Host plant density and territorial behavior of the seed bug,Neacoryphus bicrucis (Hemiptera: Lygaeidae)

Summary Adults of the seed bug, Neacoryphus bicrucis, are most numerous in large host plant patches and where host plants are most dense within patches. Males patrol small territories composed of host plants and expel other males and unreceptive females. Larger males aggressively exclude smaller males from high quality territories where both host plants and females are most dense. Consequently, large males mate more frequently than small males. Males remain for longer time in areas where host plants are more dense. However, mating experience influences tenure within a host plant patch such that males tend to remain where they have recently mated. Courtship is aggressive and male territorial behavior is similar to courtship behavior. Preference for territories in areas of high host plant density appears to have been selected to increase the rate of encounter with females which preferentially oviposit in areas of high host plant density.     

Sex ratios,mating behavior and sexual size dimorphism of the northern water snake,Nerodia sipedon

Competition among males to mate is generally associated with male-biased size dimorphism. In this study we examine mating behavior in the northern water snake (Nerodia sipedon), a species in which males are much smaller than females despite substantial competition among males to mate. Competition among males was a consequence of a male-biased operational sex ratio due to slightly higher female mortality from a birth sex ratio of 1 : 1, and, in 1 year, more synchronous and longer mating activity by males. Approximately one-third of both males and females appeared not to mate in a given year. Larger males were generally more likely to attempt mating, but size did not explain the variance in the number of aggregations in which individual males participated. Within aggregations, males that were successful at achieving intromission were larger than unsuccessful males in 1 of 2 years. Variation in condition (mass relative to length) and relative tail length were not generally useful predictors of either mating effort or success in males. Because large size was often advantageous to males, sexual size dimorphism appeared not to be a consequence of sexual selection favoring smaller males. Because sexual dimorphism was evident at birth, and both males and females matured sexually at about 4 years, sexual dimorphism was not simply a consequence of one sex growing at the maximum rate for longer. Female fecundity increased with size, and sex differences in size-fecundity relations may underly the pattern of sexual size dimorphism. However, because multiple mating by females is common, sperm competition is likely to be important in determining male reproductive success. Therefore, allocation of energy to sperm rather than growth may also prove to be an important influence on male growth rates and sexual size dimorphism.     

Reproductive success of male bank voles (<Emphasis Type="Italic">Clethrionomys glareolus</Emphasis>): the effect of operational sex ratio and body size

The operational sex ratio (OSR) may influence the intensity of competition for mates and mate choice and is therefore thought to be a major factor predicting the intensity and direction of sexual selection. We studied the opportunity for sexual selection, i.e., the variance in male reproductive success and the direction and intensity of sexual selection on male body mass in bank vole (Clethrionomys glareolus) enclosure populations with experimentally manipulated sex ratios. The opportunity for sexual selection was high among male-biased OSRs and decreased towards female-biased OSRs. Paradoxically, selection for large male body mass was strongest in female-biased OSRs and also considerable at intermediate OSRs, whereas at male-biased OSRs, only a weak relationship between male size and reproductive success was found. Litters in male-biased OSRs were more likely to be sired by multiple males than litters in female-biased OSRs. Our results suggest that the intensity and direction of sexual selection in males differs among different OSRs. Although the direction of sexual selection on male body mass was opposite than predicted, large body mass can be favored by sexual selection. Naturally varying OSRs may therefore contribute to maintain variation in male sexually selected traits.     

Mate choice,fecundity and sexual dimorphism in two pipefish species (Syngnathidae)

