Evolution of passerine incubation behavior: influence of food, temperature, and nest predation

Abstract.— Incubation behavior is one component of reproductive effort and thus influences the evolution of life-history strategies. We examined the relative importance of body mass, frequency of mate feeding, food, nest predation, and ambient temperature to explain interspecific variation in incubation behavior (nest attentiveness, on- and off-bout durations, and nest trips per hour) using comparative analyses for North American passerines in which only females incubate. Body mass and frequency of mate feeding explained little variation in incubation behavior. We were also unable to detect any influence of food; diet and foraging strategy explained little interspecific variation in incubation behavior. However, the typical temperature encountered during reproduction explained significant variation in incubation behavior: Species breeding in colder environments take shorter bouts off the nest, which prevents eggs from cooling to temperatures below the physiological zero temperature. These species must compensate for shorter off-bouts by taking more of them (thus shorter on-bouts) to obtain needed energy for incubation. Nest predation also explains significant variation in incubation behavior among passerines: Species that endure high nest predation have evolved an incubation strategy (long on- and off-bouts) that minimizes activity that could attract predators. Nest substrate explained additional variation in incubation behavior (cavity-nesting birds have shorter on-bouts and make more frequent nest trips), presumably because nest predation and/or temperature varies among nest substrates. Thus, nest predation can influence reproductive effort in a way previously not demonstrated–by placing a constraint on parental activity at the nest. Incubating birds face an ecological cost associated with reproductive effort (predation of entire brood) that should be considered in future attempts to explain avian life-history evolution.     

Patterns of nest relief during incubation,and incubation period variability in the yellow-eyed penguin (Megadyptes antipodes)

Abstract

Daily nest checks were used to determine the yellow-eyed penguin’s pattern of nest relief during incubation, and to examine the inter-pair variability in incubation period. Nests at two breeding areas on the Otago Peninsula were visited between September and December in 1985 and 1986. At each visit, status of the nest and the identity and behaviour of the sitting bird was recorded. Incubation is shared between the sexes, with males tending to sit longer than females. Incubation spells ranged from 1–7 days with a mode of 1–2 days. Records of lone incubation following mate death show the yelloweyed penguin to be capable of 15 day unrelieved spells at the nest The incubation period of the yellow-eyed penguin ranges from 39–51 days and is the most variable of all penguins. This variability is because of the failure of some individuals to adequately cover the clutch until up to 5 days after the laying of the second egg.     

Incubation temperature impacts nestling growth and survival in an open‐cup nesting passerine

For oviparous species such as birds, conditions experienced while in the egg can have long‐lasting effects on the individual. The impact of subtle changes in incubation temperature on nestling development, however, remains poorly understood, especially for open‐cup nesting species with altricial young. To investigate how incubation temperature affects nestling development and survival in such species, we artificially incubated American robin (Turdus migratorius) eggs at 36.1°C (“Low” treatment) and 37.8°C (“High” treatment). Chicks were fostered to same‐age nests upon hatching, and we measured mass, tarsus, and wing length of experimental nestlings and one randomly selected, naturally incubated (“Natural”), foster nest‐mate on days 7 and 10 posthatch. We found significant effects of incubation temperature on incubation duration, growth, and survival, in which experimentally incubated nestlings had shorter incubation periods (10.22, 11.50, and 11.95 days for High, Low, and Natural eggs, respectively), and nestlings from the Low treatment were smaller and had reduced survival compared to High and Natural nestlings. These results highlight the importance of incubation conditions during embryonic development for incubation duration, somatic development, and survival. Moreover, these findings indicate that differences in incubation temperature within the natural range of variation can have important carryover effects on growth and survival in species with altricial young.     

