Great tits provided with ad libitum food lay larger eggs when exposed to colder temperatures

The amount of nutrients deposited into a bird egg varies both between and within clutches of the same female. Larger eggs enhance offspring traits, but as a tradeoff, laying large eggs also infers energetic costs to the female. Income breeders usually lay larger eggs later in the season, when temperatures and food availability are higher. Egg size is thus affected by the daily amount of energy available to produce an egg under cold conditions, but it is less well known in how far temperature exerts direct effects on egg size. We show that great tit females Parus major with access to ad libitum food and breeding in climate‐controlled aviaries varied their egg investments. The size of an individual egg was best predicted by mean temperatures one week pre‐laying, with females laying larger, rather than smaller, eggs under colder conditions. Eggs increased in size over the season, but not significantly over the laying sequence. The degree of daily temperature fluctuation did not influence egg size. In addition to a substantial between‐female variation, sisters were more similar to each other than unrelated females, showing that egg size does also reflect heritable intrinsic female properties. Natural variation in egg size is thus not only determined by energy‐limitation, but also due to females allocating more resources to eggs laid in colder environments, thus increasing early survival of the chicks. That the positive correlation between temperature and egg investments that is found in a natural population is reversed under ad libitum food conditions demonstrates that wild great tits tradeoff own condition with survival prospects of their chicks as a function of available food, not ambient temperature.     

BREEDING OF THE ADÉLIE PENGUIN PYGOSCELIS ADELIAE AT CAPE BIRD

Observations were made during four seasons (1967–68, 1968–69, 1969–70, 1970–71) on the breeding of Adelie Penguins at Cape Bird, Ross Island, Antarctica. Breeding data from individuals were related to date of return, laying date, clutch-size, nest location, and change of mate. Some females consistently laid near the mean date of laying, while others were consistently early or late layers. Laying date may be under direct genetic control, or may reflect feeding ability. The mean clutch-size was smaller in peripheral compared to central nests, and smallest of all in isolated nests. Clutches laid late in the season were smaller than those laid near the peak date, and small, late clutches were laid in peripheral rather than central nests. These differences may reflect age and/or feeding ability. Penguins that are better able to find food will return earlier, obtain central sites, and have larger clutches than those with a lesser ability. The two main causes of egg and chick losses were predation and parental failure. Losses were highest in single-egg clutches, at isolated and peripheral nests, and among eggs laid late in the season. These results may be partly related to the age and experience of the penguins. However, regardless of age, peripheral nesting and late laying were always disadvantageous. The sex ratio of adults in the colonies was 117♂:100♀. This may be explained by the higher mortality of females. Some males could not find partners, but females that did not breed had probably been unable to obtain sufficient food for gonad development. The return of penguins, incidence of non-breeding, adult mortality, clutch-size, and breeding success at Cape Bird were all markedly different in 1968–69 compared to the other three seasons studied. This season was marked by the persistence of sea-ice along the northwestern shores of Ross Island. The low reproductive output in 1968–69 was thought to result from a shortage of food for egg-laying and incubation.     

Reduced parental effort in relation to laying date in house sparrows (Passer domesticus): a study under controlled conditions

It is known that breeding success in birds declines with an advance in the laying date. In this paper, breeding success diminished with laying date in a house sparrow (Passer domesticus) population breeding in captivity. The principal cause of the cline in breeding success with date was a decrease in the female feeding rate. The parental effort in females decreased with the advance of the laying date, although food access was ad libitum throughout the breeding season. Consequently, this cline in breeding success may be due to a reduced parental investment with the advance of the laying date or to exhaustion due to previous reproductive events in the season. Hypotheses usually used to explain the cline in breeding success with date can not explain results in this study.     

The effects of low dietary calcium during egg-laying on eggshell formation and skeletal calcium reserves in the Zebra Finch Taeniopygia guttata

