Access to mates in a territorial ungulate is determined by the size of a male's territory, but not by its habitat quality

1. Territoriality is commonly associated with resource defence polygyny, where males are expected to gain access to females by anticipating how resources will influence female distribution and competing for resource-rich sites to establish their zone of dominance. 2. We tested this hypothesis in European roe deer (Capreolus capreolus) by simultaneously assessing the influence of resources on female distribution and the influence of female distribution on male distribution and breeding success using paternity analyses. 3. Females did not fully distribute themselves among male territories in relation to resources. As a result, relative female abundance in a male's territory depended on territory size, but not on its habitat quality. In turn, relative female abundance in a male's territory determined, at least partially, his breeding success. 4. Interestingly, male territory size, and hence access to females, was partly determined by male body mass (all males) and by residual antler size (subadults only). The latter result suggests that large antlers may be important to young males for establishing their first territory, which is then usually retained for all subsequent reproductive seasons. 5. To conclude, although territoriality of male roe deer has certainly evolved as a tactic for ensuring access to mates, our results suggest that it does not really conform to a conventional resource defence polygyny strategy, as males seem to gain no obvious benefit from defending a territory in an area of high habitat quality in terms of enhanced access to mates. 6. This may explain the stability of male territories between years, suggesting that male territoriality conforms to an 'always stay' and 'low risk-low gain' mating strategy in roe deer.     

Male mating strategies and the mating system of great-tailed grackles

Great-tailed grackles (Quiscalus mexicanus) are sexually dimorphic,dichromatic, colonially nesting blackbirds. In this study, males pursued three basic types of conditional mating strategies,each of which employed a different set of mating tactics. Territorialmales defended one or more trees in which several females nested.They achieved reproductive success by siring the offspringof their social mates and through extrapair fertilization.Resident males lived in the colony but did not defend territoriesor have social mates. Transient males passed through the colony, staying no more than a few days, and probably visited more thanone colony. Residents appeared to queue for access to territories,but transients did not. Residents and transients gained allpaternity through extrapair fertilizations and provided noparental care. Territorial males sired the majority of offspring,but residents and transients also sired small numbers of nestlings. Territorial males were larger and had longer tails than nonterritorialmales. The number of social mates was related to body size,and males that sired nestlings were heavier and had longertails than males with no genetic reproductive success. Malesthat gained paternity through extrapair fertilization wereheavier and had longer tails than males that did not. The matingsystem of great-tailed grackles can best be categorized as "non-faithful-female frank polygyny."     

Male brood provisioning rates provide evidence for inter‐age competition for mates in female Cooper's Hawks Accipiter cooperii

Life history theory predicts that individuals should maximize lifetime reproductive success (LRS) by breeding as soon as they reach sexual maturity, yet many species delay breeding, either because there are insufficient available mates or breeding sites, or because delayed breeding yields higher LRS. Accipitriform species, such as Cooper's Hawk Accipiter cooperii, exhibit both delayed breeding and delayed plumage maturation. However, in certain circumstances, first‐year females in non‐definitive plumage do breed and apparently compete with older females for high‐quality breeding territories. We predicted that these young females are at a competitive disadvantage compared with older females and that older females would have both higher reproductive success and be able to acquire higher quality nesting territories. We conducted brood counts and measured prey delivery rates by male Cooper's Hawks in an expanding urban population located in Albuquerque, New Mexico (USA), to assess our prediction. We found that older females had higher reproductive success, fledging 1.6 more offspring than younger females, and that they occupied territories where males provisioned at higher rates of 0.37 more prey items per 2‐h period. Our results showed that older females fared better than first‐year females but it is unclear if this is the result of passive or active competition. Older females initiated nesting 14.3 days sooner than first‐year females and thus may have filled vacant, high‐quality territories before first‐year females began seeking mates. Additionally, first‐year females were never observed persistently to confront older females for breeding territories, but they did actively compete against each other. First‐year females may defer to older females who, in a direct competitive interaction, would be most likely to prevail. Thus, delayed plumage maturation in Cooper's Hawks may serve to focus competition for nesting territories within age classes.     

