Incubation Attendance Research Articles

High Incubation Attendance and Nesting Site Constraints of the Sclater's Monal in an Alpine Environment in Southwestern China.

Breeding in alpine environments poses significant challenges to birds, requiring specific adaptations for survival. The Sclater's monal (Lophophorus sclateri), a regionally threatened, typical alpine pheasant species, is restricted to high-elevation habitat from the East Himalayas to the mountains of west Yunnan, China. Due to its low population density and the difficulty of accessing its habitats, the breeding ecology of this species is understudied. Therefore, we aimed to understand the breeding behavior, nest site use, and life-history traits that allow this monal species to cope with the alpine environment. During our fieldwork from March to June 2015 and 2016 in the Gaoligong Mountains in western Yunnan, China, we found six cliff nesting sites ranging from 3535 m to 3892 m. Three sites were active, with one being used in both years, and the remaining three were inactive but had been used in prior years. The clutch size was 2.75 ± 0.5 (2 or 3 eggs; n = 4 nests at three nesting sites), and all 11 eggs were successfully hatched. The female solely performed incubation, spending 97.2% of its time in incubating with an average duration of 25.69 ± 13.79 h (n = 43 bouts across three females) per on-bout which indicates the female bird invested more time than other pheasants. High incubation attendance by females highlights the importance of increased parental care in ensuring reproductive success. These findings highlight that the Sclater monal exhibits specific breeding behaviors and nesting strategies that reflect adaption to harsh environments. Additionally, our observations of intense male-male interactions and reuse of nesting sites suggest that suitable nesting sites are limited, which could significantly impact population dynamics. Together, these insights are crucial for conserving this regionally threatened pheasant.

Biotic and abiotic factors affecting Atlantic ghost crab (Ocypode quadrata) spatiotemporal activity at an important shorebird nesting site in Virginia.

Atlantic ghost crabs (Ocypode quadrata) are predators of beach-nesting shorebird nests and chicks on the United States' Atlantic and Gulf coasts. Ghost crabs may also disturb birds, altering foraging, habitat use, or nest and brood attendance patterns. Shorebird conservation strategies often involve predator and disturbance management to improve reproductive success, but efforts rarely target ghost crabs. Despite the threat to shorebird reproductive success, ghost crabs are a poorly understood part of the beach ecosystem and additional knowledge about ghost crab habitat selection is needed to inform shorebird conservation. We monitored ghost crab activity, defined as burrow abundance, throughout the shorebird breeding season on Metompkin Island, Virginia, an important breeding site for piping plovers (Charadrius melodus) and American oystercatchers (Haematopus palliatus). We counted burrows at shorebird nests and random locations throughout the breeding season and investigated whether ghost crab activity was greater at nest sites relative to random locations without shorebird nests. While we observed burrows at all nest sites (n = 63 nests), we found that burrow counts were lower at piping plover nests with shell cover, relative to random locations with no shell cover. Ghost crabs may avoid piping plover nest sites due to anti-predator behaviors from incubating adults or differences in microhabitat characteristics selected by piping plovers. We also investigated the effects of habitat type, date, and air temperature on the abundance of ghost crab burrows. We found that while crab burrows were present across the barrier island landscape, there were more burrows in sandy, undisturbed habitats behind the dunes, relative to wave-disturbed beach. Additionally, ghost crab activity increased later in the shorebird breeding season. Understanding when and where ghost crabs are most likely to be active in the landscape can aid decision-making to benefit imperiled shorebird populations.

Interspecific differences in brood desertion by female diving ducks in relation to duckling age and environmental conditions

Studies on brood desertion in birds have been mainly conducted on species with biparental care, and less often on uniparental species. Females of many duck species remain with their ducklings, unassisted by males, during variable periods of time before the chicks can fledge. In this paper, we examined factors that influence brood desertion by female red-crested pochards ( Netta rufina), common pochards ( Aythya ferina) and white-headed ducks ( Oxyura leucocephala) during a 10-year period in lakes in southern Spain that experienced interannual variations in water levels. At an interspecific level, we found that brood desertion was less frequent in the species that nested earlier (red-crested pochard), while it was more frequent in the species that nested later (white-headed duck). Larger (older) ducklings were deserted more frequently than smaller ones in the three species. However, an interaction between flooding conditions in wetlands and species identity on brood desertion, after accounting for chick age, suggests that there were variations between the ducklings of the three species in their requirements for maternal care, conditioned by environmental conditions. Therefore, our study indicates that environmental conditions may affect parental care. A longer duration of brood attendance by female ducks may buffer against adverse effects of environmental conditions.