Influence of olfactory and visual cover on nest site selection and nest success for grassland‐nesting birds

Habitat selection by animals is influenced by and mitigates the effects of predation and environmental extremes. For birds, nest site selection is crucial to offspring production because nests are exposed to extreme weather and predation pressure. Predators that forage using olfaction often dominate nest predator communities; therefore, factors that influence olfactory detection (e.g., airflow and weather variables, including turbulence and moisture) should influence nest site selection and survival. However, few studies have assessed the importance of olfactory cover for habitat selection and survival. We assessed whether ground‐nesting birds select nest sites based on visual and/or olfactory cover. Additionally, we assessed the importance of visual cover and airflow and weather variables associated with olfactory cover in influencing nest survival. In managed grasslands in Oklahoma, USA, we monitored nests of Northern Bobwhite (Colinus virginianus), Eastern Meadowlark (Sturnella magna), and Grasshopper Sparrow (Ammodramus savannarum) during 2015 and 2016. To assess nest site selection, we compared cover variables between nests and random points. To assess factors influencing nest survival, we used visual cover and olfactory‐related measurements (i.e., airflow and weather variables) to model daily nest survival. For nest site selection, nest sites had greater overhead visual cover than random points, but no other significant differences were found. Weather variables hypothesized to influence olfactory detection, specifically precipitation and relative humidity, were the best predictors of and were positively related to daily nest survival. Selection for overhead cover likely contributed to mitigation of thermal extremes and possibly reduced detectability of nests. For daily nest survival, we hypothesize that major nest predators focused on prey other than the monitored species’ nests during high moisture conditions, thus increasing nest survival on these days. Our study highlights how mechanistic approaches to studying cover informs which dimensions are perceived and selected by animals and which dimensions confer fitness‐related benefits.     

Size matters: nest colonization patterns for twig‐nesting ants

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Supplemental food increases ornamentation of male nestling Eastern Bluebirds

Although the condition‐dependence and signaling function of ornamental plumage coloration among adult males is well studied, less research has focused on the information content of ornamental coloration among juvenile birds. Eastern Bluebird (Sialia sialis) nestlings grow their nuptial plumage while in the nest and dependent on parents for food, making them an ideal species for studying the development and function of elaborate plumage. Previous research suggests that plumage brightness of Eastern Bluebirds functions, in the juvenile stage, in parent–offspring interactions as a sexually selected trait in adults. Using an experimental approach, we tested the effects of supplemental food on the structural plumage coloration (i.e., tips of primary feathers) of Eastern Bluebird nestlings in Watauga County, North Carolina, during the 2011 breeding season. We provided supplemental mealworms daily to breeding pairs from the onset of incubation through the nestling period, and measured plumage brightness, UV chroma, and mass of nestlings (N = 89 males and 71 females). Male nestlings of supplementally fed parents exhibited brighter plumage. The mass and UV chroma of young bluebirds were not significantly affected by food supplementation. However, the relationship between mass and brightness differed between male nestlings in the control and supplementally fed treatments. Males reared in food‐supplemented territories exhibited a positive relationship between color and mass. Nestlings in control territories, however, exhibited a negative relationship between size and brightness, suggesting that reduced food availability results in a tradeoff between allocating resources toward somatic growth and development of bright plumage. Our results suggest that UV‐blue structural plumage in male juvenile Eastern Bluebirds is at least partially condition‐dependent and may help to explain why plumage color can influence social interactions in Eastern Bluebirds.     

Effects of environmental conditions on reproductive effort and nest success of Arctic‐breeding shorebirds

