Parental Incubation Research Articles

The effect of ambient temperature on bird embryonic development: a comparison between uniparental incubating silver‐throated tits and biparental incubating black‐throated tits

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.

The clutch size, incubation behavior of Reeves's Pheasant (Syrmaticus reevesii) and their responses to ambient temperature and precipitation

Weather conditions play a pivotal role in embryo development and parental incubation costs, potentially impacting the clutch size and incubation behavior of birds. Understanding these effects is crucial for bird conservation. Reeves's Pheasant (Syrmaticus reevesii) is a threatened species endemic to China, which is characterized by female-only incubation. However, there is a lack of information regarding the impact of weather conditions on clutch size and incubation behavior in this species. Using satellite tracking, we tracked 27 wild female Reeves's Pheasants from 2020 to 2023 in Hubei Province, China. We explored their clutch size and incubation behavior, as well as their responses to ambient temperature and precipitation. Clutch size averaged 7.75 ± 1.36, had an association with average ambient temperature and average daily precipitation during the egg-laying period, and was potentially linked to female breeding attempts. Throughout the incubation period, females took an average of 0.73 ± 0.46 recesses every 24 h, with an average recess duration of 100.80 ± 73.37 min and an average nest attendance of 92.98 ± 5.27%. They showed a unimodal recess pattern in which nest departures peaked primarily between 13:00 and 16:00. Furthermore, females rarely left nests when daily precipitation was high. Recess duration and nest attendance were influenced by the interaction between daily mean ambient temperature and daily precipitation, as well as day of incubation. Additionally, there was a positive correlation between clutch size and recess duration. These results contribute valuable insights into the life-history features of this endangered species.

Dabbling Duck Eggs Hatch after Nest Abandonment in the Wild

In most birds, parental incubation of eggs is necessary for embryo development and survival. Using a combination of weekly nest visits, temperature dataloggers, infrared video cameras, and GPS tracking of hens, we documented several instances of duck eggs hatching after being abandoned by the incubating female. Of 2826 Mallard (Anas platyrhynchos) and Gadwall (Mareca strepera) nests monitored 2015–2019 in Suisun Marsh, California, 48 (1.7%) were abandoned during late incubation (≥ 20 days). Of these, we identified six (12.5%) where at least one egg hatched 2–9 days after abandonment. In all six cases, eggshell membranes were found in the nest (indicating hatch), and ducklings were observed at three nests. Abandoned nests were unattended for an average of 5.9 days before eggs hatched; during this time, mean nest temperatures (23.6°C–29.0°C) were substantially lower than before nest abandonment (31.7°C–36.4°C). We estimated that abandonment resulted in a 9% longer time period between clutch completion and hatch (0–4 days longer) and a lower rate of egg hatching success (36%). Our results provide evidence that some older embryos (≥ 20 days) in mild climates can survive without parental incubation for several days and continue to develop (at a reduced rate) to the point of successfully hatching.

Parenting in the city: effects of urbanization on incubation behaviour and egg temperature in great tits, Parus major

With 68% of the world's population expected to live in cities by 2050, it is crucial to understand how animals respond to urbanization. Urban areas are associated with changes in microclimate, light, noise and human activity, which can affect animal physiology, behaviour and fitness. In particular, urbanization may affect how parents allocate time and energy between parental care and self-maintenance. In birds, a critical aspect of parental care is incubation, where parents must maintain egg temperatures for successful offspring development. However, incubation is energetically costly, and anthropogenic disturbances may affect parental decisions related to this cost. Determining whether urbanization affects incubation is crucial for understanding the differences between urban and nonurban populations in their reproductive performance and fitness. We measured incubation behaviour and egg temperature in one urban and one forest population of great tits, a uniparental incubator, using temperature loggers. We found that, compared to forest females, urban females spent more time incubating, took shorter off-bouts and ended their last daily off-bouts at a later hour. Furthermore, time spent off the nest, the number and duration of off-bouts and daily start of activity were related to ambient temperature; start of activity was related to clutch size; and time spent off the nest was related to the day of incubation. Despite urban females spending more time on the nest, eggs of both forest and urban females experienced relatively similar average egg temperatures, and urban eggs tended to experience greater variability in egg temperature. Furthermore, there were no differences between sites for any fitness-related offspring variable. Our results suggest that urbanization can influence the thermal environment of developing avian offspring by causing urban adults to be inefficient incubators through increased activity off the nest in the evening, differences in nest insulation and/or suboptimal incubation behaviour compared to forest counterparts. • Urbanization appears to influence parental incubation behaviour of great tits. • Urban birds spent more time warming their eggs compared to birds in the forest. • However, outcomes for offspring were similar. • Urban eggs and forest eggs experienced similar temperatures.

