Avian parents that physically incubate their eggs must balance demands of self-maintenance with providing the proper thermal environment for egg development. Low incubation temperatures can lengthen ...the incubation period and produce changes in neonate phenotype that may influence subsequent survival and reproduction. We artificially incubated wood duck (Aix sponsa) eggs at three temperature regimes (low, 35.0; mid, 35.9; and high, 37.3°C) that are within the range of temperatures of naturally-incubated nests. We tested the effect of incubation temperature on duckling body composition, fledging success, the probability that females were recruited to the breeding population, and their subsequent reproductive success. Incubation period was inversely related to incubation temperature, and body mass and lipid mass for newly-hatched ducklings incubated at the lowest temperature were lower than for ducklings produced at higher temperatures. In 2008, ducklings (n = 412) were individually marked and broods (n = 38) containing ducklings from each temperature treatment were placed with wild foster mothers within 24 hrs of hatching. Ducklings incubated at the lowest temperature were less likely to fledge from nest sites than ducklings incubated at the higher temperatures. We recaptured female ducklings as adults when they were either prospecting for nest sites (n = 171; 2009-2011) or nesting (n = 527; 2009-2012). Female ducklings incubated at the lowest temperature were less likely to survive and be recruited to the breeding population than females incubated at higher temperatures. Reproductive success of surviving females also was greater for females that had been incubated at warmer temperatures. To our knowledge, this is the first avian study to link developmental conditions experienced by neonates during incubation with their survival and recruitment to the breeding population, and subsequent reproductive success. These results advance our understanding of incubation as an important reproductive cost in birds and support the potential significance of incubation in influencing the evolution of avian life histories.
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DOBA, IZUM, KILJ, NUK, PILJ, PNG, SAZU, SIK, UILJ, UKNU, UL, UM, UPUK
Incubation starts during egg laying for many bird species and causes developmental asynchrony within clutches. Faster development of late-laid eggs can help reduce developmental differences and ...synchronize hatching, which is important for precocial species whose young must leave the nest soon after hatching. In this study, we examined the effect of egg laying sequence on length of the incubation period in Wood Ducks (Aix sponsa). Because incubation temperature strongly influences embryonic development rates, we tested the interactive effects of laying sequence and incubation temperature on the ability of late-laid eggs to accelerate development and synchronize hatching. We also examined the potential cost of faster development on duckling body condition. Fresh eggs were collected and incubated at three biologically relevant temperatures (Low: 34.9°C, Medium: 35.8°C, and High: 37.6°C), and egg laying sequences from 1 to 12 were used. Length of the incubation period declined linearly as laying sequence advanced, but the relationship was strongest at medium temperatures followed by low temperatures and high temperatures. There was little support for including fresh egg mass in models of incubation period. Estimated differences in length of the incubation period between eggs 1 and 12 were 2.7 d, 1.2 d, and 0.7 d at medium, low and high temperatures, respectively. Only at intermediate incubation temperatures did development rates of late-laid eggs increase sufficiently to completely compensate for natural levels of developmental asynchrony that have been reported in Wood Duck clutches at the start of full incubation. Body condition of ducklings was strongly affected by fresh egg mass and incubation temperature but declined only slightly as laying sequence progressed. Our findings show that laying sequence and incubation temperature play important roles in helping to shape embryo development and hatching synchrony in a precocial bird.
