Assessing mate size in the red swamp crayfish Procambarus clarkii: effects of visual versus chemical stimuli

1. Mate assessment requires an efficient system of information exchange between the sexes and often relies on several forms of communication. Both sexes of the crayfish, Procambarus clarkii , select a mate on the basis of its body size, independently of chelar size/symmetry and dominance status, large size being a proxy for a high quality partner. Here, we explored the effects that visual and chemical stimuli, emitted alone or in combination, exert on the assessment of size of a potential mate.
2. We followed a binary choice test paradigm, in which two male or female 'targets', of either large or small size, were simultaneously presented to a female or a male 'chooser'. We recorded the first target visited, the total duration of the visits per target, and the behaviour of the chooser when approaching the target area.
3. Our results show that females require a combination of visual and chemical stimuli to select the larger male. A more complex pattern was found when males were the choosers. While visual and chemical cues together rendered them willing to mate with the larger female, their initial choice and the total time spent near her depended only on smell. In mate assessment by P. clarkii , visual and chemical information seem to act as 'non-redundant signals'.     

Evidence of female cryptic choice in crayfish

To test whether male body size affects female reproductive investment in the polygamous crayfish Procambarus clarkii, we described mating behaviour of virgin females paired with either small or large males, and analysed the number, size and weight of both eggs and juveniles sired by either types of male. Along with confirming the overt selection by females of larger mates, we found that the size and weight of both the eggs and the juveniles were higher when sired by larger fathers. This suggests that P. clarkii females exert a form of cryptic choice for large males, seemingly adjusting the quantity of egg deutoplasm in function of the mate body size. The question of why females spend time and energy to brood low-fitness offspring is finally raised.     

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.     

Seasonal Variation in Mate Choice of Photinus ignitus Fireflies

Mate choice by either sex may vary with changes in the associated costs and benefits, determined by factors such as the availability of potential mates and variation in mate quality. We examined seasonal variation in operational sex ratio, courtship behavior, spermatophore mass, egg count, and the relationship between morphological traits and mating success in Photinus ignitus fireflies to determine if mate choice in either sex varied with the availability and relative reproductive investment of fertilizable females and sexually active males. Successfully mating males had larger lanterns than unsuccessful males when the operational sex ratio was male‐biased. In addition, female responsiveness to male signals increased as the number of courting males decreased, and male spermatophore mass decreased with body size across the mating season. Successfully mating females had larger body mass than unsuccessful females. Female body mass predicted egg count and female rejection by males increased as the season progressed and female size decreased. These results suggest that both male and female P. ignitus exhibit mate choice, and that such choice is influenced by seasonal variation in the abundance and quality of potential mates.     

The large-male advantage in brown antechinuses: female choice, male dominance, and delayed male death

Male-biased dimorphism in body size is usually attributed tosexual selection acting on males, through either male competitionor female choice. Brown antechinuses (Antechinus stuartii) aresexually dimorphic in size, and heavier males are known to siremore offspring in the wild. We investigated four possible mechanismsthat might explain this large-male reproductive advantage. Wetested if there is a female preference for large males, a femalepreference for dominant males, if larger males compete moreeffectively for mates, and if there is a survival advantagefor large males during the mating season. We established nestinggroups of males in captivity and conducted mate choice trialsin which males from nesting groups either could or could notinteract. We assessed male dominance rank and recorded survivaltimes after mating. Females did not prefer larger males directly.The results suggest that the other three mechanisms of sexualselection tested account for the large-male advantage: largemales competed more successfully for mates, so were sociallydominant; females rejected subordinates (males they saw losingtwice in contests to previous mates); and dominant males survivedfor longer after their first mating. Females judged male rankbased on direct observation of male competitive interactionsat the time of mating and apparently could not distinguish rankfrom male scent. Effects of size and dominance on male reproductivesuccess are not confounded by age because male antechinusesare semelparous.     

Male and Female Mate-Choice Decisions by Evarcha culicivora, An East African Jumping Spider

There has been a considerable recent interest in the criteria by which animals choose mates and in the extent to which mating systems tend to be based on mutual mate choice. In this study, we consider Evarcha culicivora, a salticid spider from East Africa. This species has some unusual characteristics, including active display by females as well as males, males that kill females more frequently than females kill males and wide intrasexual variation in body size. For females, larger males are especially dangerous. Here, we demonstrate, using two experimental designs (live‐mate choice and mount choice), that virgin males, virgin females and previously mated males prefer larger opposite‐sex individuals as potential mates, but mated females prefer smaller, safer males as potential mates.     

