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.     

Selection for male choice based primarily on mate compatibility in the oldfield mouse, Peromyscus polionotus rhoadsi

Despite the consensus that mate choice acts as a mechanism for selection of secondary sexual traits, the evolutionary forces affecting mate preferences themselves remain controversial. In this study, we first demonstrated selection acting directly on the mate preferences of monogamous male oldfield mice, Peromyscus polionotus rhoadsi. One group of male oldfield mice were allowed to express a social preference between two potential mates, and were subsequently paired with either their preferred or rejected female. Among these pairs, those containing preferred females produced more offspring than did those containing rejected females. We next demonstrated that this fitness advantage depended primarily on compatibility between the members of a mated pair. A second group of male oldfield mice were not allowed the opportunity to express a social preference between potential mates. Rather, these males were paired with females that had been either preferred or rejected by males in the first group. Among these pairs, those containing preferred females did not produce more offspring than those containing rejected females. In other words, individual mate preferences had fitness consequences only for those males that expressed them, demonstrating that these preferences were based primarily on compatibility between mates.     

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     

Paternity control for externally fertilised eggs: behavioural mechanisms in the waterfrog species complex

Male competition for mates and female mate choice are key mechanisms involved in sexual selection. Surprisingly, these mechanisms have often been investigated separately although they appear to interact in many species. Male–male competition for territories located at the best places or to establish dominance relationships often explain mating patterns. Such male behaviours may affect and sometimes even hinder female mate choice, as in the case of sexual coercion. While in many species females are able to exert cryptic control over paternity (i.e. a process allowing females to bias offspring production toward certain males after intromission), in other species external fertilisation prevents females from doing so. This is the case in the waterfrog hybridisation complex where the hybrid Pelophylax esculentus can only produce viable offspring by pairing with the parental species Pelophylax lessonae (hybridogenetic reproduction). We examined two potential processes that could enhance such mating combinations. Firstly, by monitoring male spatial distribution within six choruses, we showed that the proportion of P. lessonae males located at the edge (in the best position to grasp females arriving at the chorus) cannot explain the frequency of mating combinations observed. Secondly, an experimental approach emphasised a new way for anuran females to favour paternity of a particular male in a sexual coercion context. When females are forcefully paired with an incompatible male, they cannot remove the male grasped on their back by themselves. Nevertheless, by controlling the movement of the pair within the chorus, these females often change mates by enhancing male competition instead of laying eggs. In many species with externally fertilised eggs, it may be thus necessary to take into account this new possibility for females to control offspring paternity.     

Sex differences in partner preference in mated pairs of saddle-back tamarins (Saguinus fuscicollis)

Summary The attraction of permanently cohabiting males and females to their pair mate was assessed in a preference situation in which a subject was given a choice between its pair mate and a stranger of the mate's sex. Animals were tested in a choice apparatus, consisting of 1 cage housing the subject, 1 cage housing its pair mate, and 1 cage housing the strange stimulus animal. Wire mesh tunnels connected the subject's cage with that of each stimulus animal. Each tunnel was divided into 5 sections of equal length. During a series of 30-min choice tests, the subjects' location was scored every 10 sec as being in the home cage or in one of the sections of the tunnels leading to the cages of the other animals. In addition to these location scores, being at Zero Distance from a stimulus animal, as well as a number of behaviors directed at both stimulus animals, were recorded. Males were in closer proximity to their females than to strange females (Fig. 2), approached their females more frequently and directed more Tongue Displays and Sniffs at their mates (Figs. 3–5). Females did not prefer their males in terms of these behaviors (Figs. 2–5). Moreover, they directed a higher number of Gazes at strange males (Fig. 6). Both males and females were more frequently at Zero Distance from their pair mates than from strange stimulus animals and had more Nonaggressive Contacts with their pair mates (Figs. 7–8). These results suggest that males show a strong attachment to their female pair mates while females are attracted to their mates as well as to strange males.     