Sex Role Reversal and High Frequency of Social Polyandry in the Pheasant-Tailed Jacana (Hydrophasianus chirurgus)

In a few species, males invest more than females in parental care while the females invest in mating competition and producing multiple broods for several mates. Species in the family Jacanidae are commonly used for studying this type of breeding system (called sex-role reversal), and previous studies found discrepancies and variation between species in the expected characteristics of reversed sex roles. Yet, a better understanding of sex role differences in breeding behavior in such species is crucial for disentangling possible evolutionary mechanisms leading to this peculiar breeding system. Sex-role reversal in the pheasant-tailed jacana Hydrophasianus chirurgus has been documented long time ago. Since the very early observation of this species, however, there was no attempt to provide a comprehensive and quantitative description of their breeding. This study aims to fill these knowledge gaps by investigating the sex role differences in the breeding behavior of pheasant-tailed jacanas, by observing and monitoring a breeding population in Taiwan. We focused on three main characteristics of sex-role reversal: (1) competition between females for access to males, such as agonistic and courtship behaviors, (2) polyandrous mating, and (3) male-only care. As expected, we found that females provide most of the territory defense toward conspecifics. Males also participated in agonistic behaviors, although less frequently than females. Furthermore, contrary to what was expected, we found that males spent more time than females on courtship behavior. Polyandrous females performed mating and laying sequentially with different mates but maintained the pair bonds simultaneously with multiple males. For the first time for the species, we could estimate that the average number of mates per female (i.e., degree of polyandry) was 2.4 and that at least 81.8% of the females in the population were polyandrous. Finally, our observations corroborated that brood care is predominantly provided by males, nevertheless females were also participating to some degree in brood attendance but never in direct care (i.e., brooding). This study highlights that some aspects of polyandrous breeding might deviate from stereotyped view on sex-role reversal, and stress the importance of further within species and comparative studies in order to fully understand the mechanisms leading to sex-role reversal.

Sex bias in parental care is associated with brood age and fledglings’ growth rate in Western Bluebirds Sialia mexicana

Parental care is defined as any behavioral trait of the parents that improves the fitness of the offspring, and that probably evolved or is currently maintained by this function. There is variation in the type and amount of care allocated by parents depending on fluctuation in brood demands that respond to growth and developmental environment. Each sex, within the breeding pair, may respond differently to changing brood demands, and such variation might impact offspring development and fitness. Even though there is a plethora of studies evaluating sex differences in parental care, proximate factors influencing distribution of parental duties within the breeding pairs have received far less attention. In a wild population of Western Bluebirds, we assessed sex differences in parental care (i.e. brood attendance, food deliveries, nest cleaning) through nestling development, and evaluated whether distribution of parental care between sexes is associated with offspring growth. We found that females adjusted their parental behavior according to changes in brood demands more often than males, and fed, attended, and cleaned more than males. Moreover, a more female-biased distribution of care through nestling development was positively associated with the growth of fledglings. We conclude that sex differences in parental care may be a measure of specialization in parental duties within breeding pairs favored by natural selection. These findings are in marked contrast to earlier studies in Northern populations of Western Bluebirds that have failed to detect differences in parental care between sexes, and suggest that differences in the social environment or ecological conditions between populations of the same species might lead to differences in parental cooperation between males and females.

