A predator's perspective of nest predation: predation by red squirrels is learned,not incidental

Nest predation has been used to explain aspects of avian ecology ranging from nest site selection to population declines. Many arguments rely on specific assumptions regarding how predators find nests, yet these predatory mechanisms remain largely untested. Here we combine artificial nest experiments with behavioural observations of individual red squirrels Tamiasciurus hudsonicus to differentiate between two common hypotheses: predation is incidental versus learned. Specifically, we tested: 1) whether nest survival could be explained solely by a squirrel's activity patterns or habitat use, as predicted if predation was incidental; or 2) if predation increased as a squirrel gained experience preying on a nest, as predicted if predation was learned. We also monitored squirrel activity after predation to test for evidence of two search mechanisms: area‐restricted searching and use of microhabitat search images. Contrary to incidental predation and in support of learning, squirrels did not find nests faster in areas with high use (e.g. forest edges). Instead, survival of artificial nests was strongly related to a squirrel's prior experience preying on artificial nests. Experience reduced nest survival times by over half and increased predation rates by 150–200%. Squirrels returned to and doubled their activity at the site of a previously preyed on nest. However, neither area‐restricted searching nor microhabitat search images can explain how squirrels located artificial nests more readily with experience. Instead, squirrels likely used cues associated with the nests or eggs themselves. Learning implies that squirrels could be increasingly effective predators as the density or profitability of nests increases. Our results add support to the view that nest predation is complex and broadly influenced (e.g. by predator experience, motivation), and is unlikely to be predicted consistently by simple relationships with predator activity, abundance or habitat.     

Mortality of Wood Warbler Phylloscopus sibilatrix nests in Welsh Oakwoods: predation rates and the identification of nest predators using miniature nest cameras

Capsule Predation was the main cause of nest failure, but predation rates have remained unchanged since the 1980s. Eurasian Jays Garrullus glandarius were the most common predator.

Aims To quantify, and compare, nest predation rates for 1982–84 and 2009–11, and to identify predators of Wood Warbler Phylloscopus sibilatrix nests in Welsh oakwoods.

Methods During 2009–11, 167 Wood Warbler nests were monitored and purpose-built miniature nest cameras deployed at 73 of them. Nest predation rates were compared with 67 nests monitored during 1982–84.

Results Of 167 nests monitored from 2009 to 2011, 62 failed due to predation (32/73 camera nests, 30/94 non-camera nests), giving an overall Daily Survival Rate (DSR?±?se) of 0.979?±?0.003. This was not significantly different from the rate during 1982–84 (0.967?±?0.006). In 2009–11, the DSR of nests declined temporally during the season at both the egg and chick stages. For chick stage nests, DSR varied annually and nonlinearly with age of nestlings. There was no evidence for an effect of cameras at either stage. Of 32 camera nests lost to predation, the predator was identified from 28, resulting in 30 predators being identified. There was one case of multiple predators at a single nest. The majority of nest predation was carried out by birds (28/30), predominantly Eurasian Jays (18/28), but also Common Buzzards Buteo buteo (5/28), Great Spotted Woodpeckers Dendrocopos major (3/28) and Eurasian Sparrowhawks Accipiter nisus (2/28). There was one predation by both a Eurasian Badger Meles meles and a Red Fox Vulpes vulpes. There were no records of Grey Squirrels Sciurus carolinensis depredating nests.

Conclusions Nest predation rates were similar in both periods, suggesting that increased rates of nest predation have not been driving the decline of the Wood Warbler population in Wales. Deployment of nest cameras did not affect nest survival rates and were successful in identifying nest predators, the majority of which were avian, especially Eurasian Jays. Knowledge of the identity of nest predators can aid the development of conservation measures.     

Predators and nest success of Sky Larks Alauda arvensis in large arable fields in the Czech Republic

Capsule Nest failure owing to a range of predators was high, but the level and specificity of nest depredation cannot be generalised.

Aims To determine fates and predators of Sky Lark nests in conventionally managed arable fields in the Czech Republic.

