Nesting ecology of a naturalized population of Mallards Anas platyrhynchos in New Zealand

Investigating the reproductive ecology of naturalized species provides insights into the role of the source population's characteristics vs. post‐release adaptation that influence the success of introduction programmes. Introduced and naturalized Mallards Anas platyrhynchos are widely established in New Zealand (NZ), but little is known regarding their reproductive ecology. We evaluated the nesting ecology of female Mallards at two study sites in NZ (Southland and Waikato) in 2014–15. We radiotagged 241 pre‐breeding females with abdominal‐implant transmitters and measured breeding incidence, nesting chronology and re‐nesting propensity. We monitored 271 nests to evaluate nest survival, clutch and egg size, egg hatchability and partial clutch depredation. Breeding incidence averaged (mean ± se) 0.91 ± 0.03, clutch size averaged 9.9 ± 0.1 eggs, 94 ± 2% of eggs hatched in successful nests, partial depredation affected 6 ± 1% of eggs in clutches that were not fully destroyed by predators, and re‐nesting propensity following failure of nests or broods was 0.50 ± 0.003. Nesting season (first nest initiated to last nest hatched) lasted 4.5 months and mean initiation date of first detected nest attempts was 28 August ± 3.3 days. Smaller females were less likely to nest, but older, larger or better condition females nested earlier, re‐nested more often and laid larger clutches than did younger, smaller or poorer condition females. Younger females in Southland had higher nest survival; cumulative nest survival ranged from 0.25 ± 0.007 for adult females in Waikato to 0.50 ± 0.007 for yearling females in Southland. Compared with Mallards in their native range, the nesting season in NZ was longer, clutches and eggs were larger, and nest survival was generally greater. Different predators and climate, introgression with native heterospecifics and/or the sedentary nature of Mallards in NZ may have contributed to these differences.     

Time-dependent reproductive decisions in the blue tit

Many breeding attempts in birds do not result in any fledged young due to predation on eggs or young. Consequently, the influence of time constraints on reproductive decisions are integrated parts of the reproductive behaviour of birds breeding within short, seasonal climate zones. In this study, I mimicked nest predation by removing blue tit (Parus caeruleus) clutches shortly after completion. Around 75% of the removed clutches were followed by a repeat clutch. Females producing their first clutch early in the season and females with an early onset of incubation in the laying sequence (an indication of high parental or territory quality) were most likely to initiate a repeat clutch. A trade‐off between the benefits of a repeat clutch and survival likely stopped late females in bad condition from investing more in the current reproductive season. Females producing a repeat clutch laid fewer eggs, had an earlier onset of incubation in the laying sequence and produced larger eggs than they did when producing their original first clutch. Eggs produced after the onset of incubation were especially large in the repeat clutches. Since food availability was presumably higher when the female produced her repeat clutch compared with her first clutch, females made a strategical decision when reducing clutch size, whereas onset of incubation and egg size may have been energetically constrained when producing the first clutch. Females that produced a relatively large clutch, had a relatively early onset of incubation, and laid relatively large eggs in their first clutch also did so when producing a repeat clutch, indicating that some of the variation in breeding parameters are due to differences in parental or territory quality. Differences between years in the temperature‐dependent development rate of caterpillars seem to affect the time constraints on breeding. A year with a predicted early seasonal decline in caterpillars resulted in short intervals between removal and relaying, small clutches and an early onset of incubation.     

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.     

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.     

Breeding success of the Eurasian eagle owl (<Emphasis Type="Italic">Bubo bubo</Emphasis>) and rodent population dynamics

The dependence of the breeding success of the eagle owl on the population dynamics of rodents, which are a staple of its diet in Mordovia, was traced. At peak numbers of rodents, large clutches and a high survival rate of fledglings were observed in the breeding pairs; after a year of depression of rodent populations, in the following year, the pairs do not nest at all or their breeding success is reduced to a minimum due to the death of the clutches. In 52% of the cases, the nesting pairs laid three eggs; in 31%, two eggs; in 9%, four eggs; in 4%, one egg or five eggs. The average clutch size was 2.78 ± 0.17. The average number of chicks grown from a successful nest was 2.41 ± 0.27. The nests in Mordovia were located at a distance of 1.1–3.7 km from residential areas.     

