The effect of energy and seasonality on avian species richness and community composition

We analyzed geographic patterns of richness in both the breeding and winter season in relation to a remotely sensed index of seasonal production (normalized difference vegetation index [NDVI]) and to measures of habitat heterogeneity at four different spatial resolutions. The relationship between avian richness and NDVI was consistent between seasons, suggesting that the way in which available energy is converted to bird species is similar at these ecologically distinct times of year. The number and proportion of migrant species in breeding communities also increased predictably with the degree of seasonality. The NDVI was a much better predictor of seasonal richness at finer spatial scales, whereas habitat heterogeneity best predicted richness at coarser spatial resolutions. While we find strong support for a positive relationship between available energy and species richness, seasonal NDVI explained at most 61% of the variation in richness. Seasonal NDVI and habitat heterogeneity together explain up to 69% of the variation in richness.     

Bird community composition patterns in urban parks of Madrid: The role of age, size and isolation

This study intends to assess the influence of fragment age, size and isolation (from the regional species pool) on bird community composition patterns in urban parks in Madrid, and the role of local and regional factors on community structure. Park age was a good indicator of habitat complexity. Park age and area accounted for 62% of the variability in species richness, but two measures of isolation from the regional species pool were not included as significant factors. Species composition in urban parks showed a high degree of nestedness, which was associated with park age and area, but not with two measures of isolation from the regional species pool. The degree of nestedness increased with park age; the distribution of species varying from nested in old and mature parks to random in young parks. The incidence (% of species occurrence in parks) in young parks was correlated with regional densities, whereas in mature and old parks the incidence was correlated with local densities. In this urban landscape, species composition appears to be regulated by local factors (particularly in mature and old parks), such that species accumulate in an orderly (not random) fashion in relation to park age and area. Regional influences seem to be more pronounced only in young parks, which are mainly colonized by species from the regional species pool.     

A global model of island biogeography

Aim The goal of our study was to build a global model of island biogeography explaining bird species richness that combines MacArthur and Wilson's area–isolation theory with the species–energy theory. Location Global. Methods We assembled a global data set of 346 marine islands representing all types of climate, topography and degree of isolation on our planet, ranging in size from 10 ha to 800,000 km². We built a multiple regression model with the number of non‐marine breeding bird species as the dependent variable. Results We found that about 85–90% of the global variance in insular bird species richness can be explained by simple, contemporary abiotic factors. On a global scale, the three major predictors — area, average annual temperature and the distance separating the islands from the nearest continent — all have constraining (i.e. triangular rather than linear) relationships with insular bird species richness. We found that the slope of the species–area curve depends on both average annual temperature and total annual precipitation, but not on isolation. Insular isolation depends not only on the distance of an island from the continent, but also on the presence or absence of other neighbouring islands. Range in elevation — a surrogate for diversity of habitats — showed a positive correlation with bird diversity in warmer regions of the world, while its effect was negative in colder regions. We also propose a global statistical model to quantify the isolation‐reducing effect of neighbouring islands. Main conclusions The variation in avian richness among islands worldwide can be statistically explained by contemporary environmental variables. The equilibrium theory of island biogeography of MacArthur and Wilson and the species–energy theory are both only partly correct. Global variation in richness depends about equally upon area, climate (temperature and precipitation) and isolation. The slope of the species richness–area curve depends upon climate, but not on isolation, in contrast to MacArthur and Wilson's theory.     

Bird diversity patterns in Neotropical temperate farmlands: The role of environmental factors and trophic groups in the spring and autumn

Productivity, habitat heterogeneity and environmental similarity are of the most widely accepted hypotheses to explain spatial patterns of species richness and species composition similarity. Environmental factors may exhibit seasonal changes affecting species distributions. We explored possible changes in spatial patterns of bird species richness and species composition similarity. Feeding habits are likely to have a major influence in bird–environment associations and, given that food availability shows seasonal changes in temperate climates, we expect those associations to differ by trophic group (insectivores or granivores). We surveyed birds and estimated environmental variables along line‐transects covering an E‐W gradient of annual precipitation in the Pampas of Argentina during the autumn and the spring. We examined responses of bird species richness to spatial changes in habitat productivity and heterogeneity using regression analyses, and explored potential differences between seasons of those responses. Furthermore, we used Mantel tests to examine the relationship between species composition similarity and both the environmental similarity between sites and the geographic distance between sites, also assessing differences between seasons in those relationships. Richness of insectivorous birds was directly related to primary productivity in both seasons, whereas richness of seed‐eaters showed a positive association with habitat heterogeneity during the spring. Species composition similarity between assemblages was correlated with both productivity similarity and geographic proximity during the autumn and the spring, except for insectivore assemblages. Diversity within main trophic groups seemed to reflect differences in their spatial patterns as a response to changes between seasons in the spatial patterns of food resources. Our findings suggest that considering different seasons and functional groups in the analyses of diversity spatial pattern could contribute to better understand the determinants of biological diversity in temperate climates.     

