Fighting behaviour as a correlate of male mating success in black grouse Tetrao tetrix

Fighting is a fundamental determinant of male fitness in species where females prefer socially dominant males as mates or where dominants can prevent subordinates from mating. This in turn can lead to the evolution of honest inter- and intra-sexual cues of male dominance. Fighting as a behaviour comprises both fighting rate (number of fights per unit of time) and fighting performance (success in winning fights), but it is not always clear which of these components are important for female choice and how they link to signals of male quality. To quantify the relative importance of fighting as a cue for females, we recorded detailed behavioural data from male black grouse Tetrao tetrix at leks. We explored the relationship between phenotypic traits (body mass, eye comb size, tail (lyre) length and blue chroma colouration) and fighting performance and rates and how these were related to male mating success. In older males' pairwise fights, winners had lower blue chroma than losers, but there were no differences in other morphological traits. In yearlings, no morphological trait predicted success in pairwise contests. Both fighting rate and performance were positively related to the number of copulations acquired by a male; however, when controlled for lek centrality, fighting performance and not fighting rate was significantly related to mating success. Our results indicate that females may be using components of fighting behaviour as cues for mate choice.     

Female control of offspring paternity in a western population of red-winged blackbirds (Agelaius phoeniceus)

Extra-pair copulations, which occur when individuals that have formed social relationships to breed copulate outside their pairbond, now are recognized as an important component of reproductive success in many species. In situations where both males and females benefit from extra-pair copulations without incurring much risk, an inevitable conflict arises between pairbonded mates. In this study I investigated the conflict of interest between male and female reproductive strategies in a western population of red-winged blackbirds (Agelaius phoeniceus). Female red wings in this population initiate extra-pair copulations, which resulted in a 35% rate of extra-pair fertilization. Females initiated the majority (78%) of extra-pair copulations away from their nesting territory where pairbonded individuals typically copulate, and females that engaged in extra-pair copulations spent a significantly greater amount of time off the marsh during peak fertilization compared to females that did not. In addition, females that nested in areas with a large number of potential extra-pair partners produced significantly more extra-pair fertilized young compared to females that nested on marshes with few male neighbors. Males’ strategies to protect paternity were limited primarily to patrolling territory boundaries and to opportunistically preventing extra-pair copulations off the marsh when they were visible. In this population females appear to use behavioral means to control nestling paternity, which in turn directly affected their mate’s reproductive success, and males were restricted to using strategies that were largely ineffective at preventing the threat of extra-pair paternity. Received: 23 December 1994/Accepted after revision: 17 December 1995     

Mate choice in the yellow-shouldered widowbird: correlates of male attractiveness

Recent investigations of male ornaments in sexual selection have used experimental manipulation of tail length in three widowbird species, but only for one of these have correlates of male reproductive success been reported. I examined correlates of male attractiveness to nesting females over two breeding seasons for the polygynous yellow-shouldered widowbird, Euplectes macrourus, in order to discover which cues females may be using to select mates. The black, long-tailed ( 10 cm) males defend large territories and build nest frames, or cock's nests, which females then line and use for nesting. I examined various aspects of male morphology, five behavioral displays, territory characteristics, and the number of cock's nests that males built. Few correlates of mating success were found. The best predictor was the number of cock's nests that a male builds, though one courtship display also correlated with male mating success in 1 year, as did average grass height. Tail length did not correlate with male mating success. A partial correlation analysis confirmed that cock's nests and, in 1 year, grass height, were the primary contributers to male success. Females may choose where to nest primarily on the availability of suitable nesting sites. Long tails may be used by females seeking extra-pair copulations or in male-male competition for territories.     

