Intraspecific nest parasitism in the Spotless Starling Sturnus unicolor

Intraspecific nest parasitism in two colonies of Spotless Starling Sturnus unicolor breeding in nestboxes was studied in central Spain from 1991 to 1994. Nests were monitored regularly and three criteria were used to detect nest parasitism: the appearance of more than one egg per day during the laying period of the host; the appearance of an egg after the start of incubation; eggs with unusual shape or pigmentation. The proportion of parasitized nests in first clutches (37%) was twice that of intermediate (19%) or second (20%) clutches in colony B, whereas parasitism occurred in first (35%) and intermediate (12%) but not in second clutches in colony A. Most clutches (52–70%) were parasitized during the host's laying period and received one parasitic egg. In 10% of the parasitized clutches in colony B, one of the host's eggs disappeared on the day the parasitic egg was added, suggesting that the parasitic female removed this egg. Although parasitism increased clutch size significantly, it led to a decrease in host breeding success, mainly through the removal of eggs and the loss of host nestlings and the survival of parasitic chicks. Observations suggested that parasitic females were young individuals without their own nests and/or those whose breeding attempt had been disrupted while laying in their own nest.     

Conspecific nest parasitism in the Pied Avocet Recurvirostra avosetta

Evidence for the occurrence of conspecific nest parasitism (CNP) in Pied Avocets Recurvirostra avosetta is presented. Clutches of more than four eggs had obviously been produced by more than one female but were incubated by only one pair each. Minimum estimates for the frequencies of parasitized clutches and parasitic eggs were 3.3% and 1.3% respectively. CNP increased in frequency in colonies with higher nest densities. The frequency of CNP was unaffected by the rate of nest failures early in the season. On average, parasitic eggs were laid earlier in the season than the majority of non-parasitic eggs. Parasitic Avocets usually deposited their eggs during the laying period of host nests. Parasitized nests had significantly longer incubation periods than unparasitized nests. Hatching success in supernormal clutches was insignificantly reduced compared with four-egg clutches. The annual breeding success of individuals with parasitized clutches was considerably (but not significantly) higher than those of non-parasitized individuals. This was probably due to the fact that parasites chose to deposit their eggs in dense colonies whose members had significantly higher breeding success than the individuals in loose colonies. In colonies with a high rate of CNP, the frequency of clutches of more than five eggs increased. These clutches had little chance of survival.     

Breeding success and chronology of Wood Storks Mycteria americana in northern and central Florida, U.S.A.

The breeding success and chronology of Wood Storks Mycteria americana were studied at eight colonies in northern and central Florida during 1981–1985. Mean ± s.d. clutch size for all colony-years was 3.07 ± 0.56 (n = 2694 nests), with three-egg clutches (72%) most frequent. Mean clutch size among all colonies and years ranged from 2.73 ± 0.55 to 3.41 ± 0.61. Many colonies exhibited significant negative trends in clutch size with, hatching date because of a proportional decrease in four-egg clutches later in the season. Mean colony clutch size was not correlated with nest numbers, nesting density or mean hatching date within most years. Mean ± s.d. number of fledglings for all colonies and years was 1.29 ± 1.16 fledglings per nest (n = 2812 nests). Mean annual fledging rates in colonies ranged from 0 (colony failed) to 2.66 fledglings per nest. Most breeding failure occurred prior to egg hatching, and the second highest mortality occurred between hatching and 2 weeks of age. Four-egg clutches fledged more storks than three-egg clutches, which in turn were more successful than two-egg clutches. However, all clutch sizes showed similar fledgling per egg rates. The seasonal decline in productivity was associated proportionally with smaller clutch sizes later in the breeding season. An increase in mean hatching date was correlated with an increase in latitude. There was greater within-year breeding synchrony among colonies than interyear breeding synchrony within each colony. Breeding synchrony was not correlated with mean hatching date, latitude, longitude, nest numbers or nesting density.     