Influence of settlement time, human population, park shape and age, visitation and roads on the number of alien plant species in protected areas in the USA

Abstract. I examined a data set of 77 protected areas in the USA (including national and state parks) to determine which of the following variables most strongly influence alien plant species richness: park area, climate (temperature and precipitation), native species richness, visitation rate, local human population size, total road length, park shape and duration of European settlement. Many of these predictor variables are intercorrelated, so I used multiple regression to help separate their effects. In support of previous studies, native species richness was the best single predictor of alien species richness, probably because it was a good estimator of both park area and habitat diversity available for establishment of alien species. Other significant predictors of alien species richness were years of occupation of the area by European settlers and the human population size of adjacent counties. Climate, visitation rate, road length and park shape did not influence alien species richness. The proportion of alien species (alien richness/native richness) is inversely related to park area, in agreement with a previous study. By identifying which variables are most important in determining alien species richness, such findings suggest ways to reduce alien species establishment.     

Functional diversity of urban bird communities: effects of landscape composition,green space area and vegetation cover

In this study, we aim to gain a better insight on how habitat filtering due to urbanization shapes bird communities of Vienna city parks. This may help to derive implications for urban planning in order to promote and maintain high diversity and ecosystem function in an increasing urbanized environment. The structure of wintering bird communities of 36 Vienna city parks – surveyed once a month in January 2009, December 2009, December 2012, and January 2013 – was described by species richness and the functional diversity measurements FRic (functional richness), FEve (functional evenness), and FDiv (functional divergence). Environmental filtering was quantified by park size, canopy heterogeneity within the park, and the proportion of sealed area surrounding each park. Species richness, FRic, and FDiv increased with increasing park size. Sealed area had a strong negative effect on species richness and FDiv. Canopy heterogeneity played a minor role in explaining variance in FDiv data. FEve did not respond to any of these park parameters. Our results suggest a loss of species richness and functional diversity, hence most likely indicate a decline in ecosystem function, with decreasing park size and increasing sealed area of the surrounding urban landscape matrix.     

Breeding bird diversity in relation to environmental gradients in China

Geographic variation in species richness has been explained by different theories such as energy, productivity, energy–water balance, habitat heterogeneity, and freezing tolerance. This study determines which of these theories best account for gradients of breeding bird richness in China. In addition, we develop a best-fit model to account for the relationship between breeding bird richness and environment in China. Breeding bird species richness in 207 localities (3271 km² per locality on average) from across China was related to thirteen environmental variables after accounting for sampling area. The Akaike's information criterion (AIC) was used to evaluate model performance. We used Moran's I to determine the magnitude of spatial autocorrelation in model residuals, and used simultaneous autoregressive model to determine coefficients of determination and AIC of explanatory variables after accounting for residual spatial autocorrelation. Of all environmental variables examined, normalized difference vegetation index, a measure of plant productivity, is the best variable to explain the variance in breeding bird richness. We found that species richness of breeding birds at the scale examined is best predicted by a combination of plant productivity, elevation range, seasonal variation in potential evapotranspiration, and mean annual temperature. These variables explained 47.3% of the variance in breeding bird richness after accounting for sampling area; most of the explained variance in richness is attributable to the first two of the four variables.     

