Elevational gradients in species abundance,assemblage structure and energy use of rainforest birds in the Australian Wet Tropics bioregion

Elevational patterns of species richness, local abundance and assemblage structure of rainforest birds of north‐eastern Australia were explored using data from extensive standardized surveys throughout the region. Eighty‐two species of birds were recorded with strong turnover in assemblage structure across the elevational gradient and high levels of regional endemism in the uplands. Both species richness and bird abundance exhibited a humped‐shaped pattern with elevation with the highest values being between 600 and 800 m elevation. While much of the variability in species richness could be explained by the species–area relationship, analyses of net primary productivity (NPP) and total daily energy consumption of the bird community (energy use) suggest that ecosystem energy flow or constraints may be a significant determinant of species richness. Species richness is positively correlated with local bird abundance which itself is closely related to total energy use of the bird community. We suggest the hypothesis that lower NPP limits bird abundance and energy use in the uplands (>500 m) and that low bird energy use and species richness in the lowlands is limited by a seasonal bottleneck in available primary productivity and/or a species pool previously truncated by an extinction filter imposed by the almost complete disappearance of rainforest in the lowlands during the glacial maxima. We suggest that some of the previously predicted impacts of global warming on biodiversity in the uplands may be partially ameliorated by increases in NPP because of increasing temperatures. However, these relationships are complex and require further data specifically in regard to direct estimates of primary productivity and detailed estimates of energy flow within the assemblage.     

Breeding bird species richness in Taiwan: distribution on gradients of elevation, primary productivity and urbanization

Aim To examine the richness of breeding bird species in relation to elevation, primary productivity and urbanization. Location The island of Taiwan (120°–122° E, 22°–25° N). Methods We arranged bird species richness (BSR) data from 288 bird censuses undertaken in Taiwan into a 2 × 2 km quadrat system and calculated average values of elevation, primary productivity [surrogated by normalized difference vegetation index (NDVI)], and urbanization (surrogated by road density and percentage of built area) for each 2 × 2 km quadrat. Results Bird species richness showed a hump‐shaped relationship with elevation. It increased with elevation from sea level (10–64 species per 2 × 2 km quadrat), peaked around 2000 m (43–76 species), and then decreased with elevation towards its minimum at the highest elevation. Road density and percentage of built area decreased with elevation, and NDVI showed a hump‐shaped relationship with elevation and inverse relationships with road density and percentage of built area. BSR increased with NDVI and decreased with road density and percentage of built area. Linear and cubic terms of elevation together explained 31.3% of the variance in BSR, and road density explained additional 3.4%. The explanatory power of NDVI on BSR was insignificant after the effects of elevation and road density had been justified. Main conclusions We argue that urbanization plays an important role in the BSR of Taiwan. Urbanization might indirectly decrease BSR through decreasing primary productivity and therefore change the hypothetical inverse relationship between BSR and elevation into a hump‐shaped relationship. We also propose a time hypothesis that the biotic communities in the mid‐elevation zone of Taiwan had relatively longer periods of existence during the Pleistocene glacial cycles, which might be one underlying process of the observed hump‐shaped relationship between species diversity and elevation.     

Small mammal species richness and turnover along elevational gradient in Yulong Mountain,Yunnan, Southwest China

Understanding the species diversity patterns along elevational gradients is critical for biodiversity conservation in mountainous regions. We examined the elevational patterns of species richness and turnover, and evaluated the effects of spatial and environmental factors on nonvolant small mammals (hereafter “small mammal”) predicted a priori by alternative hypotheses (mid‐domain effect [MDE], species–area relationship [SAR], energy, environmental stability, and habitat complexity]) proposed to explain the variation of diversity. We designed a standardized sampling scheme to trap small mammals at ten elevational bands across the entire elevational gradient on Yulong Mountain, southwest China. A total of 1,808 small mammals representing 23 species were trapped. We observed the hump‐shaped distribution pattern of the overall species richness along elevational gradient. Insectivores, rodents, large‐ranged species, and endemic species richness showed the general hump‐shaped pattern but peaked at different elevations, whereas the small‐ranged species and endemic species favored the decreasing richness pattern. The MDE and the energy hypothesis were supported, whereas little support was found for the SAR, the environmental stability hypothesis, and the habitat complexity. However, the primary driver(s) for richness patterns differed among the partitioning groups, with NDVI (the normalized difference vegetation index) and MDE being the most important variables for the total richness pattern. Species turnover for all small mammal groups increased with elevation, and it supported a decrease in community similarity with elevational distance. Our results emphasized for increased conservation efforts in the higher elevation regions of the Yulong Mountain.     