Many small passerines forage intensively for calcium-rich foods during laying. Increased incidences of shell defects in eggs of small passerines have been reported, particularly in western Europe, and these have been explained in terms of declining calcium availability in soils, resulting from prolonged anthropogenic acid deposition. Studies in the field have provided laying birds, nesting in areas of low calcium availability, with calcium supplements. An alternative approach was adopted in this study by allowing captive Zebra Finches Taeniopygia guttata to lay first clutches on ad libitum calcium, switching them to a low calcium diet for 72 hours for the formation of all but the first egg of the second clutch and reinstating ad libitum calcium for the final clutch. Control females had access to ad libitum calcium for all three clutches. Clutch sizes did not vary significantly between birds on low calcium and controls. The former took over three days longer to lay clutch 3 than did controls but the difference was not statistically significant. Birds on low calcium laid eggs that declined in shell ash mass with laying sequence, indicating that birds may have been calcium-limited. Although not statistically significant, eggshell thickness also declined with laying sequence in clutches laid by females on low calcium. The remaining egg measurements (shell mass, shell surface area and volume] of clutches laid by birds on low calcium did not differ significantly from those of controls. Furthermore, females on low calcium did not resort to skeletal reserves to provide sufficient calcium for egg formation. Dietary calcium appears to be of paramount importance in providing sufficient calcium for clutch formation.     

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 breeding season of three species of kiwi (Apteryx) in captivity as determined from egg-laying dates

Records from kiwi ( Apteryx spp.) breeding centres were used to obtain laying dates of 403 North Island brown kiwi ( Apteryx australis mantelli ) eggs, 24 great spotted kiwi ( Apteryx haastii ) eggs and 25 little spotted kiwi ( Apteryx owenii ) eggs. North Island brown kiwi outdoors had an annual cycle of egg-laying with 88.5% of eggs laid from June to January. Eggs were laid in every month of the year, with fewest eggs in April (2.1 % of all eggs). The greatest number of eggs was laid in July (15.6% of all eggs), with a second peak of laying in October (13.5% of all eggs). Both the great spotted kiwi and little spotted kiwi had annual cycles of egg-laying, with the main egg-laying seasons being August-January (87.5% of eggs) for the former and July-December (96.0% of eggs) for the latter. The highest peaks of laying were in October and August, respectively. Kiwi eggs were considered to belong to the same clutch if they were laid no more than 40 days apart. Clutch size in the North Island brown and little spotted kiwi was most commonly one or two eggs. In the great spotted kiwi only clutches of one egg were found. The mean clutch size of North Island brown kiwi outdoors (1.51±0.05) did not differ from that of free-living North Island brown kiwi (1.33±0.09). Similarly, there was no difference in the mean clutch size of captive and free-living little spotted kiwi (1.39±0.14 and 1.10±0.07). The mean interval between eggs for kiwi outdoors did not differ between North Island brown kiwi and little spotted kiwi (27.4±0.5 and 30.7±1.8 days). The results indicate that all three species of kiwi maintain annual cycles of egg-laying in captivity, with the main egg-laying season being longer than in free-living birds.     

Reproduction and monogamy in captive flock of greater flamingos (Phoenicopterus Roseus)

Due to the great number of greater flamingos in captivity and their long life span, studying their behavior and welfare might be useful to improve the husbandry and breeding of this species in zoos. This study aimed to investigate factors affecting the breeding activity of captive greater flamingos (Phoenicopterus roseus) from 2012 to 2016. To estimate reproductive success, the number of pairs, eggs laid, and hatchlings were recorded. In addition, information on age, egg-laying history of the partners, and pair composition was collected. An increase in the number of pairs, eggs, and hatchlings was reported over the years. For each breeding season, there were pairs who laid more than once, especially in 2014 and 2015. Approximately 50% of pairs were monogamous between consecutive years; however, the percentage dropped gradually when comparing nonconsecutive years. Senescence and previous experience seemed to affect the reproductive success of the study flamingos. In conclusion, different factors can influence the reproduction of greater flamingos in zoos. All these factors are related to nonhuman animal welfare and need to be considered in developing and improving management practices.     

Parental age and reproduction in the Marsh Tit Paras palustris

In the Marsh Tit Parus palustris , breeding performance improved with the age of both the male and the female parent—older birds produced larger clutches earlier in the season. The effect of male age, however, was mainly due to a positive association with the age of the female. When the effect of laying date was accounted for, there was no additional effect of female age on clutch-size. However, pairs with previous breeding experience laid larger clutches than newly formed pairs of the same age. There was some evidence that young females that laid late in the season were less likely to survive to breed again, leading to an age effect on reproductive performance. There was no evidence that individual birds laid earlier or produced larger clutches when they became older.     

