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.     

ON THE BIRDS OF THE SANDVELD KALAHARI OF SOUTH WEST AFRICA

Keffen, R. H. &; Jarvis, M. J. F. 1984. Some measurements relating to Ostrich eggs. Ostrich 55:182-187.

Eggs from wild Zimbabwean Ostriches Struthio camelus australis and domesticated South African hybrid birds, were compared. Formulae were tested to determine their accuracy when used to calculate egg weights, surface area and density. Calculated values were compared with some actual measurements. The main motivation for the study was the need to calculate fresh egg weights of wild Ostriches, as an aid to determining weight loss during incubation under natural conditions. Eggs from wild birds were, on average, smaller than those from domesticated birds, even though the wild adults were larger than domesticated birds. However, newly batched chicks from hybrid eggs weighed less than those from wild eggs.     

Relative reproductive success of female moorhens using conditional strategies of brood parasitism and parental care

In a population of moorhens (Gallinula chloropus), at least27% of netting females laid one or more eggs in a neighbor'snest Females laid parasitically under three conditions: 56%of parasitic eggs were from nesting females that preceded layinga dutch in their own nest by a parasitic laying bout, 19% werefrom females whose nests were depredated before clutch completionand that laid the following egg parasiticaDy, and 25% were froma small number of females without territories, "non-nesting"parasites, that each laid a series of parasitic eggs. Clutchsizes varied greatly between females, but nesting females eachlaid a consistent clutch size both within and between seasonsfor a given mate and territory. Nesting females that employeda dual strategy of brood parasitism and parental care producedextra eggs that they laid in the nests of neighbors before layinga dutch in their own nests. Two out of ten females whose dutchesI experimentally removed during the laying period were successfullyinduced to lay their next egg in the nest of a neighbor. Nestingfemales that laid parasitically selected their hosts opportunisticallyfrom among the nests dosest to their territories. An experimentin which parasitic eggs were removed and hosts left to rearonly their own young showed that parasites did not choose hoststhat were better parents than pairs with contemporary neststhat were not parasitized. Females that only laid parasiticaDywithin a given season timed their parasitic laying bouts poorlyand achieved no reproductive success. Parasitic young rarelyfledged, and the mean seasonal reproductive success of nestingbrood parasites did not differ from that of nonparasitic females.However, the variance in reproductive success of nesting broodparasites was significantly higher than that of nonparasiticfemales.     

4. CLIMPSES OF THE NARINA TROGON

Bertram, B. C. R. &; Burger, A. E. 1981. Aspects of incubation in Ostriches. Ostrich 52:36-43.

We studied incubation in domesticated Ostriches Struthio camelus in South Africa and wild Ostriches in Kenya. Although the eggs were large, with relatively high thermal capacities, unattended eggs exposed to the sun reached dangerously high temperatures (40,5°C). Experimental exposure of fresh eggs to the sun for seven days prior to incubation greatly reduced the percentage of embryos which developed, and no embryos survived 15 days of exposure. In the wild. Ostriches frequently shade their eggs in the pre-incubation period to prevent overheating.

During natural incubation, temperatures in the eggs (range 30,8-33,8°C) and of nest-air (31,9-34,6°C) were remarkably constant, despite the daily ambient fluctuations of air temperatures (17,8-38,9°C). Similarly the humidity of the nest-air (39–52%) was lower and less variable than the ambient air (39–72%). Water loss during 42 days of incubation was 11–12% of initial egg weight and, in addition, early laid eggs lost 3–4% during the 2½-3 week pre-incubation period. The water vapour conductivity and the daily water loss of Ostrich eggs were similar to those of other birds, in proportion to epg size, despite the arid environment inhabited by Ostriches. Some of the constraints on the feeding and breeding behaviour of Ostriches imposed by the physical requirements of their eggs are discussed.     

Experimental support for the role of nest predation in the evolution of brood parasitism

In 1965, Hamilton and Orians (HO) hypothesized that the starting point for the evolution of obligate interspecific brood parasitism in birds was the facultative laying of physiologically committed eggs in neighbouring active nests of con‐ and heterospecifics, following predation of a bird’s own nest during the laying stage. We tested this prediction of the HO hypothesis by using captive pairs of zebra finches (Taeniopygia guttata), a species with evidence for intraspecific parasitism both in the wild and in captivity. As predicted, in response to experimental nest removal, subjects laid eggs parasitically in simulated active conspecific nests above chance levels. Across subsequent trials, we detected both repeatability and directional change in laying patterns, with some subjects switching from parasitism to depositing eggs in the empty nest. Taken together, these results support the assumptions and predictions of the HO hypothesis, and indicate that the zebra finch is a potential model species for future behavioural and genetic studies in captive brood parasite research.     

