Integument colouration and circulating carotenoids in relation to urbanisation in Eurasian kestrels (Falco tinnunculus).

Urbanisation is one of the biggest environmental challenges of our time, yet we still lack an integrative understanding of how cities affect behaviour, physiology and parasite susceptibility of free-living organisms. In this study, we focus on carotenoids, strictly dietary micronutrients that can either be used as yellow-red pigments, for integument colouration (signalling function), or as antioxidants, to strengthen the immune system (physiological function) in an urban predator, the Eurasian kestrel (Falco tinnunculus). Kestrels are specialised vole hunters but shift to avian prey in cities where diurnal rodents are not sufficiently available. This different foraging strategy might determine the quantity of carotenoids available. We measured integument colouration, circulating carotenoids in the blood and ectoparasite burden in kestrels along an urban gradient. Our results showed that nestlings that were raised in more urbanised areas displayed, unrelated to their ectoparasite burden, a paler integument colouration. Paler colours were furthermore associated with a lower concentration of circulating carotenoids. These findings support the hypothesis that the entire urban food web is carotenoid deprived and only prey of low quality with low carotenoid content is available (e.g. fewer carotenoids in urban trees, insects, small birds and finally kestrels). The alternative hypothesis that nestlings allocate carotenoids to reduce physiological stress and/or to cope with parasites rather than invest into colouration could not be supported. Our study adds to existing evidence that urban stressors negatively affect carotenoid production in urban areas, a deficiency that dissipate into higher trophic levels.

Land-use intensification causes multitrophic homogenization of grassland communities.

Land-use intensification is a major driver of biodiversity loss. Alongside reductions in local species diversity, biotic homogenization at larger spatial scales is of great concern for conservation. Biotic homogenization means a decrease in ß-diversity (the compositional dissimilarity between sites). Most studies have investigated losses in local (α)-diversity and neglected biodiversity loss at larger spatial scales. Studies addressing ß-diversity have focused on single or a few organism groups (for example, ref. 4), and it is thus unknown whether land-use intensification homogenizes communities at different trophic levels, above- and belowground. Here we show that even moderate increases in local land-use intensity (LUI) cause biotic homogenization across microbial, plant and animal groups, both above- and belowground, and that this is largely independent of changes in α-diversity. We analysed a unique grassland biodiversity dataset, with abundances of more than 4,000 species belonging to 12 trophic groups. LUI, and, in particular, high mowing intensity, had consistent effects on ß-diversity across groups, causing a homogenization of soil microbial, fungal pathogen, plant and arthropod communities. These effects were nonlinear and the strongest declines in ß-diversity occurred in the transition from extensively managed to intermediate intensity grassland. LUI tended to reduce local α-diversity in aboveground groups, whereas the α-diversity increased in belowground groups. Correlations between the ß-diversity of different groups, particularly between plants and their consumers, became weaker at high LUI. This suggests a loss of specialist species and is further evidence for biotic homogenization. The consistently negative effects of LUI on landscape-scale biodiversity underscore the high value of extensively managed grasslands for conserving multitrophic biodiversity and ecosystem service provision. Indeed, biotic homogenization rather than local diversity loss could prove to be the most substantial consequence of land-use intensification.

Biodiversity at multiple trophic levels is needed for ecosystem multifunctionality.

Many experiments have shown that loss of biodiversity reduces the capacity of ecosystems to provide the multiple services on which humans depend. However, experiments necessarily simplify the complexity of natural ecosystems and will normally control for other important drivers of ecosystem functioning, such as the environment or land use. In addition, existing studies typically focus on the diversity of single trophic groups, neglecting the fact that biodiversity loss occurs across many taxa and that the functional effects of any trophic group may depend on the abundance and diversity of others. Here we report analysis of the relationships between the species richness and abundance of nine trophic groups, including 4,600 above- and below-ground taxa, and 14 ecosystem services and functions and with their simultaneous provision (or multifunctionality) in 150 grasslands. We show that high species richness in multiple trophic groups (multitrophic richness) had stronger positive effects on ecosystem services than richness in any individual trophic group; this includes plant species richness, the most widely used measure of biodiversity. On average, three trophic groups influenced each ecosystem service, with each trophic group influencing at least one service. Multitrophic richness was particularly beneficial for 'regulating' and 'cultural' services, and for multifunctionality, whereas a change in the total abundance of species or biomass in multiple trophic groups (the multitrophic abundance) positively affected supporting services. Multitrophic richness and abundance drove ecosystem functioning as strongly as abiotic conditions and land-use intensity, extending previous experimental results to real-world ecosystems. Primary producers, herbivorous insects and microbial decomposers seem to be particularly important drivers of ecosystem functioning, as shown by the strong and frequent positive associations of their richness or abundance with multiple ecosystem services. Our results show that multitrophic richness and abundance support ecosystem functioning, and demonstrate that a focus on single groups has led to researchers to greatly underestimate the functional importance of biodiversity.

