A Large-scale, Multihabitat Inventory of the Phylum Tardigrada in the Great Smoky Mountains National Park, USA: A Preliminary Report

An All Taxa Biodiversity Inventory (ATBI) is underway in the Great Smoky Mountains National Park (GSMNP), with the goal of attempting to identify all species of life in the 2000 km² park. The GSMNP is a hotbed of biodiversity, a U.N. Biosphere Reserve, and one of the largest protected, deciduous forests in the temperate world. We have completed two field seasons of work on the tardigrades in the park (2001–2002). As of July 2003, we have collected 420 samples from soil/decomposed leaf litter, lichens and mosses on trees, and stream sediment and periphyton. A few samples from caves, bird nests, and lichens/mosses on rocks were also collected. Samples were taken from within permanent plots established for the ATBI, representing the major biological communities of the GSMNP. Tardigrades were extracted from samples using centrifugation with Ludox AM™, individually mounted on microscope slides in Hoyer’s medium, and studied with phase contrast and DIC microscopy. We have examined 1524 slides from 60 samples as of July 2003. Prior to our work, only three species of tardigrades had been previously reported from a few samples in the park. We have now recorded 42 species, 8 of which we believe may be new to science. Species richness estimates were calculated using EstimateS 6 software for each of the major tardigrade habitats. Overall, we predict that there are 47 to 76 species in the GSMNP, with generally similar species richness in soil, lichen, moss, and stream habitats. Species richness estimates were also used to determine that the number of tardigrade species was greater in mosses at breast height on trees than in mosses at the base of trees.     

A comparison of alpha and beta diversity patterns of ferns,bryophytes and macrolichens in tropical montane forests of southern Ecuador

We present a first comparison of patterns of alpha and beta diversity of ferns, mosses, liverworts and macrolichens in neotropical montane rainforests, and explore the question whether specific taxa may be used as surrogates for others. In three localities in southern Ecuador, we surveyed terrestrial and epiphytic species assemblages in ridge and slope forests in 28 plots of 400 m2 each. The epiphytic habitat was significantly richer in ferns, liverworts, and macrolichens than the terrestrial habitat; mosses, however, were primarily terrestrial. Alpha diversity of ferns and of liverworts was congruent in both habitats. Mosses were similar to ferns and liverworts only in the epiphytic habitat. Macrolichens did not share patterns of alpha diversity with any other group. Beta diversity of ferns, mosses and liverworts (lichens excluded due to low species richness) was similar in the terrestrial habitat, but not in the epiphytic habitat. Our results demonstrate that patterns of alpha diversity of the studied taxa cannot be used to predict patterns of beta diversity. Moreover, diversity patterns observed in epiphytes are different from terrestrial plants. We noted a general coincidence in species patterns of liverworts and ferns. Diversity patterns of macrolichens, in contrast, were completely independent from any other taxonomic group studied.     

Spatial gradients in species diversity of microscopic animals: the case of bdelloid rotifers at high altitude

Aim Organisms smaller than 2 mm appear not to follow the spatial patterns in richness and diversity commonly observed in macroscopic organisms. We describe spatial patterns in species diversity in a group of microscopic organisms, bdelloid rotifers, living in moss and lichen patches, in order to test the hypotheses of no relationship between species richness and composition and spatial gradients, suggested by previously published patterns in microscopic organisms. Location Moss and lichen patches as habitats for bdelloids, on high‐elevation peaks at altitudes between 2984 and 4527 m a.s.l. across the Italian, French and Swiss Alps, with distances among sample sites ranging from 1 m to 420 km, in comparison with lower‐elevation samples at altitudes from 850 to 1810 m a.s.l. Methods We sampled species assemblages of bdelloid rotifers living in isolated moss and lichen patches in 47 sites. We described the observed α, β and γ diversities; the heterogeneity of species assemblages; and the estimated number of species (incidence‐based coverage estimator). Patterns in species distribution were analysed at three different levels: (1) habitat, comparing species richness on moss and lichen substrates, testing differences in α diversity and heterogeneity (anova ), species composition (analysis of similarities test), and γ diversity (rarefaction curves); (2) altitude, comparing the observed richness with previously published data from locations well below 2000 m; and (3) distances between sites, correlating the matrix of Jaccard dissimilarities and the matrix of geographical distances with a Mantel test. Results Both species richness and species composition of bdelloid rotifers differed significantly between mosses and lichens at high elevations, but no difference was found in the heterogeneity of species assemblages. Alpha diversity was significantly lower at high‐elevation than at low‐elevation sites, but the estimated number of species was not reduced when compared with sites at low elevations. Geographical distance between sites had no effect on species composition in either mosses or lichens. The distribution of species was highly heterogeneous, with a low similarity among assemblages. Main conclusions As expected, bdelloids appear to occupy habitats selectively. The altitudinal gradient in species richness for bdelloid rotifers is limited to a decrease in α diversity only; such a decrease is not caused by a lower number of species (low γ diversity) being able to tolerate harsh conditions, and high‐altitude species are not a subset of species living at lower elevations. The observed values of α, β and γ diversity at high altitudes in the Alps are compatible with the scenario of a very low number of available propagules because of the low density of patches of favourable habitat. Our results suggest that the geographical distribution of animals, and therefore biodiversity patterns, may be strongly influenced by animal size, as small organisms such as bdelloids appear to show spatial patterns that differ from those known in larger animals. Differences in body size should be taken into account carefully in future studies of biodiversity patterns.     

