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.     

Absence from the nest due to human disturbance induces higher nest predation risk than natural recesses in Common Eiders Somateria mollissima

Human disturbance of nesting birds may cause reduced breeding success. It is therefore necessary to assess the impact of disturbance to identify steps that minimize negative impacts. We carried out a study of nesting success at two adjacent colonies of Common Eider Somateria mollissima on the islands of Grindøya and Håkøya in northern Norway between 2006 and 2011. Over the study period, nesting success was consistently higher on Håkøya (69–82%) than on Grindøya (35–60%). Between 2009 and 2011 we used camera monitoring of individual nests to identify determinants of nest survival and predation, focusing in particular on the effect of departures from the nest due to human disturbance, which differed between the colonies due to a long‐term research project on Grindøya. Overall, absence of Common Eiders from nests due to disturbance increased the predation risk by a factor of 6.42 for an increase of one additional daily disturbance. In contrast, absence due to natural recesses did not increase nest losses. Under high levels of human disturbance, camera monitoring indicated that the main cause of breeding failure was predation, primarily by Hooded Crows Corvus cornix, but also to some extent Great Black‐backed Gulls Larus marinus. The presence of cameras did not increase the predation risk. Both the presence of researchers and the sight of Common Eider females conspicuously departing from nests are likely to have provided cues to these predators. We suggest management trials to reduce nesting disturbance through the guarding of unoccupied nests to mitigate the effects of human disturbance on reproductive success.     

PVC nest boxes are less at risk of occupancy by feral honey bees than timber nest boxes and natural hollows

Feral European Honey Bee (Apis mellifera) has been identified as a potential nest competitor for Australian hollow nesting species, but few studies have investigated the impact of feral honey bee competition on Threatened species. Our study used data from Glossy Black‐cockatoo (Calyptorhynchus lathami halmaturinus) nests on Kangaroo Island, monitored and managed over an 11‐year period, and found 12% of nests became occupied by feral honey bees during that period. Our results indicate that feral honey bees were less likely to occupy nest boxes made of PVC (5%) compared with wooden nest boxes (24%) or natural hollows in Eucalyptus trees (14%). The removal of feral honey bee hives from nests is a priority for long‐term conservation of glossy black‐cockatoos on Kangaroo Island. We recommend that PVC nest boxes are chosen for future nesting habitat restoration, due to the more frequent use of wooden nest boxes by feral honey bees.     

Factors affecting egg predation in Black-tailed Gulls

In colonial seabirds, nesting density, egg-laying date and nest microhabitat affect the probability of eggs being taken by avian predators. Jungle Crows (Corvus macrorhynchos) are dominant predators of eggs of Black-tailed Gulls (Larus crassirostris). Factors affecting the probability of gulls allowing the crows to attack their nests or depredate their eggs and the probability of eggs being taken were studied by direct observation and egg census, respectively. The effect of vegetation heights, position in the colony, egg-laying date and neighbour nests on the probability of eggs being taken were examined at multiple spatial scales. Gull nests were depredated more easily by larger groups of crows. Nests in peripheral areas (<4 m from the edge of the colony) were also depredated more easily by the crows walking on the ground. Although the nests where eggs were laid early in the season were depredated more frequently, such nests highly synchronised in egg laying within a <2-m radius were less likely to be depredated than less-synchronised nests. The nests in tall vegetation were less likely to be depredated though those having neighbour nests in tall vegetation were not. The number of neighbour nests did not affect the probability of eggs being taken. Antipredation effects of nesting microhabitats vary with spatial scales at which the crows search and attack the nests of gulls.     

