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.

Summary 1.Maternal effects often explain a significant amount of variation in offspring phenotype, and can be important in the evolution of life histories. Incubation of eggs is an important maternal effect, and optimal growth and development of avian embryos takes place within a narrow range of incubation temperatures, but the effect of incubation microclimate on neonate phenotype remains relatively unexplored in birds. 2. In this study of Wood Ducks ( Aix sponsa Linnaeus) we examined effects of incubation temperature on the length of incubation period and neonate quality. We monitored nest temperatures and incubation periods of naturally incubated Wood Duck nests and found a strong inverse relationship between incubation period and average nest temperature. 3. Next, we collected three unincubated eggs from each of 48 nests, and randomly assigned eggs from each nest to one of three incubation temperatures (34·6, 36·0 and 37·4 ° C). Experimental incubation temperatures overlapped average nest temperatures of naturally incubated Wood Duck nests. 4. Hatching success varied with incubation temperature and was lowest for eggs incubated at the lowest temperature. Incubation period of experimental eggs decreased with increasing temperature but was not affected by fresh egg mass. 5. Wood Duck embryos catabolized an estimated 34-38% of egg lipids and 25-33% of egg protein during incubation. Percentage change of lipids increased with decreasing incubation temperature, but not significantly. Embryos incubated at lower temperatures used a greater percentage of protein than embryos incubated at higher temperatures. 6. In analyses using fresh egg mass as the covariate, we found that wet and dry mass of ducklings increased with increasing incubation temperature. Decreases in lipid content of Wood Duck neonates with decreasing incubation temperature were not significant, but eggs incubated at low temperatures produced ducklings that had reduced protein mass and that were structurally larger than ducklings from eggs incubated at high temperatures. 7. Our study illustrates the importance of incubation temperature on the development of Wood Duck embryos. Decisions made by incubating parents that influence egg temperature can modify incubation period and offspring phenotype. Investigations of incubation as a reproductive cost should consider how parental decisions influence both parents and offspring.

SUMMARYEarly developmental experiences, such as incubation conditions, can have important consequences for post-hatching fitness in birds. Although the effects of incubation temperature on phenotype of avian hatchlings are poorly understood, recent research suggests that subtle changes in incubation conditions can influence hatchling characteristics, including body size and condition. We designed an experiment to explore the effects of incubation temperature on hatching success, survival to 9 days post hatch, growth and the hypothalamo-pituitary-adrenal (HPA) axis in wood ducks (Aix sponsa). Wood duck eggs were collected from nest boxes and experimentally incubated at three temperatures (35.0, 35.9 and 37.0°C), each falling within the range of temperatures of naturally incubated wood duck nests. Survival and growth were monitored in ducklings fed ad libitum for 9 days post hatch. In addition, baseline and stress-induced plasma corticosterone concentrations were measured in 2 and 9 day old ducklings. Hatching success and survival to 9 days was greatest in ducks incubated at the intermediate temperature. Ducklings incubated at 35.9°C and 37.0°C had 43% higher growth rates than ducklings incubated at 35.0°C. In addition, ducklings incubated at 35.0°C had higher baseline (17-50%) and stress-induced (32-84%) corticosterone concentrations than ducklings incubated at 35.9°C and 37.0°C at 2 and 9 days post hatch. We also found a significant negative correlation between body size and plasma corticosterone concentrations (baseline and stress-induced) in 9 day old ducklings. To our knowledge, this is the first study to demonstrate that thermal conditions experienced during embryonic development can influence the HPA axis of young birds. Our results illustrate that subtle changes (<1.0°C) in the incubation environment can have important consequences for physiological traits important to fitness.

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.

Summary1. The developmental environment plays a key role in determining offspring phenotype, and the parents' behaviour and physiology often dictates developmental conditions. Despite the plethora of studies documenting the importance of incubation temperature on offspring phenotype in reptiles, very few studies have examined such relationships in birds. 2. Because nearly all birds physically incubate their eggs, altering the nest environment may be an important but previously overlooked way parents can influence their offspring's phenotype. Here, we tested the hypothesis that incubation temperature would affect thermoregulation in wood duck (Aix sponsa) hatchlings. 3. We show that a reduction in <1°C in incubation temperature affects the metabolic costs of thermoregulation in offspring of a non-domesticated bird, resulting in 27-40% greater increases in oxygen consumption of ducklings incubated at the lowest temperature relative to ducklings incubated at higher temperatures.4. Because we demonstrate that incubation temperature affects hatchling phenotypic quality, our findings provide novel support for newly proposed frameworks that highlight the importance of incubation temperature to the evolution of clutch size in birds.

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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