The effects of body size on mate choice in a grapsid crab,Gaetice depressus (Crustacea,Decapoda)

Male and female mate choices were investigated in the grapsid crab,Gaetice depressus (Crustacea, Decapoda) in a laboratory experiment. Males mated indiscriminately with regard to the body size of the females, and frequently copulated with the first females they encountered. In contrast, females showed mate discrimination with regard to the body size of males. The females tended to sample potential mates prior to copulation, and showed both a preference for the larger males and a tendency toward the rejection of males with body sizes smaller than their own. However, they did not discriminate between two males that were either larger or smaller than they themselves. Mate choice by the females of this species is though to be based upon threshold-criterion tactics, in which the body size of the female itself is used as a threshold value.     

Male and Female Soldier Beetles Relax Choice for Mate Quality Across Daily Courtship Periods

Mutual mate choice occurs when members of each sex will reject some potential mates in efforts to encounter better prospects later. The decision to reject may represent the interaction between mate preferences, mate availability, and temporal constraints. Theory predicts that mutual mate choice will favor relaxed choosiness as mate availability and time for courtship decline. We explored mutual mate choice in the soldier beetle, Chauliognathus pennsylvanicus (Cantharidae), where courtship consisted of males attempting to secure evasive females. We employed field observations, laboratory experiments, and stochastic simulations to investigate the decline in choosiness over the daily courtship period, during which individuals can mate at most once. We found that reproductive success of males and females increases with mate size and mating frequency. Females biased copulations toward larger mates by attempting to evade suitors, while males biased copulations by releasing the smaller females they capture. However, late in the day males and females may increase reproductive success by accepting rather than rejecting lower quality mates to maintain high mating rates. Stochastic simulations indicated that reproductive success, the product of mating frequency and mean mate size, was maximized in males and females by incrementally reducing mate standards across daily courtship periods. In the field, large males who rejected small females early in the daily courtship period rarely did so later. Large females used less effective evasive maneuvers later in the courtship period, resulting in copulations with progressively smaller males. These results support models of mutual mate choice that predict that individuals of high quality will maximize reproductive success by relaxing choosiness as the courtship period wanes.     

Effects of female and male size on female mating and remating decisions in a bean beetle

The body sizes of individuals of the choosing and chosen sexes in a mate choice may affect sequential mating of females. We examined the effects of the body sizes of females and their mates on attributes of female first mating, and the effects of body sizes of females and their previous and potential future mates on female remating in the adzuki bean beetle, Callosobruchus chinensis. Large- and small-sized adults were derived from larvae reared under conditions of low and high density in a bean, respectively. The speed of first mating of large females was not affected by the size of courting males, whereas small females initiated mating more rapidly when courted by small males. The remating probability of large females was not affected by first male size, whereas small females that mated first with smaller males were more likely to remate. These data suggest that pre- and post-copulatory female choices for male size depend on the female’s size, and the small females might be more willing to copulate with smaller males but prefer larger males to sire their offspring after copulation. A possible explanation for this preference is that small females may suffer greater harm from copulating with larger males.     

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.     

Mating and remating of least auklets (Aethia pusilla) relative to ornamental traits

We investigated mating patterns of the least auklet, a smallmonogamous seabird, at St. Paul Island, Alaska, during threebreeding seasons. Least auklets mated assortatively with respectto both plumage color, a trait important in status signaling,and tarsus length, an index of body size. Least auklets mateddisassortatively with respect to the extent of facial plumes,but neither assortatively nor disassortatively for any otherornamental trait (bill color, bill ornament size). Mate fidelitywas lower in least auklets than in some long-lived seabird species;when both members survived to a following year, only about two-thirdsof pairs reunited. Nearly half of the auklets paired in 1 yearobtained a new mate in the following year, either because ofmate disappearance or divorce. Interyear fidelity to mates wasrelated only to male ornamentation; males with larger facialplumes were more likely to reunite with their mates the nextyear than males with smaller plumes. There were no significantdifferences in the ornaments of females in reunited and divorcedpairs. Pairs that reunited also had significantly lighter plumagethan pairs that divorced, and the plumage of males reunitingwith their mates was significantly paler than that of divorcedmales. We conclude that the probability of both divorce andremating in this species is influenced by ornamental traits.Our finding that remating was related to male plumage colorand ornaments is consistent with the idea that remating is influencedby female choice. Pairs that reunited also bred earlier in theseason and had higher reproductive success than pairs with experiencedindividuals breeding together for the first time. We also foundevidence that failure to breed in a given year increased theprobability of subsequent divorce.     

