PREY TAKEN BY TAWNY OWLS DURING THE BREEDING SEASON

The diet of Tawny Owls during the breeding seasons 1949-52 in Wytham Woods, near Oxford, was determined (a) from analysis of pellets collected, (b) from observation at night, by the use of a red floodlight, of prey brought to the nest and (c) from records of prey left in the nest made during daily visits to weigh the young.
Analysis of pellets showed an increase in the proportion of moles and beetles (mainly cockchafen) in the diet after the first week of May (the time when, on average, the young owls are about half grown) and a decrease in the proportion of mice and voles.
These changes were confirmed in a more emphatic way from observations of food being brought to the nest and from records of prey left in the nest.
This greater emphasis suggests that tho food brought to the young may differ from that which the adults eat themselves.
The fact that no moles were observed being brought to nests at night, wherean many were recorded as surplus prey in the nest, showed that diurnal hunting is regular during the breeding reaaon.
A true assessment of prey taken by Tawny Owls during the breeding season should be based both on analyeis of pellets cast by the adults and on records of food brought to the nest throughout 24 hours. Such records could best be obtained with an automatic camera and flash and a design of nest-box which is described.     

Effects of mate removal on the behaviour and reproductive success of Reed Warblers Acrocephalus scirpaceus

The Reed Warbler Acrocephalus scirpaceus is a largely monogamous insectivorous passerine in which males and females provide equal care by day to eggs and chicks. Polygyny occurs occasionally, with males leaving one female unaided. When females were temporarily removed from three recently-completed clutches their males deserted and resumed the high song levels typical of unmated males. Males may desert either because they are physically incapable of incubation or because the energy expenditure needed for a male to return to an equivalent stage in the breeding cycle is much lower than for a female to do so. Widowed females ( n = 7 ), however, continued the breeding attempt alone, with similar incubation levels but higher provisioning rates than those of control females. In three out of four mid-season broods raised by lone females all fertile eggs were reared to healthy fledglings (in the fourth brood the female died), while only one of four late-season nests produced any fledglings (which were underweight). Late-season control nests were as successful as earlier ones. Loss of male help led to starvation of chicks, but caused no adverse effects during incubation. This explains the small changes in widows' sitting levels during incubation, but much greater effects after hatching. Females may need male help to rear late broods (but not early broods) as days are shorter and food is scarcer. Males may normally help at nests, even those in the mid-season, because in stressful spells (even for a few days) such help is vital for successful breeding but in good periods it costs the male little.     

Incubation prior to clutch completion accelerates embryonic development and so hatch date for eggs laid earlier in a clutch in the great tit Parus major

The ability of parents to respond to changes in food supply within a season will have a large effect on fitness through the number and quality of chicks fledged. Great tits, Parus major, attempt to synchronise their production of chicks with a seasonal food peak, but when food supply fails, hatching asynchrony of chicks provides a mechanism by which some young can be fledged because more developed chicks outcompete their less developed siblings for the reduced parental food supply. We tested whether female great tits can potentially control the degree of hatching asynchrony by using incubation before clutch completion, so that early laid eggs develop faster and hatch sooner. The temperature of an artificial egg placed in 29 nests during the laying period was measured with data loggers, and nocturnal incubation of eggs similar to incubation post clutch completion was recorded in all nests. We then demonstrated that eggs removed from the nest for 72 hour periods prior to clutch completion hatched later than eggs remaining in the nest for the entirety of the laying period. Our results show that variable pre clutch completion incubation (which was mostly nocturnal) can lead to faster embryo development and earlier hatching, so potentially providing a mechanism for adaptive female control of degree of hatching asynchrony.     

Larval sensory abilities and mechanisms of habitat selection of a coral reef fish during settlement

In most animals, siblings from a given reproductive event emerge over a very short period of time. In contrast, many species of birds hatch their young asynchronously over a period of days or weeks, handicapping last-hatched chicks with an age and size disadvantage. Numerous studies have examined the adaptive significance of this atypical hatching pattern, but few have attempted to explain the considerable intrapopulation variation that exists in hatching asynchrony. I explored proximate determinants of hatching asynchrony by monitoring 112 Burrowing Owl (Athene cunicularia) nests in the grasslands of southern Saskatchewan, Canada, over 4 years. Age disparities between first- and last-hatched siblings (i.e., hatching spans) varied considerably, ranging between 1 and 7 days (mode = 4 days). These hatching spans increased with increased hatching success. Hatching spans also increased with larger clutches, but the increase was less than predicted given the increased time required to lay more eggs. Hatching span was unrelated to number of prey cached in the nest during egg laying (an index of food availability), and was unaltered by a year of super-abundant prey. Furthermore, pairs given extra food during laying had hatching spans equal to those of unsupplemented control pairs. These results were inconsistent with both the energy constraint and facultative manipulation hypotheses, which predict that hatching asynchrony should vary with the level of food during laying, when incubation onset is determined. Burrowing Owls were apparently free of food limitation early in breeding, yet may not have been able to optimize hatching spans because food conditions during laying were largely unrelated to food conditions during brooding. Thus, one of the premises for facultative manipulation of hatching asynchrony—that laying females are able to forecast post-hatch food conditions—may not have been met for this population of Burrowing Owls.     