Light received by embryos promotes postnatal junior phenotypes in a seabird

Abstract Light is a salient and variable ecological factor that can impact developmental trajectories of vertebrate embryos, yet whether prenatal light environment can act as an anticipatory cue preparing organisms to cope with postnatal conditions is still unclear. In asynchronous birds, last-laid eggs are particularly exposed to sunlight as parental incubation behavior becomes intermittent after the hatching of senior chicks. Here, we explore whether natural variations in prenatal light exposure shape the distinctive phenotype showed by last-hatched chicks of a semi-precocial seabird, the yellow-legged gull (Larus michahellis), potentially preparing them to cope with the postnatal competitive context. To do this, we manipulated the amount of light received by last-laid eggs (within a natural range) during last stages of embryonic development. Prenatal exposure to light cues promoted the development of the resilient “junior phenotype” exhibited by last-hatched gull chicks, characterized by accelerated hatching, increased begging behavior and a slower growth rate. These developmental and behavioral adjustments were accompanied by down-regulation of genes involved in metabolism and development regulation (SOD2 and TRalpha), as well as changes in the HPA-axis functioning (lower baseline corticosterone and robust adrenocortical response). Junior chicks exposed to light cues during the embryonic development showed longer telomeres during the early postnatal period, suggesting that light-induced adjustments could allow them to buffer the competitive disadvantages associated with hatching asynchrony. Our study provides evidence that postnatal junior phenotypes are, at least in part, prenatally shaped by light cues that act during a critical temporal window of developmental sensitivity.

An analytical study on the effect of weather changes on birds in a machine learning perspective

Building applications that make use of the speed and specificity of algorithm development can significantly help attempts to quantify and analyze variations in bird behavior in response to weather changes in the age of machine learning. For both endotherms and ectotherms, temperature can affect everything from daily energy budgets to nesting behaviors. In birds, environmental temperature plays a key role in shaping parental incubation behavior and temperatures experienced by embryos. One of the main mechanisms used to mitigate the impacts of climate change on these species in the near future is to assess the current system of protected areas. It is necessary to ensure that these areas will continue being effective in conserving these species even under climate change. The purpose of this paper was to review the challenges faced by birds due to climate change, how it affects their growth, migration, breeding, etc. and how machine learning models can have considerable usage in helping to find out the problems affected by birds due to climate change.

Blue tit Cyanistes caeruleus males increase their reproductive effort when subject to a flea experimental manipulation

Parasites exert a strong selection pressure on their hosts as manifested in behavioural antiparasite traits to reduce negative impacts on fitness. The numerous nest‐dwelling ecto‐parasites residing in avian nests make altricial birds excellent model‐systems for investigating the relationship between parasites and their hosts. Here, we experimentally increased natural levels of hen fleas Ceratophyllus gallinae in blue tit Cyanistes caeruleus nests during incubation, and tested their effects on parental incubation behaviours and reproductive performance. Our experimental addition of fleas resulted in an increase in feeding effort of males to incubating females. Frequency of male feedings was also positively related to clutch size. These results suggest that males increase their reproductive effort in flea manipulated and large broods. This will, at least partly, compensate female costs in nests with high ecto‐parasite density and many nestlings. Furthermore, nestling mass at day six in experimental nests decreased with brood size, which was not the case in nests with a natural level of fleas. In line with male incubation feeding, parents may try to compensate for the costs inflicted by the fleas but can only partly compensate when brood size is large.