Different responses of avian feeding guilds to spatial and environmental factors across an elevation gradient in the central Himalaya

Although elevational patterns of species richness have been well documented, how the drivers of richness gradients vary across ecological guilds has rarely been reported. Here, we examined the effects of spatial factors (area and mid‐domain effect; MDE) and environmental factors, including metrics of climate, productivity, and plant species richness on the richness of breeding birds across different ecological guilds defined by diet and foraging strategy. We surveyed 12 elevation bands at intervals of 300 m between 1,800 and 5,400 m a.s.l using line‐transect methods throughout the wet season in the central Himalaya, China. Multiple regression models and hierarchical partitioning were used to assess the relative importance of spatial and environmental factors on overall bird richness and guild richness (i.e., the richness of species within each guild). Our results showed that richness for all birds and most guilds displayed hump‐shaped elevational trends, which peaked at an elevation of 3,300–3,600 m, although richness of ground‐feeding birds peaked at a higher elevation band (4,200–4,500 m). The Normalized Difference Vegetation Index (NDVI)—an index of primary productivity—and habitat heterogeneity were important factors in explaining overall bird richness as well as that of insectivores and omnivores, with geometric constraints (i.e., the MDE) of secondary importance. Granivore richness was not related to primary production but rather to open habitats (granivores were negatively influenced by habitat heterogeneity), where seeds might be abundant. Our findings provide direct evidence that the richness–environment relationship is often guild‐specific. Taken together, our study highlights the importance of considering how the effects of environmental and spatial factors on patterns of species richness may differ across ecological guilds, potentially leading to a deeper understanding of elevational diversity gradients and their implications for biodiversity conservation.     

Zooplankton species richness–productivity relationship: Confronting monotonic positive and hump‐shaped models from a local perspective

This study examined two models that are most frequently used to describe the relationship between species richness and productivity (SPR): monotonic positive and hump‐shaped models. We assessed zooplankton community diversity in response to algal productivity. The relationship between net primary productivity (NPP) and rarefied species richness was examined by fitting the data to two models and comparing them using the Akaike information criterion (AICc). Macrophyte banks with the highest net primary productivity had the highest zooplankton abundance. Our results pointed to a hump‐shaped model as the best fit to describe the relationship between zooplankton species richness and primary productivity (ΔAICc > 4). Thus, the diversity was lower at the extremes of productivity and higher at intermediate levels of productivity. We suggest that this relationship might occur because when the resource supply rates are low, environmental conditions are stressful, whereas a high availability of resources enhances competitive exclusion. Two observations supported this statement: (i) the total abundance of the community positively correlated with NPP (P < 0.05), indicating that less productive sites had few consumers and the raised productivity tended to favour the total abundance; (ii) NPP was negatively correlated with evenness (P < 0.05), indicating that productivity increased the dominance of certain species in the communities. Therefore, we challenged two of the models most frequently used to explain SPR, and discuss some mechanisms underlying a hump‐shaped SPR.     

Elevational gradients in bird diversity in the Eastern Himalaya: an evaluation of distribution patterns and their underlying mechanisms

Background

Understanding diversity patterns and the mechanisms underlying those patterns along elevational gradients is critically important for conservation efforts in montane ecosystems, especially those that are biodiversity hotspots. Despite recent advances, consensus on the underlying causes, or even the relative influence of a suite of factors on elevational diversity patterns has remained elusive.

