An experimental study of male adornment in the scarlet-tufted malachite sunbird: II. The role of the elongated tail in mate choice and experimental evidence for a handicap

Summary Scarlet-tufted malachite sunbirds (Neetarinia johnstoni) are endemic to the high altitude zones of East African mountains. On Mount Kenya the males are resident and territorial all year. The females arrive over an extended period at the beginning of the breeding season, after spending the non-breeding season in nomadic flocks at slightly lower elevations. Males are bright iridescent green with elongated central tail feathers, which are displayed during courtship. We examined the role of the tail in mate choice using natural variation and investigated the influence of tail length on male time budgets by experimental manipulation. Territorial males that paired had longer tails than territorial males which remained unpaired. Males with naturally long tails started breeding earlier, resulting in fledglings being produced earlier in the season when the weather was more clement. Birds which bred together in one year were paired together in the next year if they both survived. Thus the male's tail may only be involved as a cue in the formation of new pairs. Males which were already paired were subjected to one of three experimental treatments — having their tail elongated, shortened, or manipulated but kept at the same length. Males with experimentally shortened tails spent more time in flight and hawked for flying insects with a higher efficiency than control males. Both control and elongated tail males reduced the amount of time in flight and had a lower hawking efficiency after manipulation. These results suggest that the long tail of male scarlet-tufted malachite sunbirds is a handicap. The factors influencing the reproductive success of a pair are discussed. Offprint requests to: M.R. Evans     

Sexual dimorphism and the mating ecology of polar bears (Ursus maritimus) at Svalbard

We assessed the role of size, mass, and age in mating and non-mating polar bears (Ursus maritimus) at Svalbard, Norway, during the spring breeding season. The ratio of male to female mass, in male-female pairs, ranged from 1.00 to 3.02 ([`(x)] = 1.99 \overline x = 1.99 ) indicating that mating males were larger than mating females but with substantial variation. Paired males were older than unpaired males and male mass was related to age. However, males paired with females were not significantly different in body mass from those males caught alone. Wounds and scars resulting from fights between males began at about 6 years of age and peaked at about 17 and 20 years of age, respectively. The frequency of broken canines in males, presumably due to increased male-male conflicts, increased with age but showed little increase in females. The wide range of male size in male-female pairs and the age-related signs of injury suggest that male polar bears engage in both scramble competition and contest competition for access to breeding females. The mating system of polar bears is variable but is best described as female defense polygyny or serial monogamy.     

Replacement males in the western bluebird: opportunity for paternity, chick-feeding rules, and fitness consequences of male parental care

In western bluebirds (Sialia mexicana), most pairs remain together for life and share equally in post-hatching parental care. We removed resident males of socially monogamous pairs during laying and after clutch completion to examine chick-feeding rules used by replacement males and current and future fitness consequences of paternal care. Replacement males were not infanticidal and 7 (47%) fed nestlings. Feeding replacement males and the females they joined fed at rates similar to controls. Females without a feeding replacement male compensated by feeding more themselves so that overall feeding rates were not compromised, but they reduced their brooding time. Unlike assisted females, unassisted females exhibited reduced nesting success and their 14-day-old chicks weighed less than controls. Field metabolic rates of unassisted females were 17% higher than those of control females, but the difference was not statistically significant. Older females were better able to raise young without the male's help than were yearling females. Female condition was not affected by male provisioning and unassisted females were as likely to survive to the next breeding season as assisted females. We found no future benefits of provisioning by replacement males; those that fed were no more likely to breed with the female on her subsequent attempt than were males that did not feed, and subsequent clutch sizes were not reduced for females rearing young without the male's help. These experiments suggest that male parental care increases nesting success in western bluebirds and that replacement males use an all-or-none rule to determine whether or not to feed chicks: if they are present during the fertile period they feed at typical rates; if they are not, they usually do not feed at all. Because chick-feeding by males is tied to opportunity for paternity, influences success in the current nest, and does not affect the male's future breeding success, it appears to be parental rather than mating effort. Received: 8 May 1998 / Accepted after revision: 23 July 1998     

Mating patterns in squirrel monkeys (Saimiri oerstedi)

Summary The mating system of squirrel monkeys (Saimiri oerstedi) in Parque Nacional Corcovado, Costa Rica was studied and used to develop a model to interpret the evolution of seasonal sexual dimorphism in squirrel monkeys (Saimiri spp.). Adult male body weights in captivity and the wild may increase more than 20%, beginning approximately two months prior to and continuing through the annual two month, breeding season. Female inter-troop transfer was common in the study population, but male troop residence was stable. Instances of agression among adult males in the troop, even in sexual contexts, were rare. Reproductively mature males enlarged to varying degrees by the start of the breeding season and cooperated in mobbing females to olfactorily evaluate female, estrous condition. Female mate preference corresponded to a ranking based on relative male enlargement. The largest male obtained 70% of the copulations observed in the 1984 breeding season. Little evidence exists that females typically mate with more than one male during the period of peak receptivity. Seasonal enlargement in males is suggested to be the result, of both male intrasexual competition and female choice.     

