GPS-telemetry unveils the regular high-elevation crossing of the Himalayas by a migratory raptor: implications for definition of a "Central Asian Flyway".

Remote technologies are producing leapfrog advances in identifying the routes and connectivity of migratory species, which are still unknown for hundreds of taxa, especially Asian ones. Here, we used GPS-telemetry to uncover the migration routes and breeding areas of the massive population of migratory Black-eared kites wintering around the megacity of Delhi-India, which hosts the largest raptor concentration of the world. Kites migrated for 3300-4800 km along a narrow corridor, crossing the Himalayas at extremely high elevations (up to > 6500 m a.s.l.) by the K2 of the Karakoram Range and travelled long periods at elevations above 3500 m. They then crossed/circumvented the Taklamakan Desert and Tian Shan Range to reach their unknown breeding quarters at the intersection between Kazakhstan, Russia, China and Mongolia. Route configuration seemed to be shaped by dominant wind support and barrier avoidance. Wintering ranges were smaller than breeding ranges and concentrated around Delhi, likely in response to massive human food-subsidies. Our results illustrate that high-elevation crossings by soaring migrants may be more common than previously appreciated and suggest the delineation of a hitherto poorly-appreciated "Central Asian Flyway", which must funnel hundreds of thousands of migrants from central Asia into the Indian subcontinent via multiple modes of the Himalayan crossing.

The population density of an urban raptor is inextricably tied to human cultural practices.

Human socio-cultural factors are recognized as fundamental drivers of urban ecological processes, but their effect on wildlife is still poorly known. In particular, human cultural aspects may differ substantially between the extensively studied urban settings of temperate regions and the poorly studied cities of the tropics, which may offer profoundly different niches for urban wildlife. Here, we report how the population levels of a scavenging raptor which breeds in the megacity of Delhi, the black kite Milvus migrans, depend on spatial variation in human subsidies, mainly in the form of philanthropic offerings of meat given for religious purposes. This tight connection with human culture, which generated the largest raptor concentration in the world, was modulated further by breeding-site availability. The latter constrained the level of resource-tracking by the kites and their potential ecosystem service, and could be used as a density-management tool. Similar ties between animal population densities, key anthropogenic resources and human beliefs may occur in thousands of cities all over the globe and may fit poorly with our current understanding of urban ecosystem functioning. For many urban animals, key resources are inextricably linked with human culture, an aspect that has been largely overlooked.

Human-attacks by an urban raptor are tied to human subsidies and religious practices.

Growing urbanization is increasing human-wildlife interactions, including attacks towards humans by vertebrate predators, an aspect that has received extremely scarce investigation. Here, we examined the ecological, landscape and human factors that may promote human-aggression by raptorial Black kites Milvus migrans in the 16-millions inhabitants megacity of Delhi (India). Physical attacks depended on human activities such as unhygienic waste management, ritual-feeding of kites (mainly operated by Muslims), human density, and presence of a balcony near the nest, suggesting an association between aggression and frequent-close exposure to humans and derived food-rewards. Surprisingly, while more than 100,000 people could be at risk of attack in any given moment, attitudes by local inhabitants were strikingly sympathetic towards the birds, even by injured persons, likely as a result of religious empathy. These results highlight the importance of socio-cultural factors for urban biota and how these may radically differentiate the under-studied cities of developing countries from those of western nations, thus broadening our picture of human-wildlife interactions in urban environments. The rapid sprawling of urban and suburban areas with their associated food-subsidies is likely to increase proximity and exposure of large predators to humans, and vice versa, leading to heightened worldwide conflicts.

Offspring defense by an urban raptor responds to human subsidies and ritual animal-feeding practices.

