THE EVOLUTION OF CLUTCH SIZE. I. AN EQUATION FOR PREDICTING CLUTCH SIZE

We derive an equation for calculating the clutch sizes of birds and other long-lived animals from Murray's (1979) theory on the evolution of clutch size. For the Prairie Warbler (Dendroica discolor) in Indiana, this equation predicts an average clutch size of 3.49, less than half an egg smaller than the recorded average clutch size of 3.89. We attribute the discrepancy to sampling error and suggest that the equation satisfactorily identifies the important factors affecting the evolution of clutch size. The success of the equation in predicting clutch size of the Prairie Warbler provides additional support for Murray's theory on the evolution of clutch size.     

6. THE PROBLEM OF NOMENCLATURE

Hockey, P. A. R. 1983. Aspects of the breeding biology of the African Black Oystercatcher. Ostrich 54:26-35. Fifty-five pairs of African Black Oystercatchers Haematopus moquini bred at Marcus Island in 1979–1980. Sixteen pairs laid replacement clutches: the mean interval between loss of the fist clutch and laying of a replacement clutch was 22,2 days. Mean inter-nest distance was 19,4 m. The modal clutch size was 2, with a mean of 1.74. Mean dimensions of 105 eggs were 60,7 × 40,l mm and mean fresh egg mass was 55,s g. There were differences in egg mass and dimensions between eggs in one- and two-egg clutches. Rate of egg loss was high, due mainly to depredation by Kelp Gulls Larus dominicanus promoted by human disturbance. Fledging success was lower at a disturbed site than at undisturbed sites, with highest chick mortality occurring in the first week of life. Mortality of first-year birds of eight days and older was estimated at 48% and 69% in two successive seasons. All juvenile birds dispersed from the natal sites, and were resighted up to 168 km away. Dispersed juveniles were concentrated at the edge of an area of high oystercatcher density.     

SOME ASPECTS OF THE BEHAVIOUR OF THE WATTLED PLOVER AFRIBYX SENEGALLUS (LINNAEUS)

The question of how aridity might influence avian clutch size, through the influences of rainfall seasonality and environmental stochasticity (unpredictability), has received little attention. A marked east-west gradient in aridity across South Africa provides a unique opportunity to test for such influences. Using an extensive collection of nest records for 106 terrestrial bird species from the South African Nest Record Card Scheme, we tested three predictions related to rainfall seasonality and stochasticity. Analyses were conducted at two levels, the first examining each species independently, and the second grouping species into five dietary guilds. The first prediction, that clutch size should generally increase with higher rainfall seasonality (i.e. higher seasonal fluctuation of food availability), was supported, particularly in the most arid environments where food abundance is more closely linked to rainfall. Controlling for rainfall seasonality, the second prediction, that clutch size should generally decrease as a bet-hedging strategy in arid, stochastic environments, was also supported. Although the timing of the rainy season differs among regions in South Africa (winter, early summer, later summer, year-round), birds primarily nest during spring. The relative timing of rainfall and breeding is expected to have different consequences for seasonal variation in clutch size among rainfall regions. The third prediction, of different patterns of seasonal variation in clutch size between rainfall regions, was also supported. In the winter and early-summer rainfall regions, early-nesting birds (breeding with or soon after the rains) generally had a larger clutch size than late-nesting birds. In the late-summer rainfall region, early-nesting birds (breeding well before the rains) had a smaller clutch size than late-nesting birds.     

THE POPULATION,ECOLOGY AND CONSERVATION OF THE BLACK OYSTERCATCHER HAEMATOPUS MOQUINI

Summers, R. W. & Cooper, J. 1977. The population, ecology and conservation of the Black Oystercatcher Haematopus moquini. Ostrich 48:28-40.

The population of Black Oystercatchers Haematopus moquini in the southwestern Cape, South Africa, was estimated to be 2 942 birds. Birds occurred most abundantly on coastal islands and were also abundant on mixed (sandy and rocky) shores of the mainland. Sandy shores and coastal wetlands supported few birds. Black Oystercatchers bred mainly from December to February with the number of clutches present reaching a peak in the first half of January. The most frequent clutch size was two eggs, the mean clutch size was 1,81. No significant differences were found in either linear dimensions or mass between the first and second eggs. The mean proportion of juveniles in groups of birds in July was 3,6 % suggesting a low recruitment to the adult population. The breeding population at Marcus Island is apparently sedentary throughout the year. The primary moult season for adults extends from March to October (eight months). Introduced mammalian predators should be controlled on islands and important mainland breeding sites should be protected by the creation of nature reserves and restricting human access during the breeding season.     

