Focal Males Research Articles

Male fiddler crabs defend multiple burrows to attract additional females

Males of many species defend resources to attract females. Surprisingly, defense of multiple female breeding sites (e.g., nests or burrows) appears to be rare, primarily reported in fish and birds. In fiddler crabs, burrows are a vital resource for reproduction and survival. Both sexes defend individual territories centered on a single burrow. We examined burrow acquisition and defense in Uca capricornis to test whether males defend multiple burrows as a novel strategy to acquire additional mates. When crabs were experimentally forced to acquire a new burrow, females often settled into an empty burrow near resident males. We documented more empty burrows around males than expected by chance and, in addition, larger males had a greater proportion of empty burrows in their immediate vicinity. We experimentally introduced crabs into empty burrows next to focal males: newly introduced males were soon evicted, whereas females were courted and stayed. These results suggest that male U. capricornis defend empty burrows as a strategy to obtain more mates. Intriguingly, however, U. capricornis tend to occur in socially monogamous pairs. This raises the possibility of sexual conflict within social pairs over the presence of additional females and that female‐ female competition might constrain male mating success. Key words: mating systems, polygamy, resource-holding potential, resource monopolization, sexual conflict, sexual selection. [Behav Ecol]

Effects of Breeding Stage and Behavioral Context on Singing Behavior of Male Indigo Buntings

Abstract We studied the effects of breeding stage and behavioral context on the singing behavior of male Indigo Buntings (Passerina cyanea; n = 15) during the 2004 breeding season in Madison County, Kentucky, USA, to better understand how males with a single-song repertoire vary the characteristics of their song to convey different information. Playback experiments were conducted in 2005 in territories of focal males (n = 14) to further examine the possible effect of male-male interactions on singing behavior. We analyzed 10,919 songs of 15 male Indigo Buntings with songs consisting of a series of figures that were usually paired (i.e., phrases). Mean song duration was 2.30 ± 0.13 (SE) sec (range = 1.44–3.40 sec) with males varying song duration by varying the number of figures and phrases in each song. Singing rates varied significantly (P < 0.0001) among breeding stages and were highest prior to pairing, suggesting singing has a role in mate attraction. Singing rates also differed (P = 0.013) du...

Enhancing male sexual success in a lekking fly ( Anastrepha suspensa Diptera: Tephritidae) through a juvenile hormone analog has no effect on adult mortality

While defending lek-territories, male Anastrepha suspensa (Loew) produce chemical, acoustic and visual courtship signals. In the laboratory and under semi-natural conditions, topical application of the juvenile hormone analog methoprene doubles pheromone production and subsequently doubles sexual success. However, sexual signals and interactions are likely to be physiologically expensive and so result in higher male mortality. Comparison of males kept in isolation for 35 days, but provided daily with a potential mate or a rival male, revealed that both male- and female-interactors shortened focal-male lifespan. In addition, focal males were either treated with methoprene or not, then either provided with protein in their sucrose-based diet or not. Protein proved to similarly double sexual success and also resulted in longer male life spans in all of the interactor-categories. However, there was no evidence that methoprene induced hypersexuality resulted in higher rates of mortality, i.e., the longevity of males treated with methoprene did not significantly differ from untreated males in the same interactor/diet categories. This apparent lack of costs to a putatively sexually selected signal is unexpected but presents an opportunity to increase the sexual competence of sterile flies with few consequences to their survival following mass-release.

Interactions between inter- and intrasexually selected behaviors in the beaugregory damselfish (Stegastes leucostictus)

There is a trade-off between mutually incompatible behaviors involved in conspecific aggression and courtship by territorial male beaugregory damselfish (Stegastes leucostictus). This trade-off may be variable based on the proximity of each stimulus. We first tested whether presenting male and female conspecific stimuli at varying distances from the breeding site influenced the territorial male's behavioral response by placing a single female or a single male in a clear plastic bottle 1 m or 2 m from the breeding site of the focal male. We found that the focal males responded similarly to females at either distance while they decreased their response to males at the greater distance. To test how distance influences the trade-off between inter- and intrasexually selected behaviors, we simultaneously placed bottled males and females near a male's breeding site. Reducing the proximity of one stimulus resulted in an increased response to the other stimulus. We suggest that the trade-off between behaviors is variable and show that distance plays a role in territorial males' behavioral response to multiple conspecific stimuli.