Effects of forest structure on the interaction between avian hosts, dipteran vectors and haemosporidian parasites.

BACKGROUND: Forest habitats are important biodiversity refuges for a wide variety of bird species. Parasitism may modulate host species presence and abundance, and parasite effects can change according to forest management practices. Such processes are not well studied in vector-borne avian haemosporidians. We analyzed the effects of forest management on bird-dipteran-haemosporidian interactions, using seven common bird species in managed and unmanaged beech forest habitats in northeastern Germany. We assumed that forest structural heterogeneity affects parasite population parameters in avian hosts (i.e., prevalence and parasitemia), through its effect on the condition of the avian host but also through varying vector abundances. RESULTS: Parasite prevalence was high (about 80%) and homogeneous across different beech forest categories (i.e., young, old, unmanaged) and for all bird species, except Erithacus rubecula (35%). Parasitemia varied across bird species but not across forest categories within each avian species (lowest parasitemia were found in E. rubecula, Turdus merula, and Turdus philomelos). In our study system, we found that vector abundance was not the main driver of parasite dynamics. We found that forest structure affects parasite infection probability directly and potentially host condition via available resources that have to be used either to combat infections (i.e., high parasitemia) or to maintain a good body condition. CONCLUSIONS: The effects of each of the predictors were bird species-specific, and we found that Diptera vectors were not the foremost influence in our host-vector-parasite system. Effects of forest habitat variables indicated that for most bird species in this study, habitat regulation of infection probability was more likely (i.e., E. rubecula, Fringilla coelebs, Sylvia atricapilla), whereas for Parus major habitat characteristics impacted first individuals' body condition and subsequently the probability of infection. Our findings emphasize the need of species-specific analyses and to use continuous forest structural parameters (e.g., the proportion of gap, south facing aspect) to better understand habitat and land use effects on host-vector-parasite dynamics.

Specialisation and diversity of multiple trophic groups are promoted by different forest features.

While forest management strongly influences biodiversity, it remains unclear how the structural and compositional changes caused by management affect different community dimensions (e.g. richness, specialisation, abundance or completeness) and how this differs between taxa. We assessed the effects of nine forest features (representing stand structure, heterogeneity and tree composition) on thirteen above- and belowground trophic groups of plants, animals, fungi and bacteria in 150 temperate forest plots differing in their management type. Canopy cover decreased light resources, which increased community specialisation but reduced overall diversity and abundance. Features increasing resource types and diversifying microhabitats (admixing of oaks and conifers) were important and mostly affected richness. Belowground groups responded differently to those aboveground and had weaker responses to most forest features. Our results show that we need to consider forest features rather than broad management types and highlight the importance of considering several groups and community dimensions to better inform conservation.

Interannual variation in land-use intensity enhances grassland multidiversity.