Patterns of variation in size of Boat-tailed Grackle Quiscalus major eggs

Boat-tailed Grackle Quiscalus major eggs averaged 8^.09 g wet weight. Mean egg weight represented 8 05 % of female body weight. Based on egg weight, Grackle eggs have a shorter incubation period than the general passerine pattern.
Egg weight varied significantly with sequence of laying and between clutches in both two-egg and three-egg clutches. Last laid eggs weighed less than first laid eggs. The mean weight of the first two eggs laid in three-egg clutches did not differ from the mean egg weight of two-egg clutches. The average clutch of two and three eggs weighed 16.33 g and 24.16 g, respectively.
Mean egg weight varied between study colonies and with season. Grackles at East Lake laid eggs that weighed less than grackles at either Alligator Lake or North Lake. Eggs laid during March averaged less than eggs laid later in the season. The locality variation reflects the different timing of nesting between sites rather than food abundance. As nutrient provisioning increases with an increase in egg weight, the seasonal change in egg weight probably reflects improved feeding conditions for the female. This suggests that selection favours beginning laying before reserves of the female are sufficient to lay the largest egg possible.
The size of the young at hatching was correlated with egg size. Newly hate had young averaged 79.6% of fresh egg weight. Egg weight did not determine the probability of hatching or starvation of the young. Females do not appear to adjust egg size facultatively depending on the sex of the young. Eggs producing males and females did not vary significantly in weight, whether comparisons were made within or between clutches.     

Multiple maternity in black-headed gull Larus ridibundus clutches as revealed by protein fingerprinting

Social monogamy with biparental care is the norm in gulls Laridae , but egg colour variation suggests that some nests may contain mixed clutches laid by more than one female. Here we use protein fingerprinting of egg albumen to assess the occurrence of mixed maternity clutches in three colonies of black-headed gulls. Among 160 analysed clutches with >1 egg, 34% contained eggs from more than one female, and 15% of the eggs in clutches >1 came from other females than the major female (laying most eggs in nest). Among clutches with 2–3 eggs 28% were mixed, and among clutches with 4 or more eggs 89% contained eggs from two or more females. There were significantly fewer eggs from the major female in mixed nests (mean=2.06±0.63 SD) than in non-mixed nests (mean=2.82±0.43 SD). In nests without evidence of female conflict, hatching success of minority eggs was similar to that of eggs from the major female (12.5 and 8.4%, respectively). In 21% of mixed maternity nests, one or more minority eggs was buried or punctured, and 25% of eggs from major females were also found evicted, suggesting conflict between females and rejection of eggs. Intra-specific nest parasitism seems the most likely cause of mixed clutches, but there are also other possible causes.     

FACTORS AFFECTING THE EGG SIZE OF RED-BILLED GULLS LARUS NOVAEHOLLANDIAE SCOPULINUS

The factors influencing the egg size of the Red-billed Gull Larus novaehollandiae scopulinus were studied at Kaikoura, New Zealand, between 1964 and 1972. In two- and three-egg clutches there was a trend for the eggs to become smaller in the sequence of laying. Length, breadth and volume of eggs of one-, two- and three-egg clutches declined significantly as the season progressed. The size of eggs from single-egg clutches tended to be smaller than eggs from two-egg clutches laid at the same time. There were correlations between the proportions of one-egg and of three-egg clutches being laid at a given period and the mean egg volume of two-egg clutches. When the mean egg volume of two-egg clutches increased there was a corresponding increase in the proportion of two- and three-egg clutches laid. When the mean egg volume of two-egg clutches decreased there was an increase in the proportion of single-egg clutches laid. The egg size of the Red-billed Gull showed no direct correlation with the abundance or availability of food; the largest eggs were produced early in the season when food was in short supply. In spite of an increase in the food supply in the middle of the breeding season, birds laying at this time produced smaller eggs than birds which laid earlier in the season. However, early breeders which relayed at the peak in food abundance on average produced a larger replacement clutch than originals laid early in the season. It is suggested that the birds nesting early in the season are able to produce the largest eggs because they are the most efficient foragers for food, and those which nest later in the season produce smaller eggs, even at peak food abundance, because of their inefficiency or inexperience. Early breeders laying replacement clutches tended to lay larger eggs and larger clutches than birds which are producing their first clutches at the same time. Two-year-old females laid eggs which were significantly shorter than older aged birds while the breadth and volume of the egg increased with the age of the female up to the fifth year. There was a trend for females to lay larger eggs when mated with older rather than younger males. No statistical differences in egg size were detected between females changing or retaining the partner of the previous season. Female body weight and egg volume were positively correlated in females weighing less than 275 g but not for heavier females. It is suggested that the seasonal decline in egg size and clutch size results from a decrease in the availability of food and the ability of the individual to exploit the resource.     