BREEDING BIOLOGY OF THE EDIBLE-NEST SWIFTLET AERODRAMUS FUCIPHAGUS

A colony of the Edible-nest Swiftlet Aerodramus fuciphagus nesting in a Chinese shophouse in Penang, Peninsular Malaysia, was studied for seven months. Birds bred throughout, but laying was concentrated in the period October to February. Incubation and fledging periods were 23 ± 3 days and 43 + 6 days, respectively. Most young hatched during the dry season. The normal clutch size was two, with many birds laying second clutches (75%) and some third clutches (15-4%) using the same nest. Clutch size showed no variation with clutch order or month. The overall hatching success was 69-0 %, most losses caused by eggs falling from or with the nest. Fledging success was similar for broods of two (625%) and broods of one (59-4%), so that the former were twice as productive. The main loss of chicks occurred when they fell from the nest and were eaten by mammalian predators. There was a tendency for breeding success to decline with successive clutches, but not with month, being highest in November (53-6 %) and February (55-6 %). The growth rate of single chicks and successful broods of two was similar, except that the second chick was more prone to fluctuations in weight. Some second chicks showed evidence of starvation before falling from the nest. Diet was examined by analysis of foodballs regurgitated by mist-netted adults. Foodballs weighed 0–13-1 08 g and, on average, contained over 500 prey items. The main arthropods caught were Hymenoptera (40-8%), Ephemeroptera (26-4%), Homoptera (15-4%) and Diptera (7-7%). Flying ants and mayflies contributed most by weight, although figwasps and mayflies were the most numerous prey items. Only in December did the percentage of moulting mist-netted adult swiftlets fall below 70 %. It appeared that many birds were moulting and breeding simultaneously, although 21% of birds had primaries moulting in two places, suggesting arrested moult. Diurnal activity showed a normal high dawn exodus and dusk inflow of birds, although there was evidence of an increase of birds flying out prior to the dusk inflow. Nest harvesting was continued throughout the study, but most of the marked nests under study were left undisturbed.     

MATERNAL INHERITANCE OF CONDITION AND CLUTCH SIZE IN THE COLLARED FLYCATCHER

Maternal effects may strongly influence evolutionary response to natural selection but they have been little studied in the wild. We use a novel combination of experimental and statistical methods to estimate maternal effects on condition and clutch size in the collared flycatcher, where we define “condition” to be the nongenetic component of clutch size. We found evidence of two maternal effects. The first (m) was the negative effect of mother's clutch size on daughter's condition, when mother's condition was held constant. The second (M) was the positive effect of mother's condition on daughter's condition, when mother clutch size was held constant. These two effects oppose one another because mothers in good condition also lay many eggs. The maternal effects were large: Experimentally adding an egg to a mother's nest reduced clutch sizes of her daughters by 1/4 egg (i.e., m = -0.25). Measured degree of resemblance between mother and daughter clutch sizes yielded M = 0.43. The results weakly support the presence of heritable genetic variation in clutch size: additive genetic variance/total phenotypic variance = 0.33. This estimate was highly variable probably because, as we show, mother-daughter resemblance may depend hardly at all on the amount of genetic variance when maternal effects are present. Daughter-mother regression (a standard method for estimating heritability) is consequently a poor guide to the amount of genetic variance in clutch size. Our results emphasize the value of combining field experiments with observations for studying inheritance.     

THE INFLUENCE OF THE ENVIRONMENT ON BREEDING SUCCESS OF A SUBURBAN POPULATION OF CRESTED BARBETS TRACHYPHONUS VAILLANTII

Van Zyl, A.J. 1994. The influence of the environment on the breeding success of a suburban population of Crested Barbets Trachyphonus vaillantii. Ostrich 65: 291–296.

I studied the breeding biology of the Crested Barbet Trachyphonus vaillantii in Colbyn, a suburb east of Pretoria, South Africa, for nine breeding seasons from 1981 to 1989 to examine patterns in annual breeding success, breeding attempt success in multiple broods, and rainfall. The modal incubation period was 14 days and the nestling period ranged from 28 to 31 days. Average clutch size for all the years was 3,3 eggs/clutch and there was no significant difference in clutch size or number of young fledged/nest between years. On average, Crested Barbet pairs made 2,4 breeding attempts/season. There was no difference in clutch size or breeding success between the breeding attempts. Crested Barbets nesting in natural nests laid on average larger clutches than those in artificial nestboxes, but had non-signficantly lower breeding success. Failure to raise Crested Barbet chicks was attributed to parasitism by Lesser Honeyguides Indicator minor, bee swarms occupying nestboxes, and flooding of natural nests. Breeding performance was not correlated with rainfall or adult body size. The suburban environment may be less variable than a natural environment, resulting in a stable breeding Crested Barbet population.     

