Dynamic female preference for multiple signals in <Emphasis Type="Italic">Rhinogobius brunneus</Emphasis>

Many species base their choice of mates on multiple signals which provide them with different kinds of information. Choosers may assess the signals together to evaluate the overall quality of potential mates, but individuals often pay attention to different signals in different contexts. In Rhinogobius brunneus, a fish displaying exclusive male parental care, females generally prefer males showing larger first dorsal fins (FDF) and more active courtship displays as mates. Females choosing a mate usually initially assess the FDF and later utilize courtship for the final decision. In our experiments, females with different hunger states used different signals when selecting mates. Females in both hunger states preferred males with larger FDF in the first stage. In the second stage, well-fed females showed highly repeatable choice, whereas poorly fed females responded only to variation in the courtship activity of males. The males preferred by poorly fed females exhibited significantly higher offspring survival than nonpreferred males. Under conditions of food shortage, males allocate more energy to future reproduction at the expense of the present brood, and females may prioritize signals predictive of offspring survivorship over signals reflecting other aspects in male quality to minimize the losses in direct benefits. We conclude that R. brunneus females may employ information from both signals but dynamically adjust their prioritization of each signal to current conditions to ensure the choice that is currently most adaptive.     

Male-male association patterns and female proximity in the guppy,Poecilia reticulata

If males differ in their ability to attract potential mates, and are able to perceive such differences, theory predicts they should distribute themselves in a manner that increases their probability of obtaining potential matings. The relationship between male-male association patterns and the proximity of females in social groups, however, remains virtually unexplored. Experimental analysis of this relationship in the guppy, Poecilia reticulata, demonstrates that in preference tests males showed a strong tendency to associate with other males that were further away from potential mates than they were themselves. Male guppies pursue a behavioral strategy that involves categorizing other males based on their proximity (and possibly relative attractiveness) to females, remembering the identity of such individuals, and using this information when choosing between other males as associates. Such a strategy may increase a male's chances of being the individual chosen by a female assessing nearby males.     

Foraging ability in the scorpionfly <Emphasis Type="Italic">Panorpa vulgaris</Emphasis>: individual differences and heritability

According to indicator models of sexual selection, mates may obtain indirect, i.e. genetic, benefits from choosing partners indicating high overall genetic quality by honest signals. In the scorpionfly Panorpa vulgaris, both sexes show mating preferences on the basis of the condition of the potential partners. Females prefer males that produce nuptial gifts (i.e. salivary secretions) during copulation, while males invest more nuptial gifts in females of high nutritional status. Both characters, males' ability to produce nuptial gifts and high nutritional status of females, are known to be reliable indicators of foraging ability. Thus, besides possible direct benefits, both sexes might also obtain indirect benefits in terms of “good foraging genes” by their choice and thereby increase the fitness of their offspring. A prerequisite for this possibility is the heritability of the respective trait. In the present study, we estimated the repeatability and the heritability of foraging ability. Our results indicate (1) a significant repeatability of individual foraging efficiencies in males and females and (2) a heritable component of this trait by a significant parent–offspring regression. These findings suggest that genetic benefits in terms of increased offspring foraging ability might contribute to selection for mating preferences in both sexes.     

Female rock shrimp<Emphasis Type="Italic"> Rhynchocinetes typus</Emphasis> mate in rapid succession up a male dominance hierarchy

The dynamics of male-male competition for mates and patterns of female choice depend critically on the social environment. We released newly molted sexually receptive females of the rock shrimp Rhynchocinetes typus in the field and recorded their interactions with males. In the dense aggregations in which these shrimp live, most females were encountered and seized by males within 2 min. Usually, females were first seized by subordinate males, and subsequently taken over by the dominant males. Many females (17 out of 23) had multiple mates during the 10-min observation period, and most of them received spermatophores from multiple males. Males used different mating tactics in accordance with their dominance status: subordinate males often used the sneaking tactic, seizing the female and immediately transferring spermatophores. In contrast, all dominant males used the primary mating tactic; they seized and stimulated the female before transferring spermatophores. Results from previous studies had indicated that females may reduce the fertilization chances of subordinate males by delaying spawning and removing spermatophores. We suggest that this capability in combination with the observed rapid mate succession may enable females to exploit male contest behaviors.Communicated by P. Backwell     