Spatial and temporal dynamics at manakin leks: reconciling lek traditionality with male turnover

Leks, display grounds where males congregate and females visit to copulate, are typically traditional in location, despite often high turnover of individual males. How leks can persist in face of male turnover is not well understood, in part due to a lack of detailed field data allowing for a clear understanding of lek dynamics. We followed the fate of individual males at 11 to 15 leks of the blue-crowned manakin Lepidothrix coronata across four breeding seasons to gain insights on how leks are formed and changed in space and time. Between years, leks were traditional in location despite changes in territory ownership due to male disappearance and recruitment. New males were equally likely to recruit by taking over existing territories or by establishing new territories. Recruitment was influenced by age, as recruits were more likely to be adults than subadults. Lek size did not affect the probabilities of a male recruiting or persisting at a territory, and vocalization rate, a correlate of mating success in this population, did not affect male persistence. We used our field data to model changes in lek size and composition over longer periods of time (100 years) to understand how lek traditionality can be reconciled with high male turnover. Our simulations showed that leks in our population rapidly stabilize in size despite changes in territory ownership and that rates of male recruitment and disappearance compensate each other, such that leks have the potential to persist for several decades after the original males have disappeared from them.     

The reproductive behavior of Anthidium manicatum (Hymenoptera: Megachilidae) and the significance of size for territorial males

Summary Males of the wool-carder bee, Anthidium manicatum, patrol clumps of garden plants. Females of this species visit these plants for pollen, nectar, and pubescence; they also mate there. Females are polyandrous, with intervals between copulations as short as 35 s. Patrolling males defend their territories (0.1–1.3 m²) against other males and against other species of flower-visiting insects. Honey bees may be rendered unable to fly by the attacks of A. manicatum.Territory owners perform exploratory flights to other males' territories, changing territories often (median ownership 4–7 days; maximum 30 days) and flying up to 450 m to establish new territories. Territorial usurpations are nearly always by larger males.Female visitation rate is significantly correlated with number of flowers on a territory. The head size of territory-owner males shows significant correlation with territorial quality (measured by number of flowers, not area) and thus with number of female visits and copulatory opportunities. Some males fail to maintain territories and instead attempt to forage and copulate in other males' territories while the owners are otherwise occupied. Nonowner males are significantly smaller than owners, forage less often and from fewer flowers, and achieve significantly fewer copulations than owners. Females, however, do not reject smaller, nonowner males at a higher rate than they do larger, owner males; their choice for male size appears to be indirect, based instead on choice of food resource.The interval between a copulation and the male's next attempt with a different female is not shorter than that involving the same female. Males do not escort just-mated females about their teritories, as observed in Anthidium maculosum. Territorial behavior in this species most likely evolved through intrasexual competition for reproductive success which led to sexual dimorphism. The defense of a resourcebased territory is the mechanism used by a male to maximize his reproductive potential.     

Sexual selection in a monomorphic lek-breeding bird: correlates of male mating success in the great snipe Gallinago media

Summary In the lek-breeding great snipe, male morphology, behaviour, and territory features were recorded for individually marked birds on two adjacent leks. Partial correlation sshowed that male mating success, expressed as the number of female solicitations and copulations, was negatively correlated with the distance of a display territory to the lek center and positively correlated with the number of displays per unit time given by a male. No other variables were directly correlated with male matin success. Thus, central males obtain more matings than peripheral males and successful males display more per unit time than do less successful males, independently of position on the lek. Central males were found to be older than peripheral ones and were present more often on the lek. Furthermore, central males had a larger number of white tail feathers, which are usea as visual signals in the displays, but this may be explained by the fact that these males were older. It is suggested that male great snipe are subject to sexual selection mainly in behavioural and vocal cues and that this may explain the absence of size and plumage dimorphism in this species.     

Mating success on white-bearded manakin (Manacus manacus) leks: male characteristics and relatedness

Morphological, territorial, and behavioural characteristics were measured on white-bearded manakin (Manacus manacus) males from two leks in two consecutive years. These data were combined with data on marker-inferred relatedness to study possible co-variation with mating success. In one year, male size and male condition were correlated with mating success. In both years, males holding courts nearer the lek centre gained more matings. No observed male display behaviour appeared to be an independently important factor in explaining variance in male mating success. Successful males made more aggressive displays than non-successful males and more displays were between close relatives. Number of aggressive displays increased as the distance between male courts decreased. Mating success in the white-bearded manakin is most likely mediated by a combination of morphological and behavioural characteristics, influencing both male-male competition and female choice. Females could potentially use centrality on the lek as an indicator of male characteristics. However, levels of relatedness may influence spatial arrangement of males on a lek thereby affecting male-male interactions and ultimately influencing patterns of mating success.     