Ecological traits and landscape characteristics predicting bird sensitivity to urbanization in city parks

Habitat loss and fragmentation caused by urbanization often have negative impacts on wildlife in cities. There are considerable studies investigating the relationship between species traits and fragmentation vulnerability. However, so far, very few studies have examined the influence of species traits combined with landscape factors on vulnerability to urbanization in urbanized landscapes. In this study, we investigated how species traits and park characteristics influenced bird sensitivity to urbanization in the highly urbanized city of Nanjing, China. We used the line-transect method to survey birds in 37 urban parks. For each bird species, we collected data on nine life-history and ecological traits that are commonly assumed to influence urbanization vulnerability. For each park, we selected six landscape variables that are commonly considered to influence bird response to urbanization. After phylogenetic correction, the nine species traits were used separately and in combination to evaluate their associations with species abundance, an indicator of urbanization vulnerability. We then used the RLQ and fourth-corner analyses to test relationships between species traits and environmental variables. We found that the 75 species analyzed demonstrated considerable variation in vulnerability to urbanization. Using PGLS analyses and model averaging, we found that habitat specificity was the single best ecological predictor of urbanization vulnerability in birds in Nanjing city parks. The RLQ analysis showed that body size and habitat specificity were correlated with distance to city center and connectivity of the parks, reflecting strong effects of trait-mediated environmental filters that selectively benefit species with smaller body mass and lower habitat specificity in urbanized landscape. Therefore, conservation efforts giving priority to species with high habitat specificity and to parks with high connectivity and far away from the city center may prove effective for the preservation of bird diversity in our highly urbanized system. Meanwhile, preventing future habitat loss and destruction in existing city parks may also effectively conserve these vulnerable species.     

Effect of Urbanization and Seasonality in Bird Communities of Kathmandu Valley,Nepal

Kathmandu Valley, Nepal is undergoing rapid urbanization but its effects on the bird communities have not been reported till date. Kathmandu Valley was categorized into urban, sub-urban and rural to study the impact of urbanization in bird communities. By mobilizing volunteers, we monitored 24 transects each with one km long in summer and winter seasons of 2016. A total of 13,749 individuals of birds belonging to 102 species were recorded. Species richness and diversity of all birds declined from rural to urban areas and showed significant variation along urban–rural gradients. Insectivore was the most species-rich guild while nectarivore the least. The richness of insectivore, frugivore and carnivore guilds showed significant variations along the urban–rural gradients and higher preference towards the rural areas. Similarly, species richness of all birds and richness of insectivore and carnivore guilds showed significant seasonal variation and were higher in the winter season. Our results indicate that richness, diversity and feeding guilds of birds show different response towards the urbanization gradients and seasons. Sub-urban areas can function as bird refugia, however, habitat enrichments (like increasing green spaces, setting up new parks and gardens, plantation of native fruiting trees etc.) are utmost necessary to support the bird communities in urban areas of Kathmandu Valley.     

Explaining bird species composition and richness in eucalypt-dominated remnants in subhumid Tasmania

Aim To determine the factors influencing the distribution of birds in remnants in a fragmented agricultural landscape. Location Forty‐seven eucalypt remnants and six sites in continuous forest in the subhumid Midlands region of Tasmania, Australia. Methods Sites were censused over a two‐year period, and environmental data were collected for remnants. The avifauna of the sites was classified and ordinated. The abundances of bird species, and bird species composition, richness, abundance and diversity were related to environmental variables, using simple correlation and modelling. Results There were two distinct groups of sample sites, which sharply differed in species composition, richness, diversity and bird abundance, separated on the presence/absence of noisy miner (Manorina melanocephala Latham) colonies, remnant size, vegetation structural attributes and variables that reflected disturbance history. The approximate remnant size threshold for the change from one group to another was 20–30 ha. Remnant species richness and diversity were most strongly explained by remnant area and noisy miner abundance, with contributions from structural and isolation attributes in the second case. Segment richness was explained by precipitation, logging history and noisy miner abundance. Bird abundance was positively related to precipitation and negatively related to tree dieback. The 28 individual bird species models were highly individualistic, with vegetation structural variables, noisy miner abundance, climatic variables, variables related to isolation, area, variables related to floristics, disturbance variables, the nature of the matrix and remnant shape all being components in declining order of incidence. Age of the remnant did not relate to any of the dependent variables. Main conclusions Degraded and small remnants may have become more distinct in their avifaunal characteristics than might otherwise be the case, as a result of the establishment of colonies of an aggressive native bird, the noisy miner. The area, isolation and shape of remnants directly relate to the abundance of relatively few species, compared to vegetation attributes, climate and the abundance of the noisy miner. The nature of the matrix is important in the response of some species to fragmentation.     

