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The temperature experienced by avian embryos during development has important impacts on their growth and post‐hatching phenotypes. Ambient temperature can directly affect avian nest temperature and indirectly affect it through its impact on parental incubation behaviours. Because the nests of uniparental incubators are usually left unattended more frequently than the nests of biparental incubators whose nests can be attended by another bird when one bird leaves the nest, we predict that the effect of ambient temperature on nest temperature and thus on embryonic development (specifically, incubation period length and hatching success) and post‐hatching phenotype (i.e. potential carry‐over effect on nestling body mass and condition) should be greater in uniparental incubators than in biparental incubators. To test this prediction, we studied two congeneric species, the biparental incubating black‐throated tits Aegithalos concinnus and the uniparental incubating silver‐throated tits A.glaucogularis. We found that although the embryos of the two species both developed faster (shorter incubation period length) when ambient temperature was higher, the slope was significantly greater for silver‐throated tits than for black‐throated tits, consistent with our prediction of a greater effect of ambient temperature on embryonic development in silver‐throated tits. However, the result was not due to a greater effect of ambient temperature on nest temperature, because nest temperatures of the two species had positive relationships with ambient temperature in a similar way. Therefore, it implies a greater response of silver‐throated tit embryos to temperature change. In addition, ambient temperature during the incubation stage did not affect either hatching success or nestling body mass and condition in both species. Although our predictions were not fully supported, our findings highlight the different responses of embryonic development to environmental changes between a uniparental incubator and a biparental incubator, and suggest further research to explore the mechanisms. [ABSTRACT FROM AUTHOR]

The temperature experienced by avian embryos during development has important impacts on their growth and post‐hatching phenotypes. Ambient temperature can directly affect avian nest temperature and indirectly affect it through its impact on parental incubation behaviours. Because the nests of uniparental incubators are usually left unattended more frequently than the nests of biparental incubators whose nests can be attended by another bird when one bird leaves the nest, we predict that the effect of ambient temperature on nest temperature and thus on embryonic development (specifically, incubation period length and hatching success) and post‐hatching phenotype (i.e. potential carry‐over effect on nestling body mass and condition) should be greater in uniparental incubators than in biparental incubators. To test this prediction, we studied two congeneric species, the biparental incubating black‐throated tits Aegithalos concinnus and the uniparental incubating silver‐throated tits A. glaucogularis. We found that although the embryos of the two species both developed faster (shorter incubation period length) when ambient temperature was higher, the slope was significantly greater for silver‐throated tits than for black‐throated tits, consistent with our prediction of a greater effect of ambient temperature on embryonic development in silver‐throated tits. However, the result was not due to a greater effect of ambient temperature on nest temperature, because nest temperatures of the two species had positive relationships with ambient temperature in a similar way. Therefore, it implies a greater response of silver‐throated tit embryos to temperature change. In addition, ambient temperature during the incubation stage did not affect either hatching success or nestling body mass and condition in both species. Although our predictions were not fully supported, our findings highlight the different responses of embryonic development to environmental changes between a uniparental incubator and a biparental incubator, and suggest further research to explore the mechanisms.

Constant nest attentiveness, which benefits avian embryos, and body maintenance activities (e.g., foraging trips) by incubating parents are mutually exclusive activities. To balance these, parents must schedule the time spent on and off their nests according to a wide range of environmental conditions that change the costs and benefits of breaking or continuing an incubation bout. Here, we used data from 38 camera-monitored nests over two breeding seasons on Robben Island, Southern Africa to examine how African Black Oystercatchers Haematopus moquini, a biparental incubator, varied incubation bout length in relation to environmental variables that would have affected energy/water budgets: tide phase, air temperature, windspeed and time of day (day versus night). We predicted that incubation bout duration would decrease at high temperatures and wind speeds when energetic and water needs increase and increase at high tides when foraging opportunities decrease. Oystercatchers' overall nest attentiveness was high at 95%. Incubation bout length was longer at high tide and at night than at low tide and during the day. Incubation bout length decreased with increases in temperature but had a non-linear relationship with windspeed that varied by the time of day: it increased sharply with windspeed and plateaued at high windspeed during the day, but largely declined with moderate to high windspeeds at night. These results suggest that oystercatchers' energetic, thermoregulatory, and hydration needs may have increased with high heat loads during the day but decreased at night and with increased windspeed during the day, and oystercatchers responded by varying incubation bout lengths accordingly. The proximity of their nests to the intertidal zone, which means short travel time and quick access to food, water, and thermoregulatory opportunities, seems to be of vital importance to their capacity to maintain high nest attentiveness and deal with increasing heat loads. [ABSTRACT FROM AUTHOR]