South American sea lions in Peru have a lek‐like mating system

Five years of behavioral observations revealed significant effects of high air temperatures and breeding site topography on the mating system of South American sea lions in Peru. Unlike most polygynous mammals that defend females or fixed territories, male sea lions in Peru maintained positions along the shoreline where females passed each day to thermoregulate, and where most copulations occurred. Sex ratios (1 male per 17 females) and male mating success were extremely skewed (14% of males achieved 50% of the copulations, and 25% of them did not copulate at all). The mass daily movements of females toward the water and cool substrate of the shoreline, along with a highly skewed sex ratio, accentuated the difficulty for males to monopolize and restrict female movements. Females moved freely and chose their mates, unlike in temperate regions of their range where male South American sea lions control groups of females or access to tide pools. Our observations indicate that the South American sea lion in Peru has a lek‐like breeding system. This is a rare alternative to the common male strategies of defending females and resources, and is likely an evolutionary product of their highly skewed sex ratio, protracted breeding season, and the extreme subtropical climate where they breed.     

Female polyandry affects their sons’ reproductive success in the red flour beetle Tribolium castaneum

A potential benefit to females mating with multiple males is the increased probability that their sons will inherit traits enhancing their pre‐ or post‐mating ability to obtain fertilizations. We allowed red flour beetle (Tribolium castaneum) females to mate on three consecutive days either repeatedly to the same male or to three different males. This procedure was carried out in 20 replicate lines, 10 established with wild‐type, and 10 with the Chicago black morph, a partially dominant phenotypic marker. The paternity achieved by the sons of females from polyandrous vs. monandrous lines of contrasting morph was assessed in the F1, F2 and F3 generation by mating wild‐type stock females to two experimental males and assigning the progeny to either sire based on phenotype. The sons of polyandrous wild‐type females achieved significantly higher paternity when mating in the second male role than the sons of monandrous wild‐type females. By contrast, when mating in the first male role, males produced by females from polyandrous lines tended to have lower paternity than males from monandrous lines. Both effects were independent of the number of mates of the black competitor’s mother, and interacted significantly with the number of progeny laid by the female. These results provide the first evidence that manipulating the number of mates of a female can influence her sons’ mating success and suggest a potential trade‐off between offence and defence in this species.     

Costs and benefits of variable breeding plumage in the red-backed fairy-wren

The red-backed fairy-wren is a socially monogamous passerine bird which exhibits two distinct types of breeding male, bright males that breed in bright red and black plumage and dull males that breed in dull brown plumage. Most males spend their first potential breeding season in dull plumage and subsequent breeding seasons in bright plumage, but a relatively small proportion of males develop bright plumage in their first breeding season. This study quantifies morphology, behavior, and reproductive success of dull and bright males to assess the adaptive costs and benefits of bright plumage while controlling for age. Older bright males (two years of age or older) attempted to increase their reproductive success via copulations with extrapair females, whereas younger (one-year old) bright males and dull males did not. Thus, older bright males spent less time on their own territories, intruded on neighboring groups with fertile females more frequently, gave more courtship displays, and had larger sperm storage organs than did younger bright males and dull males. Microsatellite analyses of paternity indicate that the red-backed fairy-wren has extremely high levels of sexual promiscuity, and that older bright males had higher within-brood paternity than dull males or younger bright males. Regardless of age, bright males were more attractive to females in controlled mate choice trials than were dull males, and both age classes of bright males obtained higher quality mates earlier in the breeding season than did dull males, when nesting success was higher. In conclusion, although it appears that bright plumage increases access to higher quality mates, age also plays a central role in determining a male's overall reproductive success because of the high levels of sexual promiscuity exhibited by the red-backed fairy-wren.     

Effects of breeding success, mate fidelity and senescence on breeding dispersal of male and female blue-footed boobies