The Arctic is experiencing rapidly warming conditions, increasing predator abundance, and diminishing population cycles of keystone species such as lemmings. However, it is still not known how many Arctic animals will respond to a changing climate with altered trophic interactions. We studied clutch size, incubation duration and nest survival of 17 taxa of Arctic‐breeding shorebirds at 16 field sites over 7 years. We predicted that physiological benefits of higher temperatures and earlier snowmelt would increase reproductive effort and nest survival, and we expected increasing predator abundance and decreasing abundance of alternative prey (arvicoline rodents) to have a negative effect on reproduction. Although we observed wide ranges of conditions during our study, we found no effects of covariates on reproductive traits in 12 of 17 taxa. In the remaining taxa, most relationships agreed with our predictions. Earlier snowmelt increased the probability of laying a full clutch from 0.61 to 0.91 for Western Sandpipers, and shortened incubation by 1.42 days for arcticola Dunlin and 0.77 days for Red Phalaropes. Higher temperatures increased the probability of a full clutch from 0.60 to 0.93 for Western Sandpipers and from 0.76 to 0.97 for Red‐necked Phalaropes, and increased daily nest survival rates from 0.9634 to 0.9890 for Semipalmated Sandpipers and 0.9546 to 0.9880 for Western Sandpipers. Higher abundance of predators (foxes) reduced daily nest survival rates only in Western Sandpipers (0.9821–0.9031). In contrast to our predictions, the probability of a full clutch was lowest (0.83) for Semipalmated Sandpipers at moderate abundance of alternative prey, rather than low abundance (0.90). Our findings suggest that in the short‐term, climate warming may have neutral or positive effects on the nesting cycle of most Arctic‐breeding shorebirds.     

Increased nest defence of upland‐nesting ducks in response to experimentally reduced risk of nest predation

Parent birds should take greater risks defending nests that have a higher probability of success. Given high rates of mammalian nest predation, therefore, parents should risk more for nests in areas with a lower risk of mammalian predation. We tested this hypothesis using nest defence data from over 1300 nests of six species of dabbling ducks studied in an area where predation risk had been reduced through removal of mammalian predators. When predator removal reduced nest predation, the ducks increased risk taking as predicted. Also as predicted, risk taking varied inversely with body size, an index of annual survival, among species. For ducks to vary nest defence in response to variation in predation risk they must be able to assess the risk of nest predation. Because ducks modified nest defence in the breeding season immediately following predator removal, ducks may be able to assess predator abundance indirectly (e.g. by UV reflection from urine) rather than by seeing or interacting directly with the predators.     

Linking songbird nest predation to seedling density: Sugar maple masting as a resource pulse in a forest food web

The ecological literature presents considerable evidence for top‐down forcing on the maintenance of species diversity. Yet, in temperate forests, bottom‐up forces often exert a strong influence on ecosystem functioning. Here, we report on the indirect influence of a pulsed resource, sugar maple (Acer saccharum) seed production, on nest survival in a migratory songbird. We hypothesized that seed production in year t would determine daily nest survival rate in year t + 1 through its effects on seed‐eating rodents. We used the density of sugar maple seedlings (with cotyledons) in year t + 1 as a proxy for seed production in year t and predicted that it would be inversely related to songbird nest survival the same year. We estimated the density of sugar maple seedlings, eastern chipmunk (Tamias striatus) activity, and daily nest survival rate in the ovenbird (Seiurus aurocapilla) over four successive years in a northern hardwood forest of New Brunswick, Canada. Seedling density varied by two orders of magnitude between years, whereas an index of chipmunk activity changed by an order of magnitude. Both variables were positively correlated and negatively correlated to daily nest survival rate. A logistic‐exposure model including only seedling density received the greatest level of support (lowest AIC_c). Previous studies have reported the effect of sugar maple masting on seed‐eating rodent populations, but the strong link we report between seedling density and songbird nest survival is novel. A nocturnal seed‐eating nest predator, deer mouse (Peromyscus maniculatus), was not considered in our models, which may explain why chipmunk was not the best predictor of daily nest survival rate. The trophic linkages we observed are remarkably strong for a temperate forest ecosystem and might become more prevalent in northeastern North America, at least on calcium‐rich soils, with the loss of large‐diameter beech trees as a result of beech bark disease.     

Cold weather events provoke egg ejection behaviour in open‐nesting passerines

We analysed 25 conspecific egg ejection events from a large sample of nests (n = 762) of three passerine species. In two species, most of the events could not be explained by intraspecific brood parasitism or nest sanitation behaviour. At least some of these cases were behavioural responses to unfavourable weather conditions in the spring of 2017. We conclude that clutch size adjustment through egg ejection may be more widespread among passerines than previously thought.     