Condition-dependent control of paternity by female purple martins: implications for coloniality

Proposed causal links between extra-pair copulation (EPC) and colony formation in socially monogamous birds hinge on the question of which sex controls fertilizations. We examined in colonial purple martins Progne subis (1) whether EPCs were forced or accepted by females, and (2) the degree to which apparently receptive females were able to obtain EPCs against their mates’ paternity defenses. Paternity analyses of multilocus DNA fingerprinting confirmed previous findings of a marked relationship between age class and extra-pair fertilizations (EPFs), with young males losing paternity of 43% (n = 53) of their putative offspring compared to 4% (n = 85) by old males. All assignable extra-pair offspring were sired by old males, with one male obtaining most EPFs each year. Contrary to the hypothesis that EPCs are forced, EPF frequency within age class did not increase with seasonal increases in the number of males per fertile female. Whereas the male control hypothesis predicted that the male age class that mate-guarded more would be cuckolded less, the reverse was true: young males guarded significantly more intensely. The male age class difference in cuckoldry could not be explained by the possibility that young and inexperienced females (which are usually paired to young males) were more vulnerable to forced copulation because EPFs were unrelated to female age. These findings suggest that females (1) pair with old males and avoid EPCs, or (2) pursue a mixed mating strategy of pairing with young males and accepting EPCs from old males. The receptivity to EPCs by females paired to young males put them in conflict with their mates. Two factors determined the paternity achieved by young males: (1) the relative size of the male to the female, with young males achieving much higher paternity when they were larger than their mates, and (2) the intensity of mate-guarding. Both variables together explained 77% of the variance in paternity and are each aspects of male-female conflict. Given female receptivity to EPCs, mate-guarding can be viewed as male interference with female mating strategies. We conclude that EPCs are rarely or never forced, but the opportunity for females paired to young males to obtain EPCs is relative to the ability of their mates to prevent them from encountering other males. Evidence of mixed mating strategies by females, combined with other features of the martin mating system, is consistent with the female-driven “hidden lek hypothesis” of colony formation which predicts that males are drawn to colonies when females seek extra-pair copulations. Received: 23 March 1995/Accepted after revision: 14 January 1996     

The effect of chorus organization on mate localization in the Painted Reed frog (Hyperolius marmoratus)

Summary Experiments with female Painted Reed frog (Hyperolius marmoratus) indicate that temporal separation of the calls of neighbouring males does not enhance their locatability. The ability of females to locate a sound source is not impaired when identical calls are broadcast simultaneously through two spatially separated loudspeakers (n=8). We suggest that call alternation may be important in the spatial distribution of males rather than in the ability of females to locate mates.     

Density affects female and male mate searching in the fiddler crab, <Emphasis Type="Italic">Uca beebei</Emphasis>

In most species, only one sex searches for mates while the other waits. Models of sex-specific mate-searching behavior predict single-sex searching, but the factors that determine which sex searches are not understood. In this study, we examine the effects of density and predation risk on mate-searching behavior in the fiddler crab Uca beebei. U. beebei is one of the few fiddler-crab species in which both sexes search for mates. In a field experiment conducted in Panama, we manipulated crab density and perceived predation risk in replicate plots. Females searched more and males searched less at high densities. At high levels of perceived predation risk, both sexes similarly reduced their search rates. Observations of plots that naturally varied in crab density show that females were more likely to search for mates in areas of higher density, where there were more males. Females may preferentially search for mates in high-density areas because the abundance of nearby burrows, into which they can run to escape predators, decreases their costs of searching and because the abundance of males and male burrows facilitates comparisons and thus may increase their benefits from searching. Males at high densities decrease their mate-searching rate perhaps in response to the increase in female searching and to the corresponding increase in the intensity of their competitors' mate-attraction signals.     