Although temporal heterogeneity is a well-accepted driver of biodiversity, effects of interannual variation in land-use intensity (LUI) have not been addressed yet. Additionally, responses to land use can differ greatly among different organisms; therefore, overall effects of land-use on total local biodiversity are hardly known. To test for effects of LUI (quantified as the combined intensity of fertilization, grazing, and mowing) and interannual variation in LUI (SD in LUI across time), we introduce a unique measure of whole-ecosystem biodiversity, multidiversity. This synthesizes individual diversity measures across up to 49 taxonomic groups of plants, animals, fungi, and bacteria from 150 grasslands. Multidiversity declined with increasing LUI among grasslands, particularly for rarer species and aboveground organisms, whereas common species and belowground groups were less sensitive. However, a high level of interannual variation in LUI increased overall multidiversity at low LUI and was even more beneficial for rarer species because it slowed the rate at which the multidiversity of rare species declined with increasing LUI. In more intensively managed grasslands, the diversity of rarer species was, on average, 18% of the maximum diversity across all grasslands when LUI was static over time but increased to 31% of the maximum when LUI changed maximally over time. In addition to decreasing overall LUI, we suggest varying LUI across years as a complementary strategy to promote biodiversity conservation.

Land use intensification alters ecosystem multifunctionality via loss of biodiversity and changes to functional composition.

Global change, especially land-use intensification, affects human well-being by impacting the delivery of multiple ecosystem services (multifunctionality). However, whether biodiversity loss is a major component of global change effects on multifunctionality in real-world ecosystems, as in experimental ones, remains unclear. Therefore, we assessed biodiversity, functional composition and 14 ecosystem services on 150 agricultural grasslands differing in land-use intensity. We also introduce five multifunctionality measures in which ecosystem services were weighted according to realistic land-use objectives. We found that indirect land-use effects, i.e. those mediated by biodiversity loss and by changes to functional composition, were as strong as direct effects on average. Their strength varied with land-use objectives and regional context. Biodiversity loss explained indirect effects in a region of intermediate productivity and was most damaging when land-use objectives favoured supporting and cultural services. In contrast, functional composition shifts, towards fast-growing plant species, strongly increased provisioning services in more inherently unproductive grasslands.

Haemosporidian intensity and nestlings' life-history along an urban-to-rural gradient.

Urban areas, i.e. dense housing and reduced green spaces, can significantly impact avian health, through altering land use and increasing biotic and abiotic stress. This study assessed the association of urbanization on haemosporidian infections, vectors, immune response, and body condition in Parus major nestlings, across four classes of urbanization along an urban-to-rural gradient in Vienna, Austria. Contrary to our expectations, vector abundance remained consistent across the gradient, while an increase in leukocyte count is positively associated with total parasite intensity. We found that nestlings in more urbanized areas exhibited higher parasite intensity and altered immune response, as evidenced by variations in the heterophil to lymphocyte ratio and leukocyte counts. Culicidae female vectors were associated with nestlings' total parasites, scaled mass index, and industrial units. Nestlings in highly developed areas had higher infection rates than those in forests, suggesting increased exposure to infections. However, there was no clear relationship between total female vectors and total parasites. The level of urbanization negatively affected nestling body condition, with a decrease in fat deposits from forested to highly urbanized areas. Our findings highlight the complex interplay between urbanization, vector-borne parasite transmission, and host immune response, emphasizing the need for comprehensive urban planning to improve wildlife health and guarantee ecosystem functioning. Understanding how urbanization affects bird immunity and parasite infections is critical for adapting urban landscapes for wildlife health and ecosystem integrity.

Forest structure, plants, arthropods, scale, or birds' functional groups: What key factor are forest birds responding to?

Forest birds respond to a diverse set of environmental factors, including those altered by forest management intensity, such as resource and habitat availability in the form of food or nesting sites. Although resource/habitat availability and bird traits likely mediate responses of bird diversity to global change drivers, no study has assessed the direct and indirect effects of changes in forest management and traits on bird assemblages jointly at large spatial scales. In this context the questions remain whether (1) the birds' response to forest management changes through alterations in structural properties and/or food availability, or (2) if birds' eco-morphological traits act as environmental filters in response to environmental factors. We audio-visually recorded birds at 150 forest plots in three regions of Germany and assessed the forest structure (LiDAR) as well as the diversity of the herbaceous layer and diversity and biomass of arthropods. We further assessed eco-morphological traits of the birds and tested if effects on bird assemblages are mediated by changes in eco-morphological traits' composition. We found that abundance and species numbers of birds are explained best by models including the major environmental factors, forest structure, plants, and arthropods. Eco-morphological traits only increased model fit for indirect effects on abundance of birds. We found minor differences between the three regions in Germany, indicating spatial congruency of the processes at the local and regional scale. Our results suggest that most birds are not specialized on a particular food type, but that the size, diversity and species composition of arthropods are important. Our findings question the general view that bird traits adapt to the resources available.