Misleading mollies: surface- but not cave-dwelling Poecilia mexicana males deceive competitors about mating preferences

Individuals providing misleading information to conspecifics may benefit from deception at the receiver's expense. A recent study (Plath et al., Curr Biol 18:1138–1141, 2008c) suggested that male Atlantic mollies (Poecilia mexicana) deceive rival males about their preferred mate. Here, we contrasted potentially deceptive behavior in surface-dwelling P. mexicana males to males of the cave form of that species (the cave molly). Unlike many other cavefishes, cave mollies have retained functional eyes and readily respond to visual stimuli. Males could interact freely with two females (large and small), and an audience male was visually presented during the second part of the tests. When observed during mate choice, males reduced their mating activity, but this reduction was significantly weaker in cave mollies. Overall, the expression of mating preferences (determined through frequencies of nipping and thrusting) declined in front of an audience; again, this effect was significantly weaker in the cave form. Reduced sexual activity and reduced expression of mating preferences can be interpreted as an attempt of the focal male to avoid unintended interception of information by the rival male. Surface but not cave molly males directed their first sexual interaction (when being observed by the rival male) towards the initially non-preferred female, suggesting that surface-dwelling males deceive rival males about their mating preferences. Deception by the focal males may be an adaptation to avoid sperm competition, since other males in their social environment may use public information and copy the focal male's mate choice. It seems that sending deceptive signals is evolutionarily regressed in the cave molly, since mate choice copying is unlikely to occur under naturally dark conditions, and also the potential to deceive rivals about mating preferences is probably very limited.

Using logistic regression models to predict breeding success in male Adélie penguins (Pygoscelis adeliae)

Measures of breeding success are traditionally derived using the proportion of total nests that successfully reach strategic stages across a breeding season, such as pair formation, egg laying or fledging chicks. The use of logistic regression has recently become a popular tool in avian literature for identifying influential factors that predict nest and/or individual breeding success. In this study, we use logistic regression models to assess the importance of a range of factors affecting male Adelie penguin (Pygoscelis adeliae) breeding success during the 2002–2003 austral summer, when the presence of icebergs and extensive sea-ice meant overall breeding success of Adelie penguins was low (16/51 focal males had chicks reach fledging point). Logistic regression models for the early breeding season showed that good/average nest quality and central/middle nest location within the sub-colony were the best predictors of successful pair formation. Later, during incubation, the most successful males were those that not only returned earlier but were also heavier upon arrival and built nests of at least average quality. During the final stage when chicks had begun to fledge, the combined parameters of heavier male weight, early arrival time and good nest quality were the best predictors of breeding success. The logistic regression approach used here showed that the predictive ability of these parameters varied as the season progressed. However, most importantly, our logistic models fit the data well across all breeding stages.

Calling strategies of male túngara frogs in response to dynamic playback

AbstractTúngara frogs produce calls of varying complexity that consist of a whine followed by 0–7 chucks. In previous studies using static playbacks, males increase chuck number in response to calls with 1 versus 0 chucks but not in response to 2 or 3 versus 1 chuck. Here we use dynamic playback in which an automated interaction program counts the number of chucks present in a call and responds according to a specified calling strategy which determines the number of chucks in the response call. Males added progressively more chucks in response to the models' strategies of de-escalate, match or escalate; there was a significant difference in the focal males' responses to the de-escalate versus the escalate strategies. Similarly, males changed their calling strategy in response to the de-escalate strategy of the model. There was no evidence of change in calling patterns, as estimated by entropy, among treatments and between experimental and field data. Males produced significantly more chucks in these experiments than in the field, but in both contexts the mean chuck number is low and never approaches the maximum observed in nature. These data suggest that males are cost-sensitive when it comes to adding chucks and that they are more influenced by vocal competitors that de-escalate rather than escalate number of chucks. These are patterns of calling not previously revealed in studies using static playbacks, and this study is another demonstration of the usefulness of dynamic playback in studies of animal communication.

Black-capped chickadee dawn choruses are interactive communication networks

Summary The dawn chorus of songbirds provides an ideal opportunity to study communication networks because multiple singers are within signalling range of each other, permitting eavesdropping by both males and females. Using an Acoustic Location System, we examined the dawn chorus singing behaviour of male black-capped chickadees (Poecile atricapillus )i n 15 neighbourhoods to determine whether singing behaviour is consistent with the communication network model. We calculated levels of frequency matching for 19 focal males and all of their neighbours. The observed level of frequency matching was greater than expected by chance. All males were involved in multi-way matching at dawn and often matched two or three neighbours simultaneously. The identity of individuals involved in three-way matches was related to both previous winter-flock membership and the relative dominance rank of the interacting males. We show that male black-capped chickadee dawn choruses are interactive communication networks where males are involved in high levels of matching with neighbours, and they match multiple individuals both simultaneously and sequentially. Additionally, the existence of multi-way matching and the identities of individuals involved suggest that individual males may eavesdrop at dawn. This is the first study to quantify network communication during the dawn chorus in multiple neighbourhoods.