1. Understanding the effects of individual and population factors on variation in breeding dispersal (the movement of individuals between successive breeding sites) is key to identifying the strategies behind breeders' movements. Dispersal is often influenced by multiple factors and these can be confounded with each other. We used 13 years of data on the locations, mates, breeding success and ages of individuals to tease apart the factors influencing breeding dispersal in a colonially breeding long-lived seabird, the blue-footed booby Sula nebouxii. 2. Breeding dispersal varied among and within years. Males dispersed further in years of higher population density, and late breeding males and females dispersed further than early breeders. This temporal variation related to changes in competition for territory was taken into account in all tests of individual factors influencing breeding dispersal. 3. Individuals that retained their mates from the previous year dispersed shorter distances than those that changed their mates. 4. The effect of previous breeding success depended on mate fidelity. Unsuccessful breeding induced greater dispersal in birds that changed their mates but not in birds that retained their mates, indicating that breeders who change mates may take their own previous breeding experience into account during habitat selection. Faithful individuals may have to stay close to their previous sites to encounter their mates. 5. Male divorcees dispersed over shorter distances than their former mates, possibly because males contribute more than females to establishing territories. 6. Dispersal of males and females declined with increasing age over the first 10-11 years of life, then increased in old age, possibly due to senescent decay in the ability to compete for mates and territories.     

Fitness benefits of polyandry for experienced females

Females often mate with several different males, which may promote sperm competition and increase offspring viability. However, the potential benefits of polyandry remain controversial, particularly in birds where recent reviews have suggested that females gain few genetic benefits from extra‐pair mating. In tree swallows (Tachycineta bicolor), we found that females with prior breeding experience had more sires per brood when paired to genetically similar social mates, and, among experienced females, broods with more sires had higher hatching success. Individual females breeding in two consecutive years also produced broods with more sires when they were more genetically similar to their mate. Thus, experienced females were able to avoid the costs of mating with a genetically similar social mate and realize fitness benefits from mating with a relatively large number of males. This is one of the first studies to show that female breeding experience influences polyandry and female fitness in a natural population of vertebrates. Our results suggest that the benefits of polyandry may only be clear when considering both the number of mates females acquire and their ability to modify the outcome of sexual conflict.     

Reproductive consequences of bilateral asymmetry for individual male red-winged blackbirds

We used morphological and breeding data from a 2-year fieldstudy of red-winged blackbirds (Agelaius phoeniceus) to testthe hypothesis that males characterized by low levels of bilateralasymmetry (i.e., high developmental competence) realize a reproductiveadvantage. Specifically, we evaluated each of several distinctcomponents of male reproductive success relative to asymmetrymeasures made on five bilaterally paired characters. Resultsof a male removal experiment generally failed to support theprediction that symmetry would be associated with success incompetition for access to breeding territories: establishedterritory owners and nonterritorial replacement males were effectivelyindistinguishable in this regard. Similarly, there was no indicationthat symmetrical males were more likely to establish territoriesin high-quality marsh habitat than in marginal upland fieldhabitat. Finally, monitoring of breeding activity in high-qualityhabitat revealed that male symmetry was generally unrelatedto recruitment of social mates (i.e., harem size), the productivityof those mates (average female reproductive success), withinpairpaternity (assessed using DNA-based analysis of parentage),or extrapair mating success. Collectively these results indicatethat symmetry is not an important determinant of reproductivesuccess among individual male red-winged blackbirds. This observation,in combination with the results of several other recent investigations,suggests that the fitness consequences of subtle departuresfrom perfect symmetry may be less significant and/or less ubiquitousthan initially suggested.     

Correlates of reproductive success within alternative mating tactics of the common shrew

Male common shrews (Sorex araneus) adopt two discrete matingtactics. The most successful males, in terms of number of offspringfathered, are those that establish large overlapping home rangesin areas of high female density early in the breeding season.An alternative, less successful mating tactic is to travel longdistances in search of mating opportunities. This study is aninvestigation of correlates of reproductive success for malesadopting these different mating tactics. Reproductive successunder natural conditions was assessed using DNA fingerprinting,and survival of offspring was monitored in the field. The numberof offspring fathered by males with overlapping home rangeswas positively correlated with the number of female ranges overlappedduring the breeding season and with testes mass. The numberof offspring fathered by males that made long-distance movementswas positively correlated with their epididymal sperm counts.It is argued that competitively superior (overlapping) malesachieve high reproductive success by competing to maximize thenumber of females inseminated, whereas those adopting an alternativemating tactic instead compete largely via sperm competition,aiming to maximize insemination success with any particularfemale. There was no significant difference in the fluctuatingasymmetry (FA) of males adopting different mating tactics (FAwas measured as the difference in length of the paired lateralscent glands). Fluctuating asymmetry was not related to thenumber of offspring fathered by males adopting either matingtactic, but was significantly correlated with the proportionof male offspring fathered that survived to sexual maturity.Although apparently not correlated with mating success in thisspecies, FA may reflect some aspect of genetic quality thataffects offspring survival. [Behav Ecol 7: 334–340 (1996)]     