Supplemental food alters nest defence and incubation behaviour of an open‐nesting wetland songbird

Climate‐driven increases in spring temperatures are expected to result in higher prey availability earlier in the breeding season for insectivorous birds breeding in wetland habitats. Predation during the incubation phase is a major cause of nesting failure in open‐nesting altricial birds such as the Eurasian reed warbler. The nest predation rate in this species has recently been shown to be substantially reduced under conditions of experimentally elevated invertebrate prey availability. Food availability near the nest may be an important determinant of adult incubation and nest defence behaviours during the incubation period. We used two experimental studies to compare incubation behaviour and nest defence in food‐supplemented and unsupplemented adult Eurasian reed warblers during the incubation phase. In the first study we measured nest defence behavioural responses to a taxidermic mount of a native predator (stoat Mustela erminea). In the second study we used temperature loggers installed in nests to measure breaks in incubation as a measure of nest vulnerability. Food‐supplemented birds responded aggressively to the presence of a predator more quickly than those in the unsupplemented group, suggesting they are closer to their nest and can more quickly detect a predator in the vicinity. Food‐supplemented birds also had shorter breaks in incubation (both in terms of maximum and mean off‐bout durations), presumably because they were foraging for shorter periods or over shorter distances from the nest. This study therefore identifies the behavioural mechanisms by which changes in food availability may lead to changes in nest survival and thus breeding productivity, in open‐nesting insectivorous birds.     

The effects of rainfall on different components of seasonal fecundity in a tropical forest passerine

Seasonal fecundity is a composite metric that is determined by component parameters such as clutch size, nest survival and re‐nesting probability. Many of these component parameters are known to vary with environmental conditions, in particular rainfall prior to or during the breeding season. In some species, seasonal fecundity is positively related to rainfall, but little is known about which component parameters of seasonal fecundity respond most strongly to rainfall. We used intensive nest monitoring of a multi‐brooded tropical forest passerine, the Montserrat Oriole Icterus oberi, to examine the effects of rainfall during the pre‐breeding season on component parameters of annual fecundity. We monitored all nests of a total of 42 pairs over 5 years in which rainfall varied substantially. We then related clutch size, nest survival, onset and length of the breeding season, re‐nesting probability and re‐nesting interval to pre‐breeding season rainfall using generalized linear mixed models that accounted for random variation across sites and individual pairs, and incorporated other variables known to affect the response. Higher pre‐breeding season rainfall led to an increase in clutch size and a decrease in re‐nesting interval, but nest survival, re‐nesting probability and length of the breeding season were not affected by variation in rainfall. The onset of the breeding season was delayed in very dry years. We conclude that higher rainfall is likely to increase food availability and thus body condition of female Montserrat Orioles, leading to an increase in fecundity due to larger clutch sizes.     

Sex,growth rate,rank order after brood reduction,and hatching date affect first‐year survival of long‐lived Herring Gulls

Among most species of birds, survival from hatching throughout the first year of life is generally lower than subsequent survival rates. Survival of young birds during their first year may depend on a combination of selection, learning, unpredictable resources, and environmental events (i.e., post‐fledging factors). However, knowledge about post‐fledging development in long‐lived species is usually limited due to a lengthy immature stage when individuals are generally unobservable. Therefore, pre‐fledging characteristics are often used to predict the survival of young birds. We assessed effects of nestling growth rates, hatching date, hatching asynchrony, brood size and rank order after brood reduction, and sex on first‐year survival of 137 fledglings using a mark‐resighting analysis. We found that the survival probability (Φ_1yr = 0.39) of first‐year Herring Gulls (Larus argentatus) in our study colony located at the outer port of Zeebrugge (Belgium) was lower than that of older individuals (Φ_>1yr = 0.75). All 10 models best supported by our data included nestling growth rate, suggesting that variability in first‐year survival may be linked primarily to individual variation in growth. First‐year survival was negatively correlated with hatching date and rank order after brood reduction. Hence, carry‐over effects of breeding season events such as timing of breeding, early development, and social status had an influence on survival of Herring Gulls after fledging. Furthermore, we found sex‐biased mortality in first‐year Herring Gulls, with females (Φ_1yr = 0.45) surviving better than males (Φ_1yr = 0.38). Although adult survival is generally regarded as the key parameter driving population trajectories in long‐lived species, juvenile survival has recently been acknowledged as an important source of variability in population growth rates. Thus, increasing our knowledge of factors affecting age‐specific survival rates is necessary to improve our understanding of population dynamics and ultimately life‐history variation.     