Energetic costs of male reproduction in a scramble competition mating system

1. The assumption that the primary limitations on reproductive success differ between the sexes is inherent in traditional sexual selection theory. Although the energy that can be allocated to reproduction is assumed to be the main limitation to females, the ability to attract and defend oestrous females is assumed to be the primary limitation to males. 2. Estimates of the energetic costs of reproduction in male mammals are, however, limited and have largely been obtained from sexually dimorphic species exhibiting female defence mating systems. These studies often reveal that the energetic cost of male reproduction is similar to or even exceeds that of females, and therefore challenge long-held assumptions of inter-sexual reproductive limitations, but their generality is little known. 3. We coupled measurements of energy expenditure with detailed behavioural observations of reproductive male North American red squirrels (Tamiasciurus hudsonicus). This species displays minimal sexual dimorphism and exhibits a scramble competition mating system, under which sexual selection favours enhanced mate searching effort by males. 4. We conducted the study over 2 years characterized by a substantial variation in upcoming natural food availability and across two study populations that experienced either natural food abundance or an ad libitum food-supplementation to investigate the influence of resource availability on male reproductive energy expenditure. 5. Under natural conditions, mean energy expenditure of males across the 2 years was high, approximating that of females during lactation. Furthermore, in the anticipation of high upcoming natural food availability and resultant offspring survival, expenditure approximately doubled (from 290 +/- 7 to 579 +/- 73 kJ day(-1)). When current food availability (and consequently the density of receptive females) was experimentally elevated, males displayed the highest levels of energy expenditure we recorded (873 +/- 98 kJ day(-1)). 6. Our results provide compelling evidence that the energy available for reproductive allocation places a strong limitation on reproduction in male North American red squirrels and contribute to previous work suggesting that high and limiting energetic costs of male reproduction may be a general feature of mammalian reproduction.

Muscle Metabolic Capacities and Plasma Cortisol Levels of the Male Three‐Spine Stickleback Gasterosteus aculeatus: Are There “Femme Fatale” or “Macho Male” Effects?

To evaluate whether decreases in muscle metabolic capacities and increases in plasma cortisol explain the effects of neighboring conspecifics on male three-spine sticklebacks Gasterosteus aculeatus, we housed mature males alone, with a mature female, or with a rival mature male. The neighbors were separated from the focal male by a partition that allowed him to smell, see, and hear his neighbor. In the first experiment, focal males were allowed to reproduce, whereas in the second experiment, no reproduction occurred. Coloration and behaviors were monitored while the males tended their nests (or for the same period in the second experiment). The presence of a neighbor markedly affected the reproductive coloration of the focal male, with solitary males being less colorful than males housed with a rival male or a female. Solitary males showed greater aggression toward a model male stickleback than did males with neighbors. The presence of neighbors affected the anatomic and metabolic characteristics of the focal males primarily during nesting, when males housed with rival males had a lower hepatosomatic index and lower activities of mitochondrial and glycolytic enzymes in the axial muscle than did solitary males or males housed with females. Cortisol levels were highly variable in nesting males and did not differ with social condition but were higher in males that had been quick to construct their nests. On the other hand, when focal males were not provided with nesting material, solitary males tended to have lower cortisol levels than did males housed with rival males. While these results do not provide a mechanism for a "femme fatale" effect, they indicate that nesting males decrease metabolic status when housed with rival "macho males."