PARASITISM OF THE MASKED WEAVER PLOCEUS VELATUS ARUNDINACEUS

Williams, G. D., Coppinger, M. P. &; Maclean, G. L. 1989. Distribution and breeding of the Rock Pratincole on the Upper and Middle Zambezi river. Ostrich 60: 55–64.

The Rock Pratincole Glareola nuchalis was studied on the Upper and Middle Zambezi fromMayto November 1986 and from April to December 1987. Altogether 1938 Rock Pratincoles were counted; their distribution was closely associated with the availability of breeding habitat. The first birds were seen in mid-August and the species was still present in December. They nested in loose colonies on midstream rocks separated from the shore by flowing water. Sixty-four clutches (mean clutch size 1,8 eggs) were found and 44 eggs measured (mean size 29,2 x 21,5 mm). The eggs were usually laid on bare rock less than 1 m above the surface of the water and were sometimes in shade. Both sexes incubated and the frequency of nest relief was greatest during the hottest times of the day. The responses of incubating birds to overheating were bill-opening, panting, dorsal-feather erection, holding the wrists away from the body, and belly-soaking. Non-incubating birds drank often and did not seek shade. After hatching the young sought refuge in rock overhangs and crevices. The important breeding sites in the study area were identified and possible effects of future hydro-electric schemes are discussed.     

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.     

The phenology and clutch size of UK Blue Tits does not differ with woodland composition

The deciduous tree-herbivorous caterpillar-insectivorous bird food chain is a well-studied system for investigating the impacts of climate change across trophic levels. To date, across Europe, most attention has focused on the impacts of increasing spring temperature on changes to phenology in Oak-dominated (Quercus spp.) woodlands. Paridae species and Pied Flycatcher Ficedula hypoleuca are the most studied secondary consumers, all of which demonstrate an advancement in reproductive phenology with increases in spring temperature. Shifts in climate and phenology may also impact on reproductive investment in clutch size, and the effects of climate on phenology and clutch size may vary depending on woodland composition. To date, the effects of among-habitat variation in phenology and reproductive investment have received little attention. Insectivorous birds inhabiting woodlands that differ in tree composition may differ in the timing of breeding, due to local tree leafing phenology acting as a cue for egg-laying date and/or clutch size. Moreover, for most insectivorous birds, woodland composition within a territory is likely to be the main determinant of food availability for both adults and chicks. Consequently, if warming springs affect the temporal patterns of food availability differently across different woodland compositions, this may affect the optimal average local phenology for nesting birds. Here, using data from 34 long-term (mean 15 years) nest monitoring sites across the UK, we investigate the effect of woodland tree composition and temperature on Blue Tit Cyanistes caeruleus first egg date (FED) and clutch size. We supplemented the nest monitoring data by quantifying woodland composition, at a site level, through modified point counts. We predict that birds breeding in woodlands with greater proportions of late-leafing species, such as Oak and Ash Fraxinus excelsior, will breed later than those breeding in woodlands with greater proportions of early-leafing species, such as Birch Betula spp. and Beech Fagus sylvatica. We found no evidence for differences in Blue Tit FED or clutch size in relation to the proportion of any of the tree species investigated, after controlling for temperature and latitude (FED: −3.4 and 2.2, clutch size: −0.4 and − 0.2 eggs for one-unit increase in temperature and latitude, respectively). In recent decades and across all sites, clutch size has decreased as spring temperatures have increased, a strategy which could allow birds flexibly to adjust their breeding phenology such that nestling demand coincides with peak food availability. The lack of an effect of woodland composition on Blue Tit phenology suggests Blue Tits do not fine-tune their reproductive phenology to the local tree composition. Whether this lack of evidence for phenological divergence is due to an absence of divergent selection on breeding phenology and clutch size or to gene flow is not clear.     

ASPECTS OF THE BREEDING BIOLOGY OF THE FIERYNECKED NIGHTJAR

Jackson H. D. 1985. Aspects of the breeding biology of the Fierynecked Nightjar. Ostrich 56: 263–276.