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

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

Cooperative males reduce incubation in response to cues of female–female competition

Social groups of the joint‐laying Pukeko Porphyrio porphyrio melanotus typically contain one or two breeding females. Male Pukeko mated to two females father more offspring and therefore benefit from this mating arrangement; however, primary females should not prefer this system, because fewer eggs hatch per female in the larger joint clutches. Here, we investigated male response to simulated egg destruction, a common female competitive tactic observed in other joint‐laying species. In response to egg removal, males reduced the consistency of their incubation and in some cases nests were abandoned. Such decreases in paternal effort could eliminate any putative advantage gained by a female that destroys the eggs of a co‐nester. Our study demonstrates facultative adjustments in paternal care in a joint‐laying species and suggests that primary females may be limited in their ability to monopolize reproduction.     

黑头噪鸦的繁殖及生活史特征

黑头噪鸦是特产于中国青藏高原东南部边缘高山针叶林的一种珍稀鸟类。199—2002年在甘肃省卓尼县和四川省九寨沟县的野外研究中发现,黑头噪鸦繁殖时间早,一般3月开始繁殖,4月中旬雏鸟孵化。在孵卵期和育雏期的第一周,雌鸟95％的时间在巢内,每次离巢时间一般不超过15min。黑头噪鸦具有储食行为,其储藏的食物对雌鸟的孵卵以及幼雏的存活可能有重要意义。黑头噪鸦由3-4只个体组成一个繁殖单元,有证据表明帮手在育雏期向幼鸟提供食物。     

Body mass regulation during incubation in female common eiders Somateria mollissima

We investigated changes in incubation behaviour induced by body fuel depletion in incubating female common eiders, which, in contrast to pelagic seabirds, fast despite being close to marine food sources. In the Svalbard Archipelago, electronic scales were placed under eider nests and the incubation of six birds was prolonged by using wax-filled eggs. Based on changes in the rate of body mass loss in normally incubating females and in ten captive birds that did not incubate, body reserves neared depletion on average four days after hatching. During prolonged incubation, females took more frequent and longer recesses. Nest attentiveness consequently decreased, but was still high. In contrast to recesses during normal incubation, during which body mass of the birds decreased, mass remained constant during the recesses of prolonged incubation. The body stores of female eiders seemed to enable them to complete incubation with a limited safety margin. A further drop in body mass is avoided when a critical body mass is reached, because birds then start feeding enough to maintain mass while continuing incubation. Presumably, a similar mechanism will enable eiders to continue incubation when body reserves are prematurely depleted before hatching.     

Resource allocation between reproductive phases: the importance of thermal conditions in determining the cost of incubation

Changes in the resources allocated to particular stages of reproduction are expected to influence allocation to, and performance in, subsequent reproductive stages. Experimental manipulation of individual investment patterns provides important evidence that such physiological trade-offs occur, and can highlight the key environmental variables that influence reproductive costs. By temporarily altering the thermal properties of starling nests, we reduced the energetic demand of first-clutch incubation, and examined the effect of this manipulation on performance during the same and the subsequent reproductive attempts. Compared with controls, starlings investing less in incubation were more successful in fledging young, and were more likely to hatch all their eggs if a subsequent reproductive attempt was made. Our results show that incubation demands can limit reproductive success, and that resources saved during incubation can be reallocated to later stages of the same reproductive attempt and to future reproductive attempts. This study also shows that small changes in thermal environment can affect breeding success by altering the energetic demands imposed on incubating parents, independently of the effect of temperature on other environmental variables such as food supply.     

Temporal patterns of circulating LH and steroid hormones in a brood parasite, the Brown-headed cowbird, Molothrus afer.