Effects of parents and Brown‐headed Cowbirds (Molothrus ater) on nest predation risk for a songbird

Nest predation limits avian fitness, so ornithologists study nest predation, but they often only document patterns of predation rates without substantively investigating underlying mechanisms. Parental behavior and predator ecology are two fundamental drivers of predation rates and patterns, but the role of parents is less certain, particularly for songbirds. Previous work reproduced microhabitat‐predation patterns experienced by Yellow Warblers (Setophaga petechia) in the Mono Lake basin at experimental nests without parents, suggesting that these patterns were driven by predator ecology rather than predator interactions with parents. In this study, we further explored effects of post‐initiation parental behavior (nest defense and attendance) on predation risk by comparing natural versus experimental patterns related to territory density, seasonal timing of nest initiation, and nest age. Rates of parasitism by Brown‐headed Cowbirds (Molothrus ater) were high in this system (49% nests parasitized), so we also examined parasitism‐predation relationships. Natural nest predation rates (NPR) correlated negatively with breeding territory density and nonlinearly (U‐shaped relationship) with nest‐initiation timing, but experimental nests recorded no such patterns. After adjusting natural‐nest data to control for these differences from experimental nests other than the presence of parents (e.g., defining nest failure similarly and excluding nestling‐period data), we obtained similar results. Thus, parents were necessary to produce observed patterns. Lower natural NPR compared with experimental NPR suggested that parents reduced predation rates via nest defense, so this parental behavior or its consequences were likely correlated with density or seasonal timing. In contrast, daily predation rates decreased with nest age for both nest types, indicating this pattern did not involve parents. Parasitized nests suffered higher rates of partial predation but lower rates of complete predation, suggesting direct predation by cowbirds. Explicit behavioral research on parents, predators (including cowbirds), and their interactions would further illuminate mechanisms underlying the density, seasonal, and nest age patterns we observed.     

A new method for catching cavity‐nesting birds during egg laying and incubation

ABSTRACT The physiological condition of female birds during the egg‐laying and incubation periods is of considerable interest and yet is relatively understudied in wild birds, primarily due to the difficulty of catching birds during this period without causing nest desertion. We therefore developed a box‐net to capture cavity‐nesting birds using sections of a mist‐net placed around a metal cubic frame. We captured female Great Tits (Parus major) as they left nest boxes during the egg‐laying and incubation periods and measured desertion rates. Using box‐nets, we captured 108 of 119 (90%) females during egg laying and 10 of 12 (83%) during incubation. Our recapture rate over two consecutive days during incubation was 50% (5 of 10). Females not captured left nest boxes before we attempted to capture them, escaped through a hole in the mist‐net, or remained in nest boxes for more than 2 h, after which we ended capture attempts. Overall, 22% of egg‐laying females deserted, with desertion rates highest early in the egg‐laying period. Desertion rates of females captured using box‐nets did not differ from those of undisturbed females. One of 10 females captured in a box‐net deserted during the incubation period. Box‐nets are portable, can be set up and taken down quickly and easily, and could potentially be used with nest boxes or natural cavities at any height. Box‐nets are easy to construct and adaptable for use with an array of cavity‐nesting birds, and can be an important tool for studying female physiology during egg laying and incubation.     