Looking on the bright side: females prefer coloration indicative of male size and condition in the sexually dichromatic spadefoot toad, <Emphasis Type="Italic">Scaphiopus couchii</Emphasis>

Females across many taxa commonly use multiple or complex traits to choose mates. However, the functional significance of multiple or complex signals remains controversial and largely unknown. Different elements of multiple or complex signals may convey independent pieces of information about different aspects of a prospective mate (the “multiple messages” hypothesis). Alternatively, multiple or complex signals could provide redundant information about the same aspect of a prospective mate (the “redundant” or “back-up” signal hypothesis). We investigated these alternatives using spadefoot toads, Scaphiopus couchii. Spadefoot toads primarily use calls to attract their mates, but males also exhibit sexually dimorphic coloration. We investigated whether male coloration is indicative of male size, condition, or infection status by a socially transmitted monogenean flatworm. We found that male coloration and dorsal patterning predicts male size and condition but not infection status. Moreover, when we presented females with a choice between a bright male model and a dark male model, we found that females preferred the bright model. Because aspects of males’ calls are also associated with male size and condition, we conclude that coloration is a potentially redundant indicator of male phenotype. We suggest that coloration could enhance mate choice in conjunction with male calling behavior by providing females with a long distance cue that could enable them to identify prospective mates in a noisy chorus environment where the discrimination of individual calls is often difficult. Generally, such redundant signals may facilitate mate choice by enhancing the quality and accuracy of information females receive regarding prospective mates.     

Underwater oviposition in a damselfly (Odonata: Coenagrionidae) favors male vigilance,and multiple mating by females

Summary Female Enallagma hageni oviposit underwater where they are inaccessible to males. I demonstrate that males guard submerged females rather than perch sites, and are behaviorally distinct from lone males at the water. In contrast to lone males, which always attempt to copulate with females presented to them, guarding males exhibit a conditional latency to remating which corresponds closely to the time required by females to oviposit a complete clutch of eggs. By ovipositing underwater, females decrease the risk that their eggs become exposed. Risks associated with submerged oviposition favor both mate guarding, and multiple, within-clutch matings by females. Both guarding mates and lone males rescue females that float on the water surface as the result of improper resurfacing. Such behavior reduces the mortality risk to females from 0.06 to 0.02 per oviposition bout. By remating between bouts, females benefit from the additional vigilance of lone males, who rescue floating females 1.4 times as often as original mates. A second consequence of multiple mating is an increase in the selective advantage of vigilance by mates. Because receptive females become scarce by early afternoon, whereas male density remains high, a male has little (3%) chance of encountering a second receptive female that day. However, he incurs a 42% risk of losing fertilizations if he abandons a mate. For male E. hageni mate guarding functions in the context of both natural and sexual selection. It insures that a mate lives to lay a complete egg clutch in addition to protecting a male's sperm investment.     

Female-enhanced male competition determines the first mate and principal sire in the spider Linyphia litigiosa (Linyphiidae)

Summary Two to five days before sexual maturation, female sierra dome spiders (Linyphia litigiosa: Linyphiidae) undergo a transformation in their behavior toward males that visit their webs. During this latter part of their penultimate instar, females change from consistently positioning themselves far away from males to actively maintaining close proximity, reactions I call avoidant and associative behavior, respectively. Consistent associative behavior ceases after the female's first mating and thus is limited to soon-to-mature penultimate females. When a mate-seeking male fords an associative female, he attempts to guard her until she matures; this is often a multi-day affair. In contrast, males guard immature avoidant and mature mated females for only a single day. This dichotomy in male guarding times can be understood by the fact that associative behavior signals that the female will soon develop peak reproductive value. Upon completion of their final molt, 98% of females immediately mate with the current guarding male. Secondary suitors are not as likely to achieve mating. Moreover, first mates father 1.8 times more offspring, on average, than secondary mates. Whenever they meet on any female's web, males fight until one of the contestants withdraws. Fights typically are intensive, occasionally deadly, and often result in usurpation of the web by the newly arriving male. Larger males win more fights, but other qualities (e.g., vigor and persistence) appear to be important when contestants differ by less than 10–20% in body weight. Prolonged (i.e., multi-day) guarding of associative females enhances the intrasexual selection process by ensuring that every male that arrives at the web finds it already guarded. Therefore every male that finds the web becomes a participant in a series of male-male conflicts and web usurpations which span the period between the resident female's commencement of associative behavior and her sexual maturation. Since unforced male departures from the webs of associative females are rare, victors are retained on the web until they themselves lose a fight. This facilitates a steady increase in the fighting ability of sequential guards throughout the associative period, up until female maturation and mating. On my study site, first mates represented the final winners in a combative sorting process based on a minimum average of 2 fights; they were heavier and larger than secondary mates and randomly sampled males. The combination of (1) associative behavior by nearly mature females, (2) high mating propensity of newly mature females, and (3) first male sperm priority, constitutes a system whereby females enhance male-male competition and boost the expected fighting prowess of the principal sire of their progeny. Since males appear to make no material contribution toward progeny, the female's behavior probably functions to improve the genetic constitution of the offspring. In addition, the timing of associative behavior may limit prolonged guarding by food-stealing males to a period (1) encompassing the female's pre-molt fast and (2) before the heavy yolking of eggs, thereby ameliorating the nutritional costs of intrasexual selection.     