Summary In order to understand the causes of sexual dimorphism, mate choice and size-related fecundity were studied in two pipefish species, Syngnathus typhle and Nerophis ophidion. Sexual dimorphism is more pronounced in N. ophidion; females are larger, have sexual colourings, and are more active during courtship. In S. typhle the sexes are alike in all these respects. Males brood their offspring in both species. In N. ophidion fecundity was positively correlated with both body size and the amount of sexual colouring in females. In males no correlation between body size and fecundity, or between body size and embryo size existed. Predictably, in mate choice experiments with equal-sized females, males chose females with more extensive sexual colourings. We explain sexual dimorphism in this species as a consequence of both natural selection (fecundity increases with size in females but not in males) and sexual selection (males prefer larger females). We argue that sexual size dimorphism did not evolve by selection minimizing overlap in food niches between the sexes, because food production is high in the Zostera beds where the fishes live, and no size dimorphism was found in the sympatrically occurring S. typhle. Furthermore, in N. ophidion dimorphism is not greater in a particular mouth character than in overall body size. In S. typhle egg size and the average number of eggs transferred per spawning were positively correlated with female body size. Apparently more energy per offspring was provided by larger males than by smaller males, and larger males also carried more offspring. As predicted, large mates were preferred by both sexes in mate choice experiments. This is explicable in terms of both natural selection (fecundity increases with size in both sexes) and sexual selection (both sexes prefer large mates). As a consequence of selection acting in the same direction in both sexes, sexual dimorphism is absent in S. typhle.     

Female siskins choose mates by the size of the yellow wing stripe

Commonly, female birds use the brightly coloured patches on males to choose the best-quality mates. Coloured wing patches, however, have received little attention or have been previously related to social behaviour (as a signal to recruit conspecific individuals at feeding patches) or foraging (to flush prey) contexts, rather than to sexual selection. Here we provide evidence that in siskins (Carduelis spinus), wing patches function in mate choice. Mate-choice experiments showed that females were attracted by the size of the yellow wing stripe of the male, but not by the size of its black bib, body size, general plumage brightness or age. Experiments on birds with manipulated yellow wing stripes showed that females were sensitive to the size of this colour patch, irrespective of other male qualities. The preference of female siskins for males with larger wing patches when searching for a mate may be explained by the relationship of this trait to foraging ability, which would ensure females good parental investment from the chosen male.Communicated by W.A. Searcy     

Dispersal: an alternative mating tactic conditional on sex ratio and body size

Summary Small male milkweed beetles are less successful at obtaining mates than are larger males. Larger males usually win fights and prevent smaller males from obtaining mates and from choosing larger more fecund females as mates. When sex ratios are male-biased, smaller males are particularly likely to experience these mating disadvantages. It follows that smaller males should be especially responsive to their local competitive environment and behave so as to minimize the mating disadvantages of their smaller size. This paper tests the hypothesis that smaller males disperse from host plant patches with male-biased sex ratios and remain in patches with female-biased sex ratios more readily than larger males.Results show both larger and smaller males disperse from patches with male-biased sex ratios more frequently than from patches with femalebiased sex ratios. As predicted, however, small males are more likely to disperse from patches with male-biased sex ratios and remain in patches with female-biased sex ratios than are larger males.The data also show that smaller males dispersing from patches with male-biased sex ratios obtain more matings than non-dispersing males.For milkweed beetles, moving between patches can be viewed as an alternative mating tactic conditional on male body size and local sex ratio.     

The effect of size and cheliped autotomy on sexual competition between males of the mud crab <Emphasis Type="Italic">Cyrtograpsus angulatus</Emphasis> Dana

Size advantage in male–male competition over mates, combined with male preference over large females, is a common feature that can drive to size assortative mating and, eventually, sexual selection. In crabs, appendage autotomy can affect assortative mating and opportunity for sexual selection by affecting size advantage in mating contests. In this work, we evaluate the effect of size and appendage autotomy in generating assortative mating in the mud crab Cyrtograpsus angulatus. Field observations of guarding pairs in two different populations show a positive correlation between carapace width of males and females in both the populations. In one of the populations, incidence of appendage autotomy was low and the variability in the size of reproductive males was lower than the variability in the size of randomly collected males (i.e. only larger males were successful in getting a female), whereas there was no differences in the other population (i.e. most male sizes were successful) where the incidence of appendage autotomy was very high, indicating that the importance of size is higher when the incidence of autotomy is low. In this context, experiments (in both populations) show that, in contests for a female, larger males outcompete smaller ones only when they had intact appendages. When males had missing chelipeds, winning or loosing against smaller males was random. This may lead to a decrease in the importance of male size in populations with high incidence of cheliped autotomy, affecting assortative mating and opportunity for selection and, thus, affecting selective pressures.     