Wasp community responses to habitat complexity in Sydney sandstone forests

Abstract Habitat structure and complexity affect the diversity and composition of fauna in a number of systems. We investigated patterns in wasp species richness, abundance and composition and also their associations with habitat complexity in Sydney sandstone forests, Australia. Pitfall and flight‐intercept traps collected dissimilar wasp assemblages. High complexity habitats supported greater abundance and species richness and a dissimilar composition of pitfall‐trapped wasps to low complexity habitats. Soil moisture, tree canopy cover, ground herb cover and shrub canopy cover all had significant positive associations with the species richness of pitfall‐trapped wasps. Although the five most abundant families of wasps we trapped are endoparasitoids of other arthropods, they showed a variety of preferences for habitat variables. The mechanisms driving associations between habitat complexity and patterns in wasp communities may also provide a basis for understanding factors influencing the regulation of arthropod assemblages by wasps in agricultural and natural landscapes.     

Spatial structure, rock type, and local environmental conditions drive moss and lichen distribution on calcareous boulders

Although mosses and lichens are a relevant component of the biota of rock habitats targeted for biodiversity conservation in Europe, the ecological factors driving their distribution are still poorly known. In this work, we examined the epilithic moss and lichen assemblages colonizing boulders of different types of calcareous rocks co-occurring in the same area in the Italian Alps. The goals were: (1) to evaluate if and to what extent different calcareous rocks host different assemblages; (2) to identify species associated to each rock type; (3) to quantify the relative importance of rock type, local environmental factors, and habitat spatial structure in explaining species distribution. Our results demonstrated that different calcareous rocks host different moss and lichen assemblages with some typical species, indicating that each rock type contributes to the total diversity of both mosses and lichens. Local environmental conditions influenced mosses and not lichens whose distribution is mainly associated to rock type. The patterns of both organism groups were also significantly related to habitat spatial structure, species assemblages tending to have a patchy distribution, which may reflect dispersal dynamics. Our results have implications for conservation: (1) each rock type may play a relevant role in maintaining the overall diversity contributing with unique assemblages and typical species; (2) the patchy distribution of both moss and lichen assemblages should warn from considering rock patches as a monotonous repetition of the same habitat across space.     

Survival of freezing by hydrated tardigrades inhabiting terrestrial and freshwater habitats

The seasonality and unpredictability of environmental conditions at high altitudes and latitudes govern the life cycle patterns of organisms, giving rise to stresses that cause death or development of specific adaptations. Ice formation is a major variable affecting the survival of both freshwater fauna and fauna inhabiting lichens, mosses and leaf litter. Tardigrades occupy a wide range of niches in marine, freshwater and terrestrial environments. The highest number of species is found in terrestrial habitats thanks to their ability to enter anhydrobiosis and cryobiosis. The cryobiotic ability of tardigrade species from polar regions is well known. Consequently, we focused our research on the ability to survive freezing in the active hydrated state using seven tardigrade species differing in phylogenetic position and collected at various altitudes and from different habitats in a temperate area. Specimens were cooled at different cooling rates (from 0.31 °C min⁻¹ to 3.26 °C min⁻¹). Even though the final survival and the time required by animals to recover to active life were both inversely related to the cooling rate, highly significant interspecific differences were found. Species survival ability ranged from excellent to none. Species living in xeric habitats withstood freezing better than those living in hygrophilous habitats, while true limnic species did not exhibit any cryobiotic ability. The ability to withstand freezing seems linked to the anhydrobiotic ability. The differences in cryptobiotic performance among tardigrade species seem more influenced by selective pressures linked to local adaptation to habitat characteristics than by phylogenetic relationships.     