Applications of machine learning in behavioral ecology: Quantifying avian incubation behavior and nest conditions in relation to environmental temperature.

In the age of machine learning, building programs that take advantage of the speed and specificity of algorithm development can greatly aid efforts to quantify and interpret changes in animal behavior in response to abiotic environmental factors, like temperature. For both endotherms and ectotherms, temperature can affect everything from daily energy budgets to nesting behaviors. For instance, in birds environmental temperature plays a key role in shaping parental incubation behavior and temperatures experienced by embryos. Recent research indicates that temperatures experienced by embryos affect viability and are important in shaping fitness-related traits in young birds, sparking renewed interest in relationships among environmental factors, parental incubation behavior, and incubation temperature. Incubation behavior of birds can be monitored non-invasively by placing thermal probes into the nest and analyzing temperature fluctuations that occur as parents attend and leave the nest (on- and off-bouts, respectively). When other measures of temperature (e.g., ambient air or operative temperature) are collected simultaneously with incubation temperature it is possible to compare shifts in behavior with environmental changes. To improve analysis of incubation behavior using these large thermal data sets we developed a program, NestIQ, that uses machine learning to guide parameter optimization allowing it to track the behavior of diverse species. NestIQ's algorithm was tested using six species incubating in lab or field scenarios, that exhibit unique incubation patterns. This stand-alone and open source software is operated through a graphical user interface (i.e., no user programming is required) that provides important behavioral and thermal output statistics. Further, measures of environmental temperature can be imported alongside nest temperature into the program, which then reports various attributes of environmental temperature during shifts in parental behavior. This program will improve the ability of avian ecologists to interpret a critical parental care behavior that can be used across diverse incubation scenarios and species. Although specifically designed for quantifying avian incubation, NestIQ has the potential for broader applications, including basking and nesting behaviors of non-avian reptiles in relation to ambient temperature.

Prolactin is related to incubation constancy and egg temperature following a disturbance in a precocial bird

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.

Parental care in a sexually monomorphic, ground-nesting passerine, the Pink-billed Lark Spizocorys conirostris (Alaudidae)

Within the family Alaudidae, most monomorphic species exhibit some degree of biparental care, but direct paternal care in most species is confined to food provisioning to offspring during the nestling period. Uniquely, the genus Spizocorys is the only sexually monomorphic lark genus in which the majority of species exhibit biparental care in all stages of the breeding cycle, including incubation. Video observations were used to determine parental nest attentiveness, incubation and brooding duty partitioning, and food provision through the breeding stages of Pink-billed Lark Spizocorys conirostris. In this study we found that Pink-billed Larks exhibit biparental care in all stages of breeding, although the parental contributions differed subtly. Incubation nest attentiveness was on the lower limit for passerines with biparental care. Both male and female nest attentiveness were also well below the reported averages for biparental care in birds. There was no significant difference between the male and female incubation attentiveness, although the males had significantly longer recess-bouts. The results suggest that females are primarily responsible for incubation and brooding, with males fulfilling a supplementary role. Males, however, share the responsibility for food delivery and preferentially deliver invertebrate prey rich in proteins and calories. There is also a change in diet from predominantly insectivorous to a greater emphasis on granivory with progression of the nestling period.

Acute exposure to hyperthermic oscillating temperatures during pre-incubation influences northern bobwhite development, hatching, and survival.