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.     

Do female spotless starlings Sturnus unicolor adjust maternal investment according to male attractiveness?

In birds, female egg allocation patterns have a strong influence in offspring development and differential investment in egg size and composition has been shown to respond to male attractiveness. In this study we experimentally manipulated the perceived attractiveness of male starlings Sturnus unicolor by increasing the amount of green material in some nests (a male courtship display in this species). We predicted that, if female investment before laying is related to male attractiveness, experimental females would increase their reproductive investment in response to the addition of plants in their nests when compared to control females. We found that our manipulation caused variations in female reproductive investment in a way that seems to influence offspring quantity but not offspring quality: Females laid larger clutch sizes but not larger eggs when green plant material was added. However, yolk androgens contents were not related to the experimental manipulation. Contrary to expectations, females breeding in experimental nests laid eggs with smaller amounts of eggshell pigments. Interestingly, we found that eggs laid later in the sequence had higher testosterone levels and showed more intense egg colouration than eggs laid earlier in the sequence. These differences at the within-clutch level suggest that selection has favoured compensatory strategies for hatching asynchrony. Alternatively, since nest sabotages by other females are most common at the beginning of laying, this could be seen as female strategy to minimise losses due to nest sabotages. As far as we know, this is the first study to show that an external egg characteristic such as blue-green colouration reflects yolk androgen concentration.     

SOME PHYSICAL REQUIREMENTS FOR OSTRICH EGG INCUBATION

Jarvis, M. J. F., Keffen, R. H. & Jarvis, C. 1985. Some physical requirementsfor Ostrich egg incubation. Ostrich 56: 42–51.

Eggs from wild and domesticated Ostriches Struthio camelus australis were studied to determine temperature and humidity requirements for successful incubation. Wild eggs in Zimbabwe included some collected and incubated artificially, and others that were naturally incubated until just before hatching. Egg-weight losses during incubation were measured. The eggs artificially incubated at 35 °C to 36°C and 40–42% relative humidity, lost nearly the same weight as did eggs in naturally incubated nests. Comparisons were made between the wild eggs and chicks, and those from domesticated hybrid-strain Ostriches in Bophuthatswana. The discussion outlines some of the remaining problems associated with Ostrich egg incubation.     

Breeding biology of ostriches (Struthio camelus) in the Serengeti ecosystem, Tanzania

Ostrich breeding behaviour in the Serengeti ecosystem, Tanzania was investigated for differences in laying dates between low altitude western area (WA) and high altitude eastern area (EA) populations. Ostriches in WA laid eggs significantly earlier than in EA. The differences could be attributed to topography and rainfall pattern. Reliable rains in lower altitudes ensure availability of food that in turn influences the whole process of the reproductive cycle. Clutches were contributed by several females with a nest having up to 38 eggs. We also compared the frequency of observation of predators, ostriches, nests, 'singletons' (single eggs laid randomly) and broods between the two areas. There was no significant difference between WA and EA in 1) ostrich/nest ratio, indicating similar breeding densities; 2) ostrich/predator and predator/nest ratios, indicating that predation pressure was equally high; 3) nest/singleton and predator/singleton ratios, indicating that loss of nests did not vary between areas. However, there were significantly more predators, nests and ostriches compared to broods in EA than in WA, indicating a significantly lower reproductive success in EA. Using metapopulation terminology, ostriches in EA could be regarded as a 'sink' population and those in WA as a 'source' population, but investigations over longer time-periods are needed to further resolve if this is the case.     