Intraspecific nest parasitism in the European starling Sturnus vulgaris

Biochemical genetic markers were used along with conventional methods (abnormal laying sequence/clutch size, unusual egg shape/pigmentation) to identify intraspecific nest parasitism at two British nestbox colonies of the European starling. Between 11 and 37% of first clutches were parasitized during 1977–1979. Parasitic females probably comprised all of the following categories: (1) paired females contesting a nestbox occupied by another pair; (2) previously paired females who had laid a clutch but had been unsuccessful; (3) unpaired females who had copulated with males that already had a mate and nest site; and (4) ‘professional’ nest parasites who distributed at lest some of their eggs in one or more nests other than their own. Although parasitized nests had higher clutch sizes, parasitism led to fewer host young fledging per egg laid, mainly through the eviction of eggs and subsequent nest desertion. Number of parasitic young fledged per egg laid was highest when eggs were laid synchronously with the host, when host clutches were larger, or a smaller number of parasite eggs were added to a nest, thus favouring parasites that distribute their eggs amongst a number of nests. A greater pressure on nest sites may have accounted for the higher levels of parasitism at the Aberdeen colony and for the greater number of parasite eggs laid in a nest. Although most parasitic female starlings appeared to be much less successful than non-parasitic ones, nest parasitism in the starling might evolve directly when one or more of the following advantages are present. (1) There are no constraints on the number of eggs a female may lay but there are constraints on the number of young she may feed adequately. (2) Female survival is increased by having fewer or no eggs/young to care for. (3) Current feeding conditions favour the survival of more young than would be produced by the most common clutch size. Intraspecific nest parasitism is considered to be a first stage in the evolution of interspecific nest parasitism.     

Brood parasitism,female condition and clutch reduction in the common eider Somateria mollisima

Conspecific brood parasitism (CBP) is an alternative reproductive tactic found in many animals with parental care. Parasitizing females lay eggs in the nests of other females (hosts) of the same species, which incubate and raise both their own and the foreign offspring. The causes and consequences of CBP are debated. Using albumen fingerprinting of eggs for accurately detecting parasitism, we here analyse its relation to female condition and clutch size in High Arctic common eiders Somateria mollissima borealis. Among 166 clutches in a Svalbard colony, 31 (19%) contained eggs from more than one female, and 40 of 670 eggs (6%) were parasitic. In 6 cases an active nest with egg(s) was taken over by another female. Many suitable nest sites were unoccupied, indicating that CBP and nest takeover are reproductive tactics, not only consequences of nest site shortage. Similarity in body mass between female categories suggests that condition does not determine whether a nesting female becomes parasitised. There was no evidence of low condition in parasites: egg size was similar in hosts and parasites, and parasitism was equally frequent early and late in the laying season. Meta‐analysis of this and 3 other eider studies shows that there is a cost of being parasitised in this precocial species: host females laid on average 7% fewer eggs than other females.     

Incubation capacity contributes to constraints on maximal clutch size in Brent Geese Branta bernicla nigricans

Lack ( 1967 ) proposed that clutch size in species with precocial young was determined by nutrients available to females at the time of egg formation; since then others have suggested that regulation of clutch size in these species may be more complex. We tested whether incubation limitation contributes to ultimate constraints on maximal clutch size in Black Brent Geese (Black Brant) Branta bernicla nigricans. Specifically, we investigated the relationship between clutch size and duration of the nesting period (i.e. days between nest initiation and the first pipped egg) and the number of goslings leaving the nest. We used experimental clutch manipulations to assess these questions because they allowed us to create clutches that were larger than the typical maximum of five eggs in this species. We found that the per‐capita probability of egg success (i.e. the probability an egg hatched and the gosling left the nest) declined from 0.81 for two‐egg clutches to 0.50 for seven‐egg clutches. As a result of declining egg success, clutches containing more than five eggs produced, at best, only marginally more offspring. Manipulating clutch size at the beginning of incubation had no effect on the duration of the nesting period, but the nesting period increased with the number of eggs a female laid naturally prior to manipulation, from 25.4 days (95% CI 25.1–25.7) for three‐egg clutches to 27.7 days (95% CI 27.3–28.1) for six‐egg clutches. This delay in hatching may result in reduced gosling growth rates due to declining forage quality during the brood rearing period. Our results suggest that the strong right truncation of Brent clutches, which results in few clutches greater than five, is partially explained by the declining incubation capacity of females as clutch size increases and a delay in hatching with each additional egg laid. As a result, females laying clutches with more than five eggs would typically gain little fitness benefit above that associated with a five‐egg clutch.     