There is a growing interest in the behavioural and life history mechanisms that allow animal species to cope with rapidly expanding urban habitats, which impose frequent proximity to humans. A particular case of behavioral bottleneck (i.e. conflicting interests) faced by animals in urban environments is how they will modulate the defence of their offspring against the potential danger represented by humans, an aspect that has received scarce research attention. We examined the nest defense against humans by a dense breeding population of a raptor, the Black Kite Milvus migrans, within the megacity of Delhi (India). Here, kites live on a diet dominated by human waste and meat offered through religiously motivated bird feeding practices. Nest defense levels increased with the number of offspring, and with the progression of the breeding season. Defense also intensified close to ritual-feeding areas and with increasing human waste in the streets, suggesting synergistic effects of food availability, parental investment, personality-boldness and habituation to humans, with consequent attenuation of fear. Thus, the behavioural response to a perceived threat reflected the spatial mosaic of activity of humans in the city streets, their cultural practices of ritual-feeding, and their waste-management. For synurbic species, at the higher-end spectrum of adaptation to an urban life, human cultural practices and attitudes may well be the most defining dimensions of their urban niche. Our results suggest that, after initial urban colonization, animals may continue to adapt to the typically complex, heterogeneous environments of cities through fine-grained behavioural adjustments to human practices and activities.

Effects of interspecific coexistence on laying date and clutch size in two closely related species of hole-nesting birds.

Coexistence between great tits Parus major and blue tits Cyanistes caeruleus, but also other hole-nesting taxa, constitutes a classic example of species co-occurrence resulting in potential interference and exploitation competition for food and for breeding and roosting sites. However, the spatial and temporal variations in coexistence and its consequences for competition remain poorly understood. We used an extensive database on reproduction in nest boxes by great and blue tits based on 87 study plots across Europe and Northern Africa during 1957-2012 for a total of 19,075 great tit and 16,729 blue tit clutches to assess correlative evidence for a relationship between laying date and clutch size, respectively, and density consistent with effects of intraspecific and interspecific competition. In an initial set of analyses, we statistically controlled for a suite of site-specific variables. We found evidence for an effect of intraspecific competition on blue tit laying date (later laying at higher density) and clutch size (smaller clutch size at higher density), but no evidence of significant effects of intraspecific competition in great tits, nor effects of interspecific competition for either species. To further control for site-specific variation caused by a range of potentially confounding variables, we compared means and variances in laying date and clutch size of great and blue tits among three categories of difference in density between the two species. We exploited the fact that means and variances are generally positively correlated. If interspecific competition occurs, we predicted a reduction in mean and an increase in variance in clutch size in great tit and blue tit when density of heterospecifics is higher than the density of conspecifics, and for intraspecific competition, this reduction would occur when density of conspecifics is higher than the density of heterospecifics. Such comparisons of temporal patterns of means and variances revealed evidence, for both species, consistent with intraspecific competition and to a smaller extent with interspecific competition. These findings suggest that competition associated with reproductive behaviour between blue and great tits is widespread, but also varies across large spatial and temporal scales.

Recent natural selection causes adaptive evolution of an avian polygenic trait.

We used extensive data from a long-term study of great tits (Parus major) in the United Kingdom and Netherlands to better understand how genetic signatures of selection translate into variation in fitness and phenotypes. We found that genomic regions under differential selection contained candidate genes for bill morphology and used genetic architecture analyses to confirm that these genes, especially the collagen gene COL4A5, explained variation in bill length. COL4A5 variation was associated with reproductive success, which, combined with spatiotemporal patterns of bill length, suggested ongoing selection for longer bills in the United Kingdom. Last, bill length and COL4A5 variation were associated with usage of feeders, suggesting that longer bills may have evolved in the United Kingdom as a response to supplementary feeding.

Interspecific variation in the relationship between clutch size, laying date and intensity of urbanization in four species of hole-nesting birds.