Trade-off between mate guarding and mate attraction in the polygynous great reed warbler

Summary In a Swedish population of the polygynous great reed warbler Acrocephalus arundinaceus, about 40% of the males formed pairs with more than one female. Males sang two completely separated types of song: (1) long song when they tried to attract mates and (2) short shong when they guarded a fertile female. As soon as a male had attracted a female he immediately switched from long to short song and apparently guarded her for at least 3 days. Most males left their female and started singing long several days before her fertile period ended. This behavior probably increased the risk of cuckoldry. By assuming that males of polygynous species maximize their fitness over the whole breeding season (i.e., maximize fitness rate), we predicted longer mate guarding periods when the probability of attracting a second female was low and the risk of cuckoldry was high. These predictions were supported by observations. The probability of attracting a second female decreased during the breeding season and, in accordance, the number of days that males sang long song during the primary female's fertile period was negatively correlated with the time of the season. This trend held also for individual males when comparing mating periods of their primary and secondary female. The increasing rate of male intrusions with season may also have contributed to the increasing mate guarding periods. Offprint requests to: D. Hasselquist     

Bi-directional sex change: testing the growth-rate advantage model

Bi-directional sex change in coral-dwelling fishes (genera Gobiodon and Paragobiodon) has been attributed to a growth-rate advantage for females during the non-breeding season. This model predicts that the smallest individual in a newly formed pair should always be female. To determine if a growth-rate advantage exists for female Gobiodon histrio, I monitored the growth of males and females in natural pairs during the breeding and non-breeding season. I then used a manipulative field experiment to test four predictions of the growth-rate advantage model: (1) the larger individual should change sex to male in new pairs containing two females; (2) the smaller individual should change sex to female in new pairs containing two males; (3) neither individual should change sex in heterosexual pairs where the male is larger than the female; and (4) both individuals should change sex in heterosexual pairs where the female is larger than the male. A growth-rate advantage was detected for female G. histrio during the non-breeding season; however, only the first three of the predicted outcomes were observed in the manipulative experiment. Sex change did not occur in heterosexual pairs where the female was larger than the male. Furthermore, growth did not differ between sex-changed and non-sex-changed fish; therefore, the absence of sex change in these pairs is not due to a growth cost to sex change. I propose that the risk of moving among spatially isolated habitat patches and the low probability of finding a mate have been more important than sex-specific differences in growth rates to the evolution of bi-directional sex change in coral-dwelling gobies.     

Growth rate and the cost of calling activity in male carpenter frogs,Rana virgatipes

Summary Energy stress during the breeding season and relationships between calling activity and growth were investigated in male carpenter frogs, Rana virgatipes. This species has a prolonged breeding season of up to three months in Southern New Jersey. Monthly collections made in 1985 revealed that both dry mass and percent body lipid decreased throughout the breeding season but sharply increased at the end of the breeding season. Observations of free-living males showed that small males were more likely to gain mass than large males during the breeding season. All males gained mass at higher rates after the breeding season. A simultaneous record of calling activity and mass change was obtained for 42 males. Males called on 95% of nights, indicating that they rarely ceased their reproductive activities. Small males tended to have low calling efforts and high growth rates. When the effect of initial mass was removed, growth rate was negatively correlated with calling effort among small males. This is the first demonstration of a direct tradeoff between a reproductive activity and growth in an anuran.     

