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.     

Intraspecific nest parasitism in the swallow Hirundo rustica: the importance of neighbors

Summary Intraspecific nest parasitism in the swallow Hirundo rustica involves several parasite and anti-parasite tactics. Neighboring swallows breed asynchronously, perhaps because neighboring nests with overlapping egg-laying periods have higher frequency of intraspecific nest parasitims than neighboring nests with no such overlap. Since nest guarding prevents nest parasitism, the effect of nest guarding was removed by putting up old swallow nests. Such experimental nests were parasitized by swallows more often when the nests contained eggs, than when empty, and when the nests were far away from, rather than close to, an active neighboring nest. Close experimental nests with eggs were, however, parasitized less frequently if asynchronous (non-overlapping egg-laying periods), rather than synchronous, to active neighboring nests. Aggressive anti-parasite behavior of nest owners was determined from responses towards female intruders. Responses were more often aggressive before and especially during egg laying than during incubation. Nest owners also behaved aggressively more often when the intruder approached close to as compared to at some distance from their nest. Focal pairs behaved aggressively more often towards closely approaching intruders during the incubation period than towards distantly approaching intruders during the egg-laying period. Swallows reduce the frequency of intraspecific nest parasitism by nesting asynchronously with and close to a neighboring nest. Aggression by neighbors may reduce the success of potential parasitic swallows.     

Divorce in common murres (Uria aalge): relationship to parental quality

Behavioral precursors of 12 divorces were examined in 30 color-banded pairs of common murres (Uria aalge) over six breeding seasons. Common murres are long-lived seabirds that typically return each year to the same mate and nest site in dense colonies. At least one parent is present continuously from egg lay to chick fledging. Murres, therefore, have considerable opportunities to compare their mates parental behavior with that of several familiar neighbors. Previous reproductive success was lower for divorcing birds than for reuniting pairs. As predicted by the better option hypothesis, there were clear choosers (seven females and five males) that initiated divorce by moving to a new birds site or by courting a new partner at their current site. Choosers improved their reproductive success after the divorce, whereas their previous partners, the victims did not. Yearly divorce rates (average 8.2% per year) were significantly correlated with yearly mortality rates. Divorces appear to be opportunistic: pairs divorced after varying numbers of reproductive failures with the immediate precursor usually being the disappearance (death) of a murre from a successful neighboring site. In contrast to the delays experienced by victims, choosers formed new pairs quickly and laid their eggs no later than reuniting pairs. Prior to the divorce, victims fed their chicks less often than choosers, and some engaged in other behaviors that compromised egg or chick survival. These observations suggest that deficiencies in parental behavior were precursors to the divorce. This report is one of the first cases where reproductive failure of divorcing pairs has been linked to deficits in the parental behavior of the subsequent divorce victim.Communicated by C. Brown     

Sex-specific effects of yolk-androgens on growth of nestling American kestrels

Maternally derived androgen hormones concentrate in avian egg yolks as the yolks grow on the female’s ovary, possibly forming a basis for important maternal effects in birds. In the American kestrel (Falco sparverius), experimental elevation of yolk androgens in the first-laid egg of a clutch (a-egg) to the concentrations found naturally in a clutch’s later-laid eggs reduces the growth rate of a-egg nestlings compared to controls. These findings, together with discoveries from other species that the effects of yolk androgens on growth of female nestlings may differ from their effects on growth of male nestlings, raise the hypothesis that natural changes in yolk-androgen concentrations with laying order are ultimately due to a difference between the sexes in their yolk-androgen sensitivity and between early- and late-laid eggs in their sex ratio. By re-analyzing previously published data and adding to the analysis data from previously unanalyzed blood samples used for sex determination, we investigated possible sex-specific effects of yolk-androgens in the context of a potential sex-biased laying order in free-living American kestrels. We used a multi-level, mixed model with a Gompertz function to analyze growth of nestlings hatching from a-eggs that were control-treated or in which we experimentally elevated yolk-androgen concentrations shortly after laying to the higher concentrations naturally found in later-laid eggs. We discovered that male nestlings were more susceptible than female nestlings to growth inhibition by yolk-androgen elevation but did not find a bias in sex ratio with respect to laying order. Together, these findings do not support the above hypothesis. However, they are consistent with the hypothesis that sex differences in yolk-androgen sensitivity enable mothers to economically tune reproductive effort to an individual offspring’s reproductive value, which can vary more for one sex than the other.     