Pedigrees, assortative mating and speciation in Darwin's finches

Pedigree analysis is a useful tool in the study of speciation. It can reveal trans-generational influences on the choice of mates. We examined mating patterns in a population of Darwin's medium ground finches (Geospiza fortis) on Daphne Major Island to improve our understanding of how a barrier to the exchange of genes between populations arises in evolution. Body sizes of mates were weakly correlated. In one year, the smallest females were paired non-randomly with the males of similar size, and in another year the largest males were paired with the largest females. An influence of parental morphology on the choice of mates, as expected from sexual imprinting theory, was found; the body size of mates was predicted by the body sizes of both parents, and especially strongly by the father's. These associations imply that the seeds of reproductive isolation between species are present within a single variable population. The implication was subject to a natural test: two exceptionally large birds of the study species, apparently immigrants, bred with each other, as did their offspring, and not with the members of the resident population. The intense inbreeding represents incipient speciation. It parallels a similar phenomenon when another species, the large ground finch, immigrated to Daphne and established a new population without interbreeding with the resident medium ground finches.     

Relatedness,body size and paternity in the alpine newt,Triturus alpestris

Sexual selection has traditionally been investigated assuming that male quality is as skewed as patterns of male reproductive success can sometimes be. Recently, female choice has been investigated under the model of genetic compatibility, which assumes that each individual female has her own 'best' mate and there is no overall optimal choice for all females. We investigated female mate choice in the newt species Triturus alpestris, a member of a genus where female choice has been investigated only within the context of the optimal male (female choice for condition-dependent traits). We provided females with two males that differed in one condition-dependent trait (body size) and overall genetic composition. Both male body size and female body size did not influence paternity, but the degree of genetic relatedness between females and potential mates did. Two components of fitness (fecundity and hatching success) did not differ between singly and multiply sired clutches, indicating that females do not employ polyandry as a means of increasing offspring fitness through genetic bet-hedging. Instead, we hypothesize that females may mate initially for fertility assurance, but prefer less-related males as the most genetically compatible mates.     

The sex lives of parasites: Investigating the mating system and mechanisms of sexual selection of the human pathogen Schistosoma mansoni

The mating systems of internal parasites are inherently difficult to investigate although they have important implications for the evolutionary biology of the species, disease epidemiology, and are important considerations for control measures. Using parentage analyses, three topics concerning the mating biology of Schistosoma mansoni were investigated: the number of mates per adult male and female, variance in reproductive success among individuals, and the potential role for sexual selection on male body size and also mate choice for genetically dissimilar individuals. Results indicated that schistosomes were mostly monogamous, and evidence of only one mate change occurred over a period of 5-6 weeks. One male was polygynous and contained two females in its gynecophoral canal although offspring were only detected for one of the females. Even though they were primarily monogamous and the sex ratio near even, reproductive success was highly variable, indicating a potential role for sexual selection. Male body size was positively related to reproductive success, consistent with sexual selection via male-male competition and female choice for large males. However, relatedness of pairs was not associated with their reproductive success. Finally, genetically identical individuals differed significantly in their reproductive output and identical males in their body size, indicating important partner and environmental effects on these traits.     

Sexual selection for male dominance reduces opportunities for female mate choice in the European bitterling (Rhodeus sericeus)

Sexual selection involves two main mechanisms: intrasexual competition for mates and intersexual mate choice. We experimentally separated intrasexual (male-male interference competition) and intersexual (female choice) components of sexual selection in a freshwater fish, the European bitterling (Rhodeus sericeus). We compared the roles of multiple morphological and behavioural traits in male success in both components of sexual competition, and their relation to male reproductive success, measured as paternity of offspring. Body size was important for both female choice and male-male competition, though females also preferred males that courted more vigorously. However, dominant males often monopolized females regardless of female preference. Subordinate males were not excluded from reproduction and sired some offspring, possibly through sneaked ejaculations. Male dominance and a greater intensity of carotenoid-based red colouration in their iris were the best predictors of male reproductive success. The extent of red iris colouration and parasite load did not have significant effects on female choice, male dominance or male reproductive success. No effect of parasite load on the expression of red eye colouration was detected, though this may have been due to low parasite prevalence in males overall. In conclusion, we showed that even though larger body size was favoured in both intersexual and intrasexual selection, male-male interference competition reduced opportunities for female choice. Females, despite being choosy, had limited control over the paternity of their offspring. Our study highlights the need for reliable measures of male reproductive success in studies of sexual selection.     