Division of labour during incubation in a woodpecker Colaptes auratus with reversed sex roles and facultative polyandry

The contribution of males to incubation has rarely been studied in altricial birds because the pursuit of extra mating opportunities is believed to conflict with incubation. Woodpeckers show reversed sex roles in parental care with males doing most of the nest construction, incubating and brooding of the young while females may be polyandrous. I investigated incubation by each sex at 71 monogamous and four polyandrous nests of the Northern Flicker Colaptes auratus , predicting that males would contribute to incubation according to their energy reserves (body condition) whereas females would contribute based on alternate reproductive opportunities. Nest attendance was 99% with males contributing a mean of 66% of the total incubation including nocturnal incubation. The length of daytime bouts averaged about 2 h and did not differ between the sexes. Consistent with predictions of investment strategies, structurally larger males and those in poorer body condition incubated less than smaller males, perhaps because they required more recess time to forage or to conserve energy. Older females contributed less incubation than young females and polyandrous females contributed less incubation at their secondary nests than monogamous females. Incubation period, nest depredation rate and hatching success were not influenced by bout length, number of bouts or relative contribution of the sexes. Hatching success was 86% at nests of both monogamous and polyandrous females because males compensated for reduced female participation. Because incubation of the sexes is compensatory and not additive, incubation pattern did not influence short-term reproductive success. I conclude that males invest in incubation according to their energy needs, and females may adjust their contributions based on alternate reproductive tactics.     

Social polyandry, parental investment, sexual selection, and evolution of reduced female gamete size

Abstract Sexual selection in the form of sperm competition is a major explanation for small size of male gametes. Can sexual selection in polyandrous species with reversed sex roles also lead to reduced female gamete size? Comparative studies show that egg size in birds tends to decrease as a lineage evolves social polyandry. Here, a quantitative genetic model predicts that female scrambles over mates lead to evolution of reduced female gamete size. Increased female mating success drives the evolution of smaller eggs, which take less time to produce, until balanced by lowered offspring survival. Mean egg size is usually reduced and polyandry increased by increasing sex ratio (male bias) and maximum possible number of mates. Polyandry also increases with the asynchrony (variance) in female breeding start. Opportunity for sexual selection increases with the maximum number of mates but decreases with increasing sex ratio. It is well known that parental investment can affect sexual selection. The model suggests that the influence is mutual: owing to a coevolutionary feedback loop, sexual selection in females also shapes initial parental investment by reducing egg size. Feedback between sexual selection and parental investment may be common.     

EXTRAPAIR MATE CHOICE AND HONEST SIGNALING IN COOPERATIVELY BREEDING SUPERB FAIRY-WRENS

In many species of monogamous birds females copulate with males other than their social mates, resulting in extrapair fertilizations. Little is known about how females choose extrapair mates and whether the traits used to choose them are reliable indicators of male quality. Here we identify a novel male trait associated with extra-group mating success in the superb fairy-wren (Malurus cyaneus), a cooperatively breeding bird with one of the highest known frequencies of extra-group mating. Female fairy-wrens chose extra-group mates that molted earlier into breeding plumage. Males molted up to five months before the breeding season began, and only males that molted at least one month prior to its onset gained any extra-group fertilizations. This conclusion held after controlling statistically for the effect of age and social status on molt date. Once males acquired breeding plumage, they began courtship display to females on other territories. Thus, some males were displaying to females for several months before the breeding season began. This extraordinarily long period of advertisement by males may be facilitated by the long-term ownership of territories. We suggest that early acquisition of breeding plumage or the subsequent display behavior can be reliable cues for mate choice because they are costly to acquire or maintain.     