Methods and Principal Findings

We examined patterns of species richness, density and range size distribution of birds, and the suite of biotic and abiotic factors (primary productivity, habitat variables, climatic factors and geometric constraints) that governs diversity along a 4500-m elevational gradient in the Eastern Himalayan region, a biodiversity hotspot within the world''s tallest mountains. We used point count methods for sampling birds and quadrats for estimating vegetation at 22 sites along the elevational gradient. We found that species richness increased to approximately 2000 m, then declined. We found no evidence that geometric constraints influenced this pattern, whereas actual evapotranspiration (a surrogate for primary productivity) and various habitat variables (plant species richness, shrub density and basal area of trees) accounted for most of the variation in bird species richness. We also observed that ranges of most bird species were narrow along the elevation gradient. We find little evidence to support Rapoport''s rule for the birds of Sikkim region of the Himalaya.

Conclusions and Significance

This study in the Eastern Himalaya indicates that species richness of birds is highest at intermediate elevations along one of the most extensive elevational gradients ever examined. Additionally, primary productivity and factors associated with habitat accounted for most of the variation in avian species richness. The diversity peak at intermediate elevations and the narrow elevational ranges of most species suggest important conservation implications: not only should mid-elevation areas be conserved, but the entire gradient requires equal conservation attention.     

Energy, Density, and Constraints to Species Richness: Ant Assemblages along a Productivity Gradient

Species richness describes the number of species of a given taxon in a given time and space. The energy limitation hypothesis links the species richness of consumer taxa to net primary productivity (NPP) through two relationships: NPP limits a taxon's density, and taxon density limits species richness. We study both relationships with a survey of 15 ground ant assemblages, along a productivity gradient from deserts to rain forests. Ant density (colonies m-2) was a positive, decelerating function of net aboveground productivity (NAP). A stepwise regression suggests that the efficiency with which NAP is converted to ant colonies increases with maximum summer temperature and decreases with precipitation. Ant species richness was a positive decelerating function of density at three spatial scales. This supports the energy limitation hypothesis' assumption that average population densities are higher in environments that are more productive. These two nonlinear functions (NAP-density and density-species richness) combine to create, at a variety of scales, positive, decelerating, productivity-diversity curves for a common, ecologically dominant taxon across the terrestrial productivity gradient. However, variance in the density and diversity explained by NAP decreases with scale, suggesting that energy limitation of diversity predominates at small spatial scales (<1 ha).     

What drives the positive correlation between human population density and bird species richness in Australia?

Aim To test six hypotheses that could explain or mediate the positive correlation between human population density (HPD) and bird species richness while controlling for biased sampling effort. These hypotheses were labelled as follows: productivity (net primary productivity, NPP); inherent heterogeneity (diversity of vegetation types); anthropogenic heterogeneity (diversity of land uses); conservation policy (proportion of conservation land); increased productivity (human‐induced productivity increases); and the reduced‐slope hypothesis (which predicts that humans have a negative impact on species numbers across the full range of variation in HPD). Location Australia. Methods All data were collected at a spatial resolution of 1° across mainland Australia. Bird species richness was from 2007 atlas data and random subsampling was used to account for biased sampling effort. HPD was from the 2006 census. All other data were from government produced geographic information system layers. The most important biotic or abiotic factors influencing patterns in both species richness and HPD were assessed using simultaneous autoregressive models and an information theoretic approach. Results NPP appeared to be one of the main factors driving spatial congruence between bird species richness and HPD. Inherent habitat heterogeneity was weakly related to richness and HPD, although an interaction between heterogeneity and NPP indicated that the former may be an important determinant of species richness in low‐productivity regions. There was little evidence that anthropogenic landscape heterogeneity or human‐induced changes in productivity influenced the relationship between species richness and HPD, but conservation policy appeared to act as an important mediating factor and species richness was positively related to the proportion of conservation land only in regions of high HPD. Main conclusions The spatial congruence between bird species richness and HPD occurs because both respond positively to productivity and, in certain circumstances, habitat heterogeneity. Our results suggest that conservation policy could mediate this relationship, but further research is required to determine the importance of conservation reserves in supporting species in regions densely populated by humans.     