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DOBA, IZUM, KILJ, NUK, PILJ, PNG, SAZU, SIK, UILJ, UKNU, UL, UM, UPUK
The wood duck (Aix sponsa) is a common and important cavity-nesting duck in North America; however, we know very little about how changes in vital rates influence population growth rate (λ). We used ...estimates of fertility and survival of female wood ducks from our nest-box studies in South Carolina, Alabama, and Georgia, USA, to create a stage-based matrix population model. We conducted perturbation analyses and ranked elasticity values to examine the relative importance of 17 component vital rates to λ. Female survival is influenced by nest success, so we recognized this female heterogeneity in our analyses. Four vital rates showed the greatest importance to λ. Analytic elasticities were greatest for breeding season and nonbreeding season survival of females that nested successfully, followed by nest success and female recruitment to the breeding population. Differences in female quality were important to λ. Next, we used process variation of vital rates and conducted life-stage simulation analyses (LSA) followed by variance decomposition to determine the amount of variation in λ explained by each vital rate. Female recruitment to the breeding population explained 57.7% of the variation in λ followed by nest success (11.4%), and breeding and nonbreeding season survival of females that nested successfully (9.3% and 9.4%, respectively). Together these 4 vital rates explained 88% of the variation in λ. Mean asymptotic population growth rate (λ = 0.80 ± 0.08 SD) from LSA revealed a declining population. Recruitment of females hatched from nest boxes was insufficient to sustain the nest-box population. However, including yearling (SY) females that were produced outside of nest boxes (i.e., immigrants) increased recruitment rates 1.5 to 2 times more than when only SY females recruited from nest boxes were included. Future research that examines how emigration and immigration interact with survival and reproduction to influence local population dynamics of wood ducks will be important for identifying the value of nest-box programs to wood duck conservation and management.
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BFBNIB, FZAB, GIS, IJS, KILJ, NLZOH, NMLJ, NUK, OILJ, PNG, SAZU, SBCE, SBMB, UL, UM, UPUK
•Stress-induced prolactin was positively related to incubation constancy.•Baseline and stress-induced prolactin were positively related to egg temperature.•These relationships were present only after ...a nest disturbance.•Corticosterone was not related to incubation constancy or temperature.
To maximize fitness, parents may trade-off time and energy between parental care and self-maintenance. In vertebrates, prolactin and corticosterone are two important hormones that regulate parental investment because they stimulate parental care and mobilize energy, respectively. Further, concentrations of both hormones change in response to disturbances. One of the most important parental care behaviors in birds is incubation, since small changes in egg temperature have large effects on offspring. We investigated how prolactin and corticosterone may mediate parental incubation constancy (i.e., the daily amount of time spent incubating eggs) and regulation of egg temperature. We collected blood samples from female wood ducks (Aix sponsa) near the start and end of the incubation period to measure baseline and stress-induced (30 min after capture and restraint) hormone concentrations. We also quantified incubation constancy and egg temperature using artificial egg temperature loggers. As expected, prolactin decreased and corticosterone increased after 30 min of capture and restraint. Corticosterone concentrations (baseline and stress-induced) were negatively related to body mass, but were not related to incubation constancy. In contrast, prolactin concentrations (baseline and stress-induced) were higher at the end than the start of the incubation period, and stress-induced prolactin concentrations were positively related to incubation constancy following a nest disturbance (i.e., capture). Further, prolactin (baseline and stress-induced) concentrations were positively related to egg temperatures, but only after the disturbance. These results suggest that prolactin may be associated with the regulation of parental incubation constancy and resulting heat-transfer after a disturbance, which may ultimately affect offspring development.
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GEOZS, IJS, IMTLJ, KILJ, KISLJ, NLZOH, NUK, OILJ, PNG, SAZU, SBCE, SBJE, UILJ, UL, UM, UPUK, ZAGLJ, ZRSKP
Elucidating factors that limit the number of offspring produced is fundamental to understanding life‐history evolution. Here, we examine the hypothesis that parental ability to maintain an optimal ...physical developmental environment for all offspring constrains clutch size via effects on offspring quality.
Experimental laboratory studies of birds have shown that a <1°C difference in average incubation temperature has diverse effects on fitness‐related post‐hatching offspring phenotypes. Thus, the inability of parents to maintain optimal incubation temperatures could constrain clutch sizes.
A fundamental question that has not been sufficiently addressed is whether larger clutch sizes lead to within nest variation in egg temperature that is large enough to produce offspring with different phenotypes within a brood. This could lead to differential survival among offspring, and could create a trade‐off between offspring number and quality.