Advancing breeding phenology does not affect incubation schedules in chestnut‐crowned babblers: Opposing effects of temperature and wind

Projecting population responses to climate change requires an understanding of climatic impacts on key components of reproduction. Here, we investigate the associations among breeding phenology, climate and incubation schedules in the chestnut‐crowned babbler (Pomatostomus ruficeps), a 50 g passerine with female‐only, intermittent incubation that typically breeds from late winter (July) to early summer (November). During daylight hours, breeding females spent an average of 33 min on the nest incubating (hereafter on‐bouts) followed by 24‐min foraging (hereafter off‐bouts), leading to an average daytime nest attentiveness of 60%. Nest attentiveness was 25% shorter than expected from allometric calculations, largely because off‐bout durations were double the expected value for a species with 16 g clutches (4 eggs × 4 g/egg). On‐bout durations and daily attentiveness were both negatively related to ambient temperature, presumably because increasing temperatures allowed more time to be allocated to foraging with reduced detriment to egg cooling. By contrast, on‐bout durations were positively associated with wind speed, in this case because increasing wind speed exacerbated egg cooling during off‐bouts. Despite an average temperature change of 12°C across the breeding season, breeding phenology had no effect on incubation schedules. This surprising result arose because of a positive relationship between temperature and wind speed across the breeding season: Any benefit of increasing temperatures was canceled by apparently detrimental consequences of increasing wind speed on egg cooling. Our results indicate that a greater appreciation for the associations among climatic variables and their independent effects on reproductive investment are necessary to understand the effects of changing climates on breeding phenology.     

Incubation failure and nest abandonment by Leach's Storm‐Petrels detected using PIT tags and temperature loggers

ABSTRACT The nocturnal activity of burrow‐nesting seabirds, such as storm‐petrels and shearwaters, makes it difficult to study their incubation behavior. In particular, little is known about possible differences in the incubation behavior of adults at successful and unsuccessful nests. We combined the use of passive integrated transponder (PIT) technology and nest‐temperature data loggers to monitor the incubation behavior of 10 pairs of Leach's Storm‐Petrels (Oceanodroma leucorhoa). The mean incubation bout length was 3.31 ± 0.59 (SD) days for individual adults at successful nests (N= 4) and 1.84 ± 1.16 d for individuals at unsuccessful nests (N= 6). Mean bout length for pairs in successful burrows (3.51 ± 0.56 d) did not differ significantly (P= 0.07) from that for pairs in unsuccessful burrows (1.80 ± 1.20 d), perhaps due to one failed nest with a high mean bout length (4.15 d). The total number of incubation bouts per parent (4.3 ± 1.9 bouts) did not differ with hatching success. Adults whose nests failed repeatedly exhibited truncated incubation bouts (< 12 h) prior to complete nest failure and were more likely than successful parents to make brief visits to nearby, occupied nesting burrows. Our results suggest that the decision by Leach's Storm‐Petrels to abandon a nest is not an abrupt one. Rather, failed nesting attempts may be characterized by truncated incubation bouts where individuals pay the energetic cost of travel to and from the burrow, but do not remain long enough to successfully incubate the egg.     

Light affects parental provisioning behaviour in a cavity‐nesting Passerine

Nocturnal bird species possess special adaptations to maximise visual efficiency under low light levels. However, some typically diurnal species also experience low‐light environments. For example, cavity‐nesting Passerines raise broods in dark cavities and search for food in light‐abundant surroundings. It is not clear whether they possess special adaptations for low light vision or breed in cavities at the expense of impaired parental care. In this study, we tested whether light conditions affect the provisioning efficiency of great tits. We experimentally tested how the level of natural and artificially increased illumination inside nest boxes affects parental feeding duration, frequency and timing. We monitored 15‐h of provisioning activity of great tit parents when nestlings were day seven post hatch. We used traditional ‘dark’ nest boxes and ‘bright’ nest boxes with increased illumination obtained by using semi‐transparent plastic windows. The duration of single feedings were, on average, shorter in brightened nest boxes compared to dark ones. This difference tended to be higher early in the morning and in the evening, when the illumination in dark nest boxes was the lowest. Nest box type, however, did not influence feeding frequency or times of the onset and the end of feeding. Our findings provide new evidence for impaired efficiency of parental care due to lowered light conditions. Further research is needed to test whether prolonged feeding duration has negative effects on adult time budgets and nestling energy expenditures.     