Homosexual interactions in bed bugs: alarm pheromones as male recognition signals

Homosexual mounting is a common behaviour in bed bugs as male sexual interest is directed towards any newly fed individual. The only mode of copulation in the common bed bug, Cimex lectularius, is by traumatic insemination, where the male pierces the female abdomen with his needle-like penis. Homosexual mating would result in abdominal injuries in mounted males, as males lack the female counteradaptive spermalege structure. I here show that bed bug alarm pheromones, previously hypothesized to be a predator chemical defence, can be used by newly fed males to signal their sex and reduce the risk of homosexual mating. Mechanical blocking of the male pheromone glands significantly increased homosexual mounting duration compared to control males, while applying male extracts containing mainly alarm pheromone onto male-female mating pairs completely interrupted or shortened mating duration and reduced sperm transfer. Males confined with other males received piercing scars, demonstrating that homosexual mating occurs. The focal males in the all-male confinement experiment had reduced longevity compared to singly held males, but why this reduction in longevity occurred is not clear. Mounted males thus benefit from being able to discharge alarm pheromones, while mounting males consider the alarm signal a major sex identification cue, suggesting that male bed bugs use alarm pheromone communication to avoid homosexual harassment and mounting. (C) 2009 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Effects of extreme variation in female morph frequencies on the mating behaviour of male damselflies

Female-limited polymorphism is often attributed to selection to avoid excessive male mating attempts. It is encountered in various taxonomic groups, but is particularly common in damselflies, where one female morph (andromorph) typically resembles the conspecific male in colour pattern, while the other(s) (gynomorph(s)) do not. Two sets of theories have been proposed to explain the phenomenon in damselflies, which can be classified as the learned mate recognition (LMR) and male mimicry (MM) hypotheses. To test predictions of these hypotheses, we evaluated the rate of male sexual response towards female morphs and conspecific males in the damselfly Nehalennia irene. The LMR hypothesis predicts that males should respond sexually to andromorphs at greater rates in populations containing a higher relative frequency of andromorphs. The MM hypothesis predicts that males respond more often sexually to both andromorphs and males as the ratio of andromorphs to males increases. While LMR predicts that the rate of mating attempts towards gynomorphs should vary, the MM predicts that it should be relatively fixed. On experimentally presenting live specimens to focal males in five different populations with extreme variation in female morph frequencies, we observed that as the andromorph frequency and ratio of andromorphs to males increased, the proportion of male mating attempts increased on both andromorphs and males, whereas it decreased on gynomorphs. While the simplest form of the MM hypothesis is rejected, the results support specific predictions of both hypotheses and suggest that future studies should not treat these hypotheses as mutually exclusive.

Audience effects in the Atlantic molly (Poecilia mexicana)–prudent male mate choice in response to perceived sperm competition risk?

BackgroundMultidirectional interactions in social networks can have a profound effect on mate choice behavior; e.g., Poecilia mexicana males show weaker expression of mating preferences when being observed by a rival. This may be an adaptation to reduce sperm competition risk, which arises because commonly preferred female phenotypes will receive attention also from surrounding males, and/or because other males can copy the focal male's mate choice. Do P. mexicana males indeed respond to perceived sperm competition risk? We gave males a choice between two females and repeated the tests under one of the following conditions: (1) an empty transparent cylinder was presented (control); (2) another ("audience") male inside the cylinder observed the focal male throughout the 2nd part, or (3) the audience male was presented only before the tests, but could not eavesdrop during the actual choice tests (non-specific sperm competition risk treatments); (4) the focal male could see a rival male interact sexually with the previously preferred, or (5) with the non-preferred female before the 2nd part of the tests (specific sperm competition risk treatments).ResultsThe strength of individual male preferences declined slightly also during the control treatment (1). However, this decrease was more than two-fold stronger in audience treatment (2), i.e., with non-specific sperm competition risk including the possibility for visual eavesdropping by the audience male. No audience effect was found in treatments (3) and (5), but a weak effect was also observed when the focal male had seen the previously preferred female sexually interact with a rival male (treatment 4; specific sperm competition risk).ConclusionWhen comparing the two 'non-specific sperm competition risk' treatments, a very strong effect was found only when the audience male could actually observe the focal male during mate choice [treatment (2)]. This suggests that focal males indeed attempt to conceal their mating preferences so as to prevent surrounding males from copying their mate choice. When there is no potential for eavesdropping [treatment (3)], non-specific specific sperm competition risk seems to play a minor or no role. Our results also show that P. mexicana males tend to share their mating effort more equally among females when the resource value of their previously preferred mate decreases after mating with a rival male (perceived specific sperm competition risk), but this effect is comparatively weak.