Exploring anthropogenic and natural processes shaping fern species richness along elevational gradients

Aim (1) To explore the impact of land use, climate and environmental heterogeneity on fern species richness along a complete elevational gradient, and (2) to evaluate the relative importance of the three groups of variables within different elevational intervals. Location A temperate mountain region (55,507 km²) of Italy on the southern border of the European Alps divided into a regular grid of 1476 cells (grain 35.7 km²). Methods We applied multiple regression (spatial and non‐spatial) to determine the relative influence of the three groups of variables on species richness, including variation partitioning at two scales. We considered the whole gradient (all 1476 cells) to explain the overall elevational pattern of species richness, and we grouped the cells into elevational intervals of 500 m in order to evaluate the explanatory power of the predictors within different zones along the gradient. Results Species richness showed a hump‐shaped pattern with elevation, forming a plateau between 800 and 1500 m. The lowest species richness was found in warm and relatively dry disturbed lowlands. Moving upwards, the greatest species richness was found in forest‐dominated mid‐elevations with high environmental heterogeneity. At high elevations dominated by open natural habitats, where temperature and precipitation were relatively low, species richness declined but less sharply than in the lowlands. Although it was impossible to separate the effects of the three groups of predictors along the whole gradient, the analysis of separate elevational intervals shed light on their relative importance. The decline of species richness within lowlands was mainly related to a combined effect of deforestation and low environmental heterogeneity. In the middle part of the gradient, habitat heterogeneity and topographic roughness were positively associated with species richness. The richness decline within high‐elevation areas was related mostly to climatic constraints. Main conclusions Human impact due to land‐use modifications strongly affects the elevational pattern of species richness. It is therefore increasingly important to adopt a multiple‐hypothesis approach, taking anthropogenic effects explicitly into account when describing ecological processes along elevational gradients.     

The relationship among area, elevation, and regional species richness in neotropical birds

The elevational gradient of species richness is often claimed to mirror the latitudinal gradient and has traditionally been explained by assuming a decrease in productivity with elevation and more recently by Rapoport's rule. The influence of area on the pattern has rarely been considered. Analyses of all South American tropical land birds (more than one-fourth of the extant bird species on Earth) are used to examine four species richness/elevation models: null model, Rapoport's rule, and monotonic or hump-shaped productivity/species richness relationships. To quantify the area effect, species-area curves were created for seven elevational zones. Not accounting for area, species richness declined monotonically with elevation, but area accounted for 67%-91% of the variation in species richness per zone. When area was factored out, a hump-shaped pattern emerged, with more species in the 500-1,000-m (P<.005) and 1,000-1,500-m zones (P<.10) than in the 0-500-m zone. Rapoport's rule and the monotonic productivity/species richness relationship were thus not supported. Instead, elevational turnover rates and numbers of shared species between zones suggested that the hump-shaped pattern reflects geometric constraints (as predicted by the null model) imposed by the narrow span of the gradient, and it is suggested that midelevational zones may represent sink habitats.     

Biogeographical comparison of winter bird assemblages in urban environments in Finland

We studied biogeographical variation of urban bird assemblages in Finland. Winter birds were censused by single-visit study plot method from thirty-one centres of villages or towns along 950 km latitudinal extent. A total twenty-eight bird species was observed and the average density was 61.2 ind./10 ha. The number of dominant species in study areas varied between two and seven and their proportion of the whole assemblage was over 70%. Species richness, but not the density of birds, decreased northwards in pooled data. Higher species richness in south than in north was mainly due to the higher amount of delayed migratory birds (e.g. waterbirds, finches) and southerly distributed bird species. However, in heavily urbanized areas species richness did not decrease northwards. This observation disagreed with the hypothesis that species richness decreased northwards. Bird density, but not species richness, increased with urbanization. In particular, feral pigeon, hooded crow and house sparrow had highest densities in most urbanized areas. As only few bird species are adapted to live in urban areas, species composition and dominant bird species were almost the same in the south and in the north. These urban birds may effectively use energy rich food in feeding tables and overcome the problems of severe climate in the north. This may be the reason why bird species richness does not decrease northwards in urban areas.     

