Predator recognition and differential behavioural responses of adult Wood Warblers Phylloscopus sibilatrix

This is the final version of the article. Available from Springer Verlag via the DOI in this record.Birds often engage in nest defence against predators to improve breeding success, but defence efficiency requires the capability to assess the threat level posed by potential predators. For species with low breeding-site tenacity, which may encounter varying occurrence and density of predators in different areas, threat recognition could be compromised due to naivety, and so predator recognition may focus on broad key features to diminish the risk of misidentification. We experimentally tested this hypothesis by recording behavioural reactions of the nomadic wood warbler Phylloscopus sibilatrix to objects reflecting various levels of threat: least weasel and Eurasian jay taxidermy mounts, an inanimate object and an empty display mount. To assess actual nest predators, we used remote cameras to record predation events at wood warbler nests. As in other studies in Western Europe, Eurasian jay was found to be the main nest predator, with occasional predation by least weasel. The reaction of adult warblers to the models was generally to remain silent and on nests during the incubation stage presumably due to the need to maintain efficient nest camouflage and concealment. During the nestling stage, behavioural responses of adult warblers, calling and suspended feeding of young, showed the strongest effects from the jay taxidermy mount, moderate to the weasel and weakest to the inanimate object and empty mount. As the reaction of wood warblers reflected the degree of genuine threat posed by the predators depicted by the models, we conclude that predator recognition may be present in this species.We thank John Mallord, Chris Orsman and Andrew Cristinacce for monitoring nests in the wider study area as part of a project jointly funded by RSPB and Natural England within the Action for Birds in England partnership, Louise and Gary Clewey for assistance in the field and landowners Natural England, Woodland Trust, National Trust and Devon Wildlife Trust

Breeding success of the great tit Parus major in relation to attributes of natural nest cavities in a primeval forest

An overlap in attributes of nest cavities used by Great Tit Parus major across Eurasia suggests similar nest site preferences within the geographical range, although the drivers of these preferences are unclear. To determine whether preferred cavities provide conditions enhancing successful reproduction, we investigated the breeding performance of Great Tits in relation to tree cavity characteristics using data collected during 2008–2011 in primeval conditions (Białowieża National Park, Poland). Here, tree cavities are diverse and superabundant but nesting birds are at risk from a variety of predators. According to expectations, nest losses were high (60 % of Great Tit nests failed), mostly due to predation (69 % of nest failures). The risk of nest failure varied with nest cavity attributes. Compared to successful nests, failures were situated higher above the ground and placed closer to the cavity entrance. Very deep cavities with narrow entrances and strong livings walls provided effective protection against larger predators (e.g., martens, woodpeckers), unable to enter the cavity or pull out the contents. Yet, such holes were no barrier for the smallest predators (e.g., Forest Dormouse Dryomys nitedula), which were able to enter any Great Tit nest cavity and destroyed most of the nests. Avoiding small predators would give a selective advantage to the birds, but this seems hardly possible to achieve. We conclude that tree cavities preferred by the tits show a combination of properties which are a compromise for avoiding predation (the strongest selective pressure) and providing the minimum requirements (sufficient nest illumination, microclimate, protection against nest soaking) for development and growth of young

Microclimate in tree cavities and nest-boxes: implications for hole-nesting birds