Brood attendance by female Red Knots

Brood attendance by female Red Knots

Female incubation attendance and nest vigilance reflect social signaling capacity: a field experiment

Due to the reduced conspicuousness of female signals, their evolution has traditionally been interpreted as a by-product of sexual or natural selection in males. Recent studies have argued that they may be the result of sexual or social selection acting on females. Here, we explored the role of the white wing patch during the incubation period in female-female competition contests in a migratory cavity-nesting songbird, the pied flycatcher Ficedula hypoleuca. At this stage, female investment is crucial for offspring survival, while competition among females for nest cavities is still operating. We experimentally performed an extreme reduction of signaling capacity by covering the wing patch with dark paint in a group of females and compared their incubation attendance and social interaction patterns (vigilance and aggression at the nest as defense variables) during simulated territorial intrusion tests with female decoys, with those of an unmanipulated group of females. Tests were performed both before and after the manipulation. We hypothesized that these patches constitute signals of social dominance in female-female interactions that allow them to maintain high levels of incubation attendance by reducing the need for vigilance. We found a marked decrease in incubation attendance in experimental females after manipulation, a change that was not found in the control group. Moreover, vigilance decreased in the control group after the manipulation, a change not detected in manipulated females. No effects on aggressive nest defense were found. Female wing ornamental traits would act as a badge of status in social interactions allowing more intense incubation and reduced vigilance. Implications of social interactions on incubation patterns should be incorporated in future studies. Animals can transmit information to the others through different traits. Ornamental signals have been proposed as status indicators in social interactions in birds in many studies. Although some of them concern their function and evolution in females, only a few have been focused on their role during incubation. Our experimental study suggests that the female’s white wing patch in pied flycatchers acts as a badge of dominance status in social encounters. Thus, dominant females may enforce respect from competitors through signaling, thereby being able to reduce vigilance at the nest with positive effects on incubation efficiency. Implications of social status for incubation patterns should be considered in future studies as an additional factor modulating this crucial parental activity.

Parental investment in two large raptors breeding in a high prey density area

We investigated the breeding behaviour of Bearded Vulture Gypaetus barbatus and Golden Eagle Aquila chrysaetos between 2008 and 2011 in the Stelvio National Park (the Alps, Italy), which harbours one of highest densities of wild ungulates in the Alps. Parental care behaviours (incubation and chick brooding, nest attendance, changeover, food provisioning and nestling feeding) were recorded at nests of both species (four Bearded Vulture and 14 Golden Eagle pairs). Differences in investment between sexes and periods were found in both species: the time spent in incubation and nest attendance was higher in females and decreased with the progress of the breeding season (from incubation to post-hatching and to pre-fledging). A significant effect of the interaction between sex and period was also found. Compared to the literature, our results suggest a lower contribution by Bearded Vulture males, whereas Golden Eagle males spent more time in incubation and nestling brooding than reported. The higher investment shown by Golden Eagle males in our study area may be due to the high availability of live prey and ungulate carrion, which could allow males to spend less time in hunting, resulting in more time at the nest. Most changeovers for both species took place in the central hours of the day. No difference in food provisioning was observed between the sexes of the two species, consistent with information available for Bearded Vultures, but not for Golden Eagles. The patterns we found revealed a female-biased investment (unexpected for the Bearded Vulture), although males significantly contributed to nest behaviour, especially during the incubation period (unexpected for Golden Eagles). The large amounts of natural prey and carrion in the study area may contribute to the nest behaviour of these large raptors.

The effect of ambient temperature, habitat quality and individual age on incubation behaviour and incubation feeding in a socially monogamous songbird.