Nesting Habitat Creation Enhances Recruitment in a Predator‐Free Environment: Malaclemys Nesting at the Paul S. Sarbanes Ecosystem Restoration Project

Aquatic turtles worldwide are plagued with habitat loss due to development and shoreline alteration that destroys the terrestrial–aquatic linkage which they must cross to reproduce successfully. Furthermore, nesting habitat loss can concentrate nesting, increasing nest predator efficiency. We describe how the Paul S. Sarbanes Ecosystem Restoration Project at Poplar Island created nesting habitat for Malaclemys terrapin (Diamondback Terrapin), and document nesting success in response to construction progress and the absence of raccoons and foxes, the primary nest predators. We monitored terrapin nests throughout the nesting seasons from 2002 to 2011 to determine overall and within‐nest survivorship. Female terrapins began nesting on the restoration project within 1 year but planned construction during the study eliminated some nesting areas and opened previously inaccessible areas. Overall, nest survivorship was considerably higher than mainland nesting areas due to the absence of raccoons and foxes on the island and within‐nest survivorship was similar. Egg size, hatchling size, and the frequency of shell scute anomalies were similar to other terrapin populations, suggesting normal developmental conditions on the island. We documented annual variation in hatchling size that correlated negatively with mean air temperature during the incubation season. Our results indicate that restored or created isolated island habitat can be located rapidly by terrapins and can become an important source of recruitment in regions where nesting habitat is limited and predation is high. Poplar Island illustrates how habitat loss and restoration can affect turtle populations by revealing the changes in nesting patterns and success in newly created, predator‐free habitat.     

Factors associated with nest survival of Black‐throated Sparrows,desert‐breeding nest‐site generalists

Black‐throated Sparrows (Amphispiza bilineata) are common breeding birds throughout the desert regions of North America and can be considered nest‐site generalists. Information about how spatial (e.g., vegetation) and temporal factors influence nest survival of these sparrows is lacking throughout their range. Our objective was to examine the spatial and temporal factors associated with nest survival of Black‐throated Sparrows at the nest and nest‐patch scales in the predator‐rich environment of the northern Chihuahuan Desert of New Mexico. We used a logistic‐exposure model fit within a Bayesian framework to model the daily survival probability of Black‐throated Sparrow nests. Predation was the leading cause of nest failure, accounting for 86% of failed nests. We found evidence of negative associations between nest survival and both vegetative cover above nests and shrub density within 5 m of nests. We found no support for other habitat covariates, but did find strong evidence that daily survival rate was higher earlier in the breeding season and during the egg‐laying stage. A decline in nest survival later in the breeding period may be due to increased predator activity due to warmer ambient temperatures, whereas lower survival during the incubation and nestling stages could be a result of increased activity at nests. A generalist approach to nest‐site selection may be an adaptive response to the presence of a diverse assemblage of nest predators that results in the reduced influence of spatial factors on nest survival for Black‐throated Sparrows.     

Central–periphery gradient of individual quality within a colony of Black‐headed Gulls

Central nesting sites within avian colonies are often more profitable in terms of fitness, as they can offer better protection against predators compared with colony edges. Thus, central sites are expected to attract high‐quality individuals, which should produce a clear central–periphery gradient in the phenotypic quality of nesting individuals and reproductive output within colonies. The aim of this study was to assess spatial patterns of individual quality within a colony of a common larid, the Black‐headed Gull Chroicocephalus ridibundus. We found that laying was advanced in the central zone of the colony when compared with the periphery, and central pairs tended to lay larger eggs and had higher hatching success than peripheral pairs. Centrally nesting individuals had higher blood haemoglobin concentrations, size‐corrected body mass and heterozygosity (males only), indicating their better condition and higher genetic quality. In contrast, central females had lower plasma albumin concentration, which could possibly be attributed to their greater investment in egg production. Finally, pairs breeding in the central zone of the colony had an elevated heterophil/lymphocyte ratio, suggesting that social stress can be recognized as an important cost of nesting in dense aggregations. The results provide empirical support for central–periphery gradients in the individual quality of colonially nesting Black‐headed Gulls, indicating that spatial gradients in pair quality should be carefully taken into account when conducting research on colonial birds.     