Individual differences in nest defense in the colonial breeding Black-tailed Gulls

Often in colonial seabirds, all colony members are believed to defend against nest predators and experience equal nest predation risk. However, the variation of defense behavior among members and its reproductive consequences are largely unknown. We investigated (1) individual variation in the nest defense of breeding Black-tailed Gulls Larus crassirostris against a natural egg predator, the Jungle Crow Corvus macrorhynchos and (2) how this behavioral variation affects an individual’s own nest predation risk and that of their neighbors. Results were compared between 2 years where crow attack levels were manipulated to average 5 and 22 times normal rates (“low” and “high” predation risk years, respectively) by the placement of varying numbers of artificial nests containing unguarded eggs at the perimeter of the gull colony. In both years, 23–38% of parents, mostly males, showed “aggressive” defense behavior (strikes or chases) against crows and decoys. Other “non-aggressive” gulls showed no defense. In the year of low predation risk, intrusion rates by crows (landing within 0.5 m of an individual gull’s nest) were similar for aggressive and non-aggressive gulls. In the year of high predation risk, however, the rates of intrusion for aggressive gulls (4%) and for non-aggressive gulls with an aggressive neighbor (37%) were significantly lower than for non-aggressive gulls without an aggressive neighbor (76%). These results indicate that aggressive individuals reduce nest predation risk for themselves and conspecific neighbors in a colonially breeding species.     

Parental care of replacement clutches in common terns (Sterna hirundo)

In long-lived seabirds with low annual reproductive output, the renesting decision after breeding failure is critical, and the parents have to weigh benefits of replacement clutches against possible future reproductive costs. In this study, we investigated factors influencing renesting decisions in common terns (Sterna hirundo) and compared aspects of breeding biology and body mass between two breeding attempts by the same pairs in each of 4 years of heavy losses due to predation. Renesting birds were characterized by early laying dates and by a high age. Among early breeders, high egg mass reduced the probability of renesting. A long relaying interval coincided with low mass of replacement eggs in one year, and short intervals with high egg mass in another. Further, egg mass decreased and relaying intervals increased the later the predation events occurred. Evidence of high levels of parental care of replacement clutches came from body mass data: female mass increase prior to egg laying was higher in the second attempt than in the first, whereas male mass was lower during the second courtship period than during the first. Male mass also affected relaying intervals and mass of replacement eggs. We conclude that common terns expend high levels of parental care of replacement clutches. Intrinsic factors related to individual quality (age, body condition) seemed most important for renesting decisions and for the degree of parental care, but foraging conditions seemed to have modifying effects. Received: 13 August 1999 / Received in revised form: 5 February 2000 / Accepted: 13 March 2000     

Fieldfare (Turdus pilaris) breeding success in relation to colony size,nest position and association with Merlins (Falco columbarius)

Summary Clutch size, nestling production and breeding success were studied in colonial Fieldfares (Turdus pilaris) in a subalpine birch forest during ten breeding seasons. Reproductive success was highest for central pairs in large colonies; such pairs benefited most from communal defence against nest predators. Fieldfares and Merlins (Falco columbarius) usually bred in association. Fieldfares breeding away from Merlins had lower breeding success than pairs associated with Merlins, which also benefited by reduced nest predation. Fieldfares apparently chose to nest near Merlins, which had already laid eggs when the thrushes started nest-building.     

Are blue eggs a sexually selected signal of female collared flycatchers? A cross-fostering experiment

Impressive variation in egg colouration among birds has puzzled evolutionary biologists for a long time. The most frequently studied selective forces moulding egg colouration—predation and brood parasitism—have either received little empirical support or may play a role in only a minority of species. A novel hypothesis has suggested that egg colour may be significantly influenced by sexual selection. Females may deposit a blue-green pigment biliverdin into eggshells instead of using it for themselves as a powerful antioxidant. By this handicap, females may signal their quality to males, which are then hypothesized to increase their paternal effort. We experimentally tested the hypothesis in the collared flycatcher (Ficedula albicollis), a species laying blue-green eggs. We cross-fostered clutches between nests to disentangle effects of female/territory quality and egg colour on paternal effort and nestling quality. The results supported two assumptions of sexual signalling through egg colour hypothesis: Blue pigment seems to be a limited resource for females, and female quality is positively correlated with the intensity of the blue-green colour. However, we did not find support for the main prediction of the hypothesis, as male parental effort parameters (feeding frequencies to nestlings and intensity of nest defence) were unrelated to egg colour. We discuss possible reasons for the discrepancy between our results and previous correlative analyses that supported the hypothesis that blue egg colour may be a postmating, sexually selected signal in females.     