Frequency and timing of extrapair fertilisation in the polyandrous red phalarope (Phalaropus fulicarius)

In sequentially polyandrous bird species, where females mate with different males in succession during a single breeding season, sperm stored by females can occasionally lead to extrapair fertilisations (EPFs) in clutches cared for by the second and subsequent males. Thus, we predicted that in red phalaropes (Phalaropus fulicarius) – a sex-role-reversed, sequentially polyandrous, arctic breeding shorebird – EPFs would occur more frequently in clutches laid later in the breeding season. We used multilocus DNA profiling to examine the frequency and timing of EPFs in a population of red phalaropes breeding in the Canadian high arctic. Using a technique to determine parentage without maternal DNA, we inferred that 6 of 70 chicks in 18 broods resulted from EPFs – one extrapair chick in each of 6 broods. These results were supported by a further analysis using microsatellite DNA. As predicted, broods containing EPFs hatched from clutches laid significantly later in the season than did broods containing no EPFs. The difference in median hatch dates between broods with and without EPFs was 9.0 days, or 38% of the entire egg-laying period in that season. For the whole breeding season, we estimated that 6.5% of chicks were sired by extrapair males, which is similar to extrapair paternity estimates for other sex-role-reversed birds, but relatively low compared to the majority of socially monogamous species studied so far. Received: 24 August 1998 / Received in revised form: 18 January 1999 / Accepted: 24 January 1999     

Food supply during prelaying period modifies the sex-dependent investment in eggs of Eurasian kestrels

The theory of sex allocation suggests that if the reproductive value and the cost of producing/rearing offspring differ between male and female offspring, parents should invest differently in sexes depending on environmental conditions. Female parents could allocate more resources to eggs of one sex to compensate potential sex-dependent constraints later during the nestling period. In this study, we tested the influence of environmental conditions on sexual dimorphism in eggs of Eurasian kestrels (Falco tinnunculus) by experimentally manipulating food availability before laying. We found that an increase in food abundance before laying did not increase egg mass but changed sex-dependent resource distribution in eggs. In food-supplemented pairs, but not in control pairs, egg mass and hatchling mass were similar between males and females. In addition, we found, in the food-supplemented group, that the latest hatched females showed shorter hatching times than in the control group. In control pairs, female eggs, hatchlings and nestlings were heavier than males. In addition, male fledglings in the food-supplemented group gained less mass than those in the control group. As that food abundance was only increased until the onset of laying, female kestrels were expected to invest in eggs taking food abundance before egg formation as a predictor of future conditions during brood rearing. Our study shows that environmental conditions before laying promote a subtle adjustment of the resources invested in both sexes of offspring rather than in other breeding parameters. This adjustment resulted in a shortening of hatching time of the last hatched females that possibly gives them advantages in their competitive capacity with respect to male nest-mates.     

Dynamics of space use and male vigour amongst wood mice,Apodemus sylvaticus,in the cereal ecosystem

The spatial organisation of male and female wood mice,Apodemus sylvaticus, was investigated in a large-scale radio-tracking study on arable farmland near Oxford, United Kingdom, during the breeding season. Both sexes had significantly larger home ranges in the breeding season than at other times, and the breeding season home ranges of male (X = 1.44 ha) were significantly larger than those of females (X = 0.49 ha). Home range overlap was significantly greater between males, and between males and females, than it was between females. Overlap between males tended to be greatest in heavily utilised areas. Except during sexual consortship, there was minimal evidence of dynamic interaction among individuals. Home range sizes of breeding males varied widely, as did their body weights. There was no relationship between male body weight and home range size or any other movement parameter. However, males with the largest home ranges had the highest scores on all other movement parameters, indicating that they expended more energy in movement. These more vigorous males had access to the home ranges of more females than did males with small home ranges.     

Polygyny and male parental care in Savannah sparrows: effects on female fitness

Summary To determine the effects of male mating status on female fitness, we compared the reproductive success, survival, and future fecundity of female Savannah sparrows (Passerculus sandwichensis) mated to monogamous vs. polygynous males in a 5-year study on Kent Island, New Brunswick, Canada. The proportion of males with more than one mate varied from 15 to 43% between years and sites. Polygynous and monogamous males fledged young of equal size in every year of the study. Females who shared paternal care with other females laid as many eggs per clutch and clutches per season as monogamously mated females. In most years polygynously mated females showed no delay in laying a second clutch, and they suffered no reduction in fecundity the following year. Recruitment of a female's offspring into the breeding population was generally independent of her mating status. Fitness costs of being mated to a polygynous male were only apparent in one year of the study, during which females mated to polygynous males had higher over-winter mortality than those mated to monogamous males. That same year, young raised by polygynous males were only one-third as likely to survive to reproductive maturity (as inferred by returns) as those raised by monogamous males. A male's mating status had no effect on his own survivorship. A male's mating status did not necessarily reflect his contributions to raising nestlings, which may partially explain why monogamously and polygynously mated females had equal fitness. At 35 nests the proportion of food deliveries brought by individual males varied from 0 to 75%; on average, males brought fewer than 30% of all food deliveries. Yet parental care by polygynous males was no less than that of monogamous males, at least at the nests of their primary females. Secondary females tended to receive less male assistance during the nestling stage, but their reproductive success was indistinguishable from that of primary females. Females feeding young without male assistance made as many food deliveries/h as did pairs in which males brought at least 30% of all food deliveries. Unassisted females did not suffer diminished fledging success or produce smaller fledglings. The benefits of polygyny for male Savannah sparrows are clear: polygynous males recruit more surviving offspring into the breeding population than monogamous males. The fitness of females, on the other hand, appears to be unaffected by whether their mate was monogamous or polygynous except in occasional years. Polygyny may be maintained in this population by the constraints of a female-biased sex ratio, the inability of females to predict a male's paternal care based on his morphology or behavior, the poor correlation between a male's mating status and his assistance at the nest, and inconsistent natural selection against mating with a polygynous male. Correspondence to: N.T. Wheelwright     