Maternal behavior and clutch manipulation in the ring-legged earwig (Dermaptera: Carcinophoridae)

The expression and maintenance of maternal behavior in the earwig,Euborellia annulipes, was examined through manipulation of clutch size, age, and species and through observations of interactions between brooding females. Females underwent discrete gonadotrophic cycles culminating in oviposition of first clutches that were highly variable in size. Neither the head capsule width nor the age of the mother was correlated with clutch size. Maternal care extended through embryogenesis and for the week following hatching. Clutch removal significantly shortened the interclutch interval, indicating that the presence of brood inhibited the onset of the second gonadotrophic cycle. Brooding females readily accepted replacement clutches of the same age. Thus, mothers did not appear to distinguish their own eggs from those of other females. Experimental doubling of clutch size did not significantly reduce the proportion hatching or fledging. In contrast, reducing clutch size diminished the percentage successfully fledging. Manipulation of clutch age resulted in reduced hatching/fledging success. Placing two females, each with newly laid clutches, in the same cage usually resulted in egg transfer from the nest of one female to that of the other within 12 h. Nests of females with larger forceps were significantly more likely to contain both clutches. When mothers with first clutches were paired with mothers with third clutches, eggs were more likely to be transferred to the nest of the older female.E. annulipes females with newly laid clutches appeared to accept as replacement clutches eggs of the earwigDoru taeniatum. Alien clutches were maintained for the typical duration of embryogenesis; however, noD. taeniatum hatchlings were observed.     

Captive breeding for reintroduction: influence of management practices and biological factors on survival of captive kaki (black stilt)

An important component of the restoration strategy for the critically endangered kaki or black stilt (Himantopus novaezelandiae) is captive breeding for release. Since 1981 1,879 eggs were collected from wild and captive pairs, with birds laying up to four clutches. Eggs were incubated artificially and most chicks reared by hand until released as juveniles (about 60 days) or sub‐adults (9–10 months). Because survival in captivity is a significant determinant of the number of birds available for release, we wished to identify sources of variation in mortality to assess potential impacts of management on productivity. Hatchability was 78% for captive‐laid eggs and 91% for wild‐laid eggs. Survival of hatched eggs was 82% by 10 months of age for both wild and captive birds. Most egg mortality occurred early in incubation and around hatching: the timing of mortality was unaffected by whether birds were captive or wild, hybrid or pure kaki, or when eggs were laid. Heavier hatchlings showed higher initial survival, as did chicks from wild parents. Hatchlings from fourth‐laid eggs showed lowest survival, even though hatchling mass tended to increase with hatch order. Survival of chicks subjected to major health interventions was 69% after 4 months. No differences in survival were found between different genders, hybrids and pure kaki, hand‐reared or parent‐reared birds, chicks hatching early or late in the season, different seasons, different‐sized groups of chicks, chicks reared in different brooders, juveniles kept in different aviaries, and chicks from subsequent clutches. Birds subjected to minor health interventions were equally likely to survive as healthy chicks (82%). Survival was high despite aggressive management (quadruple clutching and collecting late in the season). Differences between captive and wild birds suggest further improvements could be made to captive diet. Wide variation in hatchability between parent pairs substantiates the practice of breaking up poorly performing pairs. Zoo Biol 0:1–16, 2005. © 2005 Wiley‐Liss, Inc.     

A molecular genetic analysis of the communal nesting of the ostrich (Struthio camelus)

The ostrich breeding system is complex and unique; communal clutches are laid by several females, although only one female, the major female, and the resident territorial male provide parental care. More eggs are laid in the nest than can be incubated and the major female ejects surplus eggs from the incubated central clutch. Microsatellite markers were used to analyse the parentage of communal nests in Nairobi National Park. This revealed that major females contributed a disproportionate number of fertile eggs to the central, incubated clutch and that multiple paternity and maternity within a nest were common; 68.9% of all incubated eggs on a nest were not parented by both the resident territorial male and the major female of that nest. All the males fertilized eggs on the clutches of neighbouring males. Unexpectedly, every major female with her own nest was also simultaneously a minor female with incubated eggs on neighbouring clutches. The relatedness between females laying in the same nest was not significantly different from the population average and significantly less than that between chicks hatched from the same nest.     