A slow-fast trait continuum at the whole community level in relation to land-use intensification.

Organismal functional strategies form a continuum from slow- to fast-growing organisms, in response to common drivers such as resource availability and disturbance. However, whether there is synchronisation of these strategies at the entire community level is unclear. Here, we combine trait data for >2800 above- and belowground taxa from 14 trophic guilds spanning a disturbance and resource availability gradient in German grasslands. The results indicate that most guilds consistently respond to these drivers through both direct and trophically mediated effects, resulting in a 'slow-fast' axis at the level of the entire community. Using 15 indicators of carbon and nutrient fluxes, biomass production and decomposition, we also show that fast trait communities are associated with faster rates of ecosystem functioning. These findings demonstrate that 'slow' and 'fast' strategies can be manifested at the level of whole communities, opening new avenues of ecosystem-level functional classification.

Bird diversity along an urban to rural gradient in large tropical cities peaks in mid-level urbanization.

The gradient from natural to urban areas strongly associates with the structure of avian communities over that gradient. Most research on urban birds is from temperate areas and knowledge from tropical Southeast Asia is lacking. We examined bird species diversity, relative abundance, and species composition along an urban to rural gradient in three Myanmar cities, and assessed potential environmental factors responsible for the changes. We counted birds within 40 point-count sites with 50-m fixed-radius in three large cities of Myanmar, namely Mandalay, Mawlamyine, and Myeik. We distinguished four urban habitat types (Downtown-urban, University Campus-suburban, Paddy Field-agriculture, Hill-forest). We classified all species into migrant or resident and into major feeding groups and related with several environmental parameters such as 'impervious surface'. We counted 5,423 individuals of 103 species with roughly equal species diversity between the three cities. Rock Pigeon (Columba livia) was the most frequent species. The species composition differed significantly between the four major habitat types. Omnivores were more abundant in the city center than all other functional groups. Interestingly, insectivores were also predominant in the city center. In addition, more generalist' species occurred towards the city center compared to the periphery, indicating that the periphery has increased relevance for specialized birds. We found some marked differences in species composition between the three cities of Mandalay, Mawlamyine, and Myeik. Additionally to species composition, species diversity and relative abundance differed significantly between each of the four major habitat types in all three cities.

Environmental factors affect Acidobacterial communities below the subgroup level in grassland and forest soils.

In soil, Acidobacteria constitute on average 20% of all bacteria, are highly diverse, and are physiologically active in situ. However, their individual functions and interactions with higher taxa in soil are still unknown. Here, potential effects of land use, soil properties, plant diversity, and soil nanofauna on acidobacterial community composition were studied by cultivation-independent methods in grassland and forest soils from three different regions in Germany. The analysis of 16S rRNA gene clone libraries representing all studied soils revealed that grassland soils were dominated by subgroup Gp6 and forest soils by subgroup Gp1 Acidobacteria. The analysis of a large number of sites (n = 57) by 16S rRNA gene fingerprinting methods (terminal restriction fragment length polymorphism [T-RFLP] and denaturing gradient gel electrophoresis [DGGE]) showed that Acidobacteria diversities differed between grassland and forest soils but also among the three different regions. Edaphic properties, such as pH, organic carbon, total nitrogen, C/N ratio, phosphorus, nitrate, ammonium, soil moisture, soil temperature, and soil respiration, had an impact on community composition as assessed by fingerprinting. However, interrelations with environmental parameters among subgroup terminal restriction fragments (T-RFs) differed significantly, e.g., different Gp1 T-RFs correlated positively or negatively with nitrogen content. Novel significant correlations of Acidobacteria subpopulations (i.e., individual populations within subgroups) with soil nanofauna and vascular plant diversity were revealed only by analysis of clone sequences. Thus, for detecting novel interrelations of environmental parameters with Acidobacteria, individual populations within subgroups have to be considered.