Methods Sky Lark nests in large fields (mainly Maize, Sugar Beet and Opium Poppy) were monitored by means of continuous video surveillance.

Results Primary nest fates of 42 active nests were fledging (13), depredation (22), desertion (5), nestling death (1), and flooding (1). The overall nest success (Mayfield estimate) was 17% (all mortality factors considered) or 27% (only depredation). Depredation events were caused by Marsh Harrier Circus aeruginosus (11), Hooded Crow Corvus cornix (4), Stone Marten Martes foina (3), Montagu’s Harrier Circus pygargus (2), Red Fox Vulpes vulpes (2), Hedgehog Erinaceus sp. (2) and Eurasian Jackdaw Corvus monedula (1). Successful nests were only slightly more away from field edge than depredated nests; nests taken by birds tended to be closer to field edge than those depredated by mammals. The possible reasons for the absence of a clear edge effect include comparatively large field parcels (about 50 ha) and location of nests far from field edge (median = 195 m).

Conclusion Nest survival and composition of nest predators are site‐specific and contingent upon the study method and may not be simply generalised.     

A Bayesian multinomial logistic exposure model for estimating probabilities of competing sources of nest failure

Understanding causes of nest loss is critical for the management of endangered bird populations. Available methods for estimating nest loss probabilities to competing sources do not allow for random effects and covariation among sources, and there are few data simulation methods or goodness‐of‐fit (GOF) tests for such models. We developed a Bayesian multinomial extension of the widely used logistic exposure (LE) nest survival model which can incorporate multiple random effects and fixed‐effect covariates for each nest loss category. We investigated the performance of this model and the accompanying GOF test by analysing simulated nest fate datasets with and without age‐biased discovery probability, and by comparing the estimates with those of traditional fixed‐effects estimators. We then exemplify the use of the multinomial LE model and GOF test by analysing Piping Plover Charadrius melodus nest fate data (n = 443) to explore the effects of wire cages (exclosures) constructed around nests, which are used to protect nests from predation but can lead to increased nest abandonment rates. Mean parameter estimates of the random‐effects multinomial LE model were all within 1 sd of the true values used to simulate the datasets. Age‐biased discovery probability did not result in biased parameter estimates. Traditional fixed‐effects models provided estimates with a high bias of up to 43% with a mean of 71% smaller standard deviations. The GOF test identified models that were a poor fit to the simulated data. For the Piping Plover dataset, the fixed‐effects model was less well‐supported than the random‐effects model and underestimated the risk of exclosure use by 16%. The random‐effects model estimated a range of 1–6% probability of abandonment for nests not protected by exclosures across sites and 5–41% probability of abandonment for nests with exclosures, suggesting that the magnitude of exclosure‐related abandonment is site‐specific. Our results demonstrate that unmodelled heterogeneity can result in biased estimates potentially leading to incorrect management recommendations. The Bayesian multinomial LE model offers a flexible method of incorporating random effects into an analysis of nest failure and is robust to age‐biased nest discovery probability. This model can be generalized to other staggered‐entry, time‐to‐hazard situations.     

Predation on artificial and natural nests in the lowland rainforest of Papua New Guinea

Capsule: Although survival of nests was similar between forest fragments and continuous forest, the range of predators differed. Artificial nests provide an under-estimate of nest predation by snakes.

Aims: To estimate the natural nest predation rate in continuous primary forest, compare it with predation rates in forest fragments. To assess the reliability of nest survival rates determined by the use of artificial nests with clay eggs and identify the main nest predators.

Methods: We observed survival of natural nests during the incubation period in continuous primary forest in Papua New Guinea. Some nests were monitored with infrared cameras. We also used artificial nests deployed with clay eggs to identify predators.

Results: There was a predation rate of 50% for natural nests and snakes were major predators of nest contents. Clutch daily survival rates (DSRs) differed among nest types. The DSR of artificial nests (0.977) was not significantly different to that of natural cup nests (0.969). Survival rates of artificial nests were similar in forest fragments and continuous forest. Forest fragments had, however, a higher proportion of avian predators than continuous forest.