The species richness pattern of vascular plants along a tropical elevational gradient and the test of elevational Rapoport's rule depend on different life‐forms and phytogeographic affinities

The research about species richness pattern and elevational Rapoport's rule (ERR) have been carried out mostly in the temperate regions in the recent years and scarcely in the tropical mountains; meanwhile, it is unclear whether the ERR is consistent among different life‐forms and phytogeographic affinities. Here, we compiled a database of plant species of Mount Kenya, a tropical mountain of East Africa, and divided these species into twelve groups depending on the life‐form and phytogeographic affinity of each species. We inspected the species richness pattern of each group along the elevation gradient and also tested ERR of each group using Stevens' method. Our results showed that species richness of the total species showed a positively skewed (hump‐shaped) pattern along the elevation gradient and different life‐forms and phytogeographic affinities showed similar hump‐shaped patterns as the total species. The average elevation range size of the total species and herbaceous species showed increasing patterns along the elevation gradient, while lycophytes and ferns, and woody species showed an obvious downward trend after peaking in the high elevation regions. We concluded that the widely distributed herbaceous species which also have broad elevation range sizes are more applicable to ERR, while the narrowly distributed woody species with small elevation range sizes occurring in the higher elevations could reverse ERR. Therefore, we concluded that the ERR is not consistent among different organisms in the same region.     

Avian Use of Wetlands in Reclaimed Minelands in Southwestern Indiana

We studied the use of mineland wetlands by birds and the relationship between avian communities and wetland characteristics. Data were collected from 20 wetlands in Pike County, Indiana, and included wetland size, depth, water conductivity and salinity, aquatic macroinvertebrate abundance, vegetation, and bird use. Principal component analysis showed that physical variables could be explained by two principal component scores and that wetlands could be grouped on the basis of size and conductivity. Principal component analysis could not reduce vegetation variables to fewer principal component scores, meaning that wetland vegetation characteristics were independent of one another and did not show any trend. Most wetlands had low invertebrate density, and wetlands with higher invertebrate density had low invertebrate diversity. Wetlands with similar habitat characteristics (physical, vegetative, and invertebrate) did not necessarily show similarities in bird assemblages. Bird similarity index values ranged from 0 to 59%, implying that each wetland has its own bird community. Stepwise multiple regression analysis (α= 0.05) relating bird use and habitat characteristics showed that bird species richness increased with the species richness of submergent vegetation and was correlated negatively with the species richness of emergent vegetation. There was no significant relationship between bird species richness or bird species diversity and wetland size. The number of species within different avian guilds correlated with different habitat characteristics. The species richness of submergent plants was a factor that correlated positively with the number of species of several guilds (dabblers, wading birds, and plunge divers). Wetland age was not a factor that determined bird use.     

Birds and people in Europe

At a regional scale, species richness and human population size are frequently positively correlated across space. Such patterns may arise because both species richness and human density increase with energy availability. If the species-energy relationship is generated through the 'more individuals' hypothesis, then the prediction is that areas with high human densities will also support greater numbers of individuals from other taxa. We use the unique data available for the breeding birds in Europe to test this prediction. Overall regional densities of bird species are higher in areas with more people; species of conservation concern exhibit the same pattern. Avian density also increases faster with human density than does avian biomass, indicating that areas with a higher human density have a higher proportion of small-bodied individuals. The analyses also underline the low numbers of breeding birds in Europe relative to humans, with a median of just three individual birds per person, and 4 g of bird for every kilogram of human.     