Incubation is an important aspect of avian life history. The behaviour is energetically costly, and investment in incubation strategies within species, like female nest attentiveness and the feeding by the non-incubating partner during incubation, can therefore vary depending on environmental and individual characteristics. However, little is known about the combined effect of these characteristics. We investigated the importance of ambient temperature, habitat quality, and bird age on female incubation behaviour and male feeding of the incubating female (incubation feeding) in blue tits Cyanistes caeruleus, a socially monogamous songbird. An increase in ambient temperature resulted in a higher nest temperature, and this enabled females to increase the time off the nest for self-maintenance activities. Probably as a consequence of this, an increase in ambient temperature was associated with fewer incubation feedings by the male. Moreover, in areas with more food available (more deciduous trees), females had shorter incubation recesses and males fed females less often. Additionally, males fed young females more, presumably to increase such females’ investment in their eggs, which were colder on average (despite the length of recesses and female nest attentiveness being independent of female age). Male age did not affect incubation feeding rate. In conclusion, the patterns of incubation behaviour were related to both environmental and individual characteristics, and male incubation feeding was adjusted to females’ need for food according these characteristics, which can facilitate new insights to the study of avian incubation energetics.Significance statementParents often invest a substantial amount of energy in raising offspring. How much they do so depends on several environmental factors and on the extent they cooperate to raise the offspring. In birds, males can feed incubating females, which may allow females to stay longer on the nest, which, in turn, may ultimately improve reproductive success. The interplay between environmental factors and such incubation feeding on incubation attendance has, however, received little attention. Here, we show that favourable circumstances (higher ambient temperature and food availability) allowed incubating blue tit females to increase the time off the nest to improve self-maintenance and males to feed them less, whereas males also fed inexperienced partners more often. Thus, we show a concerted effect of several environmental and intrinsic factors on parental effort during incubation, which will help to improve the general understanding of avian incubation and parental care.

Nest Defence Behaviour and Testosterone Levels in Female Pied Flycatchers

AbstractNesting holes are a scarce resource for obligated cavity‐nesting birds and an important selective force for the evolution of aggressive female behaviours, which may be mediated by testosterone (T) levels. It is known that during periods of intense intrasexual competition such as initial breeding stages, females are highly aggressive towards intruding females. Here, we studied the implications of T levels for female–female competition by comparing levels of aggressiveness towards simulated female intruders (decoys) in two populations of the pied flycatcher (Ficedula hypoleuca) with a marked difference in breeding density. To this end, we exposed free‐living females to simulated territorial intrusions during 30 min when nest construction was almost complete. T levels of females were measured at the beginning of incubation under the assumption that they are positively associated with T levels during nest building. We also related aggressiveness to T levels in both populations. Furthermore, we aimed at detecting whether variation of T levels may explain female incubation attendance. Females showed higher T levels in the populations where pied flycatchers were exposed to a higher likelihood of conspecific interactions (high breeding density) than in the population with low breeding density. Female territorial presence, vigilance at the nest box and proximity to decoys were negatively related to circulating T levels in the high‐density population, but not in the low‐density population. Differences in T levels between populations did not result in differences in female incubation attendance, but T levels were negatively related to the incubation attendance in females from the population showing high T levels. In our populations, T levels in females prior to laying reflect the need to defend nesting cavities which is higher at high breeding density and in subdominant females. High T levels are costly in terms of incubation attendance.

Plumage ornaments and reproductive investment in relation to oxidative status in the Iberian Pied Flycatcher (Ficedulahypoleuca iberiae)

A key aspect in the study of plumage traits with a potential role in communication is the cost associated with trait production and maintenance, expressed in terms of oxidative stress. In the Iberian Pied Flycatcher (Ficedula hypoleuca iberiae (Witherby, 1928)), males and some females exhibit a white forehead patch and both sexes present conspicuous white patches on the wings. We examined associations between these plumage ornaments and their ability to cope with oxidative stress. Furthermore, we explored oxidative costs of reproductive investment. Total antioxidant status (TAS) in plasma and glutathione (tGSH) levels in red blood cells, as well as a marker of oxidative damage in plasma lipids (malondialdehyde (MDA)), were assessed simultaneously for the first time in studies of avian reproduction. We found negative associations between antioxidants and ornaments in incubating females, although this relationship was positive while feeding nestlings. For males, MDA levels were negatively associated with ornaments, while TAS showed a positive relation. Female MDA showed a positive correlation with intensity of incubation attendance, while this relation was negative for tGSH levels. These results indicate that multiple achromatic plumage ornaments signal the individual capacity to cope with costs related to oxidative stress. Moreover, this study highlights the critical role of incubation for avian life histories.