Helper contributions to reproductive success in the splendid fairy-wren (Malurus splendens)

Summary In Malurus splendens, helpers were present in 65% of 226 group-years with at least one helper female in 37% of group-years. Most females helped for only one year, while many males did so for at least two years. Most were offspring of one or both present breeders, and in 53% of helper-years, helped both parents. For 159 helpers of known age and parentage, the mean coefficient of relatedness to the offspring was 0.47. Novice females with or without helpers produced fewer fledglings per season than females with one year breeding experience and the same level of help. Helpers did not affect production of fledglings per year by females with one year of experience. Females with two or more years experience and at least two helpers produced more fledglings than equivalent birds with one or no helpers. Experience and helpers have little effect on production of fledglings per nest but they lead to more females renesting after a first brood has been raised. Fewer than 20% of novices renest after fledging one brood, while for females with at least two years experience, the percent renesting after success is 40% with no help, 56% with one helper and 69% with 2 or more helpers. Experienced females begin their first clutch earlier than novices, and helpers reduce the time to renest after success from 66 days for an experienced female with no helpers to 50 days for females with at least two years experience and two or more helpers. Breeding females with helpers survive better (76%) than those with no helpers (55%), and helpers thus gain future indirect fitness. Despite their close relatedness to breeders and offspring, in only 19% of group-years did helpers increase their indirect fitness from an increase in productivity.     

Helping at the nest,allofeeding and social status in immature arabian babblers

Summary Arabian babblers, Turdoides sqamiceps, are cooperatively breeding, group territorial birds, occurring in desert habitats. Non-breeders participate in several types of cooperative behaviour, including care of eggs, nestlings and fledglings, and bringing food to one another (allofeeding). This paper reports on observations of helping with eggs and young, and peer allofeeding, performed by immature babblers, fledged the previous season or earlier in the same season. The birds were from a babbler population studied since 1971 in the Arava valley of eastern Israel. They belonged to groups in which all individuals had been colourringed and were accustomed to human observers. Individuals and nests in the territories of these groups could be watched from distances of 1–2 m without causing alarm, so fine details of behaviour could be observed in the field. In most allofeeding interactions observed, a more dominant individual brought food to a subordinate. Sometimes the subordinate bird avoided accepting the proferred food, and was then hit or chased by the bird that had attempted to feed it. Occasionally a subordinate bird attempted to feed a more dominant individual. Dominant birds always refused to take food offered by a subordinate, and hit or chased any subordinate attempting to feed them. Frequencies of visits to nests were correlated with helper rank for those visits where the incoming bird displaced a previous visiter from the nest. Frequences of visits to unattended nests, however, were not correlated with helper rank. Similarly, frequencies of feeding visits to fledglings were correlated with helper rank until incubation of the next brood began. After this, the correlation disappeared. Numerous instances of aborted nest visits were observed in which a helper, often carrying food, arrived at the nest-tree but left again without visiting the nest. In many other cases, a helper arrived at the nest-tree but delayed visiting the nest for several minutes. Aborted and delayed visits usually occurred when a more dominant bird was in the vicinity of the nest. This suggests interference between helpers. Direct interference between helpers visiting fledglings was occasionally observed. In such cases, a more dominant helper snatched food from a subordinate approaching fledglings, and then fed this food to the fledglings itself. Interference between helpers, and conflict between proferers and recipients of food during allofeeding, are not easily explained by kin selection or reciprocity. On the other hand, such behaviour is readily explained by a hypothesis which suggests that individuals may increase their social status in the group by performing cooperative behaviour. Babblers that establish status by demonstrating ability to bear the short-term costs of cooperative behaviour, rather than through direct aggression towards rivals, are likely to forge collaborative relationships with other group members. Since babblers must collaborate to establish and defend a territory, such relationships are essential to reproductive success.     