Population structure and sex-change in the coral-inhabiting snailCoralliophila violacea at Hsiao-Liuchiu,Taiwan

Surveys of the coral-inhabiting snailCoralliophila violacea (Lamarck) (=C. neritoidea Kiener) were made on shallow fringing reefs (<8 m deep) around Hsiao-Liuchiu, Taiwan, between July and October 1990. The snails were aggregated into patches on the surface of massive poritid coral colonies. Coral colonies >40 cm in diameter were more likely to bear patches of snails than smaller colonies, and also to have more snails. The coralliophilids ranged from 5 to 30 mm in aperture length. The sex ratio of the population was biased toward males (539:279), with only a few small individuals of indistinguishable sex. Snails between 6 and 10 mm were all males, while most snails with aperture lengths 20 mm were females. Judging from the distinct size ranges of males and females within patches and from the observed degeneration of the penis, the snails may have changed sex from male to female with increasing size. Sex-change may occur across a wide size range (10 to 20 mm). The correlation of smallest female size and largest male size among patches indicates that snail size at sex-change is peculiar to each individual patch. Those females in patches with a single female (but many males) were significantly smaller than females in multiple-female patches. It is likely that in the absence of females males change sex at a smaller size, whereas in the presence of large females males delay sexchange until they have reached a larger size. The plasticity of size at sex-change may be adaptive and a result of natural selection at the individual level.     

Mate replacement in experimentally widowed collared flycatchers (Ficedula albicollis): determinants and outcomes

There is increasing evidence that sexual selection may be intense even in socially monogamous birds, resulting from both mate choice and sperm competition. We studied these two modes of sexual selection experimentally by removing paired male collared flycatchers, Ficedula albicollis, from their mates for 2 days and investigating the factors that influenced the likelihood of a replacement male appearing and how the removals influenced paternity. Replacement males (usually neighbouring males) appeared at 81% (n = 37) of nests where males were removed. The likelihood of this appearance was unaffected by the probable reproductive value of the female's clutch to the replacing male. A replacement was, however, less likely when the original male had a large forehead patch, a trait previously shown to be subject to sexual selection in this population. Experimental removal of males increased the level of sperm competition: 74% of experimental broods were multiply sired, compared to 29% of unmanipulated broods in a previous study. Only two factors predicted how paternity was shared between males: removed males fathered more young if removed closer to laying, and if they had larger forehead patches. The former result is consistent with last-male sperm precedence determining paternity, whereas the latter adds to other evidence that forehead patch size is the target of female preference in this species. Our results suggest that females exert some control over male replacement, and also that they may influence the fertilisation success of males by behavioural means. Received: 15 July 1998 / Received in revised form: 16 March 1999 / Accepted: 28 March 1999     

Male lagoon gobies, <Emphasis Type="Italic">Knipowitschia panizzae</Emphasis>, prefer more ornamented to larger females

Female ornamentation may be directly sexually selected, by male choice or female competition, or occurs as the result of a genetic correlation, arising from sexual selection on males. However, increasing evidence supports the former hypothesis, suggesting that males actively choose their partner preferring traits indicative of female quality. In the lagoon goby, Knipowitschia panizzae, a polygynous species whose males perform parental care to eggs, body length and the size of a sex-specific yellow patch on the belly are known to be reliable indicators of female fecundity. In this paper, we tested, using dummies, the male’s mating preferences for female body and yellow belly patch sizes. The two experimental trials in which a single female trait was variable showed that males prefer a larger belly patch and a larger body size, indicating that both these characters are selected by male mate choice. However, when faced with dummies exhibiting an inverse combination of body and belly patch sizes (experiment 3), males significantly preferred the smaller ones with larger yellow belly patches. A calculation of dummy theoretical fecundity reveals that in the first two experiments, males would have received an immediate benefit from their choice in terms of egg number, whereas in the third one, males chose partners that would have provided them with fewer eggs. The male lagoon goby preference for females with larger belly patches, regardless of their size, suggests that this trait, in addition to indicating fecundity, conveys information about other aspects of female and/or egg quality.     