Egg patterning is not a reliable indicator of intraspecific brood parasitism in the blue tit Cyanistes caeruleus

A recent study reported the incidence of intraspecific brood parasitism (IBP) in the blue tit, based on differences in pigmentation between eggs within clutches and the appearance of multiple eggs within a nest in a single day. Here we present data from another population of blue tits Cyanistes caeruleus in which we show that such evidence is not necessarily indicative of IBP. As in many other studies of birds, we found that the patterning and degree of pigmentation on eggs is individually highly characteristic of a female. As in the previous study of blue tits, a minority of clutches (about 3% of 904 clutches) contained one, two or three eggs that were distinctly different from the other eggs in the clutch, either in size or patterning, but not both. Molecular genetic analyses of parentage in a sample of these clutches with suspected IBP demonstrated unequivocally that all eggs within each clutch had in fact been laid by a single female, in all cases the social parent. We also found no evidence of IBP from paternity analyses of a sample of 3,529 offspring from 427 broods in this population, adding to previous evidence of the absence of IBP in this species. These findings cast doubt on the utility of purely observational approaches to provide evidence of intraspecific brood parasitism; our study suggests that egg morphology and records of laying patterns are inadequate to enable the identification of IBP.     

Cuckoos versus reed warblers: Adaptations and counteradaptations

At study sites in Cambridgeshire, England, the percentage of reed warbler, Acrocephalus scirpaceus, nests parasitized by cuckoos, Cuculus canorus, in 2 years was 22·5% and 9·1%. The warblers rejected cuckoo eggs at 19% of parasitized nests. Parasitized clutches suffered less predation than unparasitized clutches, suggesting that the cuckoo itself was the major predator, plundering nests too advanced for parasitism so that the hosts would re-lay. The cuckoos laid a mimetic egg, parasitized nests in the afternoons during the host laying period, usually removed one host egg, laid a remarkably small egg and laid very quickly. Nests were experimentally parasitized with model eggs to study the significance of this procedure. Experiments showed that host discrimination selects for: (1) egg mimicry by cuckoos (poorer matching model eggs were more likely to be rejected); (2) parasitism during the laying period (mimetic eggs put in nests before host laying began were rejected); (3) afternoon laying (mimetic eggs were less likely to be accepted in the early morning than in the afternoon, when hosts were more often absent from the nest); (4) a small egg (large eggs, typical of non-parasitic cuckoos, were more likely to be rejected); (5) rapid laying (a stuffed cuckoo on the nest stimulated increased rejection of model eggs), and (6) sets a limit to host egg removal by cuckoos (if more than one or two are removed desertion may occur). Mimicry may also be selected for because it reduced the chance that second cuckoos can discriminate the first cuckoo's egg from the host's clutch. Predation did not select for mimicry; nests with a non-mimetic egg did not suffer greater predation than those with a mimetic egg. Host rejection of model eggs did not depend on: (1) stage of parasitism once host egg laying had begun (nevertheless cuckoos were more likely to lay early in the host laying period probably to increase the chance the cuckoo chick hatched); (2) removal of a host egg (however, this reduced the incidence of unhatched eggs so cuckoos may remove a host egg so as not to exceed the host incubation limit). There were two costs of rejection, an ‘ejection’ cost (own eggs ejected as well as the cuckoo egg) and, with mimetic eggs, a ‘recognition’ cost (own eggs ejected instead of the cuckoo egg). Reed warblers did not discriminate against unlike chicks (another species) and did not favour either a cuckoo chick or their own chicks when these were placed in two nests side by side. Possible reasons why the hosts discriminate against unlike eggs but not unlike chicks are discussed.     