Conclusion: Although, we observed similar survival rates in artificial and natural nests, the composition of nest predators was different between natural and artificial nests. Artificial nests were not suitable for estimating the real predation caused by reptiles. Nevertheless, we find that participation of avian nest predators can be estimated correctly with the use of artificial nests.     

Long-term trends in the nest survival and productivity of the Lesser Spotted Woodpecker Dryobates minor in Britain

ABSTRACT

Capsule: There has been no trend in nest survival of Lesser Spotted Woodpecker Dryobates minor in the 70 years since the 1940s whereas the numbers of young per nesting attempt has declined. Late nests in any year are now significantly less productive than early ones.

Aims: To test whether there has been a long-term decline in the nest survival and productivity of the Lesser Spotted Woodpecker in Britain.

Methods: Breeding data from 331 nests over the period 1949–2019 have been analysed. There were three sources – nest records submitted to the British Trust for Ornithology, a study by the Royal Society for the Protection of Birds from 2006 to 2009 and records submitted to the Woodpecker Network from 2015 to 2019. Generalized linear models were used to analyse the records for first egg date, clutch size, numbers of young fledged, and numbers of nest days for which the nest was under observation. Data were grouped into three periods reflecting population trends of the bird (pre-1980, 1980–1999, 2000–2019) and we also analysed trends with year and spring temperature.

Results: There was no trend in first egg date up to 1980 but subsequently it has advanced by 13 days. The mean clutch was 5.3 with no trend with period or year. There were no trends in nest survival during the egg stage or chick rearing. The mean number of young fledged was 2.6, with nests since 1980 fledging lower numbers of young than those pre-1980 and a decline with year. Loss of chicks, probably to starvation, was the main cause of low productivity. In the last period (2000–2019) nests started early in the season fledged more young than those started later, a trend not apparent in the earlier periods.

Conclusions: Low productivity is a widespread problem for the Lesser Spotted Woodpecker which has probably been exacerbated by the trend towards warm springs.     

Herbaceous ground cover reduces nest predation in olive groves

Capsule Bare ground increases artificial nest predation in olive groves.

Aims To assess the effect of different soil management regimes on nest predation rates in olive groves.

Methods We performed nest predation experiments with artificial nests during the breeding season in 2013, in two areas of southern Spain. Each artificial nest (n?=?300) contained three quail Coturnix eggs, two of which were unmanipulated and the third one was emptied and injected with plaster. Predators were identified by marks on eggs filled with plaster.

Results Ground nests were significantly more depredated, irrespective of the presence of ground cover; tree nests were less depredated in fields with ground cover. There was a clear difference in nest predators of ground and tree nests. Rodents were the most frequent predators of tree nests.

Conclusion Lower predation rates of tree nests in orchards with ground cover are probably linked to a change in the foraging behaviour of rodents, which in these more complex habitats might be restricted by rodents' own risk of predation. This study underscores the important role of agricultural practices in preserving farmland bird communities, particularly tree-nesting species, suggesting that for this group, implementation of ground cover in olive groves might enhance breeding success by reducing nest predation rates.     

Using artificial nests to predict nest survival at reintroduction sites

Artificial nests are frequently used to assess factors affecting survival of natural bird nests. We tested the potential for artificial nests to be used in a novel application, the prediction of nest predation rates at potential reintroduction sites where exotic predators are being controlled. We collected artificial nest data from nine sites with different predator control regimes around the North Island of New Zealand, and compared the nest survival rates with those of North Island robin (Petroica longipes) nests at the same sites. Most of the robin populations had been reintroduced in the last 10 years, and were known to vary in nest survival and status (increasing/stable or declining). We derived estimates of robin nest survival for each site based on Stanley estimates of daily survival probabilities and the known incubation and brooding periods of robins. Estimates of artificial nest survival for each site were derived using the known fate model in MARK. We identified the imprints on the clay eggs in the artificial nests, and obtained different estimates of artificial nest survival based on imprints made by different potential predators. We then compared the value of these estimates for predicting natural nest survival, assuming a relationship of the form s = αpβ, where s is natural nest survival and p is artificial nest survival. Artificial nest survival estimates based on imprints made by rats (Rattus spp.) and brushtail possums (Trichosurus vulpecula) were clearly the best predictors (based on AICc), and explained 64% of the variation in robin nest survival among sites. Inclusion of bird imprints in the artificial nest survival estimates substantially reduced their predictive value. We suggest that artificial nests may provide a useful tool for predicting the suitability of potential reintroduction sites for New Zealand forest birds as long as imprints on clay eggs are correctly identified.     