Bird diversity along elevational gradients in the Andes of Colombia: area and mass effects

Aim The decrease in species richness with increasing elevation is a widely recognized pattern. However, recent work has shown that there is variation in the shape of the curve, such that both negative monotonic or unimodal patterns occur, influenced by a variety of factors at local and regional scales. Discerning the shape of the curve may provide clues to the underlying causes of the observed pattern. At regional scales, the area of the altitudinal belts and mass effects are important determinants of species richness. This paper explores the relationship between bird species richness, elevation, mass effects and area of altitudinal zones for birds in tropical mountains. Location The three Andean ranges of Colombia and the peripheral mountain ranges of La Macarena and Santa Marta. Methods Lists of bird species were compiled for altitudinal belts in eastern and western slopes of the three Andean Cordilleras and for La Macarena and Santa Marta. The area of the altitudinal belts was computed from digital elevation models. The effect of area was analysed by testing for differences among altitudinal belts in the slopes and intercepts of the species‐area relationships. Mass effects were explored by separately analysing two sets of species: broadly distributed species, i.e. lowland species whose distributions extend into the Andes, and tropical Andean species, i.e., species that evolved in the Andes. Results Plotting total number of species in each altitudinal belt revealed a decline in species richness with elevation. In slopes with a complete elevational gradient from lowlands to mountain peaks, the decrease was monotonic. In internal Andean slopes where the lower elevational belts are truncated, there was a peak at mid elevations. There was a linear relationship between number of species and area of the altitudinal belts. When controlling for area, there were no differences in the number of species among altitudinal belts (500–2600 m), except for the two upper‐elevation zones (2600–3200 and > 3200 m), which had lower species richness. Diversity of widely distributed species declined monotonically with elevation, whereas tropical Andean species exhibited a mid‐elevation peak. Main conclusions A large proportion of the variation in species richness with elevation was explained by area of the altitudinal belts. When controlling for area, species richness remained constant up to 2600 m and then decreased. This pattern contrasts with a previously reported hump‐shaped pattern for Andean birds. Diversity patterns of widely distributed species suggested that immigration of lowland species inflates diversity of lower elevational belts through mass effects. This influence was particularly evident in slopes with complete altitudinal gradients (i.e. connected to the lowlands). Tropical Andean species, in contrast, were more diverse in mid‐elevational belts, where speciation rates are expected to be higher. The influence of these species was more prevalent in internal Andean slopes with no connection to the lowlands. The decline of species richness at high elevations may be related to higher extinction rates and lower resource levels.     

Modelling the responses of Australian subtropical rainforest birds to changes in environmental conditions along elevational gradients

Montane birds face significant threats from a warming climate, so determining the environmental factors that most strongly influence the composition of such assemblages is of critical conservation importance. Changes in temperature and other environmental conditions along elevational gradients are known to influence the species richness and abundance of bird assemblages occupying mountains. However, the role of species‐specific traits in mediating the responses of bird species to changing conditions remains poorly understood. We aimed to determine whether different bird species responded differently to changing environmental conditions in a relatively understudied biodiversity hotspot in subtropical rainforest on the east coast of Australia. We examined patterns in avian species richness and abundance along two rainforest elevational gradients using monthly point counts between September 2015 and October 2016. Environmental data on temperature, wetness, canopy cover and canopy height were collected simultaneously, and trait information on body size and feeding guild membership for each bird species was obtained from the Handbook of Australian, New Zealand and Antarctic Birds. We used a generalized linear mixed modelling (GLMM) framework to determine the drivers of species richness and abundance and to quantify species’ trait–environment interactions. GLMMs indicated that temperature alone was significantly positively correlated with species richness and abundance. Species richness declined with increasing elevation. When modelling abundance, we found that feeding guild membership did not significantly affect species’ responses to environmental conditions. In contrast, the predicted abundance of a species was found to depend on its body size, due to significant positive interactions between this trait, temperature and canopy cover. Our findings indicate that large‐bodied birds are likely to increase in abundance more rapidly than small‐bodied birds with continued climatic warming. These results underline the importance of temperature as a driving factor of avian community assembly along environmental gradients.     