Factors regulating annual mating success and reproductive success in spotted sandpipers (Actitis macularia)

Summary We investigated factors affecting annual mating success (MS) and reproductive success (RS) of spotted sandpipers (Actitis macularia) from 13 years of a 17-year study at Little Pelican Island, Leech Lake, Minnesota. Analyses were restricted to ages 1–3. Mean annual female MS varied from 1.3 to 2.7 mates, and the MS pattern was indistinguishable from random. However, female MS increased with age and was affected by arrival date, territory size, and beach size. Female RS also increased with age, and number of mates and year effects were the most significant explanatory variables in each age. Older female RS was increased by priority on a territory and presence of a previous mate. Territory size and beach size varied with population density and did not predictably affect RS. The strong year affect on RS was associated with annual variation in sex ratio and predation. Males produce only one successful clutch per year, so MS greater than one is a result of nest loss and does not increase RS. Neither male MS nor RS changed with age. Male reproductive failure rate varied by year. Given that a male produced young, the degree of RS was affected by year, arrival date, priority on a territory, territory size, and beach size. In years with early-season predation, late arrivals had higher RS; territory and beach size effects varied by year. Neither the presence, nor degree, of female care was associated with male RS. Male RS was more subject to annual environmental variability than was female RS, probably because of relatively low annual potential RS among males.Offprint requests to: L.W. Oring at the current address     

Females prefer larger leks: field experiments with ruffs (Philomachus pugnax)

Summary Preference by females for choosing mates at male aggregations has been hypothesized as the primary selective pressure favoring the formation of leks, but alternative hypotheses account for lek formation without invoking female preference. Observational studies to determine whether male mating success increases with lek size, as predicted under the female preference hypothesis, have produced inconsistent results, possibly due to covariation of lek size with other variables or to male-male or intersexual conflict over lek size. We tested whether females prefer larger leks in a field experiment with ruffs (Philomachus pugnax), a lekking sandpiper, in which male group size, composition, and location were controlled. Wild females chose the larger of two adjacent groups often enough such that males in larger groups had significantly higher per capita rates of female visitation (Table 3). Such behavior would probably lead to higher per male mating rates at larger leks, which is generally considered a necessary condition for female choice to select for lek display (Fig. 2). Lek size in nature will reflect both female preference for larger leks and competition among males, which may favor smaller lek size. All else being equal, however, female ruffs preferred to visit larger groups strongly enough to maintain lekking by males.     

Mixed reproductive strategy and mate guarding in a semi-colonial passerine,the swallow Hirundo rustica

Summary Both male and female swallows Hirundo rustica have a mixed reproductive strategy (parental care for offspring and extra-pair couplations). Mate guarding protects females from male harassment and male swallows from being cuckolded. Females hide their fertile period by copulating successfully with their mates for an extended period during first clutches. Males guard in the pre-fertile period, when many unpaired males are present. Early breeding swallows guard more than late breeders since more sexual chases of females by non-mate males take place in the early period. Solitarily breeding females experience few chases by strange males; copulation frequency, length of copulation period and mate guarding is adjusted to a lower level than among colonial birds. Male guarding activity is more intense in the fertile than in the pre-fertile period. Guarding reduces success of extra-pair copulation attempts.Three female swallows each paired and copulated with a single male and later changed to a new male before starting to breed. Extra-pair copulations most often take place between neighbouring swallows in the fertile period of the female. Many old, early breeding males and many young, late breeding females participate in extra-pair couplations. Successful extra-pair copulations peak in the fertile period contrary to success of pair copulations which does not change during the copulation period.     