Synchronous waving in an ocypodid crab, Ilyoplax pusilla : analyses of response patterns to video and real crabs

In several species of ocypodid crabs, including Ilyoplax pusilla (De Haan), waving displays tend to be synchronized among neighboring individuals. However, no quantitative study has yet been conducted. In this study, responses of I. pusilla males to waving of video and real crabs were examined. Spontaneous, periodic waving was observed even when no stimulus waves were present. When single isolated waves of the video crab were randomly delivered (single pulse perturbation), crabs waved either immediately after the stimuli or after some time had elapsed. Assuming that waves immediately after the stimuli were triggered prior to those stimuli, crabs typically responded after a roughly constant delay from the onset of each stimulus. The delay (response time) was close to, but slightly shorter than, the spontaneous wave period. Dependency of response time upon timing of the stimulus (stimulus time) was at most weak. Responses of crabs to periodic waves of the video crabs and waves of␣the real crabs were basically consistent with the results of single pulse perturbation, although there was a consistent tendency for stimulus time and response time to be negatively correlated near stimulus time = 0 s. Overall, these results demonstrate that males of I. pusilla adjust signal timing relative to a neighbor in a manner of phase delay synchrony. The similar response pattern has also been noted in synchronous flashing of fireflies and the calling of katydids. Moreover, these experiments with I.␣pusilla demonstrate the value of video playback as a research tool in this species. Received: 4 April 1997 / Accepted: 17 December 1997     

Sex-specific patterns of lifetime reproductive success in single and repeat breeding steelhead trout (Oncorhynchus mykiss)

All other things being equal, the lifetime reproductive success (LRS) of iteroparous and semelparous individuals should scale with the number of breeding seasons. Deviations from this relationship may occur for many reasons, including age- or size-related fecundity or life history trade-offs, which may differ between sexes. We used 19 brood years of DNA parentage analysis in a small (N = 4–143 year⁻¹) wild, unexploited population of steelhead trout (Oncorhynchus mykiss) to compare the LRS of individuals that spawned only once [“one time spawners” (OTS), N = 355 male, 371 female] to those spawning twice [“repeat spawners” (RPS), N = 13 male, 49 female]. Female RPS had nearly twice the LRS of female OTS (1.17 offspring per female vs 0.91 offspring per female), whereas male RPS had nearly three times the LRS of male OTS (1.54 offspring per male vs 0.57 offspring per male). Female RPS produced slightly more adult offspring during their second breeding season than their first (0.78 vs 0.82 offspring per female); however, male RPS produced all of their adult offspring in their second breeding season (0 vs 1.54 offspring per male). The additional growth in body size of males between breeding seasons may give them an advantage in their second breeding season, but the lack of offspring produced in their first season suggests a trade-off between survival and future reproduction that was not expressed in females.     

The annual reproductive cycles of Uca annulipes,Portunus pelagicus and Metapenaeus affinis (Decapoda: Crustacea) from the South-west coast of India

Employing the gonad index method, the reproductive cycles of three decapod crustaceans, Uca annulipes (Latreille), Portunus pelagicus (Linnaeus) and Metapenaeus affinis (Milne-Edwards) have been studied. In these crustaceans breeding is not continuous all the year round, but extends over several months of the year with distinct peak periods of gonadal activity. The male and female reproductive cycles are not concurrent. The peak of the reproductive cycle of males occurs slightly earlier in the breeding season than that of females. These studies indicate the possibility of production of successive broods of eggs during the same breeding season. In these species, the low saline conditions of the monsoon period are unfavourable for breeding. The medium and high saline conditions during the post-monsoon and pre-monsoon months, respectively, with plenty of planktonic food for the larvae, seem to be the favourable periods for breeding activity.     