Tactic‐specific benefits of polyandry in Chinook salmon Oncorhynchus tshawytscha

This study examined whether polyandrous female Chinook salmon Oncorhynchus tshawytscha obtain benefits compared with monandrous females through an increase in hatching success. Both of the alternative reproductive tactics present in male O. tshawytscha (large hooknoses and small, precocious jacks) were used, such that eggs were either fertilized by a single male (from each tactic) or multiple males (using two males from the same or different tactics). The results show that fertilized eggs from the polyandrous treatments had a significantly higher hatching success than those from the monandrous treatments. It is also shown that sperm speed was positively related with offspring hatching success. Finally, there were tactic‐specific effects on the benefits females received. The inclusion of jacks in any cross resulted in offspring with higher hatching success, with the cross that involved a male from each tactic providing offspring with the highest hatching success than any other cross. This study has important implications for the evolution of multiple mating and why it is so prevalent across taxa, while also providing knowledge on the evolution of mating systems, specifically those with alternative reproductive tactics.     

Do female black field crickets Teleogryllus commodus benefit from polyandry?

Female insects that mate multiply tend to have increased lifetime fitness, apparently because of greater access to male-derived resources (e.g. sperm, nuptial gifts) that elevate fertility/fecundity. Experiments that standardize the number of matings per female also show that polyandry can improve aspects of offspring performance, most notably early embryo survival (egg hatching success). This improvement is widely attributed to genetic benefits which would arise if polyandrous females skew paternity to produce fitter offspring. In two separate experiments with field crickets (Teleogryllus commodus) polyandrous females (two, three or four mates) did not have higher egg hatching success than monandrous females (effect sizes: r = 0.03 and 0.08 for the respective experiments), which is consistent with our finding of no sire effect on hatching success. Polyandry also had no effect on post-hatching offspring survival. Polyandrous females' offspring took significantly longer to mature but their sons were not heavier and their daughters were actually significantly smaller than those of monandrous females. Finally, after controlling for relative male size, monandrous females' sons were more successful when directly competing for a mate.     

Sexual selection for large male size in a polyandrous butterfly: the effect of body size on male versus female reproductive success in Pieris napi

Sexual selection theory predicts that the larger sex shouldbe that for which fitness increases at the faster rate withsize. In butterflies, as in most invertebrates, females areusually the larger sex, but previous comparative analysis hasshown that relative male size increases with female polyandryamong butterflies. In agreement with this pattern, males arelarger than females in the strongly polyandrous green-veinedwhite butterfly, Pieris napi L., and in this article we assessthe size dependence of reproductive success in both sexes. Inan experiment where virgin males and females were released inthe field, we found no strong association between size and malemating success. However, laboratory experiments showed thatthere was a strong correlation between size and the ejaculatethat the male delivered to the female at mating and that largeejaculates delayed female remating for a longer time comparedto small ejaculates. Moreover, female P. napi utilize male-derivednutrients received at mating to increase their fecundity. Hence,large males sire more offspring both by way of donating morenutrients to female egg production and by way of delaying femaleremating (given that the last male to mate with the female willfather most of the offspring). Laboratory experiments showedthat the association between size and fecundity was low, ornonexistent, among P. napi females allowed to mate only once.However, weak size dependence was found for polyandrous females.We hypothesize that size dependence of female fecundity maybe especially weak among polyandrous butterflies because a fundamentalsource of variation in fecundity relates to their ability tofind nutrient giving males, an ability which may be unrelatedto female size. According to this hypothesis there is a causalassociation between weak size dependence of female fecundityand polyandry, and a strong size dependence of male reproductivesuccess that may underlie the comparative pattern of positivecorrelation between relative male size and polyandry.     

Male mate choice, male availability and egg production as limitations on polyandry in the red-necked phalarope

In sequentially polyandrous birds, a female's second mate faces a substantial risk of cuckoldry due to rapid mate switching and stored sperm. Secondary males are potentially available to females because males arrive asynchronously and/or are recycled into the breeding pool following nest predation. In a study of red-necked phalaropes, Phalaropus lobatus, a sex-role reversed shorebird, we tested the hypotheses that the proportion of females that become polyandrous is proximately limited by: (1) the ability of females to produce eggs, (2) the availability of males as mates and (3) male mate choice. In a colour-banded population in which rates of nest loss were manipulated by researchers, females that produced second clutches required similar lengths of time to complete clutches as those contemporaneously producing first clutches, and increased their egg size relative to their first clutch, making egg limitation unlikely. There was no correlation between an annual measure of males' availability as potential mates following nest losses and the proportion of females that were polyandrous. The majority of males that lost clutches (66%) re-paired with their original female significantly more often than expected by random mate choice (P<0.0001). Although 76% of polyandrous nestings involved renesting males, only 6% (N=46) of renesting males changed mates if their original female was still available. Renesting males that changed mates did not select for or against females that had already produced clutches (NS). Our results suggest that the level of polyandry in this species is not constrained by the females' abilities to produce more eggs or by the number of males recycling back into the breeding pool. Instead, the proportion of females that become polyandrous is limited by males choosing to renest with their original females, thereby decreasing their probability of caring for eggs potentially fertilized by a female's previous mate.     