Extra-pair paternity in eastern bluebirds: effects of manipulated density and natural patterns of breeding synchrony

The causes of variation in rates of extra-pair paternity among avian populations remain unclear, but could include environmental factors such as breeding density and synchrony. By experimentally manipulating nest site availability, we tested the effects of breeding density on the frequency of extra-pair paternity in eastern bluebirds (Sialia sialis). We also examined the role of breeding synchrony on extra-pair paternity using natural timing of nests. Microsatellite analysis revealed 34 of 305 nestlings (11.2%) were the result of extra-pair fertilizations; and 21 of 79 broods (26.6%) had at least one extra-pair nestling. Several measures of breeding density had independent effects on extra-pair paternity. First, experimental plot type affected extra-pair paternity, with 28 of 34 (82.4%) extra-pair young from nests in high density areas, and only six (17.6%) from nests in low density areas. Independently of plot type, the number of breeding neighbors within a 320-m radius was a significant predictor of the likelihood of extra-pair paternity at the nest. Extra-pair paternity was associated with temporal factors such as absolute timing of breeding and natural levels of local breeding synchrony, but only in bivariate comparisons. We found a positive interaction between density of neighbors within a 320-m radius and local breeding synchrony; this term reduced the main effects of synchrony and number of neighbors, but not experimental treatment. Our results demonstrate the importance of utilizing multiple aspects of proximity in breeding density analyses and testing for interactions between ecological factors that can influence the behavioral events leading to extra-pair fertilizations.     

Tactics of parasitic American coots: host choice and the pattern of egg dispersion among host nests

Summary I examined the tactics adopted by a conspecific brood parasite, the American coot (Fulica americana), and the degree to which these tactics reflect sources of mortality for parasitic eggs. Only 8% of parasitic eggs produced independent offspring, compared to a 35% success rate for non-parasitic eggs, and most mortality was due to egg-rejection by hosts or the consequences of laying eggs too late in the host's nesting cycle. Parasites usually laid parasitically before initiating their own nests and usually parasitized immediate neighbours. Parasites did not remove host eggs before laying their own egg, and egg disappearance in general was not more common at parasitized nests. I found no evidence for non-random host choice, either on the basis of stage of the host's nesting cycle or the host's brood size. The absence of adaptive host choice is likely a consequence of the fact that, due to host limitation, only a small proportion of parasites had meaningful variation among potential hosts to choose from. The pattern of egg dispersion among host nests by individual parasites appears to be a compromise between constraints imposed by host limitation and the increased success obtained from spreading eggs among nests. Most females laying fewer than five parasitic eggs laid them in a single host nest while females laying five or more eggs normally parasitized two or more hosts. An examination of egg rejection and survival rates showed that parasites would maximize success by laying a single egg per host nest, and the pattern of laying several eggs per host nest is likely a consequence of host limitation. However, no egg that was the fifth laid, or later, parasitic egg in a host nest was ever successful and this probably explains why most females laying five or more eggs parasitized more than one host.     

Brood parasitism and cooperative nesting in the moorhen,Gallinula chloropus

Summary During a two year study of the moorhen, Gallinula chloropus, a quarter of the nests found had two or more females laying in them. This was the result of two features of the moorhens' breeding biology: (1) Cooperative nesting; two (or more) females were paired to the same male. Both laid in the same nest, the second laid a large clutch synchronously to the first, and both cooperated in parental care. (2) Intraspecific brood parasitism; some females (parasites or dumpers) laid a small number of eggs at random in the nests' of neighbours. Cooperatively nesting females were mother and daughter, while parasite and host were not. Parasitic females laid in their own nest as well, and generally did so after completing a dumping sequence. Parasitic eggs were laid at random during the host's laying and incubation period, and were about a quarter as likely to produce an independent chick as were non-parasitic eggs. Parasitic females would probably have reared more chicks by laying in their own nests straight away, instead of dumping any eggs at all, because there was a strong decline in reproductive success with season. Possible reasons why they did not do so are discussed. Parasitised pairs tended to rear fewer own chicks than non-parasitised pairs. There was no evidence that hosts selectively destroyed parasitic eggs. Four hypotheses that explain why hosts accepted dumped eggs are considered. (1) The host pair were unaware of the dump, (2) the host male had copulated with the dumping female, (3) the dumper was related to one of the host pair, and (4) there was a benefit to the host pair because they gained extra helpers. These hypotheses were not supported by the data, but further tests are needed. As the cost of desertion was greater than the cost of acceptance of parasitic eggs, hosts may have been forced to accept.     