Temperature extremes alter development, growth, hatching, and survival of eggs of ground-nesting birds, particularly during pre-incubation (egg laying) when eggs are left unattended and exposed to the environment for days or weeks before parental incubation begins. The northern bobwhite quail is a ground-nesting bird whose eggs experience high temperatures (≥45° C) during pre-incubation. It is known that chronic high temperatures during pre-incubation alter development and reduce hatching and survival of bobwhite eggs, but it is not known if acute doses of high temperatures during pre-incubation have the same effect. In this study, the 12-d pre-incubation period was divided into thirds. Fresh bobwhite eggs were exposed to either a commercial holding temperature for all 12 d (serving as a control), or a high oscillating temperature regimen for 4 d (one third of pre-incubation) either in the early, middle, or late third of pre-incubation, with a low oscillating temperature regimen during the remaining 8 d. The timing of acute exposure to high oscillating temperatures significantly affected bobwhite development. Eggs exposed in the first 2/3 of pre-incubation developed twice as much as eggs exposed late in pre-incubation, even though all eggs received the same amount of heating degree-hours. Thus, a critical window of thermal susceptibility exists for developing northern bobwhites. Acute exposure to high oscillating temperatures resulted in reduced hatchling mass, hatching success, survival, and compromised hatching synchrony. Thus, acute hyperthermic nest temperatures during pre-incubation could result in the observed reductions in the percentage of juveniles in natural populations during hot and droughty years.

Parental incubation exchange in a territorial bird species involves sex-specific signalling

BackgroundEffective 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.ResultsNorthern 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.ConclusionsOur 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.

Parental incubation patterns and the effect of group size in a Neotropical cooperative breeder

ABSTRACT In many cooperatively breeding species, young (“helpers”) from one year help other adults raise offspring the following year. In contrast to helper effects during the nestling or postfledging stages of the avian breeding cycle, potential benefits from helpers during incubation are poorly studied. We analyzed 39 clutches and recorded 6,027 off-bouts to document incubation behavior in the White-breasted Thrasher (Ramphocinclus brachyurus), an endangered Neotropical facultative cooperative breeder. Our goal was to test the prediction that cooperation confers benefits during incubation in terms of increased nest attendance and hatching success. We found that 65% of active hours (0500–2100) are spent on the nest, values somewhat lower than average for a tropical passerine with uniparental incubation. Highest incubation constancy was observed at a rare 4-egg clutch, which was attended by putative joint-nesters. Excluding this clutch, differences in incubation behavior between pair and cooperative group...

'Green incubation': avian offspring benefit from aromatic nest herbs through improved parental incubation behaviour.

Development of avian embryos requires thermal energy, usually from parents. Parents may, however, trade off catering for embryonic requirements against their own need to forage through intermittent incubation. This dynamically adjusted behaviour can be affected by properties of the nest. Here, we experimentally show a novel mechanism by which parents, through incorporation of aromatic herbs into nests, effectively modify their incubation behaviour to the benefit of their offspring. Our study species, the European starling, includes in its nest aromatic herbs which promote offspring fitness. We provided wild starlings with artificial nests including or excluding the typically selected fresh herbs and found strong support for our prediction of facilitated incubation. Herb effects were not explained by thermal changes of the nests per se, but by modified parental behaviours. Egg temperatures and nest attendance were higher in herb than herbless nests, egg temperatures dropped less frequently below critical thresholds and parents started their active day earlier. These effects were dynamic over time and particularly strong during early incubation. Incubation period was shorter in herb nests, and nestlings were heavier one week after hatching. Aromatic herbs hence influenced incubation in beneficial ways for offspring, possibly through pharmacological effects on incubating parents.

Egg turning behavior and incubation temperature in Forster's terns in relation to mercury contamination.

Egg turning behavior is an important determinant of egg hatchability, but it remains relatively understudied. Here, we examined egg turning rates and egg temperatures in Forster’s terns (Sterna forsteri). We used artificial eggs containing a data logger with a 3-D accelerometer, a magnetometer, and a temperature thermistor to monitor parental incubation behavior of 131 tern nests. Overall, adults turned their eggs an average (±SD) of 3.8 ± 0.8 turns h-1, which is nearly two times higher than that of other seabirds. Egg turning rates increased with nest initiation date. We also examined egg turning rates and egg temperatures in relation to egg mercury contamination. Mercury contamination has been shown to be associated with reduced egg hatchability, and we hypothesized that mercury may decrease egg hatchability via altered egg turning behavior by parents. Despite the high variability in egg turning rates among individuals, the rate of egg turning was not related to mercury concentrations in sibling eggs. These findings highlight the need for further study concerning the potential determinants of egg turning behavior.