Yelling for sex: harem males compete for female access in bronze-winged jacanas

Sperm competition in sex-role reversed, polyandrous jacanas is intense because females copulate with multiple male mates before laying each clutch. These males may be unable to attempt to maximize their share of copulations by mate guarding or forcing copulations. Instead, males in polyandrous harems may compete for sexual access to the female by giving a call, termed the 'yell', to attract her. Male bronze-winged jacanas, Metopidius indicus, yelled at higher rates in larger harems, and when the female was further from the yeller or on a comate's territory. Half of all yells were given at mating platforms where all copulations occurred. Males that received the clutch yelled at lower rates during the incubation and chick care periods. Yells attracted the female when she was far from the yeller or with a comate. When the yell of a polyandrous male was broadcast from his territory, the female was more likely to fly to his territory during playback than during control periods. Within polyandrous harems the males that yelled at the highest rates received the most copulations, and three out of four females gave clutches to the male that gave the longest and most frequent yells, so females may have used yells to assess male quality. Intrusions by females, but not males, increased during yell playbacks, and tended to be more frequent on the territories of males with high yell rates. Females may therefore respond to their mates' yells because yells may attract female intruders which may attempt to take over the territory. Copyright 1999 The Association for the Study of Animal Behaviour.     

Sexual selection in sticklebacks in the field: correlates of reproductive, mating, and paternal success

Male sticklebacks display multiple ornaments, and these ornamentshave been shown to be preferred by females in laboratory experiments.However, few field data exist, and it is not known whether thesepreferences are simultaneously or sequentially operative ina single population. We report correlates of reproductive successin two stickleback populations that differ in their ecology,over several periods within their breeding season. In both populationslarger males had higher reproductive success, but not in all periodsof the breeding season. Reproductive success increased withredness of the throat only in the Wohlensee population, andonly in one period that was characterized by low average success.In the Wohlensee population, the parasitic worm Pomphorhynchuslaevis is abundant, and reproductive success decreased withthe presence of the parasite. In the Roche population, maleswith nests concealed in a plant had higher mating success. Thesenests were less likely to fail, suggesting that females preferredto spawn in concealed nests because of higher offspring survivorship.The different sexual traits appear to reveal different aspectsof male quality (multiple message hypothesis): females probablyfind large males attractive because of their higher paternalquality, but it seems more likely that red males are preferred forbetter genetic qualities. Females also discriminate on territoryquality, and male traits may be important in competition forthese territories. The correlates of reproductive success werenot consistent during the season, probably due to changes inthe availability of ripe females. Such fluctuating selectionpressures will contribute to the maintenance of genetic variationin sexual traits.     

Emasculation to plug up females: the significance of pedipalp damage in Nephila fenestrata

Female multiple mating selects for male adaptations that maximizefertilization success in a context of sperm competition. Whilemale mating strategies usually reflect a trade-off between presentand future reproduction, this trade-off is largely removed insystems where the maximum number of matings for males is verysmall. Selection may then favor extreme mechanisms of paternityprotection that amount to a maximal investment in a single mating.Males in several arthropod taxa break off parts of their copulatoryorgans during mating, and it has frequently been suggested thatmutilated males can thus secure their paternity. Nevertheless,such a mechanism has rarely been confirmed directly. Here westudy the golden orb spider Nephila fenestrata, which has amating system with potentially cannibalistic, polyandrous females,and males that are often functionally sterile after mating withone female only. We show that males in this species can indeedprotect their paternity by obstructing the female's genitalopenings with fragments of their copulatory organs.