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     

Territory establishment in lekking marine iguanas, Amblyrhynchus cristatus: support for the hotshot mechanism

The territory establishment of male marine iguanas and their subsequent mating success were analysed to identify spatial spillover (hotshot) and temporal spillover effects on lek formation. Males started to establish small display territories 2 months ahead of the mating season. Males did not establish territories in temporal synchrony and did not settle at sites where the probability of encountering females was highest. However, males arriving later preferentially established their territories in the neighbourhood of already established territories independently of the density of female-sized iguanas in these territories. Although settling in close proximity, there were no fights between those males. The number of fights between territorial males increased towards, and peaked during, the mating season. Fights did not result in the transfer of space, indicating that space per se was no resource. Instead, fights were directed towards central (hotshot) males. These central males had higher mating success than marginal males. Female density during the time of territory establishment did not predict the mating success of males, because females changed their spatial preferences between early establishment and mating periods. Similarly, the areas where males achieved the highest numbers of copulations changed during 4 years of our study. Thus, there was no evidence for temporal spillover between subsequent seasons. However, most male-male interactions served to distract successful males and may lead to spatial spillover of females into territories of unsuccessful males. In marine iguanas, territorial establishment appears largely governed by hotshot processes.     

Recruitment of mates and deceptive behavior by male Tengmalm's owls

Summary Tengmalm's owl Aegolius funereus is a hole-nesting polygynous species in which female nomadism is a reaction to cyclic lows of staple prey. For 2 years during a peak in vole abundance, I examined recruitment of mates and male singing behavior in a local population. Females and about half of the breeding males seemed to arrive successively throughout the breeding period of the first year. In contrast, a majority of the breeding males and likely also females were already on the breeding grounds at the beginning of the breeding season in the second year. Before breeding in the first peak year, males were singing at up to five different nest holes. All early breeding males (67% of total number) continued to sing at secondary nest holes after attracting and installing primary females. Secondary singing locations were never closer than 300 m from primary females' nest holes, although closer locations were used for singing before mating. All bigynous males with nests within the study area were also singing at tertiary nest holes, but none successfully attracted a third female. Polyterritoriality was confirmed by a late male settling between primary and secondary nest holes of a bigynous male. Polyterritorial behavior and reduced breeding success of secondary females are in line with the deception model. Females apparently could not discriminate between paired and unpaired singing males. Females made short visits to different males before mating. During courtship, the number and quality of nest holes defended by males may have been of particular importance to female choice.     

Sexual conflict over sperm ejection in monogamous pairs of kittiwakes<Emphasis Type="Italic"> Rissa tridactyla</Emphasis>

Socially monogamous partners suffer conflicting interests concerning various aspects of reproduction such as parental care, copulation and fertilization. Female black-legged kittiwakes commonly eject their mates' sperm immediately following copulations. Because sperm ejection reduces male sperm competitiveness and paternity assurance, males and females have conflicting interests as regards sperm ejection. Males whose mates ejected their sperm at least once remained longer on their mates' backs after the last insemination which apparently prevented the females from ejecting sperm. These results suggest that compelling females to retain their sperm may be a previously unidentified tactic employed by males to assure their paternity. Females tried to prevent their mates from witnessing sperm ejection by ejecting sperm after their mates departed from the nest. Females were more likely to eject sperm when they terminated the copulations by unbalancing the male. The conflict over sperm ejection was related to the ability of the females to end the copulations which covaried with the body mass of their mates. These findings suggest that conflicts in monogamous pairs also exist over the disposition of sperm.Communicated by C.R. Brown     

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     

Sexual selection and cues for female choice in leks of Jackson's widowbird Euplectes jacksoni

Summary Sexual selection through female mate choice was investigated in the lekking Jackson's widowbird by applying multivariate selection analysis to observational data from four leks. Males perform a stereotyped jump display on small display courts (dance rings) constructed by the males in open grassland. Females visit the lek solely for mating and nest on their own, away from the lek area. Few cases of interference during courtship and absence of position effects on mating success indicated that female choice within the leks was not pre-empted by male-male competition. In a set of 11 male traits with mating success as the dependent fitness measure, significant selection differentials (covariances) were found for the length of the conspicuous tail and the rate of the jump display, suggesting sexual selection of these traits. They also showed the largest selection gradients (partial effects) and thereby seem to be the cues on which females base their choice. The success of males in obtaining copulations appears to depend on two components: display rate and lek attendance affect the number of female visits, whereas tail length seems to primarily influence the chance of copulating with a visiting female. Tail length was positively related to a measure of body condition, which is of interest with regard to the suggestions that sexual ornaments may serve as indicators of male viability.     