An adaptive annual rhythm in the sex of first pigeon eggs

When the reproductive value of male and female offspring varies differentially, parents are predicted to adjust the sex ratio of their offspring to maximize their fitness (Trivers and Willard, Science 179:90–92, 1973). Two factors have been repeatedly linked to skews in avian offspring sex ratio. First, laying date can affect offspring sex ratio when the sexes differ in age of first reproduction, such that the more slowly maturing sex is overproduced early in the season. Second, position of the egg in the laying sequence of a clutch may affect sex ratio bias since manipulating the sex of the first eggs may be least costly to the mother. We studied both factors in two non-domesticated pigeon species. Both the Wood pigeon (Columba palumbus) and the Rock pigeon (Columba livia) have long breeding seasons and lay two-egg clutches. In the field, we determined the sex of Wood pigeon nestlings. In Rock pigeons, housed in captivity outdoors, we determined embryo sex after 3 days of incubation. On the basis of their sex-specific age of first reproduction, we predicted that males, maturing at older age than females, should be produced in majority early and females later in the year. This was confirmed for both species. The bias was restricted to first eggs. Rock pigeons produced clutches throughout the year and show that the sex of the first egg followed an annual cycle. To our knowledge, this study presents the first evidence of a full annual rhythm in adaptive sex allocation in birds. We suggest that this reflects an endogenous seasonal program in primary sex ratio controlled by a preovulatory mechanism.     

Pigment allocation to eggs decreases plasma antioxidants in a songbird

Although evidence is accumulating on the adaptive function of female ornamentation, very little is known about maternal allocation decisions involving sexual signaling and other reproductive functions. Blue egg coloration has been suggested as a sexually selected signal of female quality to males, and some recent studies are in accordance with this hypothesis. Blue eggshell coloration results from the deposition of biliverdin pigment by laying females, which is a potent antioxidant. Thus, egg pigmentation should be costly in terms of antioxidants, an assumption of the signaling hypothesis that has not been tested yet. We induced increased reproductive effort in a set of female pied flycatchers Ficedula hypoleuca through nest removal and measured egg pigmentation and plasma antioxidant levels in relation with a control group. Experimental females showed a negative association between egg color and plasma antioxidant levels, while there was no relationship for control birds. This supports that egg pigmentation is costly in terms of general antioxidant defenses and suggests a tradeoff between the allocations to both traits. Furthermore, experimental females with more colorful eggs raised more fledglings, especially when breeding early. Controls did not show a relationship between egg color and reproductive success. Females laying more colorful eggs could have shifted their allocation decisions towards current reproduction, at the expense of their own antioxidant defenses. Our results highlight that blue egg coloration is a life-history trait, subject to tradeoffs with other attributes, and seems to be especially informative under harsh breeding conditions.     

Benefits of large broods by higher chick survival and better territories in a precocial shorebird

When reproductive success is constant in one breeding phase, different tactics that increase variation in reproductive success among individuals may evolve in other phases. For instance, in shorebirds, which usually have a limited clutch size of four eggs, variation in reproductive tactics among individuals is expected either before egg-laying (e.g. diverse mating systems) or after hatching of the young (e.g. diverse parental care). In this paper, I studied the pied avocet (Recurvirostra avosetta), a shorebird with a modal clutch size of four eggs, to test whether post-hatch chick adoption as an alternative tactic can be linked to increased variation in annual reproductive success. When predation was high, naturally adopting pairs produced more filial fledglings than did pairs not adopting chicks and not losing chicks to adoption. The number of filial fledglings increased with the number of adopted young, possibly through diluting the chances of predation on filial young. Experimental chick addition did not lead to more fledged young due to low brood integrity as shown by the frequent loss of chicks from some experimental broods. When predation was low, larger broods occupied feeding territories with higher prey abundance than smaller broods, possibly due to their dominance over smaller ones. Pairs that lost chicks to adoption (donors) fledged as many filial young in their broods as did non-adopters/non-donors, whereas the total number of donors’ filial fledglings, including those raised in adopting broods, approached that of adopters. These findings show, for the first time, that post-hatch alternative reproductive tactics can lead to variation in annual reproductive success and to higher success for some pairs even in species where past adaptations limit variation in reproductive success in a certain phase of reproduction.     