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     

Conspicuous displays in cryptic males of a polytypic poison-dart frog

The evolution of aposematism is linked to increased opportunities for conspicuous sexual displays since detection by potential predators is no longer disadvantageous. Therefore, phenotypic divergence in aposematic species leading to relatively cryptic forms is expected to constrain such opportunities, by restoring the trade-off between natural and sexual selection on the boldness of sexual displays. We asked if and how a derived phenotype of the poison-dart frog Oophaga granulifera that appears relatively cryptic to potential predators exhibits conspicuous sexual displays for potential mates. We used visual modeling of frog contrasts against their natural backgrounds to test if for conspecifics green frogs appear less conspicuous than red frogs as they do for birds. We conducted behavioral observations of focal red and green males to determine if green frogs adjust their display behavior to the availability of potential mates. Dorsal brightness is known to influence female preferences in at least one poison frog species. We found that, despite being less visible under some measures, green frogs may appear as bright as red frogs for conspecifics but not birds, when viewed on dark backgrounds. Additionally, green males called more intermittently than red males when advertising to distant females, but they exhibited a dramatic increase in calling activity in proximity of a female and were as active as red males in this context. Together, our results suggest that green frogs retain context-dependent conspicuousness to conspecifics despite the evolution of relative crypsis to potential predators.     

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.     

A test of male mating and hunting success in the kestrel: the advantages of smallness?

We tested female choice for male wing and tarsus length and body mass in the kestrel (Falco tinnunculus), a species in which males average about 10% smaller than females. We also studied how male characters are related to their hunting success. In the laboratory, females preferred lighter males with shorter tarsi as mates, if the difference in those characters between competing males was larger than average. Lighter and shorter-winged males seemed to be better hunters than heavier and longer-winged males. Field observations in a year in which voles were scarce suggested that shorter-winged males were also better food providers in courtship feeding than longer-winged males,although in good vole years such a relationship was not found. We argue that females may prefer to pair with smaller males, because they have higher flight performance and better hunting success than heavier males. By doing so, females may gain direct breeding advantages. We conclude that both female choosiness and the hunting efficiency of males well contribute to reversed sexual size dimorphism (RSD, females larger than males) in the kestrel. Received: 18 July 1995/Accepted after revision: 17 August 1996     

Sexual conflict over fertilizations: female bluethroats escape male paternity guards

Extra-pair copulations create a potential for sexual conflict in pair-bonding birds. Here we report an experimental study of the bluethroat, Luscinia s. svecica, in which the throat ornament of males was blackened with Nyanzol D in order to reduce their sexual attractiveness and thus increase the sexual conflict over fertilizations. In an earlier study, we showed that males blackened before pairing had a lower success in attracting social mates than controls, whereas males blackened after pairing guarded their mates more intensely and sang less than controls. Here we add behavioural data from one more year on males blackened after pairing and corroborate our previous finding that the manipulation caused males to guard their mates more intensely and advertise less for additional mates. Blackened males did not suffer more intrusions from neighbouring males than did controls. Paternity analyses of the combined data set, using multilocus DNA fingerprinting and microsatellite typing, revealed that blackened males lost significantly more paternity than controls. There was also a tendency for blackened males to show a lower success in achieving extra-pair fertilizations. These results indicate that females have the upper hand in the sexual conflict over fertilizations, as females paired with unattractive males can achieve more extra-pair paternity despite the greater constraint posed by the intensified mate guarding. Still, within the blackened group, there were some indications that males guarding more intensely and singing less had higher paternity than males guarding less and singing more, suggesting a marginal positive effect of guarding for unattractive males. Male mate guarding must nevertheless be considered a best-of-a-bad-job strategy in this species. Received: 4 December 1997 / Accepted after revision: 14 June 1998     

Great tits choose between food and proximity to a mate

Summary These experiments test for time of day differences in the attentiveness of territorial, paired, male and female great tits (Parus major) to their mates, by requiring them to choose between feeding and being in sight of their partner. There was a strong time of day effect favouring proximity to a mate early in the day for both males and females. A difference in male and female behaviour that would have suggested that this was male mate guarding was not found. Three independent measures show that this temporal effect is not owing to foraging being more important in the afternoon. There was a significant within-pair correlation in the strength of this proximity effect, which was strongest early in the day. Possible functional explanations are discussed. When females moved out of sight of males, male vocal behaviour increased and this increase was much greater when females moved out of sight earlier in the day than later on. These effects were not observed among females. This result is consistent with the hypothesis that male mate guarding is most important early in the day.     