Abstract Background Effective communication between sexual partners is essential for successful reproduction. Avian parents with biparental incubation need to know how to negotiate, when and who will incubate, and how to harmonize partner exchange at the nest. Although considerable effort has been dedicated to studies of incubation rhythms, few studies have investigated how behavioural signals serve to tighten cooperation between parents. Moreover, existing studies are almost exclusively restricted to species in which long distances between incubating and non-incubating parents prevent continuous communication during incubation. Thus, the most frequently described parental exchange system is a simple model characterized by the return of the non-incubating parent to the nest itself. Here, we propose more complex parental exchange behaviour in the Northern Lapwing (Vanellus vanellus), a territorial species capable of continuous partner communication during incubation and with a highly variable male contribution to incubation. Results Northern Lapwing females regularly vocalized shortly before departing from the nest, while males mostly left the nest quietly. Responsiveness of the male to female vocalization, perhaps in combination with her flying away from the nest, helped to synchronize incubation care by increasing the probability of exchange, and also by shortening the exchange gaps. In contrast, a male-to-female exchange gap most often occurred after the male quietly flew away from the nest. The frequency of female vocal signalling was not correlated with the male incubation effort on a between-nest scale, but the highest probability of a female-to-male exchange occurred after vocal signalling by females with the most nest-attentive males. Conversely, lowered effort by females to vocalize in the night was accompanied by lower willingness of males to incubate. Conclusions Our results suggest that (1) that the incubating parent can communicate with the non-incubating partner using sex-specific behavioural signals, and this helps to synchronize parental exchange on the nest, (2) this signalling may combine acoustic and visual cues, and (3) the efficiency of this signalling might influence the overall nest attendance. The presumption that the repertoire of behavioural signals during reproduction will be much more complex in territorial species that are capable of continuous communication between the partners during the incubation period should be further tested.

Parents make tradeoffs between care for offspring and themselves. Such a tradeoff should be reduced in biparental species, when both parents provide parental care. However, in some biparental species, the contribution of one sex varies greatly over time or between pairs. How this variation in parental care influences self-maintenance rhythms is often unclear. In this study, we used continuous video recording to investigate the daily rhythms of sleep and feather preening in incubating females of the Northern Lapwing (Vanellus vanellus), a wader with a highly variable male contribution to incubation. We found that the female's sleep frequency peaked after sunrise and before sunset but was low in the middle of the day and especially during the night. In contrast, preening frequency followed a 24-h rhythm and peaked in the middle of the day. Taken together, incubating females rarely slept or preened during the night, when the predation pressure was highest. Moreover, the sleeping and preening rhythms were modulated by the male contribution to incubation. Females that were paired with more contributing males showed a stronger sleep rhythm but also a weaker preening rhythm. If more incubating males also invest more in nest guarding and deterring daylight predators, their females may afford more sleep on the nest during the day and preen more when they are off the nest. Whether the lack of sleep in females paired with less caregiving males has fitness consequences awaits future investigation. [ABSTRACT FROM AUTHOR]