The breeding behaviour of female Brown-headed cowbirds, Molothrus ater , is remarkably distinct from that of females of most other passerine species. Cowbirds are brood parasites that do not build nests, incubate eggs or feed young. Instead, they have an extended egg-laying stage, and deposit eggs in the nests of hosts within large breeding territories that are defended from other female cowbirds. In this communication, we report the changes in plasma levels of circulating hormones and in morphological characteristics that occur in free-living female Brown-headed cowbirds over a breeding season.
Plasma LH in female cowbirds increases at the onset of breeding and remains elevated for approximately eight weeks, at which time levels decline to basal values. This pattern is similar to that of follicle development, and is consistent with the prolific reproductive output by individuals of this species. Circulating E, levels are quite variable and do not demonstrate any discernible pattern, while plasma B is elevated during the egg-laying period and declines subsequently.
Testosterone titre (T) is highest early in the breeding season, when territorial interactions are generally most frequent. T levels decline slowly thereafter, reaching their lowest values at the season's end. Plasma DHT levels are slightly higher than those for T; DHT exhibits little change until late in the breeding season, when it declines. Fat scores are highest prior to egg laying, and fat reserves diminish as the breeding season progresses. In contrast, body weight increases during the reproductive season, probably as a result of developing eggs in the ovary. These results are consistent with the hypothesis that temporal patterns of hormone secretion are correlated with the expression of social systems.     

Individual female clutch identification through yolk protein electrophoresis in the communally breeding guira cuckoo (Guira guira)

Avian communal breeding systems generate alternative behavioural strategies for females, resulting in differences in reproductive success. Identifying eggs of different females in such systems is problematic, however, due to egg destruction before incubation, difficulty of capturing adults, and/or inaccuracy of egg identification based on egg morphometry. Here, we describe a technique that uses electrophoresis of yolk proteins to determine egg ownership, which we applied to communally breeding guira cuckoos (Guira guira). Validation of the method included identical yolk protein banding patterns in all eggs of the same female, but different patterns in eggs of different females in budgerigars (Melopsittacus undulatus), and identical patterns in yolk follicles of the same females in guira cuckoos. We applied the protocol to 195 guira cuckoo eggs from 34 joint nests in 2 years. All multiple guira cuckoo eggs laid on the same day in single nests had distinct banding patterns of yolk proteins, practically eliminating the possibility of more than one female being represented by the same pattern. Some identical banding patterns were repeated in different days within a nesting bout, indicating that some females laid several eggs in shared nests. Identical patterns occasionally occurred in renestings of groups, indicating that some females lay eggs in consecutive nestings. Yolk protein electrophoresis is a useful tool to identify egg maternity in other circumstances, such as polygynous mating systems with joint nests and intraspecific parasitism. Additionally, it is an alternative method for species where electrophoresis of egg white proteins does not show sufficient polymorphism.     

Renesting intervals and duration of the incubation and nestling periods of Sprague's Pipits

ABSTRACT.   Sprague's Pipits ( Anthus spragueii ) are one of the least studied birds in North America, with most information about their breeding biology based on anecdotal observations at relatively few nests. I determined the renesting intervals of female Sprague's Pipits in south-central Saskatchewan and also summarized data about incubation and nestling periods of Sprague's Pipits in Saskatchewan and Manitoba to provide more robust estimates of these life-history parameters. Two of eight radio-marked females initiated replacement clutches after failed nesting attempts, with renesting intervals of 11–13 and 15–16 d, respectively. Of 74 banded females, 3 were subsequently recaptured at later nests during the same breeding season. Nests of two of these females failed due to inclement weather and their renesting attempts were initiated 6 and 7–11 d later, respectively. The third female successfully fledged young and then initiated another nest 17 d later. The new nests of females that renested were all located less than 150 m from previous nests. Incubation and nestling periods for known-aged nests ranged from 12 to 15 d (13.4 ± 0.3 [SE], N = 9) and 11 to 14 d (12.1 ± 0.2, N = 39), respectively. Variation in the length of both incubation and nestling periods was likely a function of prolonged periods of inclement weather. More data are required to determine the extent of renesting in different habitats and the extent to which Sprague's Pipits are capable of successfully raising multiple broods in a single breeding season.     