Contrasting responses of above- and belowground diversity to multiple components of land-use intensity.

Land-use intensification is a major driver of biodiversity loss. However, understanding how different components of land use drive biodiversity loss requires the investigation of multiple trophic levels across spatial scales. Using data from 150 agricultural grasslands in central Europe, we assess the influence of multiple components of local- and landscape-level land use on more than 4,000 above- and belowground taxa, spanning 20 trophic groups. Plot-level land-use intensity is strongly and negatively associated with aboveground trophic groups, but positively or not associated with belowground trophic groups. Meanwhile, both above- and belowground trophic groups respond to landscape-level land use, but to different drivers: aboveground diversity of grasslands is promoted by diverse surrounding land-cover, while belowground diversity is positively related to a high permanent forest cover in the surrounding landscape. These results highlight a role of landscape-level land use in shaping belowground communities, and suggest that revised agroecosystem management strategies are needed to conserve whole-ecosystem biodiversity.

Historic changes in species composition for a globally unique bird community.

Significant uncertainties remain of how global change impacts on species richness, relative abundance and species composition. Recently, a discussion emerged on the importance of detecting and understanding long-term fluctuations in species composition and relative abundance and whether deterministic or non-deterministic factors can explain any temporal change. However, currently, one of the main impediments to providing answers to these questions is the relatively short time series of species diversity datasets. Many datasets are limited to 2 years and it is rare for a few decades of data to be available. In addition, long-term data typically has standardization issues from the past and/or the methods are not comparable. We address several of these uncertainties by investigating bird diversity in a globally important mountain ecosystem of the Hkakabo Razi Landscape in northern Myanmar. The study compares bird communities in two periods (pre-1940: 1900-1939 vs. post-2000: 2001-2006). Land-cover classes have been included to provide understanding of their potential role as drivers. While species richness did not change, species composition and relative abundance differed, indicating a significant species turn over and hence temporal change. Only 19.2% of bird species occurred during both periods. Land-cover model predictors explained part of the species richness variability but not relative abundance nor species composition changes. The temporal change is likely caused by minimal methodological differences and partially by land-cover.

Hkakabo Razi landscape as one of the last exemplar of large contiguous forests.

Deforestation and forest degradation around the world endanger the functioning of ecosystems, climate stability, and conservation of biodiversity. We assessed the spatial and temporal dynamics of forest cover in Myanmar's Hkakabo Razi Landscape (HRL) to determine its integrity based on forest change and fragmentation patterns from 1989 to 2016. Over 80% of the HRL was covered by natural areas, from which forest was the most prevalent (around 60%). Between 1989 and 2016, forest cover declined at an annual rate of 0.225%. Forest degradation occurred mainly around the larger plains of Putao and Naung Mung, areas with relatively high human activity. Although the rate of forest interior loss was approximately 2 to 3 times larger than the rate of total forest loss, forest interior was prevalent with little fragmentation. Physical and environmental variables were the main predictors of either remaining in the current land-cover class or transitioning to another class, although remaining in the current land cover was more likely than land conversion. The forests of the HRL have experienced low human impact and still constitute large tracts of contiguous forest interior. To ensure the protection of these large tracts of forest, sustainable forest policies and management should be implemented.

Management of ecosystems alters vector dynamics and haemosporidian infections.

The presence of insect vectors is a key prerequisite for transmission of vector-borne disease such as avian haemosporidians. In general, the effects of land use change on Diptera vectors are not well studied; the response of vectors to forest management depends on vector species, as has been shown previously for the birds. We tested if abundance of insects from different Diptera families and haemosporidian infection are affected through alteration of habitat structural variables (measured by LiDAR) and forest management intensities. We identified higher large-scale variation of female insect abundance in northeastern than in southwestern Germany. Unmanaged forest stands had higher Diptera insect abundances. We found that abundance of female Diptera increased with the amount of forest gaps but decreased in forest plots with more south facing aspect, higher habitat structural heterogeneity, temperature and humidity. We found that haemosporidian infections in Diptera insects increased with increased management intensity and more canopy structural diversity (e.g., amount of edge habitat), but decreased with a denser shrub layer, deeper leaf litter and higher humidity (characteristics for unmanaged forest stands). Although higher forest management intensity decreased vector abundance, the haemosporidian infections in the vectors increased, indicating a significant effect of forest management on disease dynamics.