A marked population of nightjars in Zimbabwe was studied intensively for four breeding seasons. This paper covers certain aspects of the breeding biology of the Fierynecked Nightjar Caprimulgus pectoralis. The male shows strong site fidelity during the breeding season (September to December), singing, feeding and breeding within an area of about 5,8 ha. There is some evidence of site defence by the male. The female shows strong mate fidelity, resulting in a pair bond for life. Egg laying starts with full moon in September and is further stimulated by the next two full moon periods. The eggs are laid directly on dense leaf litter at a site overhung by foliage. The normal clutch is two eggs (12S % are one egg) laid on successive days during the afternoon. Incubation starts with the first egg and is by the male at night and the female by day. The incubation period is 18 days. The birds respond to undue disturbance by deserting the eggs and laying a replacement clutch. The chicks usually hatch on successive afternoons; they are mobile on the first day and will move to a parent if called. Both parents feed and brood the young during twilight and moonlight; the male broods them on dark nights and the female does so by day. The species is double-brooded when time permits, the female laying again once the first brood has reached independence; she may even lay a third clutch if the second one comes to grief. There is no evidence of adults transporting eggs or young.     

REPRODUCTIVE COST,AGE-SPECIFIC SURVIVAL AND A COMPARISON OF THE REPRODUCTIVE STRATEGY IN TWO EUROPEAN TITS (GENUS PARUS)

Theoretical analyses of optimal reproductive rates usually assume a trade-off between offspring production and parental survival. This study verified a survival cost for willow tit males; nonbreeding males survived better than males attending a brood. Theory also predicts a smaller clutch size in birds that are less successful in transforming reproductive investments into mature offspring. As predicted, we found that crested tits, suffering a higher nest predation rate, laid smaller clutches than willow tits. The generally lower survival rate of willow tit adults may largely be attributed to their higher reproductive commitment (larger willow tit clutch size), because no significant interspecific survival difference remained between nonbreeding males. Finally, in willow tits we found a positive correlation between average clutch size and juvenile survival rate (density-dependent) the ensuing year, suggesting that willow tits may adjust clutch size in response to changing survival prospects for their young by using the breeding density as a cue.     

Malarial parasites decrease reproductive success: an experimental study in a passerine bird

Malarial parasites are supposed to have strong negative fitness consequences for their hosts, but relatively little evidence supports this claim due to the difficulty of experimentally testing this. We experimentally reduced levels of infection with the blood parasite Haemoproteus prognei in its host the house martin Delichon urbica, by randomly treating adults with primaquine or a control treatment. Treated birds had significantly fewer parasites than controls. The primaquine treatment increased clutch size by 18%; hatching was 39% higher and fledging 42% higher. There were no effects of treatment on quality of offspring, measured in terms of tarsus length, body mass, haematocrit or T-cell-mediated immune response. These findings demonstrate that malarial parasites can have dramatic effects on clutch size and other demographic variables, potentially influencing the evolution of clutch size, but also the population dynamics of heavily infected populations of birds.     

THE SOCIABLE WEAVER,PART 3: BREEDING BIOLOGY AND MOULT

Maclean, G. L. 1973. The Sociable Weaver, Part 3: Breeding biology and moult. Ostrich 44: 219–240.

Rain or some associated phenomenon is the principal Zeitgeber releasing breeding in the Sociable Weaver. The species does not breed in the absence of rain. The same nest chambers are used for breeding as are used for roosting throughout the year. The Sociable Weaver is monogamous. The clutch size varies from two to six eggs, larger clutches being more common after good rains than in relatively poorer breeding periods. Food supply may therefore be the proximate factor regulating clutch size. Replacement clutches are not necessarily smaller than first clutches. The mean clutch size within a breeding period decreases with an apparent decrease in food supply. The parents share parental duties about equally. Up to four successive broods may be raised in a single breeding period; a breeding period may last up to nine months and may occur at any time of the year according to the somewhat erratic rainfall which averages about 226 mm per year in the study area.

First broods help their parents to feed later broods; fourth brood chicks may therefore be fed by as many as 11 birds (nine young and two parents). This has survival value especially toward the end of a breeding period when food is scarce. Of similar value is the habit of starting incubation with the first or second egg of the clutch; in a relatively poor season older chicks will survive while younger ones will starve, thereby effectively and quickly reducing brood size. Young birds moult into adult plumage at four months, but do not normally leave the home colony. The sexes are indistinguishable at all ages, but there is an approximate ratio of eight males to five females in the study area.

Wing moult is slow: each remex takes about a month for replacement. Body moult occurs within the space of a month, usually after rain while the birds are breeding. Primary remiges are moulted proximo-distally from 1 to 9; secondaries are moulted disto-proximally from 1 to 6. Body moult is antero-posterior with the dorsal surface slightly in advance of the ventral surface.     