Theoretical models predict that parents feeding offspring should partially compensate for the reduced care of their partner. However, for incubating birds, the level of compensation may depend on how reduced care changes the risk of entire brood failure, for example due to clutch predation, and on individual variation in the timing of depletion of energy stores. Although biparental incubation dominates in non-passerines, short-term manipulations of care during incubation are scarce. Here, we describe the response of 25 semipalmated sandpipers (Calidris pusilla) to an unexpected ~12-h absence (experimental removal) of their partner in the middle of the 21-day incubation period. During the period when the removed partner would have taken over to start its regular ~12-h incubation bout, parents compensated partially for the absence of their partner's care (mean: 59%, 95%CI: 49–70%). However, individuals varied in their response from no to full compensation, independent of parental sex. In contrast to incubation in undisturbed nests or by uniparental species, nest attendance of compensating parents tended to be higher during the warmer part of the day. Whereas compensation was unrelated to before-experimental share of incubation, parents that left the nest from a further distance upon human approach (more aware of or more “responsive” to their environment) compensated more. The quality of incubation in the after-experimental period, i.e., after return of the partner, was lower than usual, but improved quickly over time. In seven nests where the removed parent never returned, the widowed partner attended the nest for 0–10 days (median: 4), which suggests that widowed semipalmated sandpipers can adjust their incubation behavior to that observed in uniparental incubators. To conclude, our results indicate that biparental incubators are willing to tolerate a missed or irregular incubation bout of their partner. We speculate that all individuals would compensate fully, but that some fail because they deplete their energy stores, while others may be less responsive to or initially unaware of the absence of their partner.

Incubation is energetically demanding, but it is debated whether these demands constrain incubation-scheduling (i.e., the length, constancy, and timing of incubation bouts) in cases where both parents incubate. Using 2 methods, we experimentally reduced the energetic demands of incubation in the semipalmated sandpiper, a biparental shorebird breeding in the harsh conditions of the high Arctic. First, we decreased the demands of incubation for 1 parent only by exchanging 1 of the 4 eggs for an artificial egg that heated up when the focal bird incubated. Second, we reanalyzed the data from the only published experimental study that has explicitly tested energetic constraints on incubation-scheduling in a biparentally incubating species (Cresswell et al. 2003). In this experiment, the energetic demands of incubation were decreased for both parents by insulating the nest cup. We expected that the treated birds, in both experiments, would change the length of their incubation bouts, if biparental incubation-scheduling is energetically constrained. However, we found no evidence that heating or insulation of the nest affected the length of incubation bouts: the combined effect of both experiments was an increase in bout length of 3.6 min (95% CI: -33 to 40), which is equivalent to a 0.5% increase in the length of the average incubation bout. These results demonstrate that the observed biparental incubation-scheduling in semipalmated sandpipers is not primarily driven by energetic constraints and therefore by the state of the incubating bird, implying that we still do not understand the factors driving biparental incubation-scheduling. [ABSTRACT FROM AUTHOR]

Both nest survival and incubation behaviour are highly variable among shorebirds (Charadrii), and we tested whether more conspicuous incubation behaviour increased the risk of nest predation. During 2000–2006, we monitored nest fate at 901 shorebird nests at three study sites across the circumpolar Arctic. Using miniature video recorders and nest temperature sensors, we obtained 782 days of behavioural data for 161 nests of 11 species. We related nest fate to the rate and duration of adults'' nest absences or restless movements on the nest, as well as the total proportion of each day that adult birds engaged in these activities. Nest predation was positively related to the proportion of time that each species left the nest unattended. After controlling for species effects, the likelihood of a successful nesting attempt was lower for individuals that spent more time off the nest, but among failed nests, the number of days that a nest survived prior to depredation was not significantly predicted by measures of incubation behaviour. To control for weather or seasonal effects, we paired observations from nests that were ultimately depredated with observations from successful nests of the same species on the same day. In this paired sample (dominated by two species: red phalaropes, Phalaropus fulicarius, and little stints, Calidris minuta), both incubation recesses and restless movements were more numerous among failed versus successful nests. Our results suggest that more conspicuous incubation behaviour is indeed related to a higher risk of nest predation, and that this relationship may underlie patterns of nest survival within and among shorebird species. [Copyright &y& Elsevier]