Faithful females receive more help: the extent of male parental care during incubation in relation to extra‐pair paternity in songbirds

Parental care provided by males occurs in a diverse array of animals and there are large differences among species in its extent compared with female care. However, social and ecological factors responsible for interspecific differences in male's share of parental duties remain unclear. Genetic fidelity of females has been long considered important. Theory predicts that females should receive more help from their mates in raising the offspring in species with high genetic fidelity. Using avian incubation behaviour as a model system, we confirmed this prediction. The extent of male's help during incubation increased with decreasing rate of extra‐pair paternity across species (22 species of socially monogamous songbirds from 13 families; male's share of incubation ranged from 6% to 58%), even after accounting for covariates, biases in species selection and intraspecific variability. Moreover, this result was not sensitive to two different phylogenies and branch length estimates. We suggest that our findings support the notion, backed by theory, that genetic fidelity is an important factor in the evolution of male parental care. We offer several behavioural scenarios for the coevolution between male's share of parental duties and the genetic mating system.     

Losing anti‐predatory behaviour: A cost of translocation

Translocation of endangered species to habitats where exotic predators have been removed is now a common conservation practice around the world. Many of these translocated populations have thrived, and they are often used as sources for the harvesting of individuals for translocations to sites where exotic predators still exist, albeit at reduced densities. This study investigates how isolation from exotic predators affects the ability of individuals to recognize such predators using the North Island robin (Petroica longipes) as a model. The study was carried out in three robin populations in the North Island, New Zealand: a translocated population on Tiritiri Matangi Island, where exotic mammalian predators are absent; a population reintroduced from Tiritiri Matangi Island to Wenderholm Regional Park, a mainland site where these mammals are controlled to low densities; and a mainland population at Benneydale where exotic predatory mammals are common. The response intensity of robins to a model stoat was high at Benneydale and low at Tiritiri Matangi and Wenderholm. This result indicates that isolation from mammalian predators on Tiritiri Matangi has suppressed the ability of North Island robins to recognize these predators. It is possible that the low predatory mammal densities at Wenderholm have reduced robin contact with stoats, therefore reduced the opportunity for robins to learn to recognize stoats. Thus, translocation of individuals from populations without predators to places where key predators still exist could be unsuccessful if translocated individuals fail to perform appropriate anti‐predator behaviours.     

Off‐road vehicles affect nesting behaviour and reproductive success of American Oystercatchers Haematopus palliatus

As human populations and associated development increase, interactions between humans and wildlife are occurring with greater frequency. The effects of these interactions, particularly on species whose populations are declining, are of great interest to ecologists, conservationists, land managers and natural resource policy‐makers. The American Oystercatcher Haematopus palliatus, a species of conservation concern in the USA, nests on coastal beaches subject to various forms of anthropogenic disturbance, including aircraft overflights, off‐road vehicles and pedestrians. This study assessed the effects of these human disturbances on the incubation behaviour and reproductive success of nesting American Oystercatchers at Cape Lookout National Seashore, on the Atlantic coast of the USA. We expanded on‐going monitoring of Oystercatchers at Cape Lookout National Seashore by supplementing periodic visual observations with continuous 24‐h video and audio recording at nests. Aircraft overflights were not associated with changes in Oystercatcher incubation behaviour, and we found no evidence that aircraft overflights influenced Oystercatcher reproductive success. However, Oystercatchers were on their nests significantly less often during off‐road vehicle and pedestrian events than they were during control periods before the events, and an increase in the number of off‐road vehicles passing a nest during incubation was consistently associated with significant reductions in daily nest survival (6% decrease in daily nest survival for a one‐vehicle increase in the average number of vehicles passing a nest each day; odds ratio = 0.94; 95% confidence interval (CI) 0.90, 0.98) and hatching success (12% decrease in hatching success for a one‐vehicle increase in the average number of vehicles passing a nest each day; odds ratio = 0.88; 95% CI 0.76, 0.97). Management of vehicles and pedestrians in areas of Oystercatcher breeding is important for the conservation of American Oystercatchers.