Female distribution affects mate searching and sexual selection in male northern water snakes (Nerodia sipedon)

Mating systems and sexual selection are assumed to be affected by the distribution of critical resources. We use observations of 312 mating aggregations to compare mate-searching success of male northern water snakes (Nerodia sipedon) in two marshes in which differences in mating substrate availability resulted in more than fourfold differences in female dispersion. Reproductive males had significantly larger home ranges where females were dispersed than where females were clumped. The number of females encountered by males increased significantly with male home range size where females were dispersed, and decreased significantly where females were clumped. Where females were clumped, males were more likely to encounter other males when they located females. We found no evidence in either population that mate searching was energetically expensive or that males with relatively more energy had larger home ranges. However, males with greater fat reserves at the start of the season participated in more mating aggregations when females were dispersed, suggesting that fat reserves could affect a male’s willingness to attempt mating or to persist in aggregations. When females were dispersed there was weak stabilizing selection acting to maintain male body size (β=–0.14), but strong directional selection favoring larger (β=0.50) and fatter (β=0.37) males. Over 7 years, the intensity of selection favoring larger males varied substantially (β=0.14–1.15), but that variation was not related to variation in the operational sex ratio. We found no evidence of directional selection on either body size (β=0.05) or fat reserves (β=0.10) of males when females were spatially clumped. Overall, the distribution of females had a pronounced effect on male behavior, on the factors that affected male success in locating females, and probably on the extent of sperm competition once females had been located. Received: 23 November 1998 / Received in revised form: 9 August 1999 / Accepted: 18 August 1999     

The effect of perceived female parasite load on post-copulatory male choice in a sex-role-reversed pipefish

The last several decades of research in behavioral ecology have resulted in a deeper appreciation of post-mating processes and sexual conflict in sexual selection. One of the most controversial aspects of sexual selection is cryptic mate choice. Here, we take advantage of male pregnancy in a sex-role-reversed pipefish (Syngnathus typhle) to quantify cryptic choice based on perceived parasite load and other sources of variance in female fitness. Studies have shown that S. typhle males preferentially mate with females with lower parasite loads and that a male’s perception of female parasite load can be altered by tattooing females. We manipulated the apparent parasite load of females in controlled mating experiments to test the hypothesis that post-copulatory sexual selection is dependent on a male’s perception of female parasite load in pipefish. Our results provided no evidence for cryptic male choice based on perceived female parasite load. However, we found evidence that eggs from larger females were more likely to result in viable offspring than eggs from smaller females and that the first female to mate with a male transferred more eggs per copulation on average. Overall, our results show that potential for post-copulatory sexual selection does exist in pipefish, but the male’s perception of female parasite load does not play a major role in this process.     

Sexual selection and the function of a melanin-based plumage ornament in polygamous penduline tits <Emphasis Type="Italic">Remiz pendulinus</Emphasis>

Melanin-based ornaments are often involved in signaling aggression and dominance, and their role in sexual selection is increasingly recognized. We investigated the functions of a melanin-based plumage ornament (facial ‘mask’) in male Eurasian penduline tits Remiz pendulinus in the contexts of male–male aggression, mating success, and parental care. The penduline tit is a passerine bird with a unique mating system in which both sexes may mate with several mates in a breeding season, and one (or both) parent deserts the clutch. Our study revealed that mask size of males is more likely an honest signal used by females in their mate choice decisions than a trait involved in male–male competition. First, mask size increased with both age and body condition, indicating that the mask may signal male quality. Second, males with larger masks paired more quickly and had more mates over the breeding season than males with smaller masks. Third, we found no evidence that male mask size signals male–male aggression or dominance during competitive encounters. The increased mating success of large-masked males, however, did not translate into higher reproductive success, as nestling survival decreased with mask size. Therefore, we conclude that there is either no directional selection on male mask size or males with larger masks receive indirect, long-term benefits.     

Size-dependent mating and gender choice in a simultaneous hermaphrodite, Bulla gouldiana