Avifaunal responses to habitat fragmentation in the threatened littoral forests of south-eastern Madagascar

Aim Madagascar's lowland forests are both rich in endemic taxa and considered to be seriously threatened by deforestation and habitat fragmentation. However, very little is known about how these processes affect biodiversity on the island. Herein, we examine how forest bird communities and functional groups have been affected by fragmentation at both patch and landscape scales, by determining relationships between species richness and individual species abundance and patch and landscape mosaic metrics. Location Littoral forest remnants within south‐eastern Madagascar. Methods We sampled 30 littoral forest remnants in south‐eastern Madagascar, within a landscape mosaic dominated by Erica spp. heathland. We quantified bird community composition within remnants of differing size, shape and isolation, by conducting point counts in November–December in 2001 and October–November 2002. Each remnant was characterized by measures of remnant area, remnant shape, isolation, and surrounding landscape complexity. We used step‐wise regression to test the relationship between bird species richness and landscape structural elements, after correcting for sampling effort. Relationships between bird species abundances and the landscape variables were investigated with Canonical Correspondence Analysis and binomial logistic regression modelling. Results Bird species richness and forest‐dependent bird species richness were significantly (P < 0.01) explained by remnant area but not by any measure of isolation or landscape complexity. The majority of forest‐dependent species had significant relationships with remnant area. Minimum area requirements for area‐sensitive species ranged from 15 to 150 ha, with the majority of species having area requirements > 30 ha. Surprisingly, there was no relationship between bird body size and minimum area requirement. Forest‐dependent canopy insectivorous species and large canopy frugivorous species were the most sensitive functional groups, with > 90% species sensitivity within each group. The distribution of four forest‐dependent species also appeared to be related to remnant shape where remnant area was < 100 ha. Main conclusions The majority of forest‐dependent species, including many that are considered widespread and common, were found to have significant relationships with fragment size, indicating that they are sensitive to processes associated with habitat loss and fragmentation. As deforestation and habitat fragmentation remain serious problems on the island, it follows that many forest‐dependent bird species will decline in abundance and become locally extinct. At the regional scale, we urge that large (> 200 ha) blocks of littoral forest are awarded protected status to preserve their unique bird community.     

北京的公园鸟类群落结构研究

不同类型公园的鸟类群落组成有很大差异，这主要和公园的面积、植被发育程度、位于城郊区还是城区、水域环境等自然条件，以及气候的季节性变化有关。面积大、植被发育好的公园，其群落多样性优于面积小、植被发育差的公园，公园内有无水域对鸟类群落的左右不明显，气候的季节性变化主要对鸟类群落物种数有明显的影响。通过对典型公园鸟类群落的多样性、均匀性及相关性进行比较，提出有关城市园林绿地招引鸟类的生态学对策。     

Energy density and its variation in space limit species richness of boreal forest birds

Aim An area’s ability to support species may be dependent not only on the total amount of available energy it contains but also on energy density (i.e. available energy per unit area). Acknowledging these two aspects of energy availability may increase mechanistic understanding of how increased energy availability results in increased species richness. We studied the relationship between energy density, its variation in space and boreal forest bird species richness and investigated two possible mechanisms: (1) metabolic constraints of organisms, and (2) increased resource availability for specialists. Location Protected areas in Finland’s boreal forest. Methods We tested whether bird species richness was best determined by total energy availability in an area or by energy density and its variation within the area, before and after including bird abundance in the models. We evaluated two main explanatory variables: tree growth reflecting the rate of energy production and tree volume as a measure of biomass. In addition, we modelled individual species’ responses to energy density and its variation, and evaluated the prediction of the metabolic constraints hypothesis that small species are limited by energy density whereas large species are limited by total energy availability in the area. Results Energy density and its variation were good predictors of species richness: together with abundance they explained 84% of variation in species richness (compared with 74% for abundance alone). Pure metabolic constraints were unlikely to explain this relationship. Instead, the mechanism probably involved increased habitat heterogeneity benefiting specialist species. Total energy availability was also an important factor determining species richness but its effect was indirect via abundance. Main conclusions Our results corroborate the importance of energy availability as a driver of species richness in forest bird communities, and they indicate that energy density and its variation in the landscape strongly influence species richness even after accounting for abundance.     