Patterns of diversity in microscopic animals: are they comparable to those in protists or in larger animals?

Aim General patterns of biodiversity, such as latitudinal gradients and species‐area relationships, are found consistently in a wide range of organisms, but recent results for protist diversity suggest that organisms shorter than 2 mm do not display such patterns. We tested this prediction in bdelloid rotifers, pluricellular metazoans smaller than 2 mm, but with size and ecology comparable to protists. Location A single valley in northern Italy was surveyed in detail and compared to all available faunistic data on bdelloids worldwide. Methods We analysed 171 local assemblages of bdelloid rotifers living in 5 systems of dry mosses and submerged mosses in running water and in lakes. We compared patterns of alpha, beta, and gamma diversity, and nestedness of metacommunities, with those known from protists and larger organisms. Results Bdelloid rotifers showed low local species richness (alpha diversity), with strong habitat selection, as observed in larger organisms. The number of species differed among systems, with a higher number of species in dry than in aquatic mosses. There was no hierarchical structure or exclusion of species in the metacommunity pattern within each system. Local diversity for the entire valley was surprisingly high compared with worldwide bdelloid diversity, similar to observed patterns in protists. Main Conclusions Bdelloid rotifers have some of the peculiarities of protist biodiversity, although at slightly different spatial scales, thus confirming the idea of a major change in biodiversity patterns among organisms shorter than 2 mm. However, bdelloids show stronger habitat selection than protists. We suggest two possible explanations for the observed patterns: (1) dispersal is very rare, and not all bdelloid clones are arriving everywhere; and (2) dispersal is effective in displacing propagules, but environmental heterogeneity is very high and prevents many species from colonizing a given patch of moss.     

Differences in leaf‐litter invertebrate assemblages between radiata pine plantations and neighbouring native eucalypt woodland

Abstract We investigated the structure, composition and environmental correlates of leaf‐litter invertebrate assemblages in Pinus radiata plantations and in neighbouring native eucalypt woodland in the Jenolan Caves Karst Conservation Reserve, south‐east Australia. Invertebrate assemblages of plantations were compared with remnant eucalypt woodland located well away from the influence of plantations to determine the direct effects of plantations as a result of habitat‐replacement with a non‐native plantation species. We also included in our comparisons edge habitat of eucalypt woodland located immediately adjacent to plantations. This unique edge habitat is exposed to the intrusion of large volumes of pine leaf‐litter from plantations, which has the potential to affect indirectly invertebrate assemblages of surrounding woodland. We found that species richness of invertebrates was significantly lower in pine plantations compared with remnant eucalypt woodland. There was a complete absence of species from 12 invertebrate orders that were found in surrounding eucalypt woodland. A rich and abundant native plant understorey that provides increased habitat heterogeneity is the most likely explanation for the richer invertebrate assemblage found in remnant eucalypt woodland. The total abundance of all invertebrate taxa in pine plantations in winter was significantly higher than in remnant eucalypt woodland, pine‐litter edges and pine‐free edges. Plantations were characterized by particularly high abundances of species in two orders, Acari and Collembola. High abundances of acarine and collembolan species in plantations were associated with a decompositional environment represented by comparatively higher moisture contents and higher C : N ratios of both leaf‐litter and soil, higher soil conductivity and lower soil pH. We suggest that implementation of The Plantation Biodiversity Benefits Score will be a fruitful way forward to assess the environmental benefits that can be gained from pine plantations in this region of south‐eastern Australia.     

Marine dock pilings foster diverse,native cryptobenthic fish assemblages across bioregions