Reproductive assessment of the great hornbill (Buceros bicornis) by fecal hormone analysis

The population of great hornbills (Buceros bicornis) in the United States is rapidly aging, and captive breeding efforts have not met population managers' expectations for a sustainable captive group. Little is known about the reproductive physiology of these birds. This study reports the first data on the re‐productive endocrinology of the great hornbill. The hormone profiles of the only pair of these birds that hatched a chick in the 1999–2000 breeding season are compared to the profiles of six other pairs of hornbills, from different institutions in the United States, that did not reproduce successfully that season. The study investigates the estradiol, corticosterone, and testosterone profiles of these seven pairs of birds, establishing a base of knowledge from which endocrine data may be used to improve the success of captive breeding programs. The estradiol profiles from this study indicate a difference in hormonal patterns between laying and non‐laying female great hornbills. Egg‐laying females had significantly higher estradiol concentrations during the breeding season than the non‐laying females (P<0.003). Testosterone concentrations of the males were not significantly different between the mates of egg‐laying and non‐egg‐laying females. The corticosterone concentrations tended to be lower in the females that laid eggs vs. the non‐egg‐laying group. The males of the egg‐laying pairs showed a significantly lower (P<0.036) corticosterone concentration than the non‐egg‐laying male pairs. This, combined with the extremely low corticosterone levels (compared to the other birds in the study) of the pair of hornbills that hatched a chick in the 1999–2000 breeding season, suggests that adrenal activity may play a role in the reproductive failure of some captive great hornbills. Zoo Biol 22:135–145, 2003. © 2003 Wiley‐Liss, Inc.     

Errors in egg-laying by female Common Cuckoo Cuculus canorus in nests of its common host

Dozens of studies have documented that brood parasites are well adapted to a brood parasitic lifestyle but not all parasitism events are successful. Co-evolution between brood parasites and their hosts is a dynamic process so it is reasonable to expect that a female brood parasite may commit errors during egg deposition by laying her eggs outside the laying period of the host, with consequent impacts on her fitness. Using an extensive dataset from a long-term study, we evaluated egg-laying patterns and errors related to the timing of egg-laying in the Common Cuckoo Cuculus canorus (hereafter ‘Cuckoo’). Specifically, we tested whether the Cuckoo avoids laying before or on the day of host clutch initiation to reduce the risk of rejection of parasitic eggs, whether laying errors will be more frequent in periods with a lack of active host nests, and whether the laying errors will be more frequent in periods with intense Cuckoo parasitism and a consequent lack of suitable host nests. We found that about one-third of Cuckoo eggs were laid on the host clutch initiation day or 1 day before, and the percentage of Cuckoo eggs laid decreased thereafter. Surprisingly, the probability of Cuckoo egg acceptance by the hosts was not affected by the egg-laying stage of the host clutch. Errors in the timing of egg-laying with fatal consequences (i.e. those precluding Cuckoo hatching because of laying in incubated or deserted clutches) were recorded in about 5% of cases. Only laying date of a Cuckoo egg had a significant effect on the probability of errors, which increased during the breeding season. This may be related to the higher number of deserted and incubated host nests at the site at the end of the breeding season. Errors in egg-laying may be attributed to young and inexperienced females but also impaired body condition or intraspecific competition may cause this behaviour. Future studies, which will test these possible explanations, will help to understand better the mechanism of co-evolutionary arms races and differences between host specialist and generalist brood parasites in various host–parasite systems.     

Regulation of Yolk-Androgen Concentrations by Plasma Prolactin in the American Kestrel

The concentrations of maternally derived androgens in the yolks of avian eggs vary within and among clutches, but a mechanistic basis for this variation has not been elucidated. We investigated in the American kestrel, Falco sparverius, whether changes in plasma-prolactin concentrations induced by changes in photoperiod and food supply affect yolk-androgen concentrations. Over the course of a photoinduced breeding period in the laboratory, we measured concentrations of plasma immunoreactive prolactin (ir-prolactin) in female kestrels with ad libitum food availability (control) or food availability that was reduced during the early breeding period. In a second laboratory study, we administered via osmotic mini-pumps ovine prolactin (o-prolactin) to females beginning on the day they laid their first egg of a clutch (egg-day 1) to determine the effects of high prolactin concentrations on yolk-androgen concentrations. In both this study and one on free-living kestrels, we quantified changes in yolk-androgen concentrations with date of clutch initiation. Concentrations of ir-prolactin in nonlaying females rose with date, irrespective of food treatment. Egg-day 1 ir-prolactin concentrations were higher in control females laying late during the breeding phase than in those laying early. This increase was absent in food-reduced females. Yolk-androgen concentrations in eggs 3 and 4 but not eggs 1 and 2 of the clutch were higher in clutches initiated late than in clutches initiated early in the breeding phase in both the field and laboratory. o-prolactin treatment elevated yolk-testosterone but not androstenedione concentrations. These findings suggest that, in American kestrels, seasonal and laying-associated increases in plasma-prolactin concentrations elevate yolk-testosterone concentrations. Food availability and other factors may interact with date to regulate the effects of prolactin on yolk-testosterone deposition.     