We manipulated clutch size in nests of free‐living wood ducks and measured incubation temperature among and within clutches using multiple temperature loggers.
As clutch size increased, average incubation temperatures were lower and more variable, and eggs took longer to hatch. Notably, the range in average incubation temperature among eggs within nests increased with clutch size and exceeded 1°C in large clutches. Clutch size did not affect hatch success.
In conjunction with our companion laboratory studies that used artificial incubation to document the effects of temperature variation on fitness‐related traits in this species, our work suggests that suboptimal incubation temperatures could be a factor that limits clutch size through diminishing returns on post‐hatch offspring quality.
A free plain language summary can be found within the Supporting Information of this article.
A free plain language summary can be found within the Supporting Information of this article
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BFBNIB, FZAB, GIS, IJS, KILJ, NLZOH, NUK, OILJ, SAZU, SBCE, SBMB, UL, UM, UPUK
Optimal development of avian embryos occurs within a narrow range of incubation temperatures. Most parents that physically incubate their eggs through direct contact are challenged to balance their ...time on the nest with taking foraging recesses to satisfy their energetic requirements. To explore the costs and investment strategies of incubating female Wood Ducks (Aix sponsa), we manipulated the microclimate of nests by reducing down insulation from the typical 4.0 g to 0.5 g. Cooling rates of clutches during morning recesses increased when down insulation was reduced, especially at low ambient temperatures. Females with reduced down responded to increased cooling rates by shortening morning recesses and increasing daily incubation constancy, and these behavioral changes were independent of their body mass at the start of incubation. Females in both treatment groups responded similarly to changes in ambient temperature and spent less time incubating as ambient temperatures increased. Clutch temperatures at the end of morning recesses were similar for females with reduced and normal insulation. Average clutch temperatures for the full incubation period did not differ between treatments, and, correspondingly, there were no differences in length of the incubation period, hatching success, or duckling phenotype. Our results show that female Wood Ducks were sensitive to changes in both clutch temperature and ambient temperature and that they modified their time on the nest to provide developing eggs with an optimal thermal environment without negatively affecting their body mass at the end of incubation. Further examination of the limits of behavioral plasticity in incubating birds will be essential, particularly in light of future challenges presented by climate change.
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BFBNIB, NMLJ, NUK, PNG, SAZU, UL, UM, UPUK
The wood duck (Aix sponsa) is a common and important cavity-nesting duck in North America; however, we know very little about how changes in vital rates influence population growth rate (λ). We used ...estimates of fertility and survival of female wood ducks from our nest-box studies in South Carolina, Alabama, and Georgia, USA, to create a stage-based matrix population model. We conducted perturbation analyses and ranked elasticity values to examine the relative importance of 17 component vital rates to λ. Female survival is influenced by nest success, so we recognized this female heterogeneity in our analyses. Four vital rates showed the greatest importance to λ. Analytic elasticities were greatest for breeding season and nonbreeding season survival of females that nested successfully, followed by nest success and female recruitment to the breeding population. Differences in female quality were important to λ. Next, we used process variation of vital rates and conducted life-stage simulation analyses (LSA) followed by variance decomposition to determine the amount of variation in λ explained by each vital rate. Female recruitment to the breeding population explained 57.7% of the variation in λ followed by nest success (11.4%), and breeding and nonbreeding season survival of females that nested successfully (9.3% and 9.4%, respectively). Together these 4 vital rates explained 88% of the variation in λ. Mean asymptotic population growth rate (λ = 0.80 ± 0.08 SD) from LSA revealed a declining population. Recruitment of females hatched from nest boxes was insufficient to sustain the nest-box population. Furthermore, including yearling (SY) females that were produced outside of nest boxes (i.e., immigrants) increased recruitment rates 1.5 to 2 times more than when only SY females recruited from nest boxes were included. Future research that examines how emigration and immigration interact with survival and reproduction to influence local population dynamics of wood ducks will be important for identifying the value of nest-box programs to wood duck conservation and management.