Simultaneous hermaphrodites are predicted to optimally divide resources between male and female function, which can result in both size-dependent mating behaviors and conflict between potential mates. Predicted strategies include size-assortative mating, conditional exchange of gametes, and mating patterns where relative size affects investment in each sexual role. This study investigated the effect of body size on the mating strategies of a hermaphroditic opisthobranch, Bulla gouldiana. Although individuals were spatially aggregated in the field with high levels of movement and size variation, there was little evidence for predictions. Laboratory experiments, however, revealed complicated effects of mass on the probability and duration of mating, as well as gender choice. Pairs were more likely to mate if they included at least one large animal, with the larger animal typically inseminating the smaller. When both individuals were large, they were more likely to each mate in both sexual roles by switching roles once. Although B. gouldiana did not usually alternate between sexual roles multiple times within mating events, paired individuals behaved similarly (neither or both mating as sperm donors) more often than expected by chance. This suggests some level of reciprocity, which is unlikely to be conditional given rates of unilateral mating. When the larger member of the mating pair inseminated the smaller, the duration of insemination increased with the size of the smaller sperm recipient. Copulations lasted longer in pairs that switched sexual roles than in those that did not switch roles. This study suggests that variation in body size can lead to size-dependent mating patterns, but only some of the patterns in B. gouldiana support theoretical predictions. We review other studies that have addressed similar issues, providing inconsistent mating patterns in sperm-storing hermaphrodites.     

Sources of variation in male mating success and female oviposition rate in a nonterritorial dragonfly

Summary Variance in male mating success and female oviposition rate was studied on a lifetime and daily scale in a nonterritorial libellulid dragonfly. While controlling for weather, population density, and sex ratio, we investigated whether phenotypic variation could explain a significant amount of the observed variation. By collecting most of the data in a large outdoor field enclosure, we had access to (1) real lifetime data as emigration was prevented, (2) exact values of density and sex ratio, and (3) phenotypic data that are virtually unavailable under natural conditions. Variance in LMS was higher in males than variance in lifetime number of ovipositions in females. Lifespan was of major importance in explaining the variation. Males readily remated on the same day whenever possible (maximum 6 times), whereas in females each oviposition bout usually was followed by a nonreproductive period of approximately 4 days. An analysis of the sources of variance in daily reproduction rate showed that, besides the expected effects of weather, an excess of one sex resulted in a negative effect on the individual mating probability of members of that sex. Even after correction for weather and population parameters, phenotypic traits explained a significant amount of the remaining variation: males with a high mating rate were large, agonistic, active mate searching, and long-living individuals. Females were more likely to oviposit when they showed higher flight activity and when more days had elapsed since the previous oviposition. The presence of a strong size effect on male mating probability, and its absence in females, was confirmed at a pond (native habitat). We propose a model on the causal relationships between the relevant parameters and the variation in reproductive success. Finally, we briefly discuss why variance in reproductive success is not a very good measure of the opportunity for sexual selection in this species. Offprint requests to: N.K. Michiels at the present address     

Experimental assessment of ecological and phenotypic factors affecting male mating success and polyandry in northern watersnakes, Nerodia sipedon

To resolve conflicting field observations regarding the action of sexual selection, we used breeding experiments and paternity analysis of the 927 resulting offspring to assess how male size, condition, tail length, genetic similarity to the female, and variation in operational sex ratio (OSR) affected male reproductive success and the incidence of polyandry in northern watersnakes (Nerodia sipedon). Only size affected male mating success. Large males were more successful, but only when male size varied substantially and competition among males was intense (i.e., male-biased OSR). The conditional nature of the size advantage may explain why studies of free-living watersnakes have produced inconsistent results regarding the relationship between male size and mating success. Size differences between males did not affect the proportion of offspring each male sired within multiply sired litters. We found positive size-assortative mating, but only when the OSR was female biased, suggesting that smaller males had improved access to females when competition among males was reduced, but that competition with larger males still restricted mating opportunities of small males to less preferred, smaller females. Most litters (58%) were multiply sired and larger females were more likely to produce multiply sired litters, similar to free-living watersnakes. There was no association between the incidence of multiple paternity and OSR, however, suggesting that polyandry is not simply a function of opportunity, with females passively waiting for males to court them.     

The genetic mating system of a sea spider with male-biased sexual size dimorphism: evidence for paternity skew despite random mating success

Male-biased size dimorphism is usually expected to evolve in taxa with intense male–male competition for mates, and it is hence associated with high variances in male mating success. Most species of pycnogonid sea spiders exhibit female-biased size dimorphism, and are notable among arthropods for having exclusive male parental care of embryos. Relatively little, however, is known about their natural history, breeding ecology, and mating systems. Here we first show that Ammothella biunguiculata, a small intertidal sea spider, exhibits male-biased size dimorphism. Moreover, we combine genetic parentage analysis with quantitative measures of sexual selection to show that male body size does not appear to be under directional selection. Simulations of random mating revealed that mate acquisition in this species is largely driven by chance factors, although actual paternity success is likely non-randomly distributed. Finally, the opportunity for sexual selection (I _s), an indirect metric for the potential strength of sexual selection, in A. biunguiculata males was less than half of that estimated in a sea spider with female-biased size dimorphism, suggesting the direction of size dimorphism may not be a reliable predictor of the intensity of sexual selection in this group. We highlight the suitability of pycnogonids as model systems for addressing questions relating parental investment and sexual selection, as well as the current lack of basic information on their natural history and breeding ecology.     