THE OCCURRENCE AND SIGNIFICANCE OF ANOMALOUS REPRODUCTIVE ACTIVITIES IN TWO NORTH AMERICAN NON-PARASITIC CUCKOOS COCCYZUS SPP.

Among the unusual breeding habits of the non-parasitic Yellow-billed and Black-billed Cuckoos of North America are great variability in clutch size and rate of laying, initiation of incubation long before the clutch is complete, occasional laying in nests of other species, annual irregularity in the timing of the breeding season, and semi-nomadic post-migratory movements into breeding areas where food is abundant. These facts, in addition to their peculiar diet and the very large size of their eggs, suggest that cuckoos have extraordinary problems in obtaining adequate energy for reproduction. At Bloomington, Indiana (U.S.A.), during a 15-year period, anomalies in the reproductive activities of cuckoos were concentrated into two years in which food was abundant. This was particularly true of one of these years, when there was a vast emergence of periodical cicadas: the Yellow-billed Cuckoo advanced its normal schedule and bred during peak cicada abundance, laid unusually large clutches, and parasitized Black-billed Cuckoo nests. Some females may have resumed laying in nests in which, having already deposited clutches of normal size, they had been incubating for long periods; the alternative possibility is that there was intraspecific brood parasitism. The erratic egg-laying behaviour of these cuckoos is attributed to the evolution of mechanisms permitting very quick exploitation of a favourable feeding situation. It is suggested that reproductive behaviour has become so responsive to an abundance of food that normal ordering and integration of the stages of breeding have been lost in some females. Such a loss could be responsible for the laying of eggs in alien nests, and it may have been the antecedent of obligate brood parasitism in parasitic cuckoo species.     

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.     

Causes and consequences of egg mass variation between and within blue tit clutches

Causes of egg-size variation between and within clutches were studied in clutches of the blue tit ( Parus caeruleus L.). We measured the mass of each egg in the laying sequence in unmanipulated clutches, in clutches of parents experimentally supplied with extra food before egg-laying, and in clutches of parents supplemented with extra food after the start of egg-laying. Hatchlings were weighed at an age of two days and their mass was found to be positively related to egg mass. No general trend of decreasing or increasing egg mass was found within the laying sequence. Females provided with extra food before egg-laying laid clutches with significantly less variation in egg mass than did control females. The reason for this was that the first-laid egg of unmanipulated females was lighter than the rest of the eggs in the clutch. This pattern disappeared in clutches of females receiving extra food. Thus, the reduction in egg mass variation among clutches of foodsupplemented females depended on an ability of these females, in contrast to control females, to lay a first egg of the same mass as the rest of the clutch. Eggs laid after the initiation of incubation were significantly heavier than equivalent eggs in those clutches where incubation started after clutch completion. The difference was small, however, and the adaptive significance of the finding is questionable. We argue that intra-clutch variation in egg mass is connected with greater fitness consequences than in inter-clutch variation. Furthermore, our results indicate that energetical constraints on the laying female are more important as a cause of the observed intra-clutch variation in egg mass than are adaptive responses to the environment.     

Experimental dissociation of the effects of diet, age and breeding experience on primary reproductive effort in zebra finches Taeniopygia guttata

Reproductive performance varies with age in a wide range of organisms, and increasingly such patterns are interpreted in terms of state-dependent models. We sought to characterise 'state' with regards to age-related variation in clutch size, egg mass and timing of breeding in captive zebra finches Taeniopygia guttata , focusing on the roles of diet quality, age and breeding experience. Females on a high-quality diet laid larger clutches of larger eggs than did females on a low-quality diet. The effect of age on reproductive performance was examined by comparing females breeding (i.e. paired) for the first time at either 3- and/or 6-months of age. Clutch size increased with age but on the low-quality diet only, not on the high-quality diet. Furthermore, clutch size decreased between 3- and 6-months of age in birds bred first on the high-quality diet and then on the low-quality diet. Age did not affect egg mass but older birds had shorter laying intervals. Reproductive performance did not differ between females breeding at 6-months of age for the first or second time: the effects of age were not due to 'training' effects or experience specific to breeding (e.g. undergoing the physiological process of egg formation). In conclusion, nutritional condition (diet) emerged as a central component of state that could strongly influence, and even reverse, any age-dependent increase in primary reproductive performance.     