Brood parasitism,relatedness and sociality: a kinship role in female reproductive tactics

Conspecific brood parasitism (CBP) is a reproductive tactic in which parasitic females lay eggs in nests of other females of the same species that then raise the joint brood. Parasites benefit by increased reproduction, without costs of parental care for the parasitic eggs. CBP occurs in many egg‐laying animals, among birds most often in species with large clutches and self‐feeding young: two major factors facilitating successful parasitism. CBP is particularly common in waterfowl (Anatidae), a group with female‐biased natal philopatry and locally related females. Theory suggests that relatedness between host and parasite can lead to inclusive fitness benefits for both, but if host costs are high, parasites should instead target unrelated females. Pairwise relatedness (r) in host–parasite (h‐p) pairs of females has been estimated using molecular genetic methods in seven waterfowl (10 studies). In many h‐p pairs, the two females were unrelated (with low r, near the local population mean). However, close relatives (r = 0.5) were over‐represented in h‐p pairs, which in all 10 studies had higher mean relatedness than other females. In one species where this was studied, h‐p relatedness was higher than between nesting close neighbours, and hosts parasitized by non‐relatives aggressively rejected other females. In another species, birth nest‐mates (mother–daughters, sisters) associated in the breeding area as adults, and became h‐p pairs more often than expected by chance. These and other results point to recognition of birth nest‐mates and perhaps other close relatives. For small to medium host clutch sizes, addition of a few parasitic eggs need not reduce host offspring success. Estimates in two species suggest that hosts can then gain inclusive fitness if parasitized by relatives. Other evidence of female cooperation is incubation by old eider Somateria mollissima females of clutches laid by their relatives, and merging and joint care of broods of young. Merging females tended to be more closely related. Eiders associate with kin in many situations, and in some geese and swans, related females may associate over many years. Recent genetic evidence shows that also New World quails (Odontophoridae) have female‐biased natal philopatry, CBP and brood merging, inviting further study and comparison with waterfowl. Kin‐related parasitism also occurs in some insects, with revealing parallels and differences compared to birds. In hemipteran bugs, receiving extra eggs is beneficial for hosts by diluting offspring predation. In eggplant lace bugs Gargaphia solani, host and parasite are closely related, and kin selection favours egg donation to related females. Further studies of kinship in CBP, brood merging and other contexts can test if some of these species are socially more advanced than presently known.     

Cryopreservation of houbara semen: A pilot study

The potential of producing viable houbara bustard Chlamydotis undulata undulata progeny with cryopreserved semen was investigated during the 1998 breeding season. Houbara semen was diluted and rapidly frozen into pellet form in LN₂ using dimethylacetamide (DMA) as a cryoprotectant. Thawed semen exhibited 24.86± 10.61% recovery of fresh sperm motility levels and 36.44±16.17% of fresh viability counts. Two second‐year females were inseminated on two and three occasions, respectively, with thawed semen. One female laid four eggs between two clutches and the second female laid two eggs in a single clutch. All eggs were fertile; two developed to 18–21days but failed to pip; one chick died at pip and three chicks hatched. This is the first documented occasion that houbara bustard chicks were conceived with cryopreserved semen and it demonstrates the feasibility of using a rapid, simple, and relatively inexpensive methodology to achieve this result. Zoo Biol 18:147–152, 1999. © 1999 Wiley‐Liss, Inc.     

How avian incubation behaviour influences egg surface temperatures: relationships with egg position,development and clutch size

While understanding heat exchange between incubating adults and their eggs is central to the study of avian incubation energetics, current theory based on thermal measurements from dummy eggs reveals little about the mechanisms of this heat exchange or behavioural implications for the incubating bird. For example, we know little about how birds distribute their eggs based on temperature differences among egg positions within the nest cup. We studied the great tit Parus major, a species with a large clutch size, to investigate surface cooling rates of individual eggs within the nest cup across a range of ambient temperatures in a field situation. Using state‐of‐the‐art portable infrared imaging and digital photography we tested for associations between egg surface temperature (and rate of cooling) and a combination of egg specific (mass, shape, laying order, position within clutch) and incubation specific (clutch size, ambient temperature, day of incubation) variables. Egg surface temperature and cooling rates were related to the position of the eggs within the nest cup, with outer eggs being initially colder and cooling quicker than central eggs. Between foraging bouts, females moved outer eggs significantly more than centrally positioned eggs. Our results demonstrate that females are capable of responding to individual egg temperature by moving eggs around the nest cup, and that the energy cost to the female may increase as incubation proceeds. In addition, our results showing that smaller clutches experience lower initial incubation temperatures and cool quicker than larger clutches warrant further attention for optimal clutch size theory and studies of energetic constraints during incubation. Finally, researchers using dummy eggs to record egg temperature have ignored important elements of contact‐incubation, namely the complexity of how eggs cool and how females respond to these changes.     