The increase in size of human populations in urban and agricultural areas has resulted in considerable habitat conversion globally. Such anthropogenic areas have specific environmental characteristics, which influence the physiology, life history, and population dynamics of plants and animals. For example, the date of bud burst is advanced in urban compared to nearby natural areas. In some birds, breeding success is determined by synchrony between timing of breeding and peak food abundance. Pertinently, caterpillars are an important food source for the nestlings of many bird species, and their abundance is influenced by environmental factors such as temperature and date of bud burst. Higher temperatures and advanced date of bud burst in urban areas could advance peak caterpillar abundance and thus affect breeding phenology of birds. In order to test whether laying date advance and clutch sizes decrease with the intensity of urbanization, we analyzed the timing of breeding and clutch size in relation to intensity of urbanization as a measure of human impact in 199 nest box plots across Europe, North Africa, and the Middle East (i.e., the Western Palearctic) for four species of hole-nesters: blue tits (Cyanistes caeruleus), great tits (Parus major), collared flycatchers (Ficedula albicollis), and pied flycatchers (Ficedula hypoleuca). Meanwhile, we estimated the intensity of urbanization as the density of buildings surrounding study plots measured on orthophotographs. For the four study species, the intensity of urbanization was not correlated with laying date. Clutch size in blue and great tits does not seem affected by the intensity of urbanization, while in collared and pied flycatchers it decreased with increasing intensity of urbanization. This is the first large-scale study showing a species-specific major correlation between intensity of urbanization and the ecology of breeding. The underlying mechanisms for the relationships between life history and urbanization remain to be determined. We propose that effects of food abundance or quality, temperature, noise, pollution, or disturbance by humans may on their own or in combination affect laying date and/or clutch size.

Variation in clutch size in relation to nest size in birds.

Nests are structures built to support and protect eggs and/or offspring from predators, parasites, and adverse weather conditions. Nests are mainly constructed prior to egg laying, meaning that parent birds must make decisions about nest site choice and nest building behavior before the start of egg-laying. Parent birds should be selected to choose nest sites and to build optimally sized nests, yet our current understanding of clutch size-nest size relationships is limited to small-scale studies performed over short time periods. Here, we quantified the relationship between clutch size and nest size, using an exhaustive database of 116 slope estimates based on 17,472 nests of 21 species of hole and non-hole-nesting birds. There was a significant, positive relationship between clutch size and the base area of the nest box or the nest, and this relationship did not differ significantly between open nesting and hole-nesting species. The slope of the relationship showed significant intraspecific and interspecific heterogeneity among four species of secondary hole-nesting species, but also among all 116 slope estimates. The estimated relationship between clutch size and nest box base area in study sites with more than a single size of nest box was not significantly different from the relationship using studies with only a single size of nest box. The slope of the relationship between clutch size and nest base area in different species of birds was significantly negatively related to minimum base area, and less so to maximum base area in a given study. These findings are consistent with the hypothesis that bird species have a general reaction norm reflecting the relationship between nest size and clutch size. Further, they suggest that scientists may influence the clutch size decisions of hole-nesting birds through the provisioning of nest boxes of varying sizes.

Calcium effects on life-history traits in a wild population of the great tit (Parus major): analysis of long-term data at several spatial scales.

Calcium is an essential micronutrient for birds during egg formation and for skeletal development in nestlings. Habitat level studies suggest that birds breeding in low-calcium areas may be limited in the size or number of eggs they lay and in the quality of their nestlings. However, as birds forage non-randomly and may travel considerable distances to acquire calcium, describing different breeding environments in terms of their calcium availability is problematic. Here we explore the spatial relationships between 300-fold variation in soil calcium and the life-history traits of ca. 6,000 pairs of great tits breeding in a single continuous woodland over 41 years. Controlling for other habitat differences, we found strong positive associations between soil calcium, clutch size and recruitment at spatial scales of over 300 m from each nestbox, suggesting that females may have been travelling inter-territorially to acquire calcium during egg formation. Soil calcium near each nestbox (mean distance = 58 m) was a strong positive predictor of mean fledgling mass, suggesting that local calcium was more important during nestling stages. We found no effect of soil calcium on lay-date or egg mass. This study is the first to provide evidence that small woodland passerines are limited by calcium availability at several different spatial scales. However, experimental work is necessary to test the causality of these spatial patterns.

Edge effects in the great tit: analyses of long-term data with GIS techniques.