Social and ecological factors affecting paternity allocation in American robins with overlapping broods

Factors that affect extra-pair mating in birds are likely to vary across the breeding season. Changing densities of active nests may alter the opportunities for extra-pair mating, and parental duties may alter a male’s opportunity to guard his mate from extra-pair mating. The latter affects species with multiple broods, where males care for fledglings from first nests while females initiate second nests. We studied a population of multi-brooded American robins (Turdus migratorius) to assess how seasonal changes in nesting density and changes in mate-guarding opportunity influenced paternity patterns over successive breeding attempts. Extra-pair paternity (EPP) occurred in 71.9% of broods and accounted for 48.1% of young. High nesting densities in the study population may explain the high overall rate of EPP, but seasonal variation in breeding density did not explain patterns of EPP among nests. Contrary to the predictions of the mate-guarding hypothesis, EPP did not increase in the second nests that followed successful first nests, and the percentage of extra-pair young in second nests did not decline as the overlap between successive nests increased. The fact that EPP was actually lower when the interval between clutches was shorter suggests that the sooner the males can assume sole care of first broods and allow their mates to renest (indicative of superior paternal quality), the more paternity they realize in the next nest. These results suggest that mate-guarding opportunity does not influence paternity in this population of American robins and that female robins may allocate paternity based on their assessment of male parental performance at first nests.     

Seasonal anorexia in the male red-sided garter snake,<Emphasis Type="Italic"> Thamnophis sirtalis parietalis</Emphasis>

Many animals show seasonal shifts in behaviors that coincide with breeding, migration, or hibernation. These behavioral shifts provide ideal opportunities to study the regulation of behavior. The red-sided garter snake (Thamnophis sirtalis parietalis) spends 8 months of the year inactive in underground hibernacula, 1 month breeding, and 3 months feeding to build up enough energy stores to survive the following winter. Although they emerge from 8 months of hibernation with severely depleted energy reserves, they do not feed until weeks later, after the breeding season. We tested the hypothesis that this lack of feeding during the breeding season is due to a shift in behavior rather than the distribution of food and potential mates. Male garter snakes were given a series of choices between pursuing a breeding or feeding opportunity. The proportion of tests in which males selected feeding over breeding gradually increased throughout the study period, reaching almost 100% in the final tests. Males also were given opportunities to feed and court at the beginning and end of the study. Males initially refused food and courted females, but when retested at the end of the study they fed and did not court females. Thus aphagia during the breeding season is due at least in part to an endogenous shift in behavior.Communicated by P. Weatherhead     

Natal philopatry and breeding area fidelity of long-billed curlews (Numenius americanus): Patterns and evolutionary consequences

Summary Long-billed curlews (Numenius americanus) appear unique among scolopacid shorebirds so far studied in possessing a significant sex bias in natal philopatry. We resighted 9 curlews at least attempting to breed that were color-banded as chicks; 8 of these were males. Male curlews also cooperate extensively with neighbors in mobbing potential chick predators. This mutualistic behavior may have evolved through kin selection among philopatric males. If so, we would expect such an evolutionary consequence to lead to a similar sex bias in breeding area fidelity. Yet our resightings of colorbanded adults over 4 consecutive years indicate that males and females were equally likely to return to previous nesting territories. Excessive disturbance such as capture and nest loss within a single breeding season was correlated with the likelihood of breeding dispersal by females but not males. This suggests potentially stronger breeding area fidelity of males.     

Seasonal decline in reproductive performance varies with colony size in the fairy martin, <Emphasis Type="Italic">Petrochelidon ariel</Emphasis>

Reproductive success within populations often varies with the timing of breeding, typically declining over the season. This variation is usually attributed to seasonal changes in resource availability and/or differences in the quality or experience of breeders. In colonial species, the timing of breeding may be of particular importance because the costs and benefits of colonial breeding are likely to vary over the season and also with colony size. In this study, we examine the relationship between timing of breeding and reproductive performance (clutch size and nest success) both within and between variable sized colonies (n = 18) of fairy martins, Petrochelidon ariel. In four of these colonies, we also experimentally delayed laying in selected nests to disentangle the effects of laying date and individual quality/experience on reproductive success. Within colonies, later laying birds produced smaller clutches, but only in larger colonies. The general seasonal decline in nest success was also more pronounced in larger colonies. Late laying birds were generally smaller than earlier laying birds, but morphological differences were also related to colony size, suggesting optimal colony size also varies with phenotype. Experimentally delayed clutches were larger than concurrently produced non-delayed clutches, but only in larger colonies. Similarly, delayed clutches were more likely to produce fledglings, particularly later in the season and in larger colonies. We suggest that the reduced performance of late breeding pairs in larger colonies resulted primarily from inexperienced/low quality birds preferring to settle in larger colonies, possibly exacerbated by an increase in the costs of coloniality (e.g., resource depletion and ectoparasite infestations) with date and colony size. These findings highlight the importance of phenotype-related differences in settlement decisions and reproductive performance to an improved understanding of colonial breeding and variation in colony size.