Reliability of egg morphology to detect conspecific brood parasitism in goldeneyes Bucephala clangula examined using protein fingerprinting

Eadie (1989) developed a method based on variation between females in egg length, width and weight to detect conspecific brood parasitism in the field: using these three egg measures, Euclidean distance between all pairs of eggs within a clutch is calculated, and if maximum Euclidean distance (MED) between any two eggs exceeds a threshold value the nest is considered parasitized. The MED method has been tested in Finnish and Scottish common goldeneye Bucephala clangula populations but the results have been contradicting. Here we use protein fingerprinting to assess the validity of the MED method. Data comprised 35 clutches of which we knew, based on protein fingerprinting, how many different females laid the clutch (range 1–5 females). The mean MED of non-parasitized clutches (laid by 1 female only) was 1.470 (95% CI: lower 1.169, upper 1.771; n=21) and that of parasitized clutches (laid by 2 or more females) was 3.654 (95% CL: lower 3.083, upper 4.225; n=14). Using a MED>3.0 as a criterion to identify parasitized clutches 89% of all clutches were classified correctly either parasitized or non-parasitized when compared to the identification based on protein fingerprinting. Clutch size and the number of females (beyond 2 females) did not affect the clutch MED, whereas the status of parasitism did. Repeatability of egg length, width and weight were: 0.63, 0.76 and 0.80, respectively, implying that, variation in these egg measures occurs among rather than within females. Our new results confirm that the MED method is reliable enough to detect parasitism in common goldeneye.     

Intraspecific variation in size and density of Avocet colonies: effects of nest-distances on hatching and breeding success

In many colonial bird species there is considerable intraspecific variation in colony size and inter‐nest distance (colony density). Possible causes of this variation and its effects on hatching success (survival of eggs) and breeding success (probability of a pair raising chicks) were studied in 48 Avocet Recurvirostra avosetta colonies in Schleswig‐Holstein (Germany) between 1991 and 1996. Colony density was influenced by time of year and habitat (categories: island or mainland, close to or far from feeding grounds). Colonies on islands had the highest densities. When all available space at a colony site was used, colonies became very dense (mean nearest‐neighbour nest distance less than 1 m). Colony size (number of clutches) was influenced by time of year, but not by habitat. Hatching success was low in high density colonies and in very low density ‘colonies’ (single nests) and high over a broad range of intermediate nest densities. The low success rate of single nests was caused by a very high predation rate, whereas the low success rate in very dense colonies was caused by a high rate of nest abandonment. Nest abandonment in very dense colonies was associated with a high level of aggressiveness among Avocets during the egg‐laying period. Due to territorial behaviour, Avocets seemed to be expelled from the densest breeding sites. In very dense colonies, high frequencies of clutches of unusual size occurred due to conspecific nest parasitism. The number of Avocets taking part in attacks on potential egg predators was small and (in colonies of more than one clutch) depended neither on colony size nor on colony density. Despite a low hatching success in very dense colonies, individuals breeding in the densest colonies had significantly better chances of raising chicks than Avocets breeding in less dense colonies. Coloniality seemed to be obligatory for Avocets in order to ensure hatching success. The size and density of colonies seemed to be associated with the availability of suitable nesting habitats (islands).     

Temporal patterns of host availability, brown-headed cowbird brood parasitism, and parasite egg mass

I studied relationships between temporal patterns of host availability, brood parasitism, and egg mass for the parasitic brown-headed cowbird (Molothrus ater). At a study site consisting largely of edge habitat in north-eastern Illinois, I found 834 bird nests from 27 species. A total of 407 cowbird eggs and nestlings were found in these nests over three laying seasons. Nearly all (n= 379; 93%) were found in the nests of seven host species. For these species and all taken together, weekly nest availability generally decreased whereas parasitism frequency generally increased throughout the cowbird laying season, but the proportions of nests parasitized and the mean number of cowbird eggs in them did not. Additionally, no correlation was found between the proportions of nests parasitized and nest availability. Cowbird egg mass generally increased throughout the laying season, indicating that foraging conditions improved and that, early in the laying season, egg mass and quality may be less important than quantity. Consistently high weekly levels of parasitism indicate that cowbird reproduction was less limited by resources needed for egg production and more by the availability of suitable host nests. Fluctuating weekly host availabilities suggest that previously established, constant rates of cowbird egg laying would produce an excess of eggs during periods of low host availability. Further, the low frequency of parasitism (1%) of nests in stages too advanced for successful parasitism, and of abandoned nests, is consistent with the hypothesis that cowbirds' consistently high rate of egg production helps assure an egg is available when an appropriate nest is found. Frequently, nests were parasitized multiple times, raising the possibility that cowbirds were interfering with their own reproduction. A diverse host community increases the possibility that a decline of any one host species is unlikely to affect cowbird reproduction significantly. Received 11 July 1997 / Accepted: 31 March 1998     