In contemporary fragmented landscapes, edges are commonplace, and understanding the effects of edge environments is thus essential for the conservation of forest communities. The reproductive output of forest passerines is often reduced close to forest edges. Possible explanations include overcrowding by conspecifics, elevated rates of predation, and the occurrence of lower-quality habitat and/or individuals at forest edges. We attempted to separate these processes by examining edge effects in the absence of nest predation and by effectively controlling for differences in breeding density and the quality of habitats and individuals. We used an edge distance index (EDI), which accounts for the number and distribution of edges in close proximity to a breeding location, to help explain variation in breeding density, nesting success, and reproductive traits of 8308 pairs of Great Tits (Parus major) breeding between 1965 and 2005, in Wytham, near Oxford, United Kingdom. Results from linear mixed modeling confirmed higher breeding density and a higher proportion of immigrant individuals at forest edges. Nevertheless, independently of these effects, we also found that birds laying later, with smaller clutches but larger eggs, were typical of edge environments. The number of offspring recruited to the breeding offspring per breeding attempt was also reduced at edges, both directly and mediated through changes in clutch size and laying date. Edge effects on life histories were detectable within individual females and up to 500 m from the woodland edge. Woodland edges are increasingly common in contemporary fragmented landscapes. Therefore these results, which suggest a pervasive effect of edges on reproduction, are of considerable importance to the management and conservation of forest communities.

Speckled eggs: water-loss and incubation behaviour in the great tit Parus major.

Many small passerine birds worldwide lay white eggs speckled with red, brown and black protoporphyrin pigment spots (maculation). Unlike some patterns of avian eggshell pigmentation which clearly serve a crypsis or signalling function, the ubiquity of maculation among passerines suggests that its origins lie in another function, not specific to any particular ecological or behavioural group. Elsewhere, we have presented evidence that protoporphyrin pigments serve a structural function related to eggshell thickness and calcium availability: eggshell maculation in the great tit Parus major increases with decreasing soil calcium levels, pigments demarcate thinner areas of shell, and both the pigment intensity and distribution are related to shell thickness. Here we show that maculation also affects the rate of water loss from the egg during incubation (approximately Mass Loss per Day or MLD, which is critical to egg viability), but not that of unincubated eggs. We also demonstrate, both by observation and experiment, that the effect of female incubation behaviour on MLD compensates in some way for variation in egg characteristics, and that differences between females in the degree of such compensation are related to differences in clutch maculation. Our results suggest that, while a principal function of maculation in this species may be to strengthen the eggshell, it may also reduce eggshell permeability when large amounts of pigment are used, and that this necessitates a behavioural adjustment from the female during incubation. We discuss these findings and make further testable predictions from our model.

Density effects on life-history traits in a wild population of the great tit Parus major: analyses of long-term data with GIS techniques.

1. Population density often has strong effects on the population dynamics and reproductive processes of territorial animals. However, most estimates of density-dependent effects use the number of breeding pairs per unit area in a given season and look for correlations across seasons, a technique that assigns the same density score to each breeding pair, irrespective of local spatial variation. 2. In this study, we employed GIS techniques to estimate individual breeding densities for great tits breeding in Wytham Woods UK, between 1965 and 1996. We then used linear mixed modelling to analyse the effect of density on reproductive processes. 3. The areas of Thiessen polygons formed around occupied nestboxes were used to approximate territory size (necessarily inverse of breeding density). There were significant, independent and positive relationships between clutch size, fledging mass and the number of offspring recruited to the population, and territory size (all P < 0.001), but no effect of territory size on lay-date or egg mass. 4. Thiessen polygons are contiguous and cover all of the available area. Therefore, at low nest densities territory polygons were excessively oversized. Using a novel procedure to address this limitation, territory sizes were systematically capped through a range of maxima, with the greatest effect in the models when territories were capped at 0.9-2.3 ha. This figure approximates to the maximum effective territory size in our population and is in close agreement with several field-based studies. This capping refinement also revealed a significant negative relationship between lay-date and territory size capped at 0.9 ha (P < 0.001). 5. These density-dependent effects were also detected when analyses were restricted to changes within individual females, suggesting that density effects do not merely result from either increased proportions of low-quality individuals, or increased occupation of poor sites, when population density is high. 6. Overall, these results suggest that, in the current population, great tits with territories smaller than c. 2 ha independently lay smaller and later clutches, have lighter fledglings, and recruit fewer offspring to the breeding population. These analyses thus suggest a pervasive and causal role of local population density in explaining individual reproductive processes.