Song rates of dark-eyed juncos do not increase when females are fertile

Frequency of singing by birds may vary with reproductive stage in ways that reflect variation in the functions of song in intersexual and intrasexual communication. In dark-eyed juncos (Junco hyemalis) high-amplitude song is produced only by males. To investigate the function of this song, we tested whether fertility of females affected singing by their mates or by neighboring males. Using focal observations, song censuses, and radiotracking data, we determined whether song production varied between and among periods when females were fertile and non-fertile. Our findings show that males do not increase song production when their mates are fertile, nor do they increase song production when neighboring females are fertile. These results suggest that male juncos do not signal their intent to defend territories (or mates) more when females are fertile and that they do not use song to advertise to specific potential participants in extra-pair fertilizations. Received: 13 February 1997 / Accepted after revison: 2 May 1997     

Temporal constraints and female preference for burrow width in the fiddler crab, <Emphasis Type="Italic">Uca mjoebergi</Emphasis>

We studied sampling behaviour and mate choice in the fiddler crab Uca mjoebergi. Once a female selects a mate, she copulates in his burrow and remains there until releasing her aquatic larvae. U. mjoebergi occurs in habitats that are inundated only by the highest amplitude spring tides. Females can only release their larvae during these tides, and release before or after will result in complete failure of reproductive effort. Matings occur over a 5-day period near the end of neap tides. Our results suggest that within the mating period, females adjust their larval developmental rates by selecting specific burrows in which to incubate their clutches. We found that at the start of the mating period, females chose larger males as mates. Since male size was positively correlated to burrow width, females were selecting wide burrows and effectively incubating at lower temperatures. This would slow down the developmental rates of larvae. In contrast, females that mated late in the mating period selectively chose small males. By incubating in narrower, warmer burrows, these females may increase the developmental rates of larvae. We propose that females are selecting burrows to influence incubation rate and ensure timely release of their larvae. Female U. mjoebergi appear to adjust their preference for the direct benefits of mate choice to increase their reproductive success.     

Correlates of extra-pair fertilization success in hooded warblers

We examined correlates of extra-pair fertilization (EPF) success in the hooded warbler (Wilsonia citrina), a species where females pursue extra-pair matings. The good genes hypothesis predicts that females choose extra-pair mates with morphological or behavioral traits that reflect differences in male genetic quality. EPFs were common, as 35.3% (42/119) of broods contained extra-pair young (EPY) and 26.7% (95/356) of nestlings were the result of EPFs. There was a strong skew in male EPF success which increased the variance in annual male mating success 2–3 fold. However, male morphology did not predict EPF success, as extra-pair males were not older or larger than the males they cuckolded. Likewise, there were no significant correlations between the proportion of extra-pair young in a brood and male size or age. The good genes hypothesis predicts that high-quality males will be consistently preferred as genetic mates, but the number of young sired by a male with his social mate was not consistent from one year to the next. There was a significant negative correlation between female age and proportion of EPY produced, which could result if older females obtain higher-quality social mates. We found no strong evidence that females choose extra-pair mates for good genes, but females may use behavioral rather than morphological cues to assess relative male quality. Received: 31 May 1996 / Accepted after revision: 17 October 1996     