The provision of nest-boxes is widely used as a conservation intervention to increase the availability of cavities for hole-nesting birds, particularly in managed forests, but it is uncertain whether nest-boxes are an appropriate substitute for tree cavities. Tree cavities and nest-boxes may differ in many aspects, including microclimate, but there are few data with which to examine this. We measured the air temperature and relative humidity in vacant tree cavities previously used by breeding marsh tits Poecile palustris (a non-excavating forest passerine) and in nest-boxes provided for this species that had similar dimensions to natural nest sites, and we compared values from both with ambient conditions. We examined how tree cavity characteristics influenced microclimate and if similar conditions were replicated in nest-boxes. Tree cavities, particularly those in thicker parts of trees, were more efficient thermal insulators, with temperature extremes dampened to a greater extent relative to ambient values. In contrast, the nest-boxes provided poor insulation with negligible buffering against ambient temperatures. Mean daily relative humidity was high (on average c. 90%) in tree cavities, which all had walls of living wood, and this averaged 24% higher than in nest-boxes at comparable ambient conditions (mean humidity 76–78%). These results support previous studies that incorporated various types of tree cavities and nest-boxes, indicating that the environment within nest-boxes differs significantly from that of tree cavities. We conclude that providing nest-boxes may affect microclimatic conditions available for cavity-users, which may have ecological implications for nesting birds

Social structure of coal tits Periparus ater in temperate deciduous forest

Analysis of the impacts of social structure on the behaviour and life history of birds is a rapidly developing area of ornithology. Such studies commonly focus on the Paridae (tits and chickadees) as a model group, but detailed assessment of the basic social structure is lacking for many parids, particularly in the non-breeding season. Such baseline information is essential for understanding the organisation of bird populations, including associations between individuals within social and spatial networks. We assessed the non-breeding (autumn-winter) and spring (i.e. breeding season) social structure of Coal Tits Periparus ater in broadleaved forest in southern Britain. Coal Tits were found to be resident and sedentary in this habitat, with birds remaining close to their spring territories during the non-breeding season and associating in small groups averaging 3-4 individuals. Associations were variable, however, with no evidence of stable flock membership, group territoriality or strong pair bonds during autumn-winter. The non-breeding social structure did not change between years of high and low population density, being most similar to a 'basic flock' organisation. This differed from that expected of a food-hoarding parid and contradicted some earlier classifications, highlighting the potential plasticity of social behaviour between bird populations. Such variation can inform the understanding of the evolution of avian social structures and associated behavioural traits, such as food hoarding, and aid the interpretation of intraand inter-specific differences in behaviour

Multi-scale settlement patterns of a migratory songbird in a European primeval forest

The drivers of animal settlement are core topics in ecology. Studies from primaeval habitats provide valuable but rare insights into natural settlement behaviour, where species are unconstrained by habitat fragmentation and modification. We examined whether territorial male songbirds (wood warblers Phylloscopus sibilatrix) exhibited clustered distributions when settling in extensive primaeval forest. We tested whether settlement patterns were consistent between years, the influence of habitat preference and the spatial extent of these processes. Remote sensing (airborne LiDAR and satellite multi-spectral) and field survey data were combined to investigate settlement at the landscape (tens of km2), plot (46–200 ha) and microhabitat (20 m diameter) scales, by sampling across 5.9 × 4.5 km of the Białowieża National Park (Poland) during 2018–2019. We assessed bird distribution patterns using nearest neighbour analyses, and habitat preference with generalized linear mixed models. Variation in bird density between plots and years was suggestive of large-scale clustering at the landscape scale. At the local plot scale, any clustering was masked by territoriality, with birds appearing dispersed or randomly distributed. At the microhabitat scale within territories, birds preferred a tall, closed canopy forest structure and avoided a dense herb layer of wild garlic patches. Settlement appeared unconstrained by the availability of preferred habitat at the plot scale, which remained unsaturated. Wood warbler settlement apparently involved hierarchical decisions at multiple spatial scales, compatible with potential social attraction, territoriality, habitat preference or predator avoidance. Selecting an appropriate scale of analysis is an important consideration when assessing the clustering of territorial species

The red-billed leiothrix (Leiothrix lutea): a new invasive species for Britain?