THE EFFECTS OF DIURNAL AND TIDAL PERIODICITIES IN THE NUMBERS AND ACTIVITIES OF HERRING GULLS LARUS ARGENTATUS IN A COLONY

The average numbers of Herring Gulls Larus argentatus present in a breeding colony on Walney Island, Cumbria, were found to vary with the tidal cycle but to remain effectively constant with time of day through the breeding season. An activity survey, based on 50 Herring Gulls observed at half-hourly intervals during March and April 1973, showed that sleep and rest varied inversely with each other with sleep increasing to 50 per cent at midday. After a peak in the proportion of gulls asleep four hours before low tide, sleeping progressively decreased until low tide; seemingly a result of resident gulls waking and remaining more alert as others left the colony in search of food. Preening was constant throughout the day and tide cycle. Other behaviours (mostly courtship and agonistic behaviour associated with territory defence) increased slightly during low tide and were more common early and late in the day. Night observations of the gulls' activities showed that there was a peak of sleeping between midnight and 02.00 hours. It is suggested that Herring Gulls have a bimodal diel sleep pattern.     

Nesting ecology of solitary‐nesting Amur Falcons (Falco amurensis) in central Mongolia

Amur Falcons (Falco amurensis) are a migratory species that face a variety of threats across their range, but little is known about their breeding ecology. These falcons breed in forest habitats in Eastern and Central Asia using nests constructed by corvids, including Eurasian Magpies (Pica pica). We monitored nests of 21 pairs of Amur Falcons at Hustai National Park in central Mongolia in 2017. Our objectives were to describe their basic nesting ecology, estimate nest survival by modeling the daily survival rate (DSR), examine nest selection by modeling it as a function of nest and site covariates, and use a spatial simulation to test hypotheses concerning intra‐ and interspecific avoidance. Clutch sizes averaged 4.1 eggs (N = 21 nests), and incubation and nestling periods averaged 25.7 and 26.1 d, respectively. The daily survival rate was 0.98, with young in 12 nests surviving to fledging. Nest structures were more likely to be selected as percent cover of nest bowls increased, usually in the form of a dome of sticks with multiple side entrances. Closed nests likely provide increased protection from predators. In contrast to congeneric Red‐footed Falcons (F. vespertinus) that nest in large colonies, Amur Falcons nested no farther from or closer to nests of either conspecifics or congeners than expected by chance. One factor likely contributing to this difference is that Red‐footed Falcons often use the nests of colonial‐nesting Rooks (Corvus frugilegus), whereas Amur Falcons typically use the nests of non‐colonial Eurasian Magpies. The ongoing loss of deciduous trees like white birch (Betula platyphylla) across the breeding range of Amur Falcons, probably due to climate change and increased grazing pressure, is likely to reduce the availability of nesting habitat for Eurasian Magpies which, in turn, will likely reduce availability of nests for Amur Falcons and other small falcons.     

Genetic structure at three spatial scales is consistent with limited philopatry in Ricord's Rock Iguanas (Cyclura ricordii)

