Laying sequence interacts with incubation temperature to influence rate of embryonic development and hatching synchrony in a precocial bird.

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.

Ecological, evolutionary, and conservation implications of incubation temperature-dependent phenotypes in birds.

Incubation is a vital component of reproduction and parental care in birds. Maintaining temperatures within a narrow range is necessary for embryonic development and hatching of young, and exposure to both high and low temperatures can be lethal to embryos. Although it is widely recognized that temperature is important for hatching success, little is known about how variation in incubation temperature influences the post-hatching phenotypes of avian offspring. However, among reptiles it is well known that incubation temperature affects many phenotypic traits of offspring with implications for their future survival and reproduction. Although most birds, unlike reptiles, physically incubate their eggs, and thus behaviourally control nest temperatures, variation in temperature that influences embryonic development still occurs among nests within a population. Recent research in birds has primarily been limited to populations of megapodes and waterfowl; in each group, incubation temperature has substantial effects on hatchling phenotypic traits important for future development, survival, and reproduction. Such observations suggest that incubation temperature (and incubation behaviours of parents) is an important but underappreciated parental effect in birds and may represent a selective force instrumental in shaping avian reproductive ecology and life-history traits. However, much more research is needed to understand how pervasive phenotypic effects of incubation temperature are among birds, the sources of variation in incubation temperature, and how effects on phenotype arise. Such insights will not only provide foundational information regarding avian evolution and ecology, but also contribute to avian conservation.

Energetic constraints and parental care: is corticosterone indicative of energetic costs of incubation in a precocial bird?

Suppression of the adrenocortical response (e.g., corticosterone release) to an acute stressor is a physiological adjustment thought to decrease the likelihood of avian parents abandoning their nests. However, some periods of parental care, like incubation, are energetically costly, thus corticosterone could increase during these stages to allow incubating parents to utilize energy reserves. Wood ducks (Aix sponsa) have ~30 day incubation periods and only the female incubates the eggs. We hypothesized that corticosterone would be important in regulating energy availability during incubation in this species. Because resources invested in reproduction increase with clutch size, we also hypothesized that clutch size would influence plasma corticosterone during incubation. We measured baseline and stress-induced corticosterone in incubating females during early and late stages of incubation. At both stages of incubation all hens had low baseline corticosterone levels. However, we found that stress-induced corticosterone was 105% greater late in incubation than early in incubation. We also detected a significant negative correlation between female body mass and stress-induced corticosterone late in incubation, but not during the early stages of incubation. Furthermore, we found a significant positive relationship between stress-induced corticosterone and clutch size. These lines of evidence support the hypothesis that incubation in wood ducks is energetically costly and corticosterone is important in supporting the energetic demands of incubating hens. Our findings suggest that corticosterone's role in supporting parental care behaviors are dynamic and are influenced by several factors and that there is a greater physiological cost associated with incubating larger clutches.

Warm is better: incubation temperature influences apparent survival and recruitment of wood ducks (Aix sponsa).

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.

Incubation temperature affects multiple measures of immunocompetence in young wood ducks (Aix Sponsa).

Parental effects play a vital role in shaping offspring phenotype. In birds, incubation behaviour is a critical parental effect because it influences the early developmental environment and can therefore have lifelong consequences for offspring phenotype. Recent studies that manipulated incubation temperature found effects on hatchling body composition, condition and growth, suggesting that incubation temperature could also affect energetically costly physiological processes of young birds that are important to survival (e.g. immune responses). We artificially incubated wood duck (Aix sponsa) eggs at three biologically relevant temperatures. Following incubation, we used two immunoassays to measure acquired immune responses of ducklings. Ducklings incubated at the lowest temperature had reduced growth, body condition and responses to both of our immune challenges, compared with those from the higher temperatures. Our results show that incubation temperatures can be an important driver of phenotypic variation in avian populations.