Helpers and egg investment in the cooperatively breeding acorn woodpecker: testing the concealed helper effects hypothesis

In cooperatively breeding acorn woodpeckers (Melanerpes formicivorus), helper males have a large positive effect on fledging success in good acorn crop years but only a small positive effect in poor acorn crop years, while helper females exhibit the opposite pattern. Based on these findings, we tested the “concealed helper effects” hypothesis, which proposes that laying females reduce investment in eggs (with respect to their size, number, or quality) in a way that confounds helper effects and results in an absence of a relationship between helpers and breeding success. Results generally failed to support the hypothesis. Mean egg size was positively related to temperatures during the 10 days prior to egg-laying and negatively related to the food supply as indexed by the prior fall’s acorn crop, but there were no significant differences vis-à-vis helpers except for interactions with the acorn crop that only partly corresponded to those predicted. With respect to clutch size, females laid larger clutches when assisted by female helpers, opposite the pattern predicted. Although our results suggest that egg size is adjusted to particular ecological circumstances, we conclude that neither egg nor clutch size is adjusted in a way that confounds the apparent effects of helpers, as proposed by the concealed helper effects hypothesis.     

Siblicide in the cooperatively breeding laughing kookaburra (Dacelo novaeguineae)

I describe siblicide in the laughing kookaburra (Dacelo novaeguineae), a reverse size-dimorphic, cooperatively breeding kingfisher. Clutches were usually of three eggs, and nestlings hatched asynchronously, with intervals of 2–72 h between successive eggs. Siblicide occurred in two temporally and mechanistically distinct episodes. The youngest nestling died in one-third of all nests within days of hatching as a result of aggression from its elders. Kookaburra nestlings attacked each other using a hook on their upper beak – a rare example of a morphological specialisation for sibling rivalry. In one-fifth of all nests, the youngest nestling starved to death much later, without overt aggression, when nestling growth rates were highest. I examined the effects of food availability and competitive disparities between nestlings on the incidence of both types of siblicide. The probability of late, starvation-mediated mortality was negatively correlated with the number of male helpers. Early, aggressively mediated siblicide occurred in nests characterised by a suite of correlated variables that I call the ”kookaburra siblicide syndrome”: (1) no male helpers attended the nest, (2) the third-hatched nestling was much smaller than the second-hatched nestling, (3) the first and second nestling to hatch were male and female, respectively, and (4) there was a short hatch interval between the first two nestlings. The kookaburra siblicide syndrome variables could be inter-correlated if they were all related to the female’s condition at the onset of incubation. Females in poorer condition may be less likely to have male helpers, more likely to lay small third eggs, and more likely to hatch the first two eggs relatively synchronously because of nutritional constraints during the onset of incubation. These females may further promote siblicide by modifying the sexes of the first two nestlings. If a female hatches soon after an older but eventually smaller brother, dominance between the first two nestlings could be destabilised. I suggest this leads to escalated aggression in the nest and the death of the third nestling, which is least able to defend itself. Received: 17 December 1999 / Received in revised form: 8 May 2000 / Accepted: 20 May 2000     

Egg adoption can explain joint egg-laying in common eiders

Hypotheses regarding the evolution and maintenance of intraspecific nest parasitism were tested with data collected during a 3-year study of common eiders (Somateria mollissima) breeding near Churchill, Manitoba. The nest parasitism rate was highest (42.4% of nests) during the year with the highest nest density and the best environmental conditions, and lowest (20.2% of nests) in the year with the lowest nest density and the poorest environmental conditions. Over the nesting season, parasitic eggs were laid at the same time as normally laid eggs. Most parasitic eggs (>75%) were laid before the host female laid her third egg. The majority of the parasitic eggs were the first or second egg produced by the parasitic female. When a parasitic egg was laid before or on the same day as the host female initiated her clutch, the probability of her first egg being depredated before incubation was significantly lowered. First- and second-laid eggs suffered a high rate of predation probably because nesting females do not attend their clutch until their second or third egg is laid. Hypotheses that some females use intraspecific nest parasitism to parasitize the parental care of other females were inconsistent with these data. Egg adoption is a likely explanation for the prevalence of females incubating parasitic eggs in this population. Received: 30 September 1997 / Accepted after revision: 6 May 1998     