Elevational gradient in clutch size of Red‐faced Warblers

Our understanding of life history evolution has benefited from debates regarding the underlying causes, and geographic ubiquity, of spatial patterns in avian clutch sizes. Past studies have revealed that birds lay smaller clutch sizes at higher elevation. However, in most previous studies, investigators have failed to adequately control for elevational differences in breeding phenology. To better understand the elevational gradient in avian clutch size, we need to know how clutch size changes across the entire elevational breeding range of a species (i.e., the shape of the relationship between elevation and clutch size), and whether the elevational gradient in clutch size is merely an artifact of elevational gradients in breeding phenology or breeding season length. We examined the relationship between breeding elevation and clutch size of Red‐faced Warblers (Cardellina rubrifrons) along a 1000‐m elevational gradient in Arizona. Our objectives were to determine how clutch size changed with elevation, and if the relationship between clutch size and elevation merely reflected elevational changes in breeding season length or phenology. The proportion of 5‐egg clutches decreased and the proportion of 3‐ and 4‐egg clutches increased non‐linearly with increasing elevation, even after controlling for the elevational gradient in nest initiation date. Thus, average clutch size declined across the elevational breeding range of Red‐faced Warblers, but this decline was not due to elevational variation in breeding phenology. Timing of breeding changed, but the duration of the breeding season did not change appreciably across the elevational gradient. Hence, elevational differences in breeding season length or breeding phenology cannot explain why Red‐faced Warblers (and perhaps other birds) breeding at higher elevations have smaller clutches.     

SHORT NOTES

Earlé, R. A. 1981. Factors governing avian breeding in Acacia savanna, Pietermaritzburg, Part 3: Breeding success, recruitment and clutch size. Ostrich 52:235-243.

The clutch size and breeding success of eight species of birds were monitored over a two year period in Acacia savanna. The mean clutch size fluctuated within the breeding season and four patterns of clutch size variation were noted. A smaller proportion of large clutches were laid in the 1979 season when less food was available. Breeding success was higher in the dry 1979 season but fewer breeding attempts were made and the overall recruitment was thus lower. At the peak of the breeding season, in mid-November, breeding success was at its lowest. Predation on nests was the greatest factor reducing breeding success although rain and strong winds affected some species.     

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.     

Can birds count eggs in their nests?

Sensory mechanisms controlling avian clutch size have diversified into distinct types, according to the nature of the input that is used to disrupt the growth of ovarian follicles and hence halt egg‐laying. In an article on brood parasitism, Lyon (2003) claimed that female American Coots Fulica americana can reduce their clutch size on the basis of visual cues in response to eggs laid in their nests by other females; in this species, therefore, egg counting would be used to control clutch size. After a close examination of the physiological determination of clutch size in American Coots, I show that seven of 17 parasitized clutches were smaller than the range controlled through the mechanism using an input to disrupt follicular growth (7–10 eggs per clutch). My reanalysis suggests that American Coots are incapable of adjusting clutch size via counting and re‐asserts that a species that can count eggs has yet to be found among birds that rely upon their own body heat for incubation.     

Testing the relationship between clutch size and brood size in the Coot (Fulica atra)

The time between egg laying and chick fledging is of crucial importance for the survival of young birds. I analyzed breeding output at consecutive phases of growth of young Coots (Fulica atra) relative to the clutch size and laying date. Considering the specific breeding biology of the Coot, I tested whether chick survival reveals clutch size-dependent variability. Clutch size did not affect hatching success; it only affected brood size, and that merely temporarily. During the first 20 days after hatching, i.e. during the time of the highest chick mortality, birds with larger clutches lost chicks at a higher rate. As a result, the number of fledged chicks was independent of the initial number of chicks, and pairs with different clutch sizes had a similar number of fledglings. The laying date had no effect. This pattern of age-related chick survival points to the greater role of the type of chick growth (semi-precocial) and behavior in their survival.     

BREEDING SEASONS OF THE HOUSE SPARROW AND TREE SPARROW PASSER SPP. AT OXFORD

The breeding season of P. domesticus in 1961 and from 1963 to 1964. at Oxford began in April and ended in early August; that of P. montunus began slightly later and ended slightly earlier and this species also laid fewer clutches. In P. domesticus laying birds fell into two groups, namely "early-" and "late-starters", which, from a comparison with data obtained from birds of known age, appeared to be mostly females two or more years old, and females one year old, respectively. In P. montanus , however, it appeared that birds of all ages were beginning breeding at about the same time in the season. The laying of first clutches of the early-starting birds of P. domesticus and of the earlier starting members of the one group of P. montanus was correlated with the prevailing air temperature; it is estimated that P. domesticus began its laying at slightly lower air temperatures than P. montunus . It is suggested that laying is adapted to begin at a particular air temperature because at lower temperatures the adult would have insufficient energy available to produce and incubate a clutch.