Altered embryonic development in northern bobwhite quail (Colinus virginianus) induced by pre-incubation oscillatory thermal stresses mimicking global warming predictions.

Global warming is likely to alter reproductive success of ground-nesting birds that lay eggs normally left unattended for days or even weeks before actual parental incubation, especially in already warm climates. The native North American bobwhite quail (Colinus virginianus) is such a species, and pre-incubation quail eggs may experience temperatures ≥45°C. Yet, almost nothing is known about embryonic survival after such high pre-incubation temperatures. Freshly laid bobwhite quail eggs were exposed during a 12 day pre-incubation period to one of five thermal regimes: low oscillating temperatures (25–40°C, mean = 28.9°C), high oscillating temperatures (30–45°C, mean = 33.9°C), low constant temperatures (28.85°C), high constant temperatures (mean = 33.9°C), or commercially employed pre-incubation temperatures (20°C). After treatment, eggs were then incubated at a standard 37.5°C to determine subsequent effects on embryonic development rate, survival, water loss, hatching, and embryonic oxygen consumption. Both quantity of heating degree hours during pre-incubation and specific thermal regime (oscillating vs. non-oscillating) profoundly affected important aspects of embryo survival and indices of development and growth Pre-incubation quail eggs showed a remarkable tolerance to constant high temperatures (up to 45°C), surviving for 4.5±0.3 days of subsequent incubation, but high oscillating pre-incubation temperature increased embryo survival (mean survival 12.2±1.8 days) and led to more rapid development than high constant temperature (maximum 38.5°C), even though both groups experienced the same total heating degree-hours. Oxygen consumption was ~200–300 μl O2.egg.min-1 at hatching in all groups, and was not affected by pre-incubation conditions. Oscillating temperatures, which are the norm for pre-incubation quail eggs in their natural habitat, thus enhanced survival at higher temperatures. However, a 5°C increase in pre-incubation temperature, which equates to the predicted long-term increases of 5°C or more, nonetheless reduced hatching rate by approximately 50%. Thus, while pre-incubation bobwhite eggs may be resiliant to moderate oscillating temperature increases, global warming will likely severely impact wild bobwhite quail populations, especially in their strongholds in southern latitudes.

High atmospheric temperatures and 'ambient incubation' drive embryonic development and lead to earlier hatching in a passerine bird.

Tropical and subtropical species typically experience relatively high atmospheric temperatures during reproduction, and are subject to climate-related challenges that are largely unexplored, relative to more extensive work conducted in temperate regions. We studied the effects of high atmospheric and nest temperatures during reproduction in the zebra finch. We characterized the temperature within nests in a subtropical population of this species in relation to atmospheric temperature. Temperatures within nests frequently exceeded the level at which embryo’s develop optimally, even in the absence of parental incubation. We experimentally manipulated internal nest temperature to demonstrate that an average difference of 6°C in the nest temperature during the laying period reduced hatching time by an average of 3% of the total incubation time, owing to ‘ambient incubation’. Given the avian constraint of laying a single egg per day, the first eggs of a clutch are subject to prolonged effects of nest temperature relative to later laid eggs, potentially increasing hatching asynchrony. While birds may ameliorate the negative effects of ambient incubation on embryonic development by varying the location and design of their nests, high atmospheric temperatures are likely to constitute an important selective force on avian reproductive behaviour and physiology in subtropical and tropical regions, particularly in the light of predicted climate change that in many areas is leading to a higher frequency of hot days during the periods when birds breed.