Eleven years' data of grassland management in Germany.

BACKGROUND: The 150 grassland plots were located in three study regions in Germany, 50 in each region. The dataset describes the yearly grassland management for each grassland plot using 116 variables.General information includes plot identifier, study region and survey year. Additionally, grassland plot characteristics describe the presence and starting year of drainage and whether arable farming had taken place 25 years before our assessment, i.e. between 1981 and 2006. In each year, the size of the management unit is given which, in some cases, changed slightly across years.Mowing, grazing and fertilisation were systematically surveyed: Mowing is characterised by mowing frequency (i.e. number of cuts per year), dates of cutting and different technical variables, such as type of machine used or usage of conditioner.For grazing , the livestock species and age (e.g. cattle, horse, sheep), the number of animals, stocking density per hectare and total duration of grazing were recorded. As a derived variable, the mean grazing intensity was then calculated by multiplying the livestock units with the duration of grazing per hectare [LSU days/ha]. Different grazing periods during a year, partly involving different herds, were summed up to an annual grazing intensity for each grassland.For fertilisation , information on the type and amount of different types of fertilisers was recorded separately for mineral and organic fertilisers, such as solid farmland manure, slurry and mash from a bioethanol factory. Our fertilisation measures neglect dung dropped by livestock during grazing. For each type of fertiliser, we calculated its total nitrogen content, derived from chemical analyses by the producer or agricultural guidelines (Table 3).All three management types, mowing, fertilisation and grazing, were used to calculate a combined land use intensity index (LUI) which is frequently used to define a measure for the land use intensity. Here, fertilisation is expressed as total nitrogen per hectare [kg N/ha], but does not consider potassium and phosphorus.Information on additional management practices in grasslands was also recorded including levelling, to tear-up matted grass covers, rolling, to remove surface irregularities, seed addition, to close gaps in the sward. NEW INFORMATION: Investigating the relationship between human land use and biodiversity is important to understand if and how humans affect it through the way they manage the land and to develop sustainable land use strategies. Quantifying land use (the 'X' in such graphs) can be difficult as humans manage land using a multitude of actions, all of which may affect biodiversity, yet most studies use rather simple measures of land use, for example, by creating land use categories such as conventional vs. organic agriculture. Here, we provide detailed data on grassland management to allow for detailed analyses and the development of land use theory. The raw data have already been used for > 100 papers on the effect of management on biodiversity (e.g. Manning et al. 2015).

Multiple forest attributes underpin the supply of multiple ecosystem services.

Trade-offs and synergies in the supply of forest ecosystem services are common but the drivers of these relationships are poorly understood. To guide management that seeks to promote multiple services, we investigated the relationships between 12 stand-level forest attributes, including structure, composition, heterogeneity and plant diversity, plus 4 environmental factors, and proxies for 14 ecosystem services in 150 temperate forest plots. Our results show that forest attributes are the best predictors of most ecosystem services and are also good predictors of several synergies and trade-offs between services. Environmental factors also play an important role, mostly in combination with forest attributes. Our study suggests that managing forests to increase structural heterogeneity, maintain large trees, and canopy gaps would promote the supply of multiple ecosystem services. These results highlight the potential for forest management to encourage multifunctional forests and suggest that a coordinated landscape-scale strategy could help to mitigate trade-offs in human-dominated landscapes.

Modeling movement of West Nile virus in the Western hemisphere.

We modeled West Nile virus (WNV) movement rates and patterns based on a migratory bird agent (the Swainson's Thrush) and a resident bird agent (the House Sparrow), and compared the results of these models with actual movement data to investigate the likelihood that the pattern of WNV outbreaks observed in the New World was consistent with migrant bird-mediated spread, as reported from the Old World. We found that, contrary to Old World patterns, WNV activity in the Western Hemisphere does not seem consistent with movement by infected migrant birds. Instead WNV spread appears best explained by a non-directional movement, perhaps that of dispersing resident birds.