Female habitat choice as a determinant of the reproductive success of the territorial male marine iguana (Amblyrhynchus cristatus)

Summary The reproductive behaviour of marine iguanas (Amblyrhynchus cristatus) was studied to answer the following questions: Which physical factors determine the number of females found on a male's territory? Do males select territories according to preferences for physical conditions or according to the number of females there? What causes the strong correlation between the number of females on a territory and the number of copulations achieved there? Females preferred as their resting places territories with the following characteristics, in order of importance: High number of shady/shelter places, low temperature of rocks at noon, proximity to the sea, sun-basking places in the afternoon. Other factors measured (probability of being wetted by spray, sun-basking places in the morning and at noon) showed no significant correlation with the number of females. Males, however, preferred territories further from the sea (within about 7 m), with high rock temperatures and a high number of basking places in the morning (in order of importance). Males primarily followed their site preferences and not female density in their choice of a territory. The site choice of females determined the number of copulations achieved on a territory, as females copulated preferentially on their resting places. A shift of females from one territory to another within one colony between two succeeding years was correlated with the change in relative weight of territory owners. This result was interpreted as an indication of female choice. Site preferences of males and females are discussed in relation to the animals' physiological requirements. Possible influences of ecological factors on the evolution of this mating system are considered.     

Female choice contributes to offspring fitness in the field cricket,Gryllus bimaculatus (De Geer)

Summary Females of the field cricket, Gryllus bimaculatus, are known to mate non-randomly with respect to male size. The reproductive output of females allowed to choose their mates was compared with that of females allocated mates. Females allowed to choose their mates laid a greater proportion of their available eggs, thereby investing more heavily in reproduction. Female choice did not appear to influence hatching success, suggesting that no short term benefits are derived from non-random mating. However, the offspring of females allowed to choose their mates developed more rapidly and began their own reproductive output before those of females who had been allocated mates. Furthermore, the faster developing offspring of females who chose their mates tended to have a consistantly higher survival rate as well as a lower variance in survival (i.e. less risk of mortality) at adult eclosion. Female choice may therefore contribute to offspring fitness and thus to long term reproductive success. Females allocated large males as mates did not equal those of females allowed to choose their mates, in terms of reproductive output and offspring fitness. This suggests that females may choose their mates with regard to a mixture of characters of which male size is only one.     

Mate sampling behaviour of black grouse females (Tetrao tetrix)

We studied female mate sampling behaviour in lekking black grouse (Tetrao tetrix). Females mainly visited males occupying territories in the centre of the lek with relatively large territories. They were also more likely to visit males that had high attendance. The same factors were also correlated with male mating success. A multiple regression model including these factors explained more of the variance in female visits per male (53%) than in mating success (33%). The pattern of female sampling conformed with a pool comparison (best-of-n) tactic. Such a tactic is expected if the costs of sampling are low. Females of high body mass visited more males than lighter females, however, which indicates that females may vary in their search tactics and suggests that there may be search costs. The existence of costs is further suggested by the fact that if the mate from a previous year was still present, females always mated with the same male in the following year. Though search costs were not measured directly, our findings suggest that some costs are negligible (e.g. energetic exhaustion or predation) whereas others (timing of mating) may be more important.     

Female mate choice in the Pied Flycatcher Ficedula hypoleuca

Summary We have examined male and territorial factors which might influence female mate choice in the Pied Flycatcher (Ficedula hypoleuca). Mating success of males was largely determined by the order of arrival on the breeding grounds. Females actively selected those males that had been longest in the area. This is likely to be due to territorial cues, early males having the best territories. About 15% of the males become polygynous, and these are the males that arrived earliest.Male age was relatively unimportant for breeding success, but because old males tend to arrive earlier than yearlings, most polygynous males were old. On average, old males are somewhat darker than young males, but male colour also varies within males of the same age. Polygynous and monogamous males did not differ significantly in colour, and female breeding success was not correlated with the colour of their mates. We show theoretically that it does not pay for females to select young males to avoid polygyny unless polygyny frequency or the number of reliable monogamous males almost double. Hence our hypothesis of polygyny-by-deceit in the Pied Flycatcher remains realistic. Because of male polyterritoriality, females are unaware of the males' status when mating.