Different nest predator guild associated with egg size in the Patagonian temperate forest

Capsule: Studies of nest predation using artificial nests need to consider the effect of egg size on the types of predator that are detected.

Aims: To estimate the nest predation rate in the Patagonian temperate forest and evaluate the influence of egg size on predator guild.

Methods: On different plant species, we placed 108 nests each containing eggs of either Atlantic Canary Serinus canaria or Common Quail Coturnix coturnix, and a model clay egg of equal size to the real egg. Nest predators were identified from the marks left on the clay eggs or by videos recorded using camera traps.

Results: 86% of the nests were predated. Birds, mainly Chimango Caracara Milvago chimango, were the main nest predators. A marsupial, the Monito del Monte Dromiciops gliroides, and rodents also contributed to nest predation. Nest predation rates were similar for both egg sizes but the nest predator guild was different. Birds and rodents preyed on both eggs but the Monito del Monte consumed mainly small eggs.

Conclusion: Egg size did not influence the rate of nest predation but, instead, affected the nest predator guild. Consequently, in order to avoid underestimating the impacts of small predators, egg size should be considered in studies of nest predation.     

Using remote cameras to validate estimates of nest fate in shorebirds

Nest survival is a key demographic parameter, yet little effort has been made to improve the accuracy of field‐based methods for assigning nest fates to shorebird nests. We used remote cameras to validate estimates of nest fate from field methods and to assess variation in accuracy of nest‐fate assignment for Snowy Plover Charadrius nivosus in Utah, USA. We correctly identified the fates of 84% of nests in the field, and photos from camera monitoring revealed incorrect assignments for 22% of successful nests and 7% of depredated nests. Traditional field methods could be improved by checking nests more frequently when hatching date nears and spending additional time searching for eggshell evidence, especially when nests are in areas susceptible to weather disturbance.     

The influence of parent body colouration and nesting habitat on bird nest predation

ABSTRACT

Capsule: Parent body colouration and nesting habitat may influence nest predation risk and, therefore, influence parental decisions on offspring care.

Aims: To investigate the adaptiveness of bird colouration under two different habitat and light conditions.

Methods: We used painted polyurethane dummies to simulate parental presence near nests, where some dummies were red, and others were cryptic brown. We placed artificial nests on trees in open and forested areas, along with two quail eggs and parental dummies.

Results: The results showed major predation on nests of the open area, however, we found no significant effect of parent colour on nest predation.

Conclusion: We suggest that the costs involved with nest site selection may overcome the costs of colouration in species that occur in these environments.     

Correcting observer effect bias in estimates of nesting success of a coastal bird,the White-fronted Plover Charadrius marginatus

Capsule A recently developed observer-effects model gives better estimates than the survival model.

Aims To compare the performance of the traditional survival model with that of an observer-effects model in estimating nesting success.

Methods We used survival data from 654 nests in a covariate-dependent extension of a maximum-likelihood estimator model to simultaneously estimate daily survival rate and observer effect on nesting success in four different sectors of a study site. Standard likelihood-ratio tests and Akaike's information criterion differences were used to compare model performances.

Results A significant (negative) observer effect was detected in one of four sectors of the study area, and was probably attributable to small mammal predators. Despite significantly higher predation risk during the laying period, there was a significant positive observer effect during the laying period, contrasting with a weak negative effect during the incubation period, suggesting that an observer's tracks around unattended nests reduced predation risk. Likelihood-ratio tests indicated a significantly better fit using the observer effects model versus the Mayfield survival model in four of six scenarios.