Female reactions to male absence after pairing in the pied flycatcher

Mating with an already mated pied flycatcher (Ficedula hypoleuca) male is costly for a female. Two hypotheses explain why some females still mate with already mated males. The deception hypothesis suggests that some females mate with already mated males since it is difficult to assess perfectly the mating status of a male with separate territories (polyterritoriality). The search cost hypothesis states that females are aware of male mating status but the costs of searching for an unmated male exceed costs associated with the status of secondary female. One potential cue that could disclose a male's mating status is the existence of brief visits to the primary territory by polyterritorial males. To mimic such visits I removed the male from the territory for 60 min soon after female settlement. Only few females abandoned their mates as a consequence of male removal even though they had available unmated males close by. This result is most consistent with the deception hypothesis. Females did not use occasional male absence as a cue to avoid presumably polyterritorial males and in this respect they were not perfect in assessing males.     

No evidence for acoustic mate-guarding in duetting buff-breasted wrens (Thryothorus leucotis)

There are few empirical tests of the acoustic mate-guarding hypothesis for the function of duetting in birds. This hypothesis states that when females are fertile, males initiate many songs or answer most of their mates solo songs to form duets and repel rival males seeking extra-pair copulations. We tested the hypothesis by comparing song initiation and answer rates of males and females in socially monogamous buff-breasted wrens (Throthorus leucotis) during pre-fertile and fertile periods. During pre-fertile periods, males often sang for short periods before being answered by their mates, yet first duets were formed earlier relative to dawn and more duets were given during the dawn chorus on pre-fertile than fertile mornings. Males initiated more songs during pre-fertile than fertile periods, whereas there was no difference between stages in female song initiation rates. The proportion of songs answered by individuals of both sexes did not differ between breeding stages. Other mate-guarding behaviours, such as frequent copulation and maintaining close proximity to mates when fertile, did not appear to be important in this species, as no copulations were observed and there was no difference in the time pairs spent in close proximity when females were fertile or not. Parentage analysis revealed that only 3% of 31 broods had young that were likely the result of extra-pair paternity. These findings do not support the acoustic mate-guarding hypothesis, and suggest that the low rate of extra-pair paternity in buff-breasted wrens was maintained without the use of acoustic or traditional paternity guards.Communicated by R. Gibson     

Sex ratio and the sexual conflict about brood care in a biparental mouthbrooder

The pay-off of deserting and leaving a mate to care for the offspring alone is generally assumed to depend mainly on the availability of alternative mating partners and on the potential spawning rate of males and females. Eretmodus cyanostictus is a monogamous mouthbrooding cichlid in which the clutch is successively incubated first by the female and then by the male. It has been suggested that parents are constrained to monogamy due to low remating probabilities for both sexes. We tested this hypothesis by varying the sex ratio experimentally. Mate desertion by either sex was not significantly higher when additional potential mates were present (males: 8.3%, females: 0%) than when there were no other same-sex conspecifics present (males: 0%, females: 0%). Males lost their mate to a male intruder during their incubation in 26.7% of cases. Pair members were more active and showed more aggression when same-sex conspecifics were present. Behavioural differences between treatments were strongest during the incubation period of a given sex. If no desertion takes place, sexual conflict may be expressed also on a second level, the amount of parental care each parent provides. Indeed, males took the offspring later when additional females were present, although male incubation time did not differ between treatments. A hitherto undescribed display behaviour of females was clear evidence of a conflict about the timing of shift of young. In conclusion, offering alternative mating opportunities did not strongly favour mate desertion in E. cyanostictus. It rather revealed a conflict between mates about when to shift the young.Communicated by M. Abrahams     

