Wood warbler population dynamics in response to mast seeding regimes in Europe.

Mast seeding is the episodic, massive production of plant seeds synchronized over large areas. The resulting superabundance of seeds represents a resource pulse that can profoundly affect animal populations across trophic levels. Following years of high seed production, the abundance of both seed consumers and their predators increase. Higher predator abundance leads to increased predation pressure across the trophic web, impacting nonseed consumers such as the wood warbler Phylloscopus sibilatrix through increased nest predation after tree mast years. Over the past 30 years, the frequency of tree seed masts has increased, while wood warbler populations have declined in several regions of Europe. We hypothesized that increasing mast frequencies may have contributed to the observed population declines by creating suboptimal breeding conditions in years after masting. We measured reproductive output in four study areas in central Europe, which was between 0.61 and 1.24 fledglings lower in the years following masting than nonmasting. For each study area, we used matrix population models to predict population trends based on the estimated reproductive output and the local mast frequencies. We then compared the predicted with the observed population trends to assess if the frequency of mast years had contributed to the population dynamics. In Wielkopolska National Park (PL) and Hessen (DE), masting occurred on average only every 4 years and populations were stable or nearly so, whereas in Jura (CH) and Bialowieza National Park (PL), masting occurred every 2 and 2.5 years, respectively, and populations were declining. The simple matrix population models predicted the relative difference among local population trends over the past 10-20 years well, suggesting that the masting frequency may partly explain regional variation in population trends. Simulations suggest that further increases in mast frequency will lead to further declines in wood warbler populations. We show that changes in a natural process, such as mast seeding, may contribute to the decline in animal populations through cascading effects.

Songbird nests on the ground as islands of diversity of ptyctimous mites (Acari: Oribatida) in the primeval Bialowieza Forest (Poland).

Due to specific microclimatic conditions and accumulation of organic matter, bird nests are microhabitats that are often inhabited by various invertebrates, including mites (Acari). We tested whether nests of the ground-nesting passerine Phylloscopus sibilatrix (Bechstein) (Passeriformes: Phylloscopidae) [wood warbler] were associated with an increased local diversity of ptyctimous mites (Acari: Oribatida) on the forest floor in the Bialowieza National Park, East Poland. In 2019-2020, we analysed 150 warbler nests shortly after they had been vacated by the birds, and additionally we sampled mites in leaf litter at 1 and 6 m distances from 21 of the collected nests. We found on average more mite species in nests sampled in 2020 than in nests sampled in 2019. Although the species composition largely overlapped between bird nests and the litter, bird nests contained a greater average number of mite species than litter samples, including species found only - Microtritia minima (Berlese) and Phthiracarus crenophilus Willmann - or mostly - Euphthiracarus cribrarius (Berlese) and Phthiracarus globosus (C.L. Koch) - in bird nests. The results suggest that the presence of bird nests may increase the local diversity of the invertebrate species assemblage.

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.

Slow development of woodland vegetation and bird communities during 33 years of passive rewilding in open farmland.

Passive rewilding is a potential tool for expanding woodland cover and restoring biodiversity by abandoning land management and allowing natural vegetation succession to occur. Land can be abandoned to passive rewilding deliberately or due to socio-economic change. Despite abandonment being a major driver of land use change, few have studied the long-term outcomes for vegetation and biodiversity in Western Europe. Studies are also biased towards sites that are close to seed sources and favourable to woodland colonisation. In this case-study, we reconstruct a time series of passive rewilding over 33 years on 25 ha of former farmland that had been subject to soil tipping, far from woodland seed sources. Natural colonisation by shrubs and trees was surveyed at three points during the time series, using field mapping and lidar. Breeding birds were surveyed at three time points, and compared with surveys from nearby farmland. Results showed that natural colonisation of woody vegetation was slow, with open grassland dominating the old fields for two decades, and small wetlands developing spontaneously. After 33 years, thorny shrub thickets covered 53% of the site and former hedgerows became subsumed or degraded, but trees remained scarce. However, the resulting habitat mosaic of shrubland, grassland and wetland supported a locally distinctive bird community. Farmland bird species declined as passive rewilding progressed, but this was countered by relatively more wetland birds and an increase in woodland birds, particularly songbirds, compared to nearby farmland. Alongside biodiversity benefits, shrubland establishment by passive rewilding could potentially provide ecosystem services via abundant blossom resources for pollinators, and recreation and berry-gathering opportunities for people. Although closed-canopy woodland remained a distant prospect even after 33 years, the habitat mosaic arising from passive rewilding could be considered a valuable outcome, which could contribute to nature recovery and provision of ecosystem services.