Females Paired with More Attractive Males Show Reduced Oxidative Damage: Possible Direct Benefits of Mate Choice in Pied Flycatchers

AbstractDirect benefits of female mate choice may concern female fertility and fecundity but also physiological status. In birds with biparental care, males may contribute to improve the condition and health of their pair‐mates through help in constructing nests, incubation or incubation feeding and nestling provisioning. They may also reduce harassment of females by non‐pair males. A consequence of these male activities could be expressed in terms of oxidative damage, which may depend on metabolic effort and social stress. Here, we have related male contribution to parental and territorial duties to female oxidative status in the pied flycatcher Ficedula hypoleuca, a species where preferred males present darker dorsal plumage and, in Iberian populations, a large white forehead patch. Darker males were paired with females with high incubation attendance and reduced nestling provisioning rates, which may lead to reduced female exertion. These males owned nest boxes at which there were fewer visits by non‐pair males. Although females paired with dark mates worked less hard, they were able to raise more fledglings. Female oxidative damage measured as malondialdehyde (MDA) level in plasma declined with increasing incubation attendance and male incubation feeding. Moreover, levels of MDA in females declined with both darkness of male dorsal plumage and male forehead patch size when controlling for female forehead patch size and male age. The effect of male plumage darkness was especially strong. Females paired with middle‐aged males (2–3 yr) showed reduced levels of MDA compared with those paired with 1‐yr‐old and more than 3‐yr‐old males. Male age could not explain the effects of male attractiveness. Females paired with attractive males were more successful in reproduction while suffering reduced oxidative damage, possibly mediated by help during incubation and nestling rearing from their pair‐mates. Although correlative, the evidence suggests direct benefits of females paired with more attractive males.

Oxidative damage in relation to a female plumage badge: evidence for signalling costs

Badges of status may be controlled by costs derived from increased aggression from dominant individuals. This cost could be translated into elevated metabolic levels and a concomitant disruption of oxidative balance. Some females in Iberian pied flycatcher Ficedula hypoleuca populations exhibit a white forehead patch similar to that exhibited by all males in this species, functioning in aggressive interactions between females when competing for breeding sites. To test if social stress imposes costs on signalling, we painted white patches on females without natural patches (NP) and compared them with females with natural control (NU). We also over-painted the natural patch in other females (FP) and compared to females with control natural patches (FU). We obtained for the whole sample of females data on reproductive investment, morphology and oxidative damage measured by blood malondialdehydes (MDA), and in a subsample of females variables related to parental care during incubation and the early nestling stage. FP and FU did not differ significantly in any variable which negates an effect of paint itself. However, NP females showed significant higher levels of MDA than NU females when controlling for breeding success for the whole sample, and for female incubation attendance for the parental care subsample. When including the four treatments, there was a significant interaction between the paint treatment and the presence/absence of badges before the experiment when controlling for the significant negative effect of incubation attendance on MDA. Addition of a badge to females without one leads to increased oxidative damage possibly mediated by social control. Badges of status in female pied flycatchers may operate as badges of oxidative status.

Evolution of sexual dichromatism in relation to nesting habits in European passerines: a test of Wallace’s hypothesis

Wallace proposed in 1868 that natural rather than sexual selection could explain the striking differences in avian plumage dichromatism. Thus, he predicted that nesting habits, through their association with nest predation, could drive changes in sexual dichromatism by enabling females in cavity nesters to become as conspicuous as males, whereas Darwin (1871, The Descent of Man and Selection in Relation to Sex, John Murray, London) argued that sexual selection was the sole explanation for dichromatism. Sexual dichromatism is currently used as indicating the strength of sexual selection, and therefore testing Wallace's claim with modern phylogentically controlled methodologies is of prime interest for comparing the roles of natural and sexual selection in affecting the evolution of avian coloration. Here, we have related information on nest attendance, sexual dichromatism and nesting habits (open and cavity nesting) to male and female plumage conspicuousness in European passerines. Nest incubation attendance does not explain male or female plumage conspicuousness but nest type does. Moreover, although females of monochromatic and cavity nesting species are more conspicuous than females of other species, males of monochromatic and open nesting species are those with more cryptic plumage. Finally, analyses of character evolution suggest that changes in nesting habits influence the probability of changes in both dichromatism and plumage conspicuousness of males but do not significantly affect those in females. These results strongly suggest a role of nesting habits in the evolution of plumage conspicuousness of males, and a role for sexual selection also in females, both factors affecting the evolution of sexual dichromatism. We discuss our findings in relation to the debate that Darwin and Wallace maintained more than one century ago on the importance of natural and sexual selection in driving the evolution of plumage conspicuousness and sexual dichromatism in birds, and conclude that our results partly support the evolutionary scenarios envisaged by both extraordinary scientists.