基于物种落差分析的公园鸟类多样性提升设计——以湖南常德螺湾湿地公园为例

随着城市化发展,许多公园鸟类栖息环境遭受破坏,导致当前的鸟类多样性与历史上的和周边的都存在较大差异,为此,我们从项目区域的当前-周边-历史三者间在鸟类物种及其栖息地上的落差分析入手,以此确定目标物种及其栖息地,并加以归类,从而有针对性地进行相关设计。2016年5月至2017年3月在湖南常德柳叶湖螺湾湿地公园开展的鸟类多样性提升设计中,通过实地调查并结合文献查阅,记录到项目区域16种,周边79种,历史上146种鸟类,运用三者间的物种落差分析法确定主要的21种可恢复目标物种和4种栖息地类型,结合项目区域及周边空间特点,将项目区域划分为4个区域,在各区域内设计并营建相应的栖息地,再配套采用多种鸟类招引措施。至2017年3月,项目区鸟类已可实地观察到39种,并于2017年12月吸引到超过2000只的野鸭群来此越冬。以上实际效果表明,设计有效地提升了湿地公园鸟类多样性且增加了景观要素,可充分发挥生物保护功能。     

Richness and composition of plants and birds on land‐bridge islands: effects of island attributes and differential responses of species groups

Aim To test relationships between the richness and composition of vascular plants and birds and attributes of habitat fragments using a model land‐bridge island system, and to investigate whether the effects of fragmentation differ depending on species natural history traits. Location Thousand Island Lake, China. Methods We compiled presence/absence data of vascular plant and bird species through exhaustive surveys of 41 islands. Plant species were assigned to two categories: shade‐intolerant and shade‐tolerant species; bird species were assigned to three categories: edge, interior, and generalist species. We analysed the relationships between island attributes (area, isolation, elevation, shape complexity, and perimeter to area ratio) and species richness using generalized linear models (GLMs). We also investigated patterns of composition in relation to island attributes using ordination (redundancy analysis). Results We found that island area explained a high degree of variation in the species richness of all species groups. The slope of the species–area relationship (z) was 0.16 for all plant species and 0.11 for all bird species. The lowest z‐value was for generalist birds (0.04). The species richness of the three plant species groups was associated with island area per se, while that of all, generalist, and interior birds was explained mainly by elevation, and that of edge bird species was associated primarily with island shape. Patterns of species composition were most strongly related to elevation, island shape complexity, and perimeter to area ratio rather than to island area per se. Species richness had no significant relationship with isolation, but species composition did. We also found differential responses among the species groups to changes in island attributes. Main conclusions Within the Thousand Island Lake system, the effects of fragmentation on both bird and plant species appear to be scale‐dependent and taxon‐specific. The number of plant species occurring on an island is strongly correlated with island area, and the richness of birds and the species composition of plants and birds are associated with variables related to habitat heterogeneity. We conclude that the effects of fragmentation on species diversity and composition depend not only on the degree of habitat loss but also on the specific patterns of habitat fragmentation.     

Macro‐scale bird species richness patterns of the East Asian mainland and islands: energy,area and isolation

Aim To create a map of bird species richness (BSR) in East Asia and to examine the effect of area, isolation, primary productivity, topographic heterogeneity, and human population density on BSR. Location East Asia (from 70° E to 180° E longitude), including the eastern half of the Palaearctic Region, the entire Oriental Region, and the entire Wallacea Subregion. Methods The breeding ranges of 2406 terrestrial bird species were mapped and overlaid to create a species richness map. The BSR map was transformed into a 100 × 100 km quadrat system, and BSR was analysed in relation to land area, average normalized difference vegetation index (NDVI), elevation range, and average population density. Results In general, BSR declined from the Tropics to the Arctic. In mainland East Asia, however, BSR was highest around the Tropic of Cancer, and fluctuated between 30° and 50° N. Islands had lower BSR than adjacent mainland areas. The NDVI was strongly positively correlated with BSR in mainland areas and on islands. For mainland areas, NDVI explained 65% of the BSR variation, and topographic heterogeneity explained an additional 6% in ordinary least‐squares regression. On islands, NDVI explained 66% of BSR variation, island area explained 13%, and distance to mainland accounted for 1%. Main conclusions In East Asia, we suggest that primary productivity is the key factor underpinning patterns of BSR. Primary productivity sets the upper limits of the capacity of habitats to support bird species. In isolated areas such as islands and peninsulas, however, BSR might not reach the richness limits set by primary productivity because the degree of isolation and area size also can affect species richness. Other factors, such as spatial heterogeneity, biotic interactions, and perturbations, may also affect species richness. However, their effects are secondary and are not as strong as primary productivity, isolation, and area size.     