Colony kin structure and host‐parasite relatedness in the barnacle goose

Conspecific brood parasitism (CBP), females laying eggs in the nest of other ‘host’ females of the same species, is a common alternative reproductive tactic among birds. For hosts there are likely costs of incubating and rearing foreign offspring, but costs may be low in species with precocial chicks such as waterfowl, among which CBP is common. Waterfowl show strong female natal philopatry, and spatial relatedness among females may influence the evolution of CBP. Here we investigate fine‐scale kin structure in a Baltic colony of barnacle geese, Branta leucopsis, estimating female spatial relatedness using protein fingerprints of egg albumen, and testing the performance of this estimator in known mother‐daughter pairs. Relatedness was significantly higher between neighbour females (nesting ≤ 40 metres from each other) than between females nesting farther apart, but there was no further distance trend in relatedness. This pattern may be explained by earlier observations of females nesting close to their mother or brood sisters, even when far from the birth nest. Hosts and parasites were on average not more closely related than neighbour females. In 25 of 35 sampled parasitized nests, parasitic eggs were laid after the host female finished laying, too late to develop and hatch. Timely parasites, laying eggs in the host’s laying sequence, had similar relatedness to hosts as that between neighbours. Females laying late parasitic eggs tended to be less related to the host, but not significantly so. Our results suggest that CBP in barnacle geese might represent different tactical life‐history responses.     

Brood-parasitism by the Shining Cuckoo Chrysococcyx lucidus at Kaikoura, New Zealand

For three seasons starting in 1976 I studied the breeding of Shining Cuckoos Chrysococcyx lucidus in forest near Kaikoura, New Zealand. There is no evidence that the cuckoo parasitizes any host on mainland New Zealand other than the Grey Warbler Gerygone igata. A nestling cuckoo returned to within 1 km of its natal site in a subsequent breeding season, presumably after migrating beyond New Zealand. Empirical and theoretical estimates of the area occupied by Shining Cuckoos while breeding are given. Cuckoos near Kaikoura laid during ten weeks, the modal week of laying following seven weeks after the presumed peak of arrival of birds in New Zealand. First clutches of the host escaped parasitism because they were laid before most cuckoos arrived. Parasitized clutches received one cuckoo egg which replaced a host's egg. It was laid before, just after or long after the host began incubating, and mimicry was lacking. Cuckoo eggs, which were about 8% of the adult cuckoo's weight, hatched in 14–17 days. The frequency of parasitism near Kaikoura was 55% of late clutches (n = 40).
At 3–7 days old, nestling Shining Cuckoos evicted from the nest all other contents. The nestling period was at least 19 days. Growth in weight followed a logistic curve and the equation is given. Just over half the cuckoo eggs produced fledglings. The effect of brood-parasitism on the Grey Warbler's productivity was small. Only 17% of late warbler eggs, and late eggs only, were prevented by parasitism from yielding fledglings. Late laying by some Shining Cuckoos (relative to the host's incubational cycle), and late eviction, often led to brief inter-specific competition among nestlings for food. The brief coexistence of young Warblers and Cuckoos in the nest may explain the apparent mimicry by newly-hatched Shining Cuckoos of the host's young.     

Nest destruction elicits indiscriminate con‐ versus heterospecific brood parasitism in a captive bird

Following nest destruction, the laying of physiologically committed eggs (eggs that are ovulated, yolked, and making their way through the oviduct) in the nests of other birds is considered a viable pathway for the evolution of obligate interspecific brood parasitism. While intraspecific brood parasitism in response to nest predation has been experimentally demonstrated, this pathway has yet to be evaluated in an interspecific context. We studied patterns of egg laying following experimental nest destruction in captive zebra finches, Taeniopygia guttata, a frequent intraspecific brood parasite. We found that zebra finches laid physiologically committed eggs indiscriminately between nests containing conspecific eggs and nests containing heterospecific eggs (of Bengalese finches, Lonchura striata vars. domestica), despite the con‐ and heterospecific eggs differing in both size and coloration. This is the first experimental evidence that nest destruction may provide a pathway for the evolution of interspecific brood parasitism in birds.     

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.     

ON MULTIPLE BROODS AND THE BREEDING STRATEGY OF ARCTIC SANDERLINGS

Sanderlings on Bathurst Island in the Canadian arctic have two patterns of incubation. At some nests the eggs are covered soon after the fourth egg has been laid and at others incubation is delayed for 5–6 days. Because the delay is about the same time required to lay a second clutch and because a single individual alone incubates at any one nest, we suspected that Sanderlings might normally lay two clutches in a season, the male caring for one brood and the female for the other.
Histological and gross examination of the ovaries of two females taken as the birds began incubation showed eight freshly ovulated follicles in each female. The size gradation and histological appearance of the follicles indicated that two clutches of four eggs each had been laid within 8–10 days by a single female. The ovary of one female had additional large yolky follicles, suggesting a physiological capability of further ovulations.
Field conditions in the arctic summer are highly variable, and the small eggs and the rapid sequence of broods of Sanderlings may be breeding adaptations that permit them to multiply the traditional wader clutch of four eggs by 2 or 3 in favourable years. Selection for mating systems characterised by brief pair bonds and by polyandry is expected in precocial birds where some broods are incubated and cared for by the male, but further field work is required to determine the precise mating system of Sanderlings.     