Costs of brood parasitism and the lack of defenses on the yellow-winged blackbird - shiny cowbird system

The shiny cowbird (Molothrus bonariensis) is a generalist brood parasite that lays either white-immaculate or spotted egg morphs in eastern Argentina and Uruguay. Some hosts accept both morphs, others accept spotted eggs and reject the white morph, but no host has been found to accept white eggs and reject spotted ones. It has been suggested that the yellow-winged blackbird (Agelaius thilius) may be that type of host. The finding of a white acceptor-spotted rejector species would help to explain the occurrence and maintenance of the parasite egg polymorphism. We studied the incidence of shiny cowbird parasitism on this host, its costs for their reproductive success and the presence of antiparasitic defenses in the yellow-winged blackbird - shiny cowbird system. The parasite affected the reproductive success of the host in two ways. Cowbirds punctured host eggs causing a reduction in clutch size, and yellow-winged blackbirds deserted their nests whenever they suffered high egg loss. In addition, parasitized nests suffered higher predation during the nestling stage, but not during egg stages, indicating that the difference found was related to the presence of the cowbird chick, and not to higher exposure of parasitized nests to both parasites␣and predators. Despite the costs imposed by the parasite, yellow-winged blackbirds have not evolved antiparasitic defenses. This host did not reject any egg morph of the shiny cowbird nor desert parasitized nests unless it had suffered high egg loss. Current explanations for the host lack of defenses, the “time lag” and the “equilibrium” hypothesis, are discussed. Received: 29 August 1997 / Accepted after revision: 10 January 1998     

Primary sex ratios in zebra finches: no evidence for adaptive manipulation in wild and semi-domesticated populations.

When breeding diet is restricted, domesticated zebra finches,Taeniopygia guttata, produce male-biased primary and secondary sex ratios, but unexpectedly produce unbiased ratios when food is unrestricted. We investigated the primary sex ratios (at laying) of wild zebra finches in southeastern Australia in response to food supplementation and environmental factors predicted to enhance female breeding condition and to bias the primary sex ratio towards daughters. Molecular sexing of all nestlings in 54 complete broods where every egg hatched, failed to show any significant biases from random. Time of egg laying (month, season) and environmental conditions (rainfall, temperature) did not significantly predict variation in the primary sex ratio, but time of breeding did affect clutch size. Wild zebra finches at our colony did not bias their sex allocation as there were no differences in the primary sex ratio and no differences in the numbers and mass of sons and daughters at the end of parental care (day 35–40 post-hatch). Biases in primary sex ratio of our wild population are probably weak or non-existent possibly due to the unpredictable environment and/or multiple contrary selective forces acting on sex ratios. We also investigated the effects of photoperiod, biases in the adult sex ratio, and parental attractiveness on primary sex ratios of semi-domesticated, laboratory zebra finches. Molecular sexing of three-day old embryos from complete clutches, failed to reveal significant biases from random. In contrast to previous studies, sex of eggs did not correlate with laying order and egg mass declined with order, rather than increased. Domestication may be responsible for these differences.     

Breeding synchrony and paternity in the barn swallow (Hirundo rustica)

The barn swallow (Hirundo rustica) is a socially monogamous passerine which usually breeds in colonies where extra-pair copulations are frequent. Males intensively guard their mates during the female fertile period. Since males are more likely to be available for extra-pair copulations when their mate is not fertile, synchrony in timing of breeding may affect paternity of individual males. In this study, we analysed the change in mate-guarding rate by males in relation to the fertility condition of the female, and the relationships between breeding synchrony and density with paternity in first broods of 52 male barn swallows. Paternity (proportion of nestlings fathered in own brood) was assessed by typing of three highly polymorphic microsatellite loci. Mate guarding by males peaked during the fertile period of their mates. Paternity increased as breeding synchrony in the colony increased. Paternity of barn swallows is positively associated with the degree of exaggeration of male tail ornaments. The relationship between male ornamentation and paternity was partly mediated by an effect of ornament size on breeding synchrony. We suggest that females might delay breeding with low-quality males to enhance their opportunities for being fertilised by high-quality extra-pair males. Received: 16 January 1998 / Accepted after revision: 25 October 1998