Invasive non-native species (INNS) are one of the major threats to global biodiversity. Climate change and garden bird-feeding may facilitate the establishment of INNS. The Red-billed Leiothrix Leiothrix lutea is a small woodland passerine, native to subtropical Asia, that is an INNS in several parts of the world, including Europe, following escapes from the aviculture trade. Recently, populations of Red-billed Leiothrix have established in Spain, Portugal, Italy and France. Previous studies have shown that much of Europe is suitable for the species, including southern Britain. The Red-billed Leiothrix has not previously been considered as at high risk of establishment in Britain, but we document recent records, including a cluster in southern England, suggesting that establishment may already be underway. We discuss the potential negative impacts of this invasive species on local ecosystems, and how a warming climate and garden bird-feeding might assist their establishment in temperate Europe. We also highlight the importance of improved recording to monitor the potential establishment of INNS, such as the Red-billed Leiothrix, in Britain

An efficient survey method for estimating populations of marsh tits Poecile palustris, a low-density woodland passerine

Capsule: A playback survey comprising two visits to woodland in early spring can reliably detect Marsh Tits Poecile palustris and permit reasonable estimation of the number of territories. Aims: To assess the efficacy of an efficient survey method for detecting and estimating populations of Marsh Tits. Methods: Detection probability of colour-marked Marsh Tits, surveyed using playback, was assessed with Cormack–Jolly–Seber models. Reliability of territory estimates was compared between colour-marked and largely or wholly unmarked populations. Results: Playback surveys over two visits in early spring were highly effective in detecting individual Marsh Tits. Territory estimates were similar for woods in years where less than 20% of the population was marked compared to years when a higher proportion of birds were colour-marked, although territories may be underestimated in larger woods with unmarked populations. Conclusion: A playback survey comprising two visits in early spring is recommended as an efficient method of surveying Marsh Tit populations and locating individuals. A survey protocol is suggested

Temporal avoidance as a means of reducing competition between sympatric species

Human activity has modified the availability of natural resources and the abundance of species that rely on them, potentially changing interspecific competition dynamics. Here, we use large-scale automated data collection to quantify spatio-temporal competition among species with contrasting population trends. We focus on the spatial and temporal foraging behaviour of subordinate marsh tits Poecile palustris among groups of socially and numerically dominant blue tits Cyanistes caeruleus and great tits Parus major. The three species exploit similar food resources in mixed groups during autumn–winter. Using 421 077 winter recordings of individually marked birds at 65 automated feeding stations in Wytham Woods (Oxfordshire, UK), we found that marsh tits were less likely to join larger groups of heterospecifics, and they accessed food less frequently in larger groups than in smaller ones. Marsh tit numbers within groups declined throughout the diurnal and winter periods, while the number of blue and great tits increased. However, sites that attracted larger groups of these heterospecifics also attracted more marsh tits. The results suggest that subordinate species exhibit temporal avoidance of socially and numerically dominant heterospecifics, but have limited ability for spatial avoidance, indicating that behavioural plasticity enables only a partial reduction of interspecific competition

Thermal ecosystem engineering by songbirds promotes a symbiotic relationship with ants

Nesting birds can act as thermal ecosystem engineers by providing warm habitats that may attract arthropods to colonise the nest structure. This cohabitation of birds and nest-dwelling invertebrates may foster symbiotic relationships between them, but evidence is lacking. We investigated whether ants are attracted to bird nests by the heat generated by the hosts, and/or the nests’ structural insulation properties, to raise their broods (larvae and/or pupae) in advantageous thermal conditions. We found that the endothermic activity of birds within their nests created ‘heat islands’, with thermal conditions potentially promoting the survival and development of ant larvae in cool environments. We experimentally confirmed that the presence of heat within bird nests, and not the structure itself, attracted the ants to colonise the nests. As ants might benefit from exploiting warm bird nests, this may be a previously overlooked commensal, mutualistic or parasitic relationship which may be ecologically significant and globally widespread among various nesting birds and reproducing ants. Similar interspecific interactions may exist with other arthropods that reproduce in avian and mammalian nests. Further research is needed to reveal the nature of these relationships between such taxa, and to understand the role of warm-blooded animals as thermal ecosystem engineers