Weather impacts on interactions between nesting birds, nest-dwelling ectoparasites and ants.

Weather has a dominant impact on organisms, including their life histories and interspecific interactions. Yet, for nesting birds, and the arthropods inhabiting bird nests, the direct and cascading effects of weather are poorly known. We explored the influence of ambient temperatures and rainfall on the cohabitation of dome-shaped bird nests by Wood Warblers Phylloscopus sibilatrix, their blowfly Protocalliphora azurea ectoparasites, and predatory Myrmica and Lasius ants that may provide nest sanitation. We sampled blowflies and ants in 129 nests, and measured warbler nestlings during 2018-2020 in the primeval Bialowieza Forest, eastern Poland. The probability of ectoparasites occurring in nests increased with increasing ambient temperatures and declining precipitation in the early nestling stage, when adult blowflies are ovipositing. Where present, the number of ectoparasites was greater if higher ambient temperatures had prevailed in the late nestling stage, but only when ants were absent from nests. However, the nestling growth was unrelated to ectoparasite abundance or ant presence within bird nests, although it was lower at high rainfall. The results suggest that weather can have conflicting impacts on interactions between nesting birds and nest-dwelling arthropods, but birds can mostly compensate for any related costs in old-growth forest, where food is generally abundant.

Interspecific attraction between ground-nesting songbirds and ants: the role of nest-site selection.

BACKGROUND: Interspecific interactions within ecological networks can influence animal fitness and behaviour, including nest-site selection of birds and ants. Previous studies revealed that nesting birds and ants may benefit from cohabitation, with interspecific attraction through their nest-site choice, but mutual interactions have not yet been tested. We explored a previously undescribed ecological link between ground-nesting birds and ants raising their own broods (larvae and pupae) within the birds' nests in a temperate primeval forest of lowland Europe. We tested whether the occurrence of ant broods within bird nests resulted from a mutual or one-sided interspecific attraction that operated through nest-site choice and was modified by weather conditions. RESULTS: We found a non-random occupation of bird nests by ants raising their own broods within them, which indicated interspecific attraction driven solely by the ants. The birds' preference to nest near tussocks of vegetation showed little overlap with the most frequent placement of ant colonies among fallen deciduous tree-leaves, dead wood and moss. Additionally, birds did not appear to select forest localities with high densities of ant colonies. The occurrence of ant broods within bird nests was also unrelated to bird nest placement near to specific habitat features. The attractiveness of bird nests to ants appeared to increase with the thermal activity of the birds warming their nests, and also during cool and wet weather when the occurrence of ant broods within bird nests was most frequent. Ants often remained in the nests after the birds had vacated them, with only a slight reduction in the probability of ant brood occurrence over time. CONCLUSIONS: The natural patterns of bird nest colonisation by ants support the hypothesis of ants' attraction to warm nests of birds to raise their broods under advantageous thermal conditions. Similar relationships may occur between other warm-blooded, nest-building vertebrates and nest-dwelling invertebrates, which depend on ambient temperatures. The findings advance our understanding of these poorly recognised interspecific interactions, and can inform future studies of ecological networks.

Long-term woodland restoration on lowland farmland through passive rewilding.