Reproductive performance changes with age and laying experience in the Red-legged Partridge Alectoris rufa

Most studies of birds have reported positive effects of ageing on reproductive traits during the first years of life followed by a decline. We examined the effects of age and laying experience, controlling for confounding factors such as food availability/quality and timing of pair formation in a captive population of Red-legged Partridge Alectoris rufa . We found that all reproductive traits, except for the period between eggs, were highly repeatable within females both within and between years. The repeatability of egg mass was lower than that of clutch size, laying onset or chick traits other than the mass and size increase rates, perhaps suggesting a lower heritability value. As females aged, they showed a tendency to produce marginally lighter eggs (1.3%) and larger clutches, although this latter trend was not significant. We suggest that the senescence process implies the involvement of hormonal changes that could affect the reduction of egg mass (e.g. low levels of oestradiol). Younger females tended to start laying later than older females, which could be due to a low responsiveness to photoperiodic stimulation of the reproductive system, which would indicate a delay in gametogenesis and steroidogenesis. The mass of last-laid eggs decreased with age of the female. We suggest that physiological changes through the laying sequence could vary with age. Experienced breeders laid larger clutches linked with the advancement of laying onset. For a given age, females with more experience fared better than those with less experience.     

Egg variability and hatching success in the Cape petrel Daption capense at Nelson Island, South Shetland Islands, Antarctica

Eggs of the Cape petrel Daption capense at Nelson Island, South Shetland Islands, Antarctica, exhibited high variability in volume between females (up to 48%), while for each individual female, both volume and shape of eggs were highly correlated over two seasons. Both female size and body-condition at laying were unrelated to egg-volume in the 'good'season 1991, but larger females produced larger eggs in the 'bad'season 1990, when eggs were smaller on average. In 1991, females in better body-condition, but not larger, started laying earlier. In spite of synchronized laying (75% of eggs laid within five days), egg-volume decreased with laying date. Egg-volume differed significantly between the two years, which could not be explained by changes in the breeding population or timing of breeding. The laying of relatively small eggs and low overall breeding success in 1990 most probably reflected changes in food availability. Overall hatchability was 88% and did not differ significantly between disturbed and control colonies in 1991. Larger and more rounded eggs showed better hatching success, but hatchability was related more to egg-volume whereas hatching rate was related more to egg-shape. As the majority of egg losses were attributable to predation, the hatching rate was influenced by parental performance relatively more than by hatchability. Hence the relationship between hatching rate and egg-shape probably reflects improvement in hatching success with female age/experience, whereas the relationship between egg-size and hatchability suggests effect of egg-size independently of parental traits.     

Managing the social environments of flamingos for reproductive success

Recent research has shown that large flamingo flocks are more likely to experience reproductive success than small flocks and that there is a positive relationship between behavioral stimulation from group displays and reproductive success. This study compares the group displays and reproductive success in a captive flock of Caribbean flamingos over 6 years at the National Zoological Park. Birds were added to the flock in two different years. In each case, the flock produced fertile eggs during the breeding season following the addition of new birds. Furthermore, the addition of new birds in 1988, and then again in 1991, significantly increased group display activity over the previous year when the flock size was lower. That the addition of birds stimulated group display activity suggests an alternative management strategy for promoting reproduction in captivity: separate birds in a flock and then reunite them prior to the breeding season. © 1994 Wiley-Liss, Inc.     

Interactive effects of food and age on breeding in the Montagu's Harrier Circus pygargus

We evaluated the direct and interactive effects of food and age on reproduction in the Montagu's Harrier Circus pygargus , to test whether variation in food supply was likely to affect age-specific breeding probability or success. Younger females were more frequently non-breeders than older females. When breeding, older females laid earlier, produced larger clutches, failed less often and had higher number of fledglings than younger females. Probability of breeding was higher, laying was earlier, and clutch size and number of fledglings per pair increased with increasing food abundance. A significant interaction between food and age was observed in both breeding probability and breeding performance: older females were more likely to breed than younger females when food abundance was low, and younger females performed less well in good food conditions than older females. Overall, differences between age groups were most marked in extreme food conditions, regardless of the quality of the conditions.     

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.