Does size really matter? Species–area relationships in human settlements

Aim To describe species–area relationships in human settlements and compare them with those from a non‐urban habitat. Location West‐central Mexico. Methods We surveyed breeding birds in 13 human settlements and five shrubland patches. We estimated bird species richness using an abundance‐based coverage estimator with equal sample sizes to eliminate biases related to sampling effort differences. To assess species–area relationships, we performed log–log linear regressions between the size of the studied patches and their estimated bird richness. We also used a logarithmic approach to determine how the species–area relationship asymptoted and made use of the Michaelis–Menten model to identify the size at which the studied patches reached their maximum species richness. We also investigated (1) possible relationships among the estimated bird richness and other variables known to affect urban‐dwelling birds (built cover, plant species richness, tree cover or human population density) and (2) changes in bird community composition related to the size of the studied human settlements. Results Species–area relationships exhibited different patterns among the studied habitats. The log–log regression slope was steeper in human settlements, while the intercept was higher in shrublands. The maximum number of species was more than twofold higher in shrublands. Human settlement patch size was the only variable significantly related to bird richness. Our community composition results show that two main bird groups are related to human settlement size, and that as the size of human settlements increases, bird community similarity in relation to the largest city increases. Main conclusions Human settlements act as ecological islands, with pronounced species–area relationships. Our results suggest that an important threshold for bird species richness and community composition is reached in human settlements > 10.2 km². This threshold is unlikely to be generalizable among bio‐regions, and thus should be quantified and considered when studying, managing and/or planning urban systems.     

Anthropogenic noise reduces bird species richness and diversity in urban parks

Anthropogenic noise is becoming more prevalent in the world and has been shown to affect many animal species, including birds. The impact of such noise was measured in Neotropical urban parks to assess how the noise affects avifauna diversity and species richness. We sampled bird species, and concurrently measured sound pressure (noise) levels (L_eq, equivalent noise levels) in eight urban green areas or parks located in a large city (Belo Horizonte) in south‐eastern Brazil over a 1‐year period. The diversity of sampled points was measured by means of total species richness, Fisher's alpha and Shannon–Wiener diversity indices. Noise levels within all parks were greater than those in natural areas. We found that an increase in noise levels and the area of open habitats surrounding sampling points were negatively related to species richness. Social factors reflecting increased urbanization, such as higher incomes, were also negatively correlated with bird species richness. However, noise was the factor that explained most of the variance. These results suggest that anthropogenic noise can have a significant negative impact on the conservation value of urban parks for bird species.     

Factors influencing geographic patterns in diversity of forest bird communities of eastern Connecticut,USA

At regional scales, the most important variables associated with diversity are latitudinally‐based temperature and net primary productivity, although diversity is also influenced by habitat. We examined bird species richness, community density and community evenness in forests of eastern Connecticut to determine whether: 1) spatial and seasonal patterns exist in diversity, 2) energy explains the greatest proportion of variation in diversity parameters, 3) variation in habitat explains remaining diversity variance, and 4) seasonal shifts in diversity provide clues about how environmental variables shape communities. We sought to discover if our data supported predictions of the species–energy hypothesis. We used the variable circular plot technique to estimate bird populations and quantified the location, elevation, forest type, vegetation type, canopy cover, moisture regime, understory density and primary production for the study sites. We found that 1) summer richness and population densities are roughly equal in northeastern and southeastern Connecticut, whereas in winter both concentrate toward the coast, 2) variables linked with temperature explained much of the patterns in winter diversity, but energy‐related variables showed little relationship to summer diversity, 3) the effect of habitat variables on diversity parameters predominated in summer, although their effect was weak, 4) contrary to theory, evenness increased from summer to winter, and 5) support for predictions of species–energy theory was primarily restricted to winter data. Although energy and habitat played a role in explaining community patterns, they left much of the variance in regional diversity unexplained, suggesting that a large stochastic component to diversity also may exist.