Intraspecific nest parasitism and anti-parasite behavior in the grey starling,Sturnus cineraceus

I studied intraspecific nest parasitism in the grey starlingSturnus cineraceus in 1992 and 1993. The population in this study consisted of 290 nests (157 nests in 1992 and 133 nests in 1993) in which the clutches were completed before May 10 in the year studied. Twenty-nine nests in 1992 and 32 nests in 1993 contained at least 1 parasitic egg. Hatching success per nest of parasitized nests was slightly higher than that of non-parasitized nests. However, fledging success per nest of parasitized nests was significantly lower than that of non-parasitized nests. Thus parasitism appeared to reduce the reproductive success of hosts. Hosts exhibited a few behaviors that minimized the potential cost of brood parasitism. These behaviors included throwing out the parasitic egg and nest guarding. Hosts threw out parasitic eggs before the onset of laying, but they never did so to parasitic eggs laid after that period. The nest guarding level was low during the hosts’ laying periods, and one observed nest was parasitized during this time. Thus, nest-guarding behavior was not effective as an anti-parasite behavior. Grey starlings do not appear to adopt strategies effective in reducing parasitism.     

Brood parasitism and egg matching in the Red-chested Cuckoo Cuculus solitarius in southern Africa

Host usage and relative rates of egg matching were investigated in the Red-chested Cuckoo Cuculus solitarius in southern Africa, using nest record cards and museum collections. Eighteen host species were found to be parasitized at varying degrees of intensity (0.14–12.5%). The most commonly recorded parasitized host, the Cape Robin Cossypha caffra , had a relatively low rate of parasitism (2.46%). The host species experiencing the most recorded pressure from parasitism was the Bearded Robin Erythropygia quadrivirgata , with 12.5% parasitism. Human perception of cuckoo/host egg matching was assessed for parasitized clutches of host species in museum egg collections. Eggs of three different cuckoo egg morphs were scored as matching those of the host species on a 1–5 scale. Perfect/good matching was recorded for eggs found in Chorister Cossypha dichroa , Heuglin's Cossypha heuglini and Natal Robins' Cossypha natalensis clutches. Poor and very poor matching was evident for cuckoo eggs found in four host species' clutches: the Cape Robin, Stonechat Saxicola torquata , Cape Rockthrush Monticola rupestris and Black Flycatcher Melaenornis pammelaina . Available evidence suggests that the Red-chested Cuckoo parasitizes hosts in a particular environment (low vegetation and trees). Good to intermediate matching was recorded with only 47% of host eggs, and with only 28.5% of Cape Robin clutches. A relatively high degree of host specificity, however, is suggested by the nest record card data, which indicate that species with large numbers of records are not those with the highest rates of parasitism.     

Relationship between clutch size,egg volume and hatching success in a Yellow-legged Gull Larus michahellis colony in south-eastern Tunisia

This study aimed to test the hypothesis that clutch size covaries with egg volume and hatching success in the Yellow-legged Gull Larus michahellis. We determined clutch size and egg volume in a sample of 131 nests, and we used the data to check whether egg volume varied among nests according to clutch size, while taking into account the effects of egg laying order. We also estimated hatching success rate and investigated the relationship between hatching success and clutch size. Egg volume varied among clutches according to clutch size, with eggs being larger in three-egg clutches than in two-egg clutches. Moreover, three-egg clutches showed higher daily survival rates, and hence hatching success, than two-egg clutches. Overall, our results suggest that in the Yellow-legged Gull clutch size covaries with egg volume and hatching success, which could possibly reflect an age effect through different mechanisms. Indeed, older females could be hypothesised to exhibit greater breeding performance than younger females because of their higher experience in tapping energy resources for egg formation and defending nests from dangers. Moreover, due to their age, older females are likely to have lower residual reproductive potential and should invest more heavily in current breeding attempts.     