Breeding seasons and laying patterns of the southern African Ostrich Struthio camelus

Wild Zimbabwe Ostriches Struthio camelus were studied during four successive years. Information on breeding seasons and laying patterns was compared with that of domesticated South African hybrid Ostriches in Bophuthatswana. In wild populations laying occurred mainly from July to December or early January, while domesticated birds continued until at least the end of February. Domesticated birds normally laid about 16 eggs in succession, one every second day. There was marked synchronisation of laying and the middle of each successive peak in egg production was about 6 weeks from the preceding peak. Wild birds laid up to eight eggs in any one nest, and normally clutches were contributed by three females, the average combined clutch being 12 or 13 per nest. Circumstantial evidence suggests that individual females may lay in more than one nest during a single laying sequence. Comparisons between rainfall patterns and laying rhythms proved inconclusive.     

Breeding biology of the Barn Owl Tyto alba in central Mali

Data were obtained on 178 clutches of African Barn Owls in central Mali from four breeding seasons during 1979–1983. Significantly more clutches were laid in 1979–1980 and significantly fewer in 1980– 1981 than the average for the 4 years and there were significantly more clutches laid in the middle period of the annual breeding season. The egg volume was significantly smaller at the beginning of the breeding season and significantly larger in the middle than the overall mean with eggs of second clutches being larger than those of first clutches. The clutch size was 605 eggs of which 479 hatched. The number of young fledged per successful nest was 319 and was 1 83 for all nesting attempts. The month was the only variable shown to affect significantly the clutch size, eggs hatched and fledging rate, the highest success rates being associated with the middle of the breeding period. The average interval between the hatching of eggs was 2–31 days. Survival rates (47'1%) to fledging were significantly affected by year (1981–1982 being the least) and month (mid-season birds the best). The order of hatching significantly affected age at death or disappearance, the first-hatched birds surviving the longest. The year significantly affected age at fledging, the young from the year in which most clutches were laid leaving the nest at the youngest age and those associated with the year having the least number of clutches remaining in the nest the longest. The month of hatching also affected fledging age, birds at the extremes of the breeding season fledging at older ages. The discussion compares these data with those from elsewhere.     

Proceedings of the New Zealand Society for Parasitology Inc.

Abstract

A study of the nesting habits and breeding biology of blue penguin Eudyptula minor was undertaken over the 1995–96 and 1996–97 breeding seasons on Matiu‐Somes Island, Wellington, New Zealand. Male and female blue penguins tended to be faithful to both mates and nest sites, although there was insufficient evidence to detect any association between a bird's breeding success in 1995 and a subsequent change of mate or nest in 1996. Over the 1995 and 1996 seasons the recorded hatching success (0.51 ±0.11 and 0.63 ± 0.10 respectively), fledging success (0.81 ±0.12 and 0.85 ±0.10 respectively) and reproductive success (0.41 + 0.11 and 0.54 ± 0.11 respectively) were similar each season. There was no significant difference between the proportion of eggs laid, or eggs hatched and chicks fledged, between the two seasons. The mean number of chicks raised over the two seasons was 0.94 ± 0.05 per nest. Replacement clutches were laid by 11 per cent of failed breeders in each season, but only in 1996 were they successful in fledging chicks.

No significant difference was found between the breeding success of the Matiu‐Somes Island blue penguin colony recorded during this study and a previous study undertaken on the island 40 years ago.     

Reproductive parameters in captive hand‐reared black‐bellied sandgrouse

Flock breeders of black‐bellied sandgrouse originated from wild‐laid eggs collected in west central Morocco in 2003 and 2004, were hatched, and hand‐reared in captivity in the framework of a reinforcement population program. Three to five pairs of different ages were housed in an aviary. Egg‐pulling procedure was used and eggs incubated artificially. Breeding parameters, hatchability, and posthatching mortality were recorded. The black‐bellied sandgrouse showed a seasonal breeding pattern with a laying period extending from 7 to12 weeks. The clutch frequency varied from 3 to 7 clutches per female per season. The mean clutch size was 2.66±0.47 eggs, and the mean interclutch interval was 10±2.7 days. The average total egg production was 12±5.83 eggs per female varying with age from 8 to18 eggs per female. Egg hatchability of incubated eggs increased with age and varied from 37.5 to 72.2%. Chick mortality occurred only in the first week after hatching, averaging 60.5%. The obtained results showed that black‐bellied sandgrouse can be successfully bred in captivity and opened the possibility of controlling, managing, and maximizing their production for the reinforcement of the local declining wild populations. Zoo Biol 27:269–281, 2008. © 2008 Wiley‐Liss, Inc.     