Cyclura ricordii is an endemic iguana from Hispaniola Island and is threatened on the IUCN Red List. The main threats are predation by introduced mammals, habitat destruction, and hunting pressure. The present study focused on two nesting sites from Pedernales Province in the Dominican Republic. The hypothesis that natal philopatry influences dispersal and nest‐site selection was tested. Monitoring and sampling took place in 2012 and 2013. Polymorphic markers were used to evaluate whether natal philopatry limits dispersal at multiple spatial scales. Ripley's K revealed that nests were significantly clustered at multiple scales, when both nesting sites were considered and within each nesting site. This suggests a patchy, nonrandom distribution of nests within nest sites. Hierarchical AMOVA revealed that nest‐site aggregations did not explain a significant portion of genetic variation within nesting sites. However, a small but positive correlation between geographic and genetic distance was detected using a Mantel's test. Hence, the relationship between geographic distance and genetic distance among hatchlings within nest sites, while detectable, was not strong enough to have a marked effect on fine‐scale genetic structure. Spatial and genetic data combined determined that the nesting sites included nesting females from multiple locations, and the hypothesis of “natal philopatry” was not supported because females nesting in the same cluster were no more closely related to each other than to other females from the same nesting site. These findings imply that nesting aggregations are more likely associated with cryptic habitat variables contributing to optimal nesting conditions.     

All you can eat: is food supply unlimited in a colonially breeding bird?

Food availability is generally considered to determine breeding site selection and therefore plays an important role in hypotheses explaining the evolution of colony formation. Hypotheses trying to explain why birds join a colony usually assume that food is not limited, whereas those explaining variation in colony size suggest that food is under constraint. In this study, we investigate the composition and amount of food items not eaten by the nestlings and found in nest burrows of colonially nesting European bee‐eaters (Merops apiaster). We aimed to determine whether this unconsumed food is an indicator of unlimited food supply, the result of mistakes during food transfer between parents and chicks or foraging selectivity of chicks. Therefore, we investigated the amount of dropped food for each nest in relation to reproductive performance and parameters reflecting parental quality. Our data suggest that parents carry more food to the nest than chicks can eat and, hence, food is not limited. This assumption is supported by the facts that there is a positive relationship between dropped food found in a nest and the number of fledglings, nestling age, and chick health condition and that the amount of dropped food is independent of colony size. There is variation in the amount of dropped food within colonies, suggesting that parent foraging efficiency may also be an important determinant. Pairs nesting in the center of a colony performed better than those nesting on the edge, which supports the assumption that quality differences between parents are important as well. However, dropped food cannot be used as an indicator of local food availability as (1) within‐colony variation in dropped food is larger than between colony variation and, (2) the average amount of dropped food is not related to colony size.     

Predation risk of eggs and nestlings relative to nest‐site characteristics of the Bull‐headed Shrike Lanius bucephalus

Appropriate nest‐site selection is one of the most important ways to minimize loss of reproductive investment due to predation. We determined the environmental characteristics associated with nest predation during the incubation and nestling periods of arboreal nesting Bull‐headed Shrikes on the oceanic Minami‐Daito Island where the predator community has low species diversity and includes only three introduced mammals: Ship Rat Rattus rattus, Japanese Weasel Mustela itatsi and Feral Cat Felis catus. Egg predation declined with increasing grassland cover around nests, whereas nestling predation declined with increasing nest concealment and nest height. Our results suggest that effective nest‐site characteristics for avoiding nest predation differ during the incubation and nestling periods and are dependent on the predator species and their search strategies, at least in habitats with low predator species diversity.     

Communal nesting under climate change: fitness consequences of higher incubation temperatures for a nocturnal lizard