The effect of food on laying date and clutch-size in Tengmalm's Owl Aegolius funereus

The breeding of Tengmalm's Owl Aegolius funereus was studied at Umeå, Sweden, during the 1984–85. Mean clutch-size was one egg larger in 1984 than in 1985 despite the later laying in 1984. The difference in clutch-size was related to a better food supply in 1984. Daily weight increase of females during the prelaying period showed a high negative correlation with laying date in 1985, and a high positive correlation with clutch-size independently of laying date in 1984–85. This suggested that food eaten before and during laying had a great and direct influence on both laying date and clutch-size. Many females increased in weight during laying and most others decreased only moderately (relative to egg weight), suggesting that body reserves were not a main source for egg production.
Late breeding females were provided with extra food during the prelaying and laying periods in 1985. Fed females weighed more, bred eight days earlier and laid one more egg than controls. At the same laying dates in mid season, and after heavy snow-fall, clutch-size and female weight were larger in the fed birds than in controls, but this was not so near the end of the laying season. Although the earliest of the fed late breeders weighed more, and probably were less restricted by food availability just before or during laying, they did not lay more eggs than did early breeders. This result suggested some limitation on clutch-size that could not be overcome by the supplementary feeding. Weights of females during laying did not show any consistent relationship with clutch-size during successive laying date intervals, suggesting that clutch-size was not directly related to body condition.     

Egg characteristics are unreliable in determining maternity in communal clutches of guira cuckoos Guira guira

Egg characteristics have been used to determine egg maternity for birds in situations where two or more females lay eggs in a single nest (as in communal breeders and intraspecific parasites). We assessed the applicability of egg morphometry and eggshell appearance in ascribing egg ownership in communal clutches of guira cuckoos Guira guira , a species where up to seven females may lay eggs in a joint nest. We used both combined variables (including egg mass, length, width, shape and two eggshell variables), and a shape index to test whether eggs laid by each female were similar but different from eggs laid by other females. Also, we conducted discriminant function analyses to verify if eggs could be correctly classified to their mothers based on their characteristics. The correct maternity was determined by yolk protein electrophoresis of freshly-laid eggs. Individual female chutches were separated through egg characteristics or shape alone in 29% and 41% of the groups tested, respectively. Differences were mostly due to a single female that differed from her nest-mates in a unique egg variable. On average, 55% of the eggs analyzed were not assigned to the correct mother using egg dimensions and eggshell speckling pattern. In conclusion, the egg characteristics used do not reliably indicate maternity in guira cuckoo communal clutches. We therefore recommend a protein-based verification of egg appearance characteristics for assigning maternity in other species in which multiple female laying occurs.     

Annual variation in breeding biology of gentoo penguins, Pygoscelis papua, at Bird Island, South Georgia

The breeding biology of the gentoo penguin, Pygoscelis papua , was studied over a three-year period (1986–1988) at Bird Island, South Georgia, with particular reference to birds of known age or breeding experience. Laying date varied significantly between all three years, being three weeks later in 1987, when the breeding population decreased markedly. Factors involved in the timing of breeding are discussed. Within years egg-laying was highly synchronous: 95% of clutches were initiated in 14·5 days or less. The incubation period was 35 days and the laying interval, between the two eggs, 3·3–3·4 days. Chicks creched when 25–30 days old, and this varied between years, possibly related to food supply and chick growth. Chicks left the colony for the first time between 75 and 85 days of age. The breeding population at Bird Island decreased by 20% and increased by 84% in successive years during the study period. Breeding success (chicks fledged per egg laid) varied between 0·33 and 0·65 within colonies, but for the whole island was very consistent over the three years: 0·45, 0·51 and 0·47. Overall, colony differences were not correlated between years. Disturbance from Antarctic fur seals, Arctocephalus gazella , is suggested as the cause of consistently lower breeding success at one colony. Mean egg weight varied annually, and with age of the breeding bird, nest location and, in one year, with laying date. Young, first-time breeders laid smaller eggs and had lower breeding success compared to older, experienced birds, similar to other seabirds. However, they differed from other species in laying on average earlier than older birds. The relationship between age, egg weight, laying date and breeding success is discussed in relation to predation and seasonal food supply.     