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     

Sperm competition risk affects male mate choice copying

Mate choice copying was mostly described as a strategy employed by females to assess the quality of potential mates, but also males can copy other males’ mate choice. An open question in this context is whether and how copying males evaluate sperm competition risk, as mating with a female that has already copulated with another male obviously sets the stage for intense sperm competition (i.e., in species with internal fertilization). Using the livebearing Atlantic molly (Poecilia mexicana) as a model, we asked (a) whether males of that species indeed copy other males’ choices, and if they do so, (b) whether copying males strategically adjust their behavior to sperm competition risk. We used an approach where focal males could first choose to associate with a large or a small stimulus female. Mate choice tests were then repeated after an “observation phase” during which either no model male was present (treatment 1, control) or the previously non-preferred female could be seen associating (treatment 2) or physically interacting (treatment 3) with a model male. We found that, after the observation phase, males spent considerably more time with the previously non-preferred female in treatment (2), i.e., they copied the model male’s choice. This effect was much weaker during treatment (3) where sexual interactions between the model male and the formerly non-preferred female were allowed. Males, therefore, seem to adjust their copying behavior strategically to the perceived risk of sperm competition.     

Inter-sexual contests in the hermit crab <Emphasis Type="Italic">Pagurus bernhardus</Emphasis>: females fight harder but males win more encounters

Contests often occur between members of the same sex when they compete for access to mates, but inter-sexual contests may occur over access to other essential resources such as food or shelter. Despite the possibility that such contests are common, most studies focus on male fighting, and very few have analysed fights between males and females. Because males and females differ physically and physiologically, fighting ability or resource-holding potential (RHP) may also be subject to inter-sexual variation. In this study, we investigate size-controlled inter-sexual contests over the ownership of empty gastropod shells in the hermit crab Pagurus bernhardus. During these fights, there are two roles, attacker and defender, and we use a fully orthogonal experimental design to compare the performance of males and females in each role, when fighting either a male or female opponent. Although females fight more intensely, male attackers have an advantage when compared to females playing the attacker role, as they are more likely to evict the defender from its shell and thus win the resource. Further, in the defender role, male defenders are subject to shorter attacks than female defenders. The differences in agonistic performance could not be attributed to differences in perceived resource value between the sexes or to differences in body or weapon size. There are clear differences in the agonistic behaviour of males and females, and this possibility should be incorporated into models of contest behaviour. In particular, evolutionarily stable strategies may be expected to vary with sex ratios.     

Sexual selection in a socially monogamous bird: male color predicts paternity success in the mountain bluebird, <Emphasis Type="Italic">Sialia currucoides</Emphasis>

Ornamental traits are thought to evolve because they give individuals an advantage in securing multiple mates. Thus, the presence of ornamentation among males in many monogamous bird species presents something of a conundrum. Under certain conditions, extra-pair paternity can increase the variance in reproductive success among males, thus increasing the potential for sexual selection to act. We addressed this possibility in the mountain bluebird (Sialia currucoides), a socially monogamous songbird in which males possess brilliant ultraviolet (UV)-blue plumage. Specifically, we asked whether a male’s success at siring offspring within his own nest and within the nests of other males was related to his coloration. In pairwise comparisons, males that sired extra-pair offspring were not more colorful than the males that they cuckolded. However, males that sired at least one extra-pair offspring were, on average, brighter and more UV-blue than males that did not sire extra-pair offspring. Brighter, more UV-blue males sired more offspring both with their own mate and tended to sire more offspring with extra-pair mates and thus sired more offspring overall. Our results support the hypothesis that the brilliant UV-blue ornamental plumage of male mountain bluebirds evolved at least in part because it provides males with an advantage in fertilizing the eggs of multiple females.     

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.     

Mate guarding in the swallowHirundo rustica

Summary The importance of mate guarding by males in the monogamous swallowHirundo rustica was studied by temporarily detaining the males. Mate guarding reduced the frequency of extra-pair copulations and of sexual chases involving female mates. Males participated in sexual chases more frequently if they had a non-fertile female. Neighbouring males of ‘widowed’ females increased their own mate guarding presumably in response to the experimentally increased rate of sexual chases. Neighbouring males with a fertile female increased their mate guarding more than did males with a non-fertile female. Addition of eggs to swallow nests in the post-fledging period of the first brood induced mate guarding by male nest owners. These males also copulated more frequently with their mates than did control males. Neighbouring male swallows responded to the increased mate guarding by showing sexual interest in the guarded females. removal of eggs from swallow nests during the laying period, leaving only one egg in the nest, resulted in reduced nest attendance by females. Male mates responded by increasing their mate guarding intensity as compared to controls, and neighbouring males showed an increased sexual interest in these females.