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BFBNIB, FZAB, GIS, IJS, KILJ, NLZOH, NMLJ, NUK, OILJ, PNG, SAZU, SBCE, SBMB, UL, UM, UPUK
The environment that animals experience during development shapes phenotypic expression. In birds, two important aspects of the early‐developmental environment are lay‐order sequence and incubation. ...Later‐laid eggs tend to produce weaker offspring, sometimes with compensatory mechanisms to accelerate their growth rate to catch‐up to their siblings. Further, small decreases in incubation temperature slow down embryonic growth rates and lead to wide‐ranging negative effects on many posthatch traits. Recently, telomeres, noncoding DNA sequences at the end of chromosomes, have been recognized as a potential proxy for fitness because longer telomeres are positively related to lifespan and individual quality in many animals, including birds. Although telomeres appear to be mechanistically linked to growth rate, little is known about how incubation temperature and lay‐order may influence telomere length. We incubated wood duck (Aix sponsa) eggs at two ecologically‐relevant temperatures (34.9°C and 36.2°C) and measured telomere length at hatch and 1 week after. We found that ducklings incubated at the lower temperature had longer telomeres than those incubated at the higher temperature both at hatch and 1 week later. Further, we found that later‐laid eggs produced ducklings with shorter telomeres than those laid early in the lay‐sequence, although lay‐order was not related to embryonic developmental rate. This study contributes to our broader understanding of how parental effects can affect telomere length early in life. More work is needed to determine if these effects on telomere length persist until adulthood, and if they are associated with effects on fitness in this precocial species.
Wood duck eggs incubated at a slightly warmer temperature and laid later in the lay‐sequence produced ducklings with shorter telomeres at hatch and 1 week later, compared to those incubated at a colder temperature or laid earlier in the sequence.
Research highlights
Wood duck eggs incubated at a slightly warmer temperature and laid later in the lay‐sequence produced ducklings with shorter telomeres at hatch and 1 week later, compared to those incubated at a colder temperature or laid earlier in the sequence.
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FZAB, GIS, IJS, KILJ, NLZOH, NUK, OILJ, SAZU, SBCE, SBMB, UL, UM, UPUK
Abstract
The environments that animals experience during development have important fitness consequences. In birds, parents influence the developmental environment of their offspring through ...incubation. Subtle changes in incubation temperature affect offspring morphology and physiology, such as growth, immune function, and thermoregulation, yet little is known about how it may affect critical early-life behaviors. Because expression of behavior can be influenced by the social environment, the effect of incubation temperature on behavior may be context-dependent. We investigated whether incubation temperature and social context influence a critical early-life task in wood ducks (Aix sponsa). Wood ducks nest in tree cavities and, shortly after hatching, ducklings must jump and climb out of the cavity. Failure to exit the nest is fatal. In 2 experiments, we incubated eggs at different mean temperatures and examined the nest exodus of ducklings individually and in mixed-incubation temperature pairs. When tested individually, ducklings incubated at 35.8 °C and 37.0 °C were ~2.5 times more successful at exiting the nest, and jumped and climbed more often, than those incubated at 35.0 °C. However, in an experiment conducted the following year, we found that social interactions mitigated these effects and there was no difference in nest exodus success when ducklings incubated at 35.0 °C and 36.0 °C were tested together in pairs. This may be because, when in pairs, ducklings incubated at the low-temperature experience social enhancement whereas those incubated at the high temperature maintain similar behaviors. These results advance our understanding of how parental effects influence offspring behaviors and performance within different social contexts.
Poor conditions during development lead to suboptimal offspring behaviors, but this can be mitigated by social interactions. Shortly after wood duck ducklings hatch, they must jump out of their nest in a tree cavity. Failure to exit is fatal. Ducklings that hatch from eggs incubated at low temperatures are less successful at exiting the nest than those incubated at higher temperatures when tested individually, but success is similar when ducklings experience social interactions in pairs.
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