Determination of clutch-size in the Kentish Plover Charadrius alexandrinus

We studied the clutch-size distribution of Kentish Plover Charadrius alexandrinus in 1991 and 1992 and tested three hypotheses for determination of clutch-size: egg-formation ability of females, incubation ability of parents and nest predation. Variation in clutch-size was small: 71 out of 74 clutches had three eggs (coeff. var. = 7%). Females spent more time foraging (51 ± 6%) pre-laying and during egg-laying than their mates (39 ± 5%). However, we concluded that egg-formation was not constrained by food availability because 77% and 100% of clutches were initiated before the peaks of prey density and of prey mass, respectively. Furthermore, the number of clutches initiated and completed over time was unrelated to prey density. By experimentally reducing and enlarging clutches, we found that enlarged clutches of four eggs took longer to hatch (24.8 ± 0.9 days) than control clutches of three (21.6 ± 0.7 days). Eggs of enlarged clutches also lost weight more slowly during incubation in both years compared with control clutches. No difference was found in the incubation behaviour or weight loss of parents between reduced, control and enlarged clutches. We found no evidence to support the nest predation hypothesis, since neither the proportion of nests predated nor the number of chicks hatched was different between reduced, control and enlarged clutches. The results of this study are most consistent with the incubation ability hypothesis, although the parental ability hypothesis has remained untested.     

Assessment of territory quality and its effects on breeding success in a migrant passerine, the Wheatear Oenanthe oenanthe

This paper examines how a returning migrant assesses the quality of an area as a breeding territory before the period of peak food demand and how effective the assessment is in terms of breeding success. Male Wheatears Oenanthe oenanthe return from Africa to choose territories in the Breckland of eastern England about March, females arriving shortly after males. The food supply was predictable: prey densities during the breeding period (egg-laying to chick independence) were strongly correlated with prey densities at the same sites during the period of arrival and territory establishment. Prey densities were also related to vegetation structure, averaging highest on short turf. Male arrival date and territory size were not significantly related to prey density but were strongly related to vegetation structure, implying that birds used vegetation as an indirect clue to prey availability. Neither territory size nor nest spacing appeared to affect nest losses caused by predators. The major variations in number of young fledged (other than predation) were caused by the number of nestlings hatched and presence of a second brood. Both early arrival and an early first brood improved first-brood success and were necessary for a second brood. Not all birds which arrived early bred early enough for a second brood. First-brood hatching date was strongly negatively correlated with pre-breeding prey availability but not significantly related to vegetation structure. Thus by using vegetation as a clue to habitat quality, some pairs suffered reduced breeding success. This result implies that birds may not be able adequately to assess prey density directly at the time of territory establishment. The critical period for food availability may not be the period of peak demand (nestling period), when food is relatively abundant, but is probably the pre-breeding period when females must accumulate reserves for eggs and when the food supply is poor. Food supply during this period may determine the timing of breeding and the ability to rear a second brood, and may thus have a greater effect on breeding success.     

Division of labour during incubation in a woodpecker Colaptes auratus with reversed sex roles and facultative polyandry

The contribution of males to incubation has rarely been studied in altricial birds because the pursuit of extra mating opportunities is believed to conflict with incubation. Woodpeckers show reversed sex roles in parental care with males doing most of the nest construction, incubating and brooding of the young while females may be polyandrous. I investigated incubation by each sex at 71 monogamous and four polyandrous nests of the Northern Flicker Colaptes auratus , predicting that males would contribute to incubation according to their energy reserves (body condition) whereas females would contribute based on alternate reproductive opportunities. Nest attendance was 99% with males contributing a mean of 66% of the total incubation including nocturnal incubation. The length of daytime bouts averaged about 2 h and did not differ between the sexes. Consistent with predictions of investment strategies, structurally larger males and those in poorer body condition incubated less than smaller males, perhaps because they required more recess time to forage or to conserve energy. Older females contributed less incubation than young females and polyandrous females contributed less incubation at their secondary nests than monogamous females. Incubation period, nest depredation rate and hatching success were not influenced by bout length, number of bouts or relative contribution of the sexes. Hatching success was 86% at nests of both monogamous and polyandrous females because males compensated for reduced female participation. Because incubation of the sexes is compensatory and not additive, incubation pattern did not influence short-term reproductive success. I conclude that males invest in incubation according to their energy needs, and females may adjust their contributions based on alternate reproductive tactics.     