Food conditions affect yolk testosterone deposition but not incubation attendance

In many bird species with hatching asynchrony, yolk androgens increase across the laying sequence. This has been hypothesized to represent a compensatory mechanism for disadvantages of later-hatching chicks – via positive effects of yolk androgens on early competitiveness and growth. However, the costs and benefits of this compensatory strategy probably depend on environmental factors determining the survival chances of the chicks such as the food conditions, which should, therefore, influence maternal yolk androgen deposition.We studied the consequences of manipulated food conditions on the expected level of hatching asynchrony in canaries (Serinus canaria) assigning females to either a low (=LQ) or high quality (=HQ) diet. We measured the incubation behaviour (as incubation attendance) and the yolk androgen deposition in order to investigate whether and how females modulate hatching asynchrony in relation to the food conditions.Females on a HQ diet laid larger and heavier clutches, showed a stronger increase in yolk testosterone content towards the last-laid eggs, but did not alter their incubation attendance. Thus, females on a HQ diet seem to favour the survival of later hatching chicks, as indicated by their yolk testosterone deposition pattern. However, females on a HQ diet laid larger clutches and might need to compensate more in order to achieve a similar degree of hatching asynchrony than females on a LQ diet, given the lack of plasticity in incubation attendance. This suggests that canary females respond to food manipulations mainly via changes in clutch size rather than by altering the degree of hatching asynchrony.

Handicapped females receive more feedings during incubation from their mates: support for the female nutrition hypothesis

The female nutrition hypothesis posits that provisioning intensity of incubating females by their mates may depend on female needs and ensure proper incubation and a corresponding high hatching and breeding success of breeding pairs. Here, we have handicapped female pied flycatchers Ficedula hypoleuca at the beginning of incubation by clipping two primaries on each wing and filmed nests during incubation and later nestling provisioning to estimate male involvement in incubation feeding at the nest and in offspring care. Incubation feeding was more frequent at late nests. Correcting for this seasonal effect, incubation feeding was significantly affected by treatment and twice as high at experimental as at control nests. There was no effect of the experiment on female incubation attendance. The handicap did not result in any effect on hatching and breeding success, nestling growth and male or female provisioning and mass at the end of the nestling period. Males adjust their incubation feeding activity at the nest to female energetic requirements during incubation.

Behavioural effects of cadmium (Cd) in free-living willow ptarmigan (Lagopus lagopus)

The objective of this study was to examine effects of sublethal cadmium (Cd) contamination on parental behaviour and chick survival in willow ptarmigan (Lagopus lagopus) in an area that does not have high environmental levels of Cd due to geology but have high environmental levels of Cd due to anthropogenic deposition and acidification. One group of willow ptarmigan hens were exposed to Cd (83.2 mg Cd/L) after egg-laying using osmotic minipumps; whereas, another group given saline served as a control. No statistically significant difference in Cd concentration between the two groups was found. There was a significant correlation between the Cd concentrations in the liver and kidney of the birds and between Cd and metallothionein (MT) concentration in the kidney. No significant differences were found between the control hens and the Cd-contaminated hens with respect to any of the behaviour parameters. Age seemed to be more important than Cd treatment as adult hens showed more distraction display, had lower flushing distance, and higher brood attendance than juveniles. There were no difference in chick survival between Cd-contaminated and control hens, and there was no correlation between Cd concentrations and chick survival. The non-significant effects of Cd concentrations on parental behaviour may be due high environmental levels of Cd caused by anthropogenic deposition and acidification. We suggest that willow ptarmigan in the area have gradually evolved abilities to cope with high environmental levels of Cd, for instance, by binding Cd to MT in kidney and liver. This may suggest that Cd exerts a strong selection pressure on birds causing a relatively rapid evolution of protection mechanisms.