Multiple responses to increasing spring temperatures in the breeding cycle of blue and great tits (Cyanistes caeruleus,Parus major)

Many bird species start laying their eggs earlier in response to increasing spring temperatures, but the causes of variation between and within species have not been fully explained. Moreover, synchronization of the nestling period with the food supply not only depends on first‐egg dates but also on additional reproductive parameters including laying interruptions, incubation time and nestling growth rate. We studied the breeding cycle of two sympatric and closely related species, the blue tit Cyanistes caeruleus and the great tit Parus major in a rich oak‐beech forest, and found that both advanced their mean first‐egg dates by 11–12 days over the last three decades. In addition, the time from first egg to fledging has shortened by 2–3 days, through a decrease in laying interruptions, incubation time (not statistically significant) and nestling development time. This decrease is correlated with a gradual increase of temperatures during laying, suggesting a major effect of the reduction in laying interruptions. In both species, the occurrence of second clutches has strongly decreased over time. As a consequence, the average time of fledging (all broods combined) has advanced by 15.4 and 18.6 days for blue and great tits, respectively, and variance in fledging dates has decreased by 70–75%. Indirect estimates of the food peak suggest that both species have maintained synchronization with the food supply. We found consistent selection for large clutch size, early laying and short nest time (laying to fledging), but no consistent changes in selection over time. Analyses of within‐individual variation show that most of the change can be explained by individual plasticity in laying date, fledging date and nest time. This study highlights the importance of studying all components of the reproductive cycle, including second clutches, in order to assess how natural populations respond to climate change.     

THE BIRDS OF LANGEBAAN LAGOON

Data are presented on breeding success of Red Bishops (Euplectes orix) collected over four breeding seasons at a colony in the Addo Elephant National Park, Eastern Cape, South Africa. Overall hatching and fledging success were 53.8% and 26.0% of all eggs laid, respectively, and the overall mean number of fledglings per breeding attempt was 0.77. Hatching and fledging success varied significantly among seasons, with both clutch and brood losses due to predation being the main reason for the observed differences. Hatching success also differed significantly among clutch sizes, being highest for four-egg clutches (63.2%), intermediate for three-egg clutches (55.5%) and lowest for two-egg clutches and five-egg clutches (33.2% and 34.3%, respectively). However, fledging success was not significantly different among clutch sizes. The mean number of fledglings per breeding attempt was 0.44 for two-egg clutches, 0.80 for three-egg clutches, 1.10 for four-egg clutches, and 0.57 for five-egg clutches. The height of accepted nests (i.e.nests in which at least one egg was laid) was significantly lower than the height of nests not accepted. In addition, accepted nests in which eggs hatched and young fledged were significantly lower than accepted nests in which no eggs hatched and no young fledged. These overall effects of nest height on nest acceptance and hatching and fledging success were, however, due only to nests built above water, since no such effects were found when nests built above ground (i.e.on dry land) were analysed separately. I detected no effect of nest coverage on the probability of a nest being accepted, nor was there any effect of nest coverage on hatching or fledging success. Nests above water were significantly more likely to be accepted than nests above ground; however, hatching and fledging success of nests that were accepted did not differ significantly between nests built above water and those built above ground.     

Cuckoo parasitism and productivity in different magpie subpopulations predict frequencies of the 457bp allele: a mosaic of coevolution at a small geographic scale

The level of defense against great spotted cuckoo (Clamator glandarius) parasitism in different European populations of magpie (Pica pica) depends on selection pressures due to parasitism and gene flow between populations, which suggests the existence of coevolutionary hot spots within a European metapopulation. A mosaic of coevolution is theoretically possible at small geographical scales and with strong gene flow, because, among other reasons, plots may differ in productivity (i.e., reproductive success of hosts in the absence of parasitism) and defensive genotypes theoretically should be more common in plots of high productivity. Here, we tested this prediction by exploring the relationship between parasitism rate, level of defense against parasitism (estimated as both rejection rate and the frequency of the 457bp microsatellite allele associated with foreign egg rejection in magpies), and some variables related to the productivity (average laying date, clutch size, and number of hatchlings per nest) of magpies breeding in different subpopulations. We found that both estimates of defensive ability (egg rejection rate and frequency of the 457bp allele) covaried significantly with between-plot differences in probability of parasitism, laying date, and number of hatchlings per nest. Moreover, the parasitism rate was larger in more productive plots. These results confirm the existence of a mosaic of coevolution at a very local geographical scale, and the association between laying date and number of hatchlings with variables related to defensive ability and the selection pressure arising from parasitism supports the prediction of coevolutionary gradients in relation to host productivity.