Body mass variation in breeding Florida Scrub‐Jays

ABSTRACT Many birds lose mass when feeding dependent young and multiple hypotheses have been proposed to explain this loss. The reproductive‐stress hypothesis suggests that mass loss results from an energy deficit. The flight‐efficiency hypothesis suggests that breeders lose mass in advance of feeding young to save energy during flight. The reserve‐mobilization hypothesis suggests that female breeders accumulate energy reserves during egg production and incubation and mobilize those reserves to meet their own energy needs after eggs hatch. Finally, birds may lose mass due to gonadal regression. From 1999 to 2001, we attracted Florida Scrub‐Jays (Aphelocoma coerulescens), sedentary cooperative breeders, to a portable electronic balance. Our objective was to determine which hypotheses might best explain mass variation during breeding. Both male and female Florida Scrub‐Jays lost mass during the period of nestling care (males, 3.2%; females, 6.5%), but not when feeding fledglings, despite this being the period of peak effort. Such results are consistent with both the flight‐efficiency and reserve‐mobilization (females only) hypotheses. We also found a significant negative influence of brood size on mass change in males, providing support for the reproductive‐stress hypothesis, and we conclude that, for males, both the flight‐efficiency and reproductive‐stress hypotheses apply. For female scrub‐jays, our results were consistent with the flight‐efficiency and energy‐reserve mobilization hypotheses, both of which view mass loss as beneficial.     

Incubation feeding as a male tactic for early hatching

Male marsh tits, Parus palustris, regularly feed their mates from the beginning of nest building until hatching. Over three periods (the 15 days preceding egg formation, egg formation/laying and incubation) the number of food passes by the male to the female increased significantly. There was a significant negative relationship between the frequency with which the male fed the female in the nest during incubation and the length of the incubation period. Female blue tits, Parus caeruleus, experimentally supplied with food in the nestbox during incubation had a significantly shorter incubation period than control females. Clutches of experimentally fed females also tended to hatch more successfully. It is concluded that feeding of the female by the male is a nutritional contribution and that the shorter incubation period and increased hatching success enhance the fitness of both parents. However, the male should balance the benefits against the costs in time and energy and therefore not necessarily work at a maximal level. In accordance with this is the finding that the male's provisioning rate increased when ambient temperatures decreased. Adverse weather may jeopardize the whole or large proportions of the clutch, thereby significantly reducing the benefit from the current breeding attempt.     

Prolactin and parental behavior in Adélie penguins: effects of absence from nest, incubation length, and nest failure

Adélie penguin (Pygoscelis adeliae) males and females, nesting in Antarctica, alternate attendance at the nest with absences of many days to forage at sea. We investigated the importance of tactile input from egg and chicks on prolactin levels by observing nest attendance patterns and obtaining blood samples (1) during the first nest exchange of the incubation stage, (2) from birds whose incubation period was artificially increased or decreased by about 10 days, and (3) from birds whose nests had failed. Prolactin levels in females after 8 to 11 days of absence from the breeding colony did not differ from those in incubating males and did not change after females resumed incubation. Moving eggs between nests resulted in nests in which chicks hatched after about 26, 36 (normal), or 46 days. Duration of incubation did not affect prolactin levels in the parents measured during incubation, at the pip stage, hatch stage, or early brood stage. Adults first left their chicks unguarded on about the same calendar date, regardless of chick age. However, chicks from long incubation nests averaged 8 days younger when they were left unguarded than chicks from control or short-incubation nests. In females, there was no effect of nest failure on prolactin levels. In males, prolactin levels were slightly lower after nest failure than in males tending nests. Testosterone was significantly higher in males after nest failure than in males still tending nests. Prolactin is elevated in Adélie penguins as part of the program of cyclical hormonal changes that accompany the lengthy reproductive season and is relatively independent of tactile input. Sustained prolactin secretion is probably required for the maintenance of parental behavior in offshore feeding species that must be absent from the nest for many days at a time.     

NESTING OF THE PURPLE-THROATED CARIB HUMMINGBIRD

We present data on certain parts of the nesting biology of the Purple-throated Carib Humming- bird Eulampis jugularis on the island of Dominica, British West Indies. We watched two nests with eggs and young and a single nest under construction. Incubation was in short periods averaging about six minutes. Females at both nests with eggs continued adding material to the nest until the end of incubation. We suggest that this common practice among hummingbirds may insure that a limited amount of energy at the time of initial nest-building and egg-production is optimally apportioned between the two activities. Brooding ceased in one nest when the young were about 13 days old. Feeding rates stayed relatively constant at one to two per hour throughout the nearly complete nestling period in one nest. Average time per feeding declined, apparently as the female-young interaction became more efficient with maturation of the young. Nesting females foraged on both insects and nectar. We could not be sure what they were feeding the young. Defence of the nest varied with the position of the female and the type of intruder. The aggressive behaviour associated with defence varied according to the size and type of intruder. We concluded that for the one nest of Eulampis for which we had the most complete data, had there been three young, the female would have been unable to provide sufficient food to nourish them. However, the universal clutch size of two and nearly universal promiscuity in humming- birds is strong evidence that factors other than the ability of adults to feed more than two young are important in determining clutch size.