EFFECTS OF FERTILIZATION DISTANCE ON MALE GAIN CURVES IN A FREE-SPAWNING MARINE INVERTEBRATE: A COMBINED EMPIRICAL AND THEORETICAL APPROACH

Male gain curves describe the relationship between allocation to sperm production and male reproductive success and are central to models of sex allocation in hermaphrodites. Sperm competition is expected to result in more linear gains and select for increased allocation. We hypothesized that high sperm production in passively mating systems may also be the result of selection to enhance the ability to fertilize distant ova. Consequently, we explored the effect of distance on male gain curves in a free-spawning colonial ascidian. The performance of focal males that varied in sperm production was assayed at three distances via microsatellite markers. An advection-diffusion model was used to estimate sperm concentration gradients, to predict male reproductive gain integrated across multiple downstream females, and explore effects of hydrodynamic conditions. As distance increased, male reproductive success decreased and empirical gain curves became increasingly linear. Our model predicted that the expected net gain curve is relatively insensitive to variation in flow regime and will saturate much more slowly than if only a single, nearby distance is considered. Thus, high levels of sperm production may enhance fitness both in competitive situations and with increasing fertilization distance, highlighting the need to consider distance effects when evaluating gain curves.

Group unity of chimpanzees elucidated by comparison of sex differences in short-range interactions in Mahale Mountains National Park, Tanzania

Chimpanzees (Pan troglodytes) form multi-male and multi-female unit groups with fission-fusion grouping patterns. Short-range interaction (SRI) plays an important role in the unity of these groups and in maintaining social bonds among members. This study evaluated three models of chimpanzee social structure that differed according to the emphasis each placed on social bonds between the sexes, i.e., the male-only, the bisexual, and the male-bonded unit-group model. I investigated differences in SRI between the sexes among group members in well-habituated wild chimpanzees in Mahale Mountains National Park, Tanzania. I followed six focal adult males and six females, and quantified their respective SRI with other chimpanzees. Except between subordinate males and adult females, adults in general engaged in SRI with about 60-90% of the individuals with whom they made visual contact each day, whether in large or small parties. Although the number of social grooming (SGR) partners was limited, male-male SGR networks were wider than were either male-female or female-female SGR networks among adults. The number of contact-seeking behavior (CSB) partners was also limited, but dominant males had more CSB partners. Adult females mainly interacted by pant-grunt greeting (PGG) with adult males, but tended to do so mainly with the highest-ranking male(s) within visual contact. These results indicated that the social bonds among adult males were essential to group unity. Because of clear male dominance, adult females established peaceful coexistence with all group members despite less frequent SRI with subordinate males by maintaining affiliative social bonds with dominant males, thereby supporting the male-bonded unit-group model. Adult females had many female SRI partners, but these interactions did not involve performing conspicuous behaviors, suggesting that females maintain social bonds with other females in ways that differ from how such bonds are maintained with and between adult males.

Adjustment of brood size and androgen levels in a teleost species with exclusive male parental care

In maximizing reproductive success, individuals face a trade-off between parental care for their current offspring and investing in androgen-dependent sexual traits to produce further offspring. It has thus been proposed that parental effort would suppress androgen levels. Here, we studied parental effort by manipulating brood size in the peacock blenny, Salaria pavo, a littoral fish species in which males show long periods of paternal care. We focused on the effects of brood size on female spawning rate (measured as number of eggs received in the nest by focal males) and androgen levels. We found a positive linear relationship between brood size and the number of eggs received subsequently. Accordingly, spawning rate increased for males with experimentally enlarged broods while it decreased for males in which brood size was reduced. However, over a longer time interval, brood sizes of both treatments returned to the nonmanipulated brood size, suggesting an effect of additional factors such as male quality. Brood size did not show the expected negative relationship with levels of testosterone and 11-ketotestosterone. On the contrary, testosterone levels were positively correlated with brood size. However, in agreement with the prediction, changes in the level of parental care by manipulation of brood size showed an inverse relationship with testosterone levels. As with brood size, given time, testosterone levels tended to return to nonmanipulated levels. Such changes suggest that males adjust their brood size to an individual value through androgen modulation of courtship or other traits influencing female spawning rate.