Conclusion The observer-effects model accounts for bias in estimates of nest survival rates, and is preferred over the survival model because the inclusion of observer effect as a variable does not over-parameterize the model.     

The effect of climate on nest orientation in the Rufous Hornero Furnarius rufus

ABSTRACT

Capsule: Within breeding season changes in nest orientation of the Rufous Hornero Furnarius rufus may reflect responses to the local climate.

Aims: To explore the effect of climate on the orientation of closed-cup nests of the Rufous Hornero at a subtropical urban site throughout its breeding period.

Methods: The breeding season was divided into two periods: a cold period with low rainfall and a warm, humid period. For each period, the mean nest entrance orientation and distribution were estimated and compared against a random orientation model.

Results: A total of 42 nests were found during the first period of reproduction and 32 during the second period. In both cases, the mean orientation was non-random and significantly different between the two periods: nests were more likely to be orientated towards the northwest in the first period and to the east in the second period.

Conclusions: Climate variation throughout the breeding season was reflected in differences in nest entrance orientation. This could be because this species takes advantage of solar radiation to increase the temperature within the nest during the first cold months of the breeding season and reduces the humidity of the nest, generated by intense rain, as the season progresses.     

Nest site selection and patterns of nest re-use in the Hooded Crow Corvus cornix

Capsule: Hooded Crows Corvus cornix selected nesting trees based on species, height, grouping and distance from an occupied house. Nest re-use was common and pairs that re-used old nests produced more fledglings than those that built a new nest.

Aims: To determine the features of trees that influenced whether they were used by Hooded Crows as nest sites, to establish what factors influenced nest re-use between years and to explore potential costs or benefits of nest re-use.

Methods: In a large area of Orkney, Scotland, the features of trees that contained a Hooded Crow nest were compared to those of trees where nests were absent. Patterns of nest re-use between years were examined in relation to the availability of alternative sites, previous nesting success and the number of equivalent options to the tree used previously within 200?m of this site.

Results: Hooded Crows favoured spruce and pine trees as nest sites, above the most locally abundant tree species, elder and willow. Preference for trees increased with tree height, local tree density and distance from occupied houses. Over half of the crows studied re-used an old nest when one was available and crows that re-used an old nest fledged more offspring than those that built a new nest. The likelihood of a new nest being built increased as the number of potential locations to build increased. Territories where a nest survived the winter were more likely to be reoccupied the following year than those where nests fell, while territories with fewer trees around the old site were most likely to be abandoned, suggesting that those were territories of lower quality.

Conclusions: Hooded Crows displayed strong preferences for nest sites that might favour nesting success by offering concealment, shelter and protection from ground-based predators. Nest re-use was common, especially when alternative sites were scarce, and appeared to facilitate greater reproductive output.     

Effects of prescribed burning on avian nest survival in the southern Great Plains

Shrubs, such as mesquite (Prosopis spp.) and cholla (Opuntia spp.), now dominate fire-suppressed grasslands in southwestern North America. Responses of birds to prescribed burning of the shortgrass prairie in this region are poorly understood. We examined daily survival rates of mourning dove (Zenaida macroura) and lark sparrow (Chondestes grammacus) nests in an experimental landscape (4,811 ha) of spatially replicated, inter-annual fire frequencies (burning every 2 yr, 4 yr, or 10 yr) near Amarillo, Texas. Herbaceous habitat structure was most developed in infrequently burned plots, but shrub densities were less variable among the burn treatments. We modeled daily nest survival (DSR) against burn frequency, shrub density at nest sites, and nest stage (incubation or nestling). Daily survival of mourning dove nests was not well-related to any measured covariate, but lark sparrow DSR was negatively related to the duration of inter-annual burn frequency. In semiarid grasslands heavily inundated with shrubs, prescribed burning may positively influence the nest success of some bird species. © 2011 The Wildlife Society.     