Mate choice by female white-crowned sparrows in a mixed-dialect population

Summary It has been argued that the song dialects of white-crowned sparrows, Zonotrichia leucophrys, may function to restrict gene flow between dialect populations, thus promoting adaptation to local environmental conditions. One proposed mechanism whereby this may occur is if a female mates preferentially with males singing her natal dialect, typically the song type of her father. Previous field and laboratory studies of the role played by song differences in mate choice by female white-crowned sparrows have produced equivocal results. These investigations were conducted in populations in which a single song dialect was predominant. We examined mate choice in a population in which two song dialects are equally common. We found that songs of mates of individual females in successive years were no more likely to be of the same dialect than expected by chance. In addition, individual females did not mate preferentially with males whose songs matched those of their fathers. We conclude that, in this population, females choose mates on the basis of characteristics other than song type.     

Experimental manipulation of mate choice by male katydids: the effect of female encounter rate

Summary This study tests the general prediction that discrimination among potential mates increases with the availability of potential mates. Specifically, we conducted two experiments that examined mate choice by male zaprochiline katydids in relation to their prior encounter rate with females. The probability of mate acceptance or rejection was measured for males given either frequent or no contact with females in the laboratory (experiment 1) and males taken directly from natural areas of either high or low female abundance (experiment 2). In both experiments, males with low female encounter rates were more likely to mate than males with high female encounter rates. In both cases, the decreased mating probability of males in the high encounter treatment resulted from their tendency to reject lighter (and less fecund) females. Despite the presumed advantage to males of selecting heavier females, field data indicate that, unlike females, males do not aggregate in rich food patches. Possible explanations for this finding are discussed. Offprint requests to: T.E. Shelly at the present address     

Light environment and mating behavior in Trinidadian guppies (<Emphasis Type="Italic">Poecilia reticulata</Emphasis>)

Male guppies, Poecilia reticulata, have color patterns that result from a balance between natural selection for crypsis to avoid predators and sexual selection for bright, complex patterns that attract females. Males use displays to show off these patterns to potential mates, but their conspicuousness also depends on the light environment in which they are viewed. We investigated variation in natural underwater guppy light environments in Trinidad, West Indies, and found that mating behavior is correlated with both the ‘quantity’ (total irradiance) and ‘quality’ (spectral composition) of light: light intensity and the proportion of ultraviolet light were negatively related to display rates. Experimental manipulation of light environment to mimic natural daily changes demonstrated that these relationships are causal and are independent of time of day effects. At lower light levels, when guppies are less detectable by visually hunting predators, females had more opportunity for active mate choice, because males displayed more. However, these light conditions may reduce the ability of females to accurately discriminate between males. Guppy mating behavior is therefore strongly affected by light environment, and this may have important effects on sexual selection.     

Identification of sex, age and species-specific proteins on the surface of the harpacticoid copepod Tigriopus japonicus

 Behavioral experiments have shown that male copepods of the species Tigriopus japonicus (Nori) can distinguish species, sex, and developmental stage of potential mates using contact chemoreception. Lectin-binding patterns on the body surface of females have indicated that surface-bound glycoproteins may be important signals in mate choice. In the present study, the proteolytic enzyme trypsin was used to cleave surface proteins from females, reducing their attractiveness to males. The protein fragments released were used to make monoclonal antibodies. Three levels of screening were used to identify monoclonal antibodies that recognized proteins involved in mate recognition. One monoclonal antibody bound to the terminal urosome and lateral prosome of CV females, and its binding significantly decreased female attractiveness to males. Western blotting showed that this antibody bound the trypsin-cleaved fragment and proteins of homogenized CV females and virgin adult females, but did not bind proteins of homogenized males, CIII females, or females of T. californicus or T. fulvus. This antibody recognized proteins on the surface of females that may enable males to discriminate conspecifics, sex, and age. It is likely that this molecule has a central role in the evolution of reproductive isolation in this group. Received: 22 July 1999 / Accepted: 21 March 2000     