Communal nesting lizards may be vulnerable to climate warming, particularly if air temperatures regulate nest temperatures. In southeastern Australia, velvet geckos Oedura lesueurii lay eggs communally inside rock crevices. We investigated whether increases in air temperatures could elevate nest temperatures, and if so, how this could influence hatching phenotypes, survival, and population dynamics. In natural nests, maximum daily air temperature influenced mean and maximum daily nest temperatures, implying that nest temperatures will increase under climate warming. To determine whether hotter nests influence hatchling phenotypes, we incubated eggs under two fluctuating temperature regimes to mimic current ‘cold’ nests (mean = 23.2 °C, range 10–33 °C) and future ‘hot’ nests (27.0 °C, 14–37 °C). ‘Hot’ incubation temperatures produced smaller hatchlings than did cold temperature incubation. We released individually marked hatchlings into the wild in 2014 and 2015, and monitored their survival over 10 months. In 2014 and 2015, hot‐incubated hatchlings had higher annual mortality (99%, 97%) than cold‐incubated (11%, 58%) or wild‐born hatchlings (78%, 22%). To determine future trajectories of velvet gecko populations under climate warming, we ran population viability analyses in Vortex and varied annual rates of hatchling mortality within the range 78– 96%. Hatchling mortality strongly influenced the probability of extinction and the mean time to extinction. When hatchling mortality was >86%, populations had a higher probability of extinction (PE: range 0.52– 1.0) with mean times to extinction of 18–44 years. Whether future changes in hatchling survival translate into reduced population viability will depend on the ability of females to modify their nest‐site choices. Over the period 1992–2015, females used the same communal nests annually, suggesting that there may be little plasticity in maternal nest‐site selection. The impacts of climate change may therefore be especially severe on communal nesting species, particularly if such species occupy thermally challenging environments.     

Nesting Success in Crimson Finches: Chance or Choice?

In avian systems, nest predation is one of the most significant influences on reproductive success. Selection for mechanisms and behaviours to minimise predation rates should be favoured. To avoid predation, breeding birds can often deter predators through active nest defence or by modifying behaviours around the nest (e.g. reducing feeding rates and vocalisations). Birds might also benefit from concealing nests or placing them in inaccessible locations. The relative importance of these strategies (behaviour vs. site selection) can be difficult to disentangle and may differ according to life history. Tropical birds are thought to experience higher rates of predation than temperate birds and invest less energy in nest defence. We monitored a population of crimson finches (Neochmia phaeton), in the Australian tropics, over two breeding seasons. We found no relationship between adult nest defence behaviour (towards a model reptile predator) and the likelihood of nest success. However, nest success was strongly related to the visibility of the nest and the structure of the vegetation. We found no evidence that adult nest building decisions were influenced by predation risk; individuals that re‐nested after a predation event did not build their nest in a more concealed location. Therefore, predator avoidance, and hence nest success, appears to be largely due to chance rather than due to the behaviour of the birds or their choice of nesting sites. To escape high predation pressures, multiple nesting attempts both within and between seasons may be necessary to increase reproductive success. Alternatively, birds may be limited in their nest‐site options; that is, high‐quality individuals dominate quality nest sites.     

Do trade‐offs between predation pressures on females versus nests drive nest‐site choice in painted turtles?

Predation strongly influences reproductive behaviours because reproducing individuals must balance mortality risks to themselves and to their offspring. In many freshwater turtles, the nest predation risk decreases with nest distance from water, whereas the predation risk to females increases farther from water. To determine whether predation pressure influences the distance from water at which female turtles nest, we measured predation pressure on nesting females and on nests, as well as the distances of nests to water, in two populations of painted turtles. Using models, we found that female survival in both populations was high and did not vary with distance from water. Nest survival was also uncorrelated with nest distance to water, although it was significantly lower than adult survival in both populations and was only 1.2% in one population. Our results suggest that nest sites are not predictably safe from predators. Instead, turtles may hedge their bets by nesting over a wide range of distances from water because any distance is risky for nests and no distance is particularly risky for the nesting female. We suggest that other factors, such as suitable incubation conditions and/or post‐emergence hatchling survival, probably play a larger role than predation in driving nest‐site choice in painted turtles.     

Power lines,roads, and avian nest survival: effects on predator identity and predation intensity

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HARTLAUB'S GULL: EGGS AND INCUBATION

A.J. Williams. 1990. Hartlaub's Gull: eggs and incubation. Ostrich 61:93-96.

At Marcus Island, South Africa in 1979 Hartlaub's Gulls Larus hartlaubii laid eggs between 31 January and 28 February. Clutches were of one to three eggs; and 49% consisted of two eggs. Egg composition, and intra-clutch variation in egg size and weight, are reported. The laying interval was two days, the incubation period 25 days, and the hatching interval 0,8 days.