Reproductive behavior and spawning success of female Amblyglyphidodon leucogaster (Pisces: Pomacentridae) from the Red Sea

The reproductive behavior of female whitebelly damselfish, Amblyglyphidodon leucogaster, was investigated in the Gulf of Aqaba, Red Sea over two breeding seasons. Females were promiscuous, mating with 7–10 different males throughout the season. Females lay eggs in distinct batches, defined as the total number of eggs laid in a day. Generally females deposit a batch of eggs with one male (87.2%) and are capable of laying a new batch every other day. Egg batch size averaged 4009 eggs and females laid from 2 to 22 egg batches per season. The variation in spawning success was not correlated to body size. Females preferred to deposit eggs in nests that already contained early stage eggs (0–2 days old). Within a nest, females chose to lay eggs contiguous to the youngest egg batch, regardless if the nest contained either a single batch or multiple batches of different ages. Female within-nest spawning patterns appear to be a consequence of between nest preferences for nests with young eggs. It is proposed that the strong within-nest preference is a consequence of mate selection where females may use new egg batches as a visual cue as part of a copying style. Such a style may reduce the risk of predation and increase feeding opportunities, because less time is expended in mate selection, which would provide additional resources for egg production and ultimately increase female spawning success over the breeding season. This revised version was published online in July 2006 with corrections to the Cover Date.     

Breeding orange-winged Amazon parrots in captivity

Captive propagation of parrots for the companion bird trade provides a potential means of reducing the economic incentive to capture these birds from the wild. To test whether environmental and social manipulations might stimulate reproduction in captive, wild-caught orange-winged Amazon parrots (Amazona amazonica), we compared reproductive performance in pairs presented with nest boxes (controls) or nest boxes plus additional environmental and social manipulations (enriched group) consisting of misting, fruit supplementation, nest hole restriction, enlarged nest boxes, and pair separation/reunification. In the first breeding trial, in 1990, enriched pairs were more likely to lay eggs (six of seven enriched pairs vs. one of eight control pairs; P < 0.09). When treatments were reversed the following years (e.g., 1990 controls were exposed to enriched conditions in 1991), reproductive performance was not different between the groups (four of seven enriched pairs laid eggs vs. five of seven control pairs), although three pairs in the 1991 enriched group laid eggs which had not laid eggs before. These results are consistent with the idea that while enriched environments may enhance the probability of a first episode of egg laying in captivity, once pairs have laid eggs in captivity, little stimulation beyond that provided by nest box presentation is required to reinitiate egg laying. The environmental and social manipulations of the enriched conditions could be useful in increasing reproductive performance of captive Amazon parrots for the companion bird trade. © 1995 Wiley-Liss, Inc.     

Leafing phenology and timing of egg laying in great tits Parus major and blue tits P. caeruleus

To evaluate the importance of tree leafing for the start of laying and clutch size of birds, we compared the breeding phenology of great tits Parus major and blue tits P. caeruleus between one coastal and two inland sites in the same geographical region. Because of the cooling influence of the sea, trees at the coastal site were known to initiate budburst about a week later than at the inland sites. During 5 years, breeding by the tits and the leaf phenology of birch Betula pendula , and oak Quercus robur were monitored. The leaf phenology of birch and oak explained a significant part of the between-year variation in the start of egg laying in blue and great tits, respectively. The tits started laying earlier at the sites with an early budburst, i.e. normally inland. However, leaf phenology was not an absolute cue to the start of laying, since blue tits laid earlier relative to leafing at the inland site than at the coastal site, and both tit species laid eggs earlier relative to leafing during late springs. In neither species was clutch size affected by leafing phenology. However, great tit females at the coastal site consistently produced fewer eggs than did those at the inland site. No such difference was found in the blue tits. Although leafing phenology may predict the start of laying in tits, other factors also influence its timing. These factors might include other cues, or differing life-history trade-offs depending on site or general climatic factors during the spring.