Calling,Courtship, and Condition in the Fall Field Cricket,Gryllus pennsylvanicus

Theoretically, sexual signals should provide honest information about mating benefits and many sexually reproducing species use honest signals when signalling to potential mates. Male crickets produce two types of acoustic mating signals: a long-distance mate attraction call and a short-range courtship call. We tested whether wild-caught fall field cricket (Gryllus pennsylvanicus) males in high condition (high residual mass or large body size) produce higher effort calls (in support of the honest signalling hypothesis). We also tested an alternative hypothesis, whether low condition males produce higher effort calls (in support of the terminal investment hypothesis). Several components of long-distance mate attraction calls honestly reflected male body size, with larger males producing louder mate attraction calls at lower carrier frequencies. Long-distance mate attraction chirp rate dishonestly signalled body size, with small males producing faster chirp rates. Short-range courtship calls dishonestly reflected male residual mass, as chirp rate and pulse rate were best explained by a curvilinear function of residual mass. By producing long-distance mate attraction calls and courtship calls with similar or higher effort compared to high condition males, low condition males (low residual mass or small body size) may increase their effort in current reproductive success at the expense of their future reproductive success, suggesting that not all sexual signals are honest.     

Sex recognition and mate choice by male Bufo gargarizans in central China

Mate choice is important for successful reproduction, and consequently species have evolved various ways to choose potential high-quality mates. Anuran mate choice and underlying processes have been the subject of several recent investigations, however we are far from a complete understanding of mate choice in this system. In the present study, when given a simultaneous choice between a male and a female of identical size, males did not discriminate between the sexes, and attempted to clasp a male or a female with equal frequency. Test males only released the stimulus toad when a release call was emitted by the stimulus male. When two males with distinct size differences were provided with a male, the male chose the larger one. Moreover, males discriminated between gravid females that differed in body size, choosing larger gravid females over smaller ones. These results suggest that male Bufo gargarizans can discriminate between the sexes, probably based on male release calls, and prefer to mate with larger individual using visual cues.     

Evidence for adaptive male mate choice in the fruit fly Drosophila melanogaster

Theory predicts that males will benefit when they bias their mating effort towards females of higher reproductive potential, and that this discrimination will increase as males become more resource limited. We conducted a series of experiments to test these predictions in a laboratory population of the fruitfly, Drosophila melanogaster. In this species, courtship and copulation have significant costs to males, and females vary greatly in fecundity, which is positively associated with body size. When given a simultaneous choice between small and large virgin females, males preferentially mated with larger, more fecund, females. Moreover, after males had recently mated they showed a stronger preference for larger females. These results suggest that male D. melanogaster adaptively allocate their mating effort in response to variation in female quality and provide some of the first support for the theoretical prediction that male stringency in mate choice increases as resources become more limiting.     

Is Self‐Grooming by Male Prairie Voles a Predictor of Mate Choice?

Self-grooming by mammals is a form of scent dissemination in which individuals anoint themselves with salivary, anogenital, and other body odours. Self-grooming has been proposed to be a sexually selected trait favoured in reproductive competition and sexual attraction. We tested the hypothesis that females would show a mating preference for males that self-groomed more than a reproductive competitor that groomed less. In mate-choice experiments in which females had a choice of two tethered males, non pair-bonded females did not choose males based on their frequency of self-grooming. In a second experiment in which pair-bonded females in postpartum oestrus had access to their current mate and two strange males, strange males groomed significantly more than pair-bonded mates, yet attained the fewest copulations. Non pair-bonded females and pair-bonded males and females groomed significantly less often than did non pair-bonded males. Self-grooming behaviour was consistent with the sexual attraction hypothesis, but the frequency of self-grooming did not increase a male's mating success. We conclude that the frequency and time spent self grooming are not good predictors of mating success.     

Reproductive effort of both male and female Bar‐throated Apalis Apalis thoracica is predicted by ornamentation of self and mate

Melanin‐based plumage ornaments have been shown to play an important role in male–male competition, but also influence inter‐sexual communication. Consequently, ornaments may be associated with reproductive effort of both males and females. Females mated to males with larger melanin ornaments may acquire access to better territories or benefit from increased paternal care. Here we investigated whether the melanin‐based breast‐band of male and female Bar‐throated Apalis Apalis thoracica is a signal of information about its bearer and is associated with male and female reproductive effort. Breast‐band size was a highly variable morphometric trait in both sexes, but only in males was it associated with body mass. We then assessed whether male and female breast‐band size predicted maternal and paternal investment. Egg mass increased with male breast‐band size, but decreased with female breast‐band size. Whether females adjust maternal hormone allocation in response to their partner's ornamentation remains a contentious issue. We found that yolk testosterone and androstenedione concentrations were not predicted by male ornamentation or body mass. Finally, males with larger breast‐bands provided their mates with more food, allowing those females to spend more time incubating. Reproductive effort of both parents is therefore predicted by their own and their mate's ornamentation in Bar‐throated Apalis, and thus breast‐band size potentially acts as a signal of reproductive performance in both sexes. These results highlight the need for more comprehensive analyses of a relationship between melanin‐based ornaments and fitness, incorporating multiple behavioural variables associated with reproductive effort.