Non-aggressive mate guarding by the blue-footed booby: a balance of female and male control

Thirteen pairs of blue-footed boobies (Sula nebouxii) were observed on their colony. Pairs courted frequently and, on average, copulated 24 times during the 30 days before laying, with 38% of those copulations occurring in the last 5 days (presumed fertile period). Males and females increased attendance at the nest site as laying approached. Seven females performed an average seven extra-pair copulations, with 1–2 paired male neighbors, but these were less concentrated in the presumed fertile period than within-pair copulations, and the last two copulations of all 7 females were with their social mates. Rates of female extra-pair copulations were six times lower when their social mate was present, and during the presumed fertile period, no female performed an extra-pair copulation in the presence of her mate. Males did not respond to infidelity of social mates with aggression, prompt copulation, retaliatory copulation, or increase in copulation. Seven of 13 males performed an average of five extra-pair copulations, with 1–3 paired female neighbors, before their own mates began egg-laying. The males' extra-pair copulations represented only 4% of their total copulations during their own mates' presumed fertile periods. Females, the larger sex, apparently control sexual access and copulate with extra males to achieve extra-pair fertilization. Males pursue a mixed strategy: they copulate with extra females, mostly outside their own mate's presumed fertile period, and they copulate increasingly with their social mate as laying approaches, probably assuring some paternity by mate guarding, involving attendance and courtship. Behavior of males and females is also consistent with other hypotheses for extensive joint nest site attendance: pairbonding, copulation access, and territory acquisition. Received: 14 November 1997 / Accepted after revision: 16 May 1998     

Female mate choice determines reproductive isolation between sympatric butterflies

Animal courtship rituals are important for species recognition, and a variety of cues might be utilized to recognize conspecific mates. In this paper, we investigate different species-recognition mechanisms between two sympatric butterfly sister species: the wood white (Leptidea sinapis) and Real’s wood white (Leptidea reali). We show that males of both species frequently court heterospecific females both under laboratory and field conditions. The long-lasting elaborate courtships impose energetic costs, since the second courtship of males that were introduced to two subsequent conspecific females lasted on average only one fourth as long as the first courtship. In this paper, we demonstrate that premating reproductive isolation is dependent on female unwillingness to accept heterospecific mates. We studied female and male courtship behavior, chemical signaling, and the morphology of the sexually dimorphic antennae, one of the few male traits visible for females during courtship. We found no differences in ultraviolet (UV) reflectance and only small differences in longer wavelengths and brightness, significant between-species differences, but strongly overlapping distributions of male L. sinapis and L. reali antennal morphology and chemical signals and minor differences in courtship behavior. The lack of clear-cut between-species differences further explains the lack of male species recognition, and the overall similarity might have caused the long-lasting elaborate courtships, if females need prolonged male courtships to distinguish between con- and heterospecific suitors.     

Indirect evidence indicates female semiochemicals release male precopulatory behaviour in the snow crab,Chionoecetes opilio (Brachyura: Majidae)

Summary Experiments were conducted in aquaria to test the hypothesis that females of the snow crab,Chionoecetes opilio, release a sex pheromone to attract mates. Males exhibited significantly increased activity to water from a source aquarium containing recently-moulted pubescent females, egg-stripped multiparous females and recently-moulted immature females than to water from an aquarium containing berried multiparous females, eggs alone, adolescent males or an empty aquarium. Males without their cephalic chemoreceptors maintained low activity levels in the presence of recently-moulted pubescent females, whereas maxilla-ablated males reacted as strongly as intact males. These findings are discussed within the context of the ecdysteroid-sex pheromone hypothesis, proposed by Kittredgeet al. (1971) and Kittredge & Takahashi (1972), but subsequently rejected. We propose that ecdysteroids from both pubescent and multiparous females may elicit male search-and-clasp behaviour inC. opilio and that the reproductive biology of species used to refute the hypothesis was inappropriate to test the role of ecdysteroids as a cue in the mating process.