Differences in size between first and replacement clutches match the seasonal decline in single clutches in Tree Swallows Tachycineta bicolor

The seasonal decline in clutch size in birds can be a response to the environmentally conditioned decrease in prospects for offspring or a consequence of a lower physical ability of late‐breeding females. To find out which of the explanations apply in Tree Swallows Tachycineta bicolor, we assessed whether replacement clutch size in this species is affected by an individual female's ability to lay a certain number of eggs. To do this, we measured the decline in clutch size as a function of laying date between first and replacement clutches in individuals that re‐nested following natural failure, and compared this with the rate of decline in clutch size with laying date for Tree Swallows that laid only a single clutch in that season. Additionally, we assessed whether the clutch size and the rate of its seasonal decline varied across years. We accounted for the truncated and under‐dispersed nature of clutch size data by using a Bayesian approach in the analysis. We found little variation in the rate of clutch size decline across years at our breeding site. Accounting for this seasonal decline in clutch size, mean clutch size was similar between single‐time breeding females and those that laid replacement clutches, implying that the number of eggs laid on the second attempt by female Tree Swallows is determined by laying date, rather than by the female's physical ability to produce a clutch of a certain size.     

Manipulation of laying effort reveals habitat-specific variation in egg production constraints in Great Tits (Parus major)

It has been suggested that a bird's clutch size is not limited by the amount of resources available at the time of laying but that differences in the availability of food for nestlings is the ultimate underlying factor determining spatio-temporal variations in clutch size. However, habitat-related variations in egg production ability has yet to be investigated explicitly. We studied the breeding of Great Tits Parus major in deciduous and coniferous forests in the same area. The sizes of both the clutches and the eggs were, on average, larger in the former habitat than in the latter. A number of females were induced to lay more eggs than usual by removing four eggs from designated experimental clutches early in the laying period. These manipulated females laid approximately one egg more than control females, with the number of additional eggs laid not differing between the habitats. However, in both study years the relative size of the extra eggs – relative to the mean size of earlier laid eggs of the same clutch – was smaller in the coniferous habitat than in the deciduous habitat, while there was no habitat-related difference in the relative size of the last-laid eggs of control clutches. This result indicates that some form of proximate limitation during egg-laying period can contribute to the relatively small clutches and eggs in the coniferous habitat. Our results emphasize the need to take egg production costs into account when attempting to account for spatial variation in the reproductive behaviour of birds.     

Consequences of a female-biased sex-ratio in a socially monogamous bird: female-female pairs in the Roseate Tern Sterna dougallii

In the socially monogamous gulls and terns, female-biased sex ratios are sometimes revealed by the occurrence of ‘supernormal clutches’, which are usually attended by female-female pairs or other multi-female associations. We studied these phenomena in the endangered Roseate Tern Sterna dougallii at Bird Island, USA, from 1970 to 1995. DNA-techniques were used to sex breeding adults in 1992–94. Supernormal clutches (with three or four eggs) have comprised 1–7% of all Roseate Tern clutches at Bird Island since at least 1970, probably increasing in frequency since 1980. Supernormal clutches were spatially clustered; most were laid late in the peak period of nesting during each season. More than 80% of supernormal clutches and at least 7% of normal clutches were attended by multi-female associations; most of these were female-female pairs, with a few trios (male + two females, or three females) and one quartet (four females). More than half of the multi-female associations attended normal clutches. Some female-female pairs were maintained for up to five years. The age-distribution of females mated to females did not differ significantly from that of females mated to males. Females mated together usually laid eggs synchronously (±2 days). Such females laid fewer eggs than females mated to males (means 1.20 versus 1.73), and had lower fertility and hatching success (about 46% versus 98%); they were less successful in raising young from eggs that did hatch (means 58% versus 73%), but this difference was not significant. Their overall breeding success was much lower (about 0.34 fledglings per female versus 1.35). The sex-ratio of breeders was about 127 females to 100 males; about 20% of breeding females did not have male mates. Female Roseate Terns that do not obtain male mates appear to be of low phenotypic ‘quality’ - based on late laying, small clutches and small eggs. Our data support the hypothesis that such females have a higher fitness if they mate with each other and raise a few young than if they do not breed at all.     

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.     

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.