The effects of heterospecifics and climatic conditions on incubation behavior within a mixed‐species colony

Parental incubation behavior largely influences nest survival, a critical demographic process in avian population dynamics, and behaviors vary across species with different life history breeding strategies. Although research has identified nest survival advantages of mixing colonies, behavioral mechanisms that might explain these effects is largely lacking. We examined parental incubation behavior using video‐monitoring techniques on Alcatraz Island, California, of black‐crowned night‐heron Nycticorax nycticorax (hereinafter, night‐heron) in a mixed‐species colony with California gulls Larus californicus and western gulls L. occidentalis. We first quantified general nesting behaviors (i.e. incubation constancy, and nest attendance), and a suite of specific nesting behaviors (i.e. inactivity, vigilance, preening, and nest maintenance) with respect to six different daily time periods. We employed linear mixed effects models to investigate environmental and temporal factors as sources of variation in incubation constancy and nest attendance using 211 nest days across three nesting seasons (2010–2012). We found incubation constancy (percent of time on the eggs) and nest attendance (percent of time at the nest) were lower for nests that were located < 3 m from one or more gull nest, which indirectly supports the predator protection hypothesis, whereby heterospecifics provide protection allowing more time for foraging and other self‐maintenance activities. To our knowledge, this is the first empirical evidence of the influence of one nesting species on the incubation behavior of another. We also identified distinct differences between incubation constancy and nest attentiveness, indicating that these biparental incubating species do not share similar energetic constraints as those that are observed for uniparental species. Additionally, we found that variation in incubation behavior was a function of temperature and precipitation, where the strength of these effects was dependent on the time of day. Overall, these findings strengthen our understanding of incubation behavior and nest ecology of a colonial‐nesting species.

Incubation temperature influences survival in a small passerine bird

In birds parental incubation behaviour is an important factor shaping the environmental conditions under which the embryos develop, and sub‐optimal incubation temperatures are known to negatively affect early growth and development. It is less well known if variation in incubation temperature can impose life‐long differences in individual performance and survival. In the present study we investigated the effects of incubation temperature on long‐term survival in a small passerine bird. Using our captive population of the zebra finchTaeniopygia guttatawe artificially incubated eggs at three biologically relevant temperatures (35.9, 37.0 and 37.9°C) for two‐thirds of the incubation period and then monitored individual lifespan of the hatched chicks for two and a half years. We found that individuals from eggs incubated under the lowest temperature exhibited significantly lower long‐term survival compared to those which had been incubated at the highest temperature. Our results show that incubation temperature in birds, and thus parental incubation behaviour, play an important role in shaping the life‐history trajectories of offspring.

Effects of inbreeding and parental incubation on captive breeding success in Hawaiian crows

Inbreeding can reduce individual or population fitness and is predicted to have particularly severe effects on early life traits. Detecting inbreeding depression remains a challenge, however, especially in endangered species or long-lived taxa. Ample and detailed data are needed to study the effects of inbreeding at various life stages. We used 17years of captive breeding records of the Hawaiian crow (Corvus hawaiiensis) to study the effects of individual and parental level of inbreeding on survival through early life. Using pedigree data to calculate inbreeding coefficients for over 300 eggs, we find that the individual’s inbreeding coefficient, but not the parents’, has a very strong effect on embryo and chick survival. Inbreeding levels of chicks that survived to 2years (reproductive age) are 31% lower than those of embryos and chicks that died. Highly inbred embryos have a>5 times reduced survival probability as a consequence of individuals carrying on average 6.9 haploid lethal equivalents for survival to 2years. Further, offspring survival is greatly improved by longer parental incubation. Individuals surviving to 2years had, on average, twice as many days of parental incubation prior to artificial incubation (four vs. two). Our study contributes to evidence that the strength of inbreeding depression is particularly severe in early life traits. It shows that the negative effects of inbreeding on reproductive success should be accounted for even in benign captive environments where survival is maximized and suggests that parental incubation should be favored over artificial incubation in avian captive breeding programs.