No evidence for sperm priming responses under varying sperm competition risk or intensity in guppies

Sperm competition theory predicts that males should tailor their investment in ejaculates according to the number of rival males competing to fertilize a female's eggs. Research spanning several taxa supports this prediction by showing that males are often sensitive to the level of sperm competition and adjust their investment in sperm numbers accordingly. More recent work has revealed that males may also tailor the quality of sperm according to the number of males competing for fertilization. Here I test for both effects in guppies (Poecilia reticulata) in an experiment that simultaneously evaluates the risk and intensity models of sperm competition. The experiment determined whether male guppies adjust the number (stripped ejaculate size) and quality (sperm velocity and viability) of sperm that are primed over a 3-day period according to experimental changes in the perceived level of sperm competition. A total of 136 focal males were initially stripped of all retrievable sperm and assayed for these sperm traits before being allocated at random to one of four treatments simulating different levels of sperm competition risk and intensity. During the 3-day treatment phase, focal males had visual and olfactory access to a sexually receptive (initially virgin) female maintained with different numbers of stimulus males to simulate variation in the risk and intensity of sperm competition. Following this, males were assayed again for the sperm traits. Contrary to predictions, there was no significant change in any of the measured variables among treatments, although qualitatively the patterns for sperm velocity and viability did conform to expectation. The lack of any trend for the number of sperm primed was unequivocal and future work examining the effects of sperm competition on sperm production should focus on whether males differentially allocate sperm numbers among matings that differ in the level of sperm competition.

Presence of rivals reduces mating probability but does not affect ejaculate size in the freshwater crayfish Austropotamobius italicus

Sperm competition models predict that males should adjust their sperm expenditure according to the risk and intensity of sperm competition, increasing it when competing with at least one other ejaculate and decreasing it when the number of competing ejaculates rises above two. However, when the raffle to fertilize eggs is strongly loaded due to sperm removal by subsequent rivals, it could be hypothesized that first-mating males progressively decrease their sperm expenditure and/or defer copulation with an increasing number of nearby competitors. Here, by varying experimentally the number of competitors around a mating pair, we analysed mating behaviour of both sexes and sperm expenditure in relation to variation in the immediate risk of sperm competition in the freshwater crayfish Austropotamobius italicus , a species where males remove all or part and feed on sperm laid by previous mating males. We found that male mating behaviour varied significantly with increasing number of rivals, because interactions between focal males and competitors, as well as male refusals to copulate, increased with number of the latter. As a consequence, the probability to reach an effective copulation decreased with increasing number of competitors. However, males released similar amounts of sperm independently of the number of surrounding competitors. Thus, the observed lack of variation in sperm expenditure in relation to the number of surrounding rivals suggests that freshwater crayfish males are unable to adjust the size of their ejaculates to short-term variation in the immediate risk of sperm competition.

Male salamanders remember individuals based on chemical or visual cues

AbstractThe use of multiple cues can enhance the detection, recognition, discrimination, and memorability of individuals by receivers. We conducted two experiments, using only males, to test whether territorial red-backed salamanders, Plethodon cinereus, could use only chemical or only visual cues to remember familiar conspecifics. In both experiments, focal males spent significantly more time threatening unfamiliar than familiar male intruders. They also chemoinvestigated the filter paper containing chemical cues of unfamiliar intruders more often than that of familiar intruders. These results suggest that red-backed salamanders can use both chemical and visual cues to recognize familiar individuals, allowing them to distinguish between less threatening neighbours and more threatening intruders in the heterogeneous forest floor habitat, where visual cues alone would not always be available.

Individual differences in protandry, sexual selection, and fitness

Protandry is the difference in arrival date between males and females, with competition among males for access to preferred territories (the rank advantage hypothesis) or mating success (the mate opportunity hypothesis) supposedly driving the evolution of protandry. The fitness costs and benefits of protandry accruing to individuals differing in degree of protandry (arrival date of a male relative to the arrival date of his partner) have never been quantified. We analyzed the fitness consequences of sex differences in arrival date in the barn swallow Hirundo rustica, in which arrival date can be precisely estimated and the fitness of pairs differing in degree of individual protandry assessed. Early arriving males had greater mating success than late arriving males. The number of extrapair offspring in own nests decreased with increasing degree of individual protandry, whereas the number of offspring fathered by a focal male was unrelated to individual protandry. There was directional selection on individual protandry as shown by pairs with a larger than average degree of protandry reproducing early and, hence, supposedly producing more recruits. There was also stabilizing selection on individual protandry as shown by pairs with an intermediate degree of protandry reproducing early. Annual production of fledglings increased with early arrival of males, but not with early arrival of females, once the effect of laying date had been considered, with no additional effect of individual protandry. Neither male nor female survival was significantly related to degree of individual protandry. These findings are consistent not only with the mate opportunity hypothesis but also with a sexual conflict hypothesis, suggesting that males and females differ in their optimal timing of arrival due to sex-specific fitness costs and benefits. Key words: barn swallow, extrapair paternity, Hirundo rustica, optimal arrival time, sex ratio. [Behav Ecol]