Natural succession of vegetation on abandoned farmland provides opportunities for passive rewilding to re-establish native woodlands, but in Western Europe the patterns and outcomes of vegetation colonisation are poorly known. We combine time series of field surveys and remote sensing (lidar and photogrammetry) to study woodland development on two farmland fields in England over 24 and 59 years respectively: the New Wilderness (2.1 ha) abandoned in 1996, and the Old Wilderness (3.9 ha) abandoned in 1961, both adjacent to ancient woodland. Woody vegetation colonisation of the New Wilderness was rapid, with 86% vegetation cover averaging 2.9 m tall after 23 years post-abandonment. The Old Wilderness had 100% woody cover averaging 13.1 m tall after 53 years, with an overstorey tree-canopy (≥ 8 m tall) covering 91%. By this stage, the structural characteristics of the Old Wilderness were approaching those of neighbouring ancient woodlands. The woody species composition of both Wildernesses differed from ancient woodland, being dominated by animal-dispersed pedunculate oak Quercus robur and berry-bearing shrubs. Tree colonisation was spatially clustered, with wind-dispersed common ash Fraxinus excelsior mostly occurring near seed sources in adjacent woodland and hedgerows, and clusters of oaks probably resulting from acorn hoarding by birds and rodents. After 24 years the density of live trees in the New Wilderness was 132/ha (57% oak), with 390/ha (52% oak) in the Old Wilderness after 59 years; deadwood accounted for 8% of tree stems in the former and 14% in the latter. Passive rewilding of these 'Wilderness' sites shows that closed-canopy woodland readily re-established on abandoned farmland close to existing woodland, it was resilient to the presence of herbivores and variable weather, and approached the height structure of older woods within approximately 50 years. This study provides valuable long-term reference data in temperate Europe, helping to inform predictions of the potential outcomes of widespread abandonment of agricultural land in this region.

Lack of specialist nidicoles as a characteristic of mite assemblages inhabiting nests of the ground-nesting wood warbler, Phylloscopus sibilatrix (Aves: Passeriformes).

Bird and mammal nests provide microhabitats that support a range of other species, including invertebrates. However, the variation between communities of nest-dwelling invertebrates in different nests is poorly understood. The major aim of this study was to analyze the assemblage structure of mites from the suborder Uropodina (Acari: Mesostigmata) and from superfamily Crotonioidea (Acari: Oribatida) inhabiting nests of the wood warbler, Phylloscopus sibilatrix (Aves: Passeriformes), located on a forest floor in Bialowieza Forest, in eastern Poland. We also assessed the correlation between the nest material used by the birds with the assemblage structure of Uropodina mites, and compared the results with published studies of the nests of other birds and a mammal (common mole, Talpa europaea), and also with communities of mites inhabiting the soil. The field research was conducted in the strict nature reserve of the Bialowieza National Park, a near-primeval European temperate forest. In 2019, immediately after the breeding period, 69 wood warbler nests and 439 soil samples were collected. Analyses revealed assemblages of Uropodina mites inhabiting the nests that consisted of 14 species, mostly common soil species. Only five species of oribatid mites from superfamily Crotonioidea were present in the nest material. Analyzed nests had a high percentage of tree leaves and grass blades, whereas moss was the least frequent component of the nest material. The Uropodina mites were more abundant in the nests that had greater amounts of grass blades, but similar relationships were insignificant for the nests with varying amounts of tree leaves or moss. The assemblages of Uropodina mites inhabiting wood warbler nests were very similar to those found in soil and nests of the common mole, but they lacked typical nest-dwelling species of Uropodina (i.e., specialized nidicoles).

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.

Breeding birds actively modify the initial microclimate of occupied tree cavities.

The microclimate of cavities used by endothermic animals may depend on dynamic relationships between a cavity's physical properties and the heating activity of cavity users, but the rudiments of these relationships are unclear. I compared the temperature and relative humidity of active tree cavities that were occupied by nesting marsh tits Poecile palustris with the conditions in vacant tree cavities previously used for breeding by this species. I tested how presence of active nests modified initial cavity microclimate, and if this modification changed with nest progression or cavity insulation. In 2013-2014, mean daily internal-ambient temperature differences averaged 1.5-4.1 °C higher and relative humidity 8-10% lower, in active cavities relative to vacant sites, with greatest differences in the late nestling period. Compared to vacant cavities and relative to respective ambient values, the greatest daily minimum temperature increase was in active cavities located in the thinnest trees, which insulated least efficiently. As daily minimum temperatures were elevated to a similar level relative to outside within all active cavities, birds appeared to compensate for heat loss from cavities by warming the air within in a homeostatic manner. Similar to vacant cavities, the differences between daily maximum internal and ambient temperatures decreased with tree girth in active cavities, indicating that daily temperature maxima were systematically moderated in the thickest trees. The study demonstrates the modifying effect of birds' breeding activity on tree-cavity microclimate and highlights the role of a cavity's thermal properties in reducing the energy expenditure and risk of overheating for cavity users.

Predator recognition and differential behavioural responses of adult wood warblers Phylloscopus sibilatrix.

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.