Energy availability, abundance, energy-use and species richness in forest bird communities: a test of the species–energy theory

Aim To test the ‘more individuals hypothesis’ as a mechanism for the positive association between energy availability and species richness. This hypothesis predicts that total density and energy use in communities is linearly related to energy availability, and that species richness is a positive function of increased density. We also evaluate whether similar energy–density patterns apply to different migratory groups (residents, short‐distance migrants and tropical migrants) separately. Location European and North American forest bird communities. Methods We collected published breeding bird census data from Europe and North America (n = 187). From each census data we calculated bird density (pairs 10 ha^?1), energy use by the community (the sum of metabolic needs of individuals, Watts 10 ha^?1) and geographical location with an accuracy of 0.5°. For each bird census data coordinate we extracted the corresponding monthly values of actual evapotranspiration (AET). From these values we calculated corresponding AET values that we expected to explain the density energy use of forest birds: total annual, breeding season (June) and winter AET. We used general linear modelling to analyse these data controlling for the area of census plots, forest type and census method. Results Total density and energy use in European and North American forest bird communities were linear functions of annual productivity, and increased density and energy use then translated into more species. Also resident bird density and energy consumption were positive functions of annual productivity, but the relationship between productivity and density as well as between productivity and energy use was weaker for migrants. Main conclusions Our results are consistent with the more individuals hypothesis that density and energy use in breeding forest bird communities is coupled tightly with the productivity of the environment, and that increased density and energy consumption results in more species. However, not all community members (migratory groups) are limited by productivity on the breeding grounds.     

The role of environment and mid-domain effect on moth species richness along a tropical elevational gradient

Aim The biodiversity of geometrid moths (Lepidoptera) along a complete tropical elevational gradient was studied for the first time. The patterns are described, and the role of geometric constraints and environmental factors is explored. Location The study was carried out along the Barva Transect (10° N, 84° W), a complete elevational gradient ranging from 40 to 2730 m a.s.l. in Braulio Carrillo National Park, Costa Rica, and adjacent areas. Methods Moths were sampled manually in 2003 and 2004 at 12 rain forest sites using light ‘towers’, each with two 15 W ultraviolet fluorescent tubes. We used abundance‐based rarefaction, statistical estimation of true richness (Chao 1), geographically interpolated observed richness and Fisher's alpha as measures of local diversity. Results A total of 13,765 specimens representing 739 species were analysed. All four measures showed a hump‐shaped pattern with maxima between 500 and 2100 m elevation. The two subfamilies showed richness and diversity maxima at either lower (Ennominae) or higher (Larentiinae) elevation than Geometridae as a whole. Among the four environmental factors tested, relative humidity yielded the highest correlation over the transect with the rarefaction‐based richness estimates as well as with estimated true species richness of Geometridae as a whole and of Larentiinae, while rainfall explained the greatest variation of Ennominae richness. The elevational pattern of moth richness was discordant with both temperature and with tree species richness. A combination of all environmental factors in a stepwise multiple regression produced high values of r² in Geometridae. The potential effects of geometric constraints (mid‐domain effect, MDE) were investigated by comparing them with observed, interpolated richness. Overall, models fitted very well for Geometridae as a whole and for Ennominae, but less well for Larentiinae. Small‐ranged species showed stronger deviations from model predictions than large‐ranged species, and differed strikingly between the two subfamilies, suggesting that environmental factors play a more pronounced role for small‐ranged species. We hypothesize that small‐ranged species (at least of the Ennominae) may tend to be host specialists, whereas large‐ranged species tend to be polyphagous. Based on interpolated ranges, mean elevational range for these moths was larger with increasing elevation, in accordance with Rapoport's elevational rule, although sampling effects may have exaggerated this pattern. The underlying mechanism remains unknown because Rapoport's ‘rescue’ hypothesis could not explain the observed pattern. Conclusions The results clearly show that moth diversity shows a hump‐shaped pattern. However, remarkable variation exists with regard to taxon and range size. Both environmental and geometric factors are likely to contribute to the observed patterns.     