Ecological factors regulating brood attendance patterns of the Western Sandpiper Calidris mauri

Parental brood attendance patterns vary greatly among shorebird species. For monogamous calidridine species, biparental care with female‐first brood departure is most common. It is believed that adult sandpipers balance potential individual survival costs associated with extended parental care against the benefit gained by their brood of prolonged parental care. These costs and benefits are difficult to quantify and factors affecting the termination of parental brood attendance are unclear. We compared clutch size, nesting phenology, and parental attendance patterns of Western Sandpipers Calidris mauri at Nome and Kanaryarmiut, Alaska, sites separated by three degrees of latitude. The sites differed in breeding density and duration of breeding season, but the distribution of clutch sizes did not differ between sites or between nesting attempts. Parental attendance patterns were similar between sites, suggesting that parental attendance is a highly conserved life‐history trait in Western Sandpipers. Male Western Sandpipers attended broods longer than females, and the duration of parental attendance decreased at a similar rate for both sexes as the season progressed. Male and female Western Sandpipers undertake differential migrations to their non‐breeding grounds, with males typically settling at more northerly locations and females at more southerly sites, a migration pattern shared by certain other monogamous calidridine species. These same species exhibit similar parental brood attendance patterns, suggesting the strong role of overall migration distance in shaping the expression of parental attendance behaviours. A contrast of more geographically disjunct sites coupled with a better understanding of the migratory connectivity between Western Sandpiper breeding and non‐breeding populations would elucidate the role of cross‐seasonal effects on parental brood attendance decisions.

Solar incubation cuts down parental care in a burrow nesting tropical shorebird, the crab plover Dromas ardeola

We describe an unknown mode of solar‐assisted egg development in the crab plover Dromas ardeola, a shorebird that breeds in self‐excavated burrows. The insulating properties of the nest burrow and the intense solar radiation allowed egg development at near‐optimal temperature (35.2°C±0.2) and humidity (60.2%±4.4), allowing a very low incubation attendance by the parent birds (28.3% of time, with recesses lasting up to 58 h). Crab plovers did not abandon completely parental incubation, possibly because of the need to turn their egg, and because the slight warming provided by parents (0.8°C) may improve hatching. This is the first case of solar assisted incubation in a species unrelated to the Megapodiidae, the only birds known to develop their eggs without contact incubation.

The energetic costs of egg heating constrain incubation attendance but do not determine daily energy expenditure in the pectoral sandpiper

Heating eggs during incubation may be relatively energetically costly, affecting the outcome or number of breeding attempts. We determined the effect of reduced egg heating costs on nest attendance, change in body mass, and daily energy expenditure (DEE using the doubly labeled water technique) by heating nests of pectoral sandpipers. We also considered ground temperature, which may influence overall incubation costs, and mass reserves and stage of incubation, which may influence an individual’s ability to respond to changes in overall incubation cost. The total proportion of time spent in attending the eggs was significantly greater in nests that were experimentally heated (3.6% or 52 min daily), and this effect was significantly greater at low ground temperatures (14.7% or 211.7 min daily). Mass change was independent of experimental heating when controlling for attendance, although mass loss rate was greater for birds that attended more (for every 10% increase in daily proportion of attendance 0.12 extra grams of body mass were lost per hour), and overall daily attendance increased by 0.5% for every extra 1 g of body mass. DEE was greater for birds that had the higher rates of mass gain (for every 0.1 g of mass gained per hour, DEE increased by 20.5 kJ per day) but was independent of experimental heating when controlling for attendance. Overall, the results suggest that females are constrained from attending more by their energy reserve levels being depleted at least partly by the costs of egg heating, but these costs probably do not determine DEE, as costs off the nest may far exceed those incurred while sitting. Breeding in the arctic is clearly energetically demanding: pectoral sandpipers had an average DEE of 361.1 6 8.9 kjd � 1 , a mean power output of 4.1 W, equivalent to 6.1 times basal metabolic rate (n ¼ 24 birds). Key words: daily energy expenditure, incubation, nest attendance, pectoral sandpiper. [Behav Ecol 15:498–507 (2004)]