Sexual selection and species recognition in the pygmy swordtail, Xiphophorus pygmaeus: conflicting preferences

Sexual selection and species recognition play important roles in mate choice; however, sexual selection preferences may overlap with traits found in heterospecifics, producing a conflict between sexual selection and species recognition. We examined female preferences in Xiphophorus pygmaeus for male traits that could provide both types of information to determine how females use multiple cues when preferences for these cues would conflict. We also examined X. pygmaeus behavior in the field to determine if females have the opportunity to choose mates. As no male-male competition was observed in the field, and females occasionally chased males from feeding areas, females apparently have the opportunity to exercise mate choice in their natural habitat. In the laboratory, female X. pygmaeus used body size as a sexual selection cue, preferring large heterospecifics (X. cortezi) to small conspecifics. Females also preferred barless X. cortezi over barred X. cortezi when males were size matched. Because X. pygmaeus males do not have bars, this preference suggests that X. pygmaeus females use vertical bars in species recognition, and that large body size and vertical bars are conflicting cues. However, X. pygmaeus females did not have a preference for males of either species when sexual selection and species recognition cues were presented concurrently. This result was surprising, because preferences for species recognition cues are often assumed to be stronger than sexual selection cues. We suggest that females may be using additional species-specific cues in mate choice to prevent hybridization.     

The lexicon of love: what cues cause size-assortative courtship by male garter snakes?

Cues that females use to select potential mates have attracted substantial research effort, but the criteria for male mate choice remain very poorly known. Red-sided garter snakes ( Thamnophis sirtalis parietalis) court and mate in large aggregations around overwintering dens in southern Manitoba, Canada. Both courtship and mating are size-assortative: small male snakes court small as well as large females, whereas larger males court only large females. This system provides a unique opportunity to assess the cues that males use in selecting mates, and in particular the mechanisms that generate a size-related shift in mate preference. Experiments in which we manipulated body sizes and scents showed that both vision and scent (sex pheromones) were important. Large males directed intense courtship only when the stimulus provided both visual and chemical (skin lipid) evidence of large body size. Small males were much less discriminating in both respects. Thus, size-assortative mating in this system is generated not by larger males excluding their smaller rivals from the largest females (as has been reported in other reptile species), but by a size-related shift in the visual and pheromonal cues that elicit courtship. Males of some species may thus show complex patterns of mate choice, with the cues that stimulate courtship differing even among males within a single population based on traits such as age or body size.     

Allocation in reproduction is not tailored to the probable number of matings in common toad (<Emphasis Type="Italic">Bufo bufo</Emphasis>) males

The theory of life history evolution assumes trade-offs between competing fitness traits such as reproduction, somatic growth, and maintenance. One prediction of this theory is that if large individuals have a higher reproductive success, small/young individuals should invest less in reproduction and allocate more resources in growth than large/old individuals. We tested this prediction using the common toad (Bufo bufo), a species where mating success of males is positively related to their body size. We measured testes mass, soma mass, and sperm stock size in males of varying sizes that were either (1) re-hibernated at the start of the breeding season, (2) kept without females throughout the breeding season, or (3) repeatedly provided with gravid females. In the latter group, we also estimated fertilization success and readiness to re-mate. Contrary to our predictions, the relationship between testes mass and soma mass was isometric, sperm stock size relative to testes mass was unrelated to male size, fertilization success was not higher in matings with larger males, and smaller males were not less likely to engage in repeated matings than larger males. These results consistently suggest that smaller males did not invest less in reproduction to be able to allocate more in growth than larger males. Causes for this unexpected result may include relatively low year-to-year survival, unpredictable between-year variation in the strength of sexual selection and low return rates of lowered reproductive investment.