The influence of a native tree species mix component on bird communities in non‐native coniferous plantations in Ireland

Capsule Norway Spruce plantations with Scots Pine as a secondary tree species had higher bird densities than pure Norway Spruce. Shrub cover was the most important structural variable, influencing bird density, species richness and Simpson’s diversity.

Aims To investigate whether incorporating a native tree component into non‐native coniferous plantations had any effect on bird communities or vegetation structure.

Methods Birds were surveyed in plantations of Norway Spruce mixed with Oak and Scots Pine, each paired with a plantation of pure Norway Spruce. distance was used to generate bird densities. Bird density, species richness and Simpson’s diversity were compared between each mix type and pure Norway Spruce. glms were used to investigate relationships between structural components of plantations and bird data.

Results Bird communities of mixed plantations differed only slightly in their composition from pure Norway Spruce. Bird density was significantly higher in Scots Pine mixes than in Oak mixes or pure Norway Spruce. Neither species richness nor Simpson’s diversity differed significantly between the plantation types. Some vegetation components differed between the plantations and shrub cover was positively associated with bird density, species richness and Simpson’s diversity. The presence of rides also increased bird density.

Conclusions There is a positive effect on bird communities of including a native tree species in non‐native coniferous plantations, but the magnitude of the effect is small. The influence of shrub cover on birds suggests that forest management may play an important role in determining the utility of plantations for birds. We recommend the establishment of mixed tree species plantations where possible, although, in the case of Oak mixes, the Norway Spruce appeared to suppress growth of the Oak and thus may be restricting its effect on birds. Changes in management, such as planting Oaks in clumps or heavier thinning of the coniferous component, could address this problem.     

Ambient, productive and wind energy, and ocean extent predict global species richness of procellariiform seabirds

Aims Tests of the energy hypothesis for the large‐scale distribution of species richness have largely been concerned with the influence of two alternative forms of environmental energy, temperature and energy from primary productivity, both of which (at least in terrestrial systems) peak within the tropics. Taxa showing extra‐tropical diversity peaks present a potential challenge to the generality of species–energy theory. One such group are pelagic seabirds of the order Procellariiformes that show not only an extra‐tropical diversity peak but one confined to the Southern Ocean, hence a highly asymmetric one. They are distinct in being exceptionally adapted to take advantage of wind energy, which they may rely on for long‐distance ocean foraging for the patchy resources needed to meet their energetic needs. Wind represents a readily available source of kinetic energy, shows a strong latitudinal gradient, and has been largely omitted from species–energy theory. Moreover, maximal benefits of wind are likely to be afforded in areas of greatest available contiguous ocean extent. We compare the relative importance of wind speed, ocean productivity (chlorophyll concentration), air temperature and available ocean extent (distance) in explaining large‐scale global distribution of procellariiform species richness across the world's oceans. Location Global, oceanic. Methods Hierarchical partitioning, model selection, ordinary least squares (OLS) and spatial generalized least squares (GLS) regression. Results Hierarchical partitioning of non‐spatial regression models indicates that ocean distance is the most important predictor of procellariiform species richness followed by wind speed and then temperature. In contrast, that of spatial regression models indicates the roughly equal importance of ocean distance and temperature, followed by wind speed. Although contributing additional model fit, ocean productivity is consistently the weakest predictor. Best‐fit models include all four predictors and explain 67% of observed variation. The species–productivity relationship is negative overall, while the species–temperature relationship is hump‐shaped. In contrast, ocean distance and wind speed are positively associated with species richness. Conclusions Large‐scale procellariiform species richness distribution may represent a trade‐off in the use of different energy forms, being highest in Southern Ocean areas where productive energy and temperature are relatively low, but where available ocean foraging extent and wind energy required to utilize it are near‐maximal.