Observation of nest visitation by multiple males in the European Nightjar Caprimulgus europeaus

ABSTRACT

We report observations of nest visitation of multiple males to three nests of European Nightjars Caprimulgus europeaus. In one case, we report the direct observation of two males apparently exhibiting parental behaviour at the same nest. In two other cases, second males visited nests. We consider the potential functions of nest visits by extra-pair males in the Nightjar.     

Nest timing,nest site selection and nest structure in a high latitude population of Bearded Reedlings Panurus biarmicus

Capsule: Bearded Reedlings Panurus biarmicus show consistent nest timing, select old, compacted areas of reed when positioning their nests, and may adjust nest structure in relation to local reed characteristics and temperature.

Aims: To investigate the nest timing, nest site selection and nest structure of a rare and elusive passerine, the Bearded Reedling at the northern limit of this species’ range.

Methods: A sample of Bearded Reedling nests from the Tay Reedbeds in Scotland were located and monitored with regard to the timing of nesting, fate, fine-scale habitat characteristics and nest structure.

Results: First egg dates and brood sizes were consistent between years of the study despite variation in spring temperatures. Bearded Reedlings nested within unmanaged patches of reed, positively selecting deep leaf litters and compacted reed. Attributes of nest structure, namely internal and external diameter, were influenced by nest site characteristics and local temperature.

Conclusion: Despite a limited sample size, the study suggests that reedbed management should ensure adequate areas of old, dry and unmanaged reed are available when aiming to encourage breeding Bearded Reedlings. Additionally, the apparently flexible structure of the nest may assist this species when coping with changeable climatic conditions.     

Using nest captures and video cameras to estimate survival and abundance of breeding Piping Plovers Charadrius melodus

The estimation of abundance is fundamental to ecology and conservation but often is difficult or impossible to accomplish reliably. Recent improvements in wildlife cameras and ecological modelling have allowed for improved accuracy in estimates of abundance. In this study, we paired nest captures and high-definition nest video camera monitoring with modelling for a novel approach to estimate survival and abundance of threatened Piping Plovers Charadrius melodus breeding on Missouri River sandbars. From 2005 to 2014, we captured individuals on nests and uniquely marked them and recaptured previously marked individuals. In 2015–2017, we resighted marked individuals using small, high-definition video cameras deployed at nests, and counted the number of marked and unmarked breeding individuals associated with nests. We estimated apparent survival and derived estimates of the abundance of breeding individuals and population growth each year using a state-space Jolly–Seber superpopulation model with the addition of a binomial band ratio model for data collected using nest video cameras. Apparent survival averaged 0.73 ± 0.03 (mean ± sd) throughout the study. The number of breeding individuals varied, with the population increasing from 2012 to 2017 following a major habitat creation event. This study provides one of the few examples of camera data being used to produce demographic parameter and abundance estimates for an avian species. The camera and modelling methods described in this study may be applicable to other avian species in which some portion of the breeding population is uniquely marked.     

Nest-site selection and characteristics in a mixed-species colony of Avocets Recurvirostra avosetta and Black-winged Stilts Himantopus himantopus

Capsule Causes of breeding failure determined optimal nest location, Black-winged Stilts being better adapted than Avocets to nest close to water.

Aims To compare nest size and location between the two species and to test whether there were inter-species differences that might suggest specific nesting adaptations, and to investigate possible correlates between hatching success and nest location.

Methods Nest-site selection, characteristics and hatching success were studied during 1989 in a large mixed-species colony located in southern Spain.

Results Black-winged Stilt nests differed in composition and size depending on distance to water (nests touching the water were larger and included mud), but Avocet nests did not. Avocets nests were more aggregated and central within the colony than those of Black-winged Stilts. Causes of breeding failure were flooding and rat predation. Successful or unsuccessful Black-winged Stilt nests did not differ significantly in characteristics or location. Successful Avocet nests were further from water than flooded nests and further from dry land than depredated ones. Distance to the centre of the colony did not affect hatching success.

Conclusion Black-winged Stilts were better adapted than Avocets to nest near water, due to their greater plasticity in nest-building behaviour. Causes of breeding failure determined optimal nest location.