjusted their nest structure in response to changing conditions. Female mate choice for extended phenotype (male nests) was environmentally dependent (females chose more compact nests in high DO conditions), while female choice for male phenotype was not (females chose large, vigorous males regardless of DO level). Examining mate choice in this dynamic context suggests that females evaluate the reliability of multiple sexual cues, taking into account environmental heterogeneity.     

Increased performance of offspring sired by parasitic males in bluegill sunfish

Males of many species are characterized by alternative matingtactics. In bluegill sunfish (Lepomis macrochirus), some malesdelay maturation and become "parentals" while other males matureprecociously and become "cuckolders." Parentals use an overt,territorial mating tactic, defending a nest and courting females.Cuckolders instead use a sneaking tactic to parasitize parentals.It has previously been shown that parentals that are heavilycuckolded provide less care to their young, yet females do notappear to discriminate against cuckolders, and they may actuallyrelease more eggs when a cuckolder is present than when spawningonly with a parental. Here I examined growth rate of fry ofknown paternity through the yolk-sac stage of development usingcomplementary laboratory and field studies to assess a potentialindirect benefit for females that mate with cuckolders. Comparisonof maternal half-siblings sired in vitro shows that cuckolderoffspring grow faster and to a larger size than parental offspringwhile feeding endogenously on their yolk sac. Because both foodresource and maternal genes are equivalent across treatments,these data indicate a genetic difference in growth between thetwo male life histories. In the field, fry from nests that haveproportionately more cuckolder offspring are larger when theyemerge from the nest. This increased size can lead to threefoldhigher survivorship for cuckolder offspring than parental offspringfrom Hydra canadensis predation, a major predator of bluegillfry. These results are discussed in the context of mate choicefor direct and indirect benefits and in the context of the evolutionof alternative mating tactics.     

Mate preferences in female canaries (Serinus canaria) within a breeding season

Divorce and remating in birds can be described as strategies used to enhance reproductive success. Mate switching often occurs because pairs failed to brood at least one chick during the previous breeding season. In the present study, we evaluated the influence of reproductive success on female preferences in domesticated canaries (Serinus canaria). For that purpose, females previously paired and having reared young were placed in a choice test situation: They were allowed to choose between their previous mate and a familiar male (a male neighbor during the breeding period). During these choice tests, females tended to stay near their previous mate longer than near a male neighbor when their reproductive success was “good” (at least two chicks). On the other hand, females with “poor” reproductive success (one chick) did not show a preference for their previous mate. Furthermore, in the present study, we observed that during choice tests males reacted to the presence of their previous mate in a particular way, by gathering nest material. This behavior was more scarcely observed in neighbor males which, on the contrary, sang significantly more than previous mates did.     

Variation between and within the sexes in body size preferences

Although it is often assumed that males and females have mating preferences for larger individuals of the other sex, potential underlying differences between male and female preferences for body size are not commonly investigated. Here, sexual differences in body size preferences are examined in the poeciliid fish, Brachyrhaphis rhabdophora. Females preferred larger males to smaller males, but preference did not appear to be affected by female size. One population-level analysis for males did not indicate an overall preference for larger females. A closer examination, however, revealed an effect of male size on preference; larger males preferred larger females, while smaller males preferred smaller females. It appears then that females, regardless of size, share a preference for large males, but males differ in their behaviour, depending on their body size. In addition, while the degree of difference in size between paired females did not appear to affect male preference, the degree of difference in size between paired males strongly affected female preference; the greater the difference, the more strongly females preferred the larger male. Thus, intersexual selection is found to operate in both sexes, but how it operates appears to differ. Intrasexual and intersexual differences in mating behaviour may be missed when evaluating population-wide preferences. That is, there can be underlying differences in how the sexes respond and the consequences of such differences should be considered when investigating mate choice. The results are considered in terms of the evolution of mating preferences, alternative mating strategies, assortative mating, the maintenance of trait variation in a population, and current methods to evaluate mating preferences.     

Effects of breeding site density on competition and sexual selection in the European lobster

The availability of breeding sites has been predicted to affectthe intensity of sexual selection, including mate competition,mate choice and ultimately, variation in mating success. Wetested the hypothesis that reduced density of shelters wouldcause an increase in the intensity of sexual selection in Europeanlobsters, Homarus gammarus. However, we found little supportfor our predictions. For example, within-sex competition bymales and by females was not more intense when shelters werescarce. Indeed, females attempted to evict one another fromshelters significantly more often when shelters were common.When shelters were abundant, shelter-holding males had greatermating success than males without shelters, yet females didnot show more interest towards these males during courtshipencounters. Mate attraction was more strongly related to largemale body size when shelters were scarce. Overall, the resultssuggest that reduced shelter density does not lead to more overtwithin-sex aggression in this species. Instead, we suggest thatimpacts of breeding resource availability on sexual selectionmay depend on the range over which resources are measured, withextreme scarcity of shelters rendering overt competition uneconomical.Furthermore, females may become more selective of male traitssuch as large size, which enhance male control of breeding sitesand hence protection of females.