Anthropogenic habitats are increasingly prevalent in coastal marine environments. Previous research on sessile epifauna suggests that artificial habitats act as a refuge for nonindigenous species, which results in highly homogenous communities across locations. However, vertebrate assemblages that live in association with artificial habitats are poorly understood. Here, we quantify the biodiversity of small, cryptic (henceforth “cryptobenthic”) fishes from marine dock pilings across six locations over 35° of latitude from Maine to Panama. We also compare assemblages from dock pilings to natural habitats in the two southernmost locations (Panama and Belize). Our results suggest that the biodiversity patterns of cryptobenthic fishes from dock pilings follow a Latitudinal Diversity Gradient (LDG), with average local and regional diversity declining sharply with increasing latitude. Furthermore, a strong correlation between community composition and spatial distance suggests distinct regional assemblages of cryptobenthic fishes. Cryptobenthic fish assemblages from dock pilings in Belize and Panama were less diverse and had lower densities than nearby reef habitats. However, dock pilings harbored almost exclusively native species, including two species of conservation concern absent from nearby natural habitats. Our results suggest that, in contrast to sessile epifaunal assemblages on artificial substrates, artificial marine habitats can harbor diverse, regionally characteristic assemblages of vertebrates that follow macroecological patterns that are well documented for natural habitats. We therefore posit that, although dock pilings cannot function as a replacement for natural habitats, dock pilings may provide cost‐effective means to preserve native vertebrate biodiversity, and provide a habitat that can be relatively easily monitored to track the status and trends of fish biodiversity in highly urbanized coastal marine environments.     

The Tardigrades of Emilia (Italy). III. Piane di Mocogno (Northern Apennines)

Samples of leaf litter, mosses and lichens were collected in a beech forest at Piane di Mocogno (Emilia, Italy), 1200 m asl. The tardigrade community of the leaf litter was similar to that found in this substrate in Italy and in the U.S.A. At Piane di Mocogno the species association within the leaf litter was different from that in mosses and lichens (which have similar fauna), confirming that very different communities characterise different substrates such as leaf litter and mosses. During this faunal analysis, six species of tardigrades were found for the first time in this area. The presence of Microhypsibius bertolanii (first record in leaf litter and in Italy) and Eohypsibius nadjae are note worthy. Ornamented eggs in an exuvium of Hypsibius cf.scabropygus were also recorded for the first time.     

Dung beetle assemblages in primary forest and disturbed habitats in a tropical dry forest landscape in western Mexico

Dung beetles (Scarabaeinae) are conspicuous components in most terrestrial ecosystems, performing important ecological functions and services. Being sensitive to several types of disturbance, they have been successfully used as indicators of habitat change. Dung beetle communities in tropical rainforests have been well studied, but considerably less information is available for tropical dry forests. In this study I sampled dung beetles in two undisturbed habitats, deciduous forest and semideciduous forest, and two disturbed habitats, secondary forest and open area habitat, in the Chamela-Cuixmala region of western Mexico. Dung beetle species with high indicator value for each habitat were identified. Beetle abundance, observed species richness and estimated species richness were similar in the three forest habitats, but significantly lower in the open area habitat. A more detailed analysis of species-specific abundances in the three forest habitats revealed some differences. Transects of one of the undisturbed habitats, the deciduous forest, were more similar to the non-adjacent transects of disturbed secondary forest, than to the adjacent undisturbed semideciduous forest transects. Unlike studies in other tropical sites that have found a decrease in equitability in Scarabaeinae assemblages between undisturbed forest and disturbed habitat (particularly open habitats), in the Chamela-Cuixmala region all four habitats showed similar low equitability in community structure, with two or three very dominant species.     

Declines in littoral species richness across both spatial and temporal nutrient gradients: a palaeolimnological study of two taxonomic groups

1. Using a palaeolimnological approach in shallow lakes, we quantified the species richness responses of diatoms and Cladocera to phosphorus enrichment. We also examined differences in species richness responses between littoral and pelagic assemblages of our focal communities. To address both spatial and temporal relationships, our study includes an analysis of both surface sediments from 40 lakes and of a lake sediment record spanning c. 120 years. The objective of our study was to determine whether similar species richness patterns occurred across trophic levels, as well as along spatial and temporal gradients. 2. We found that both diatom and Cladocera species richness estimates significantly declined with increasing phosphorus across space and through time. When the assemblages were subdivided according to known habitat preferences, littoral biodiversity maintained a negative trend, whereas pelagic species richness tended to show no relationship with phosphorus. 3. Negative productivity–diversity patterns have been observed across almost all palaeolimnological studies that span large productivity gradients. This congruence in patterns is most likely due to the similarity in data collection methods and in focal communities studied. The contrasting responses between littoral and pelagic assemblages may be explained by the differences in physical habitat and the pool of taxa in each of these environments. Consistent with the literature, we found statistical support for the idea that littoral diversity declines could be explained by an interaction between macrophytes and nutrients along strong trophic gradients. The general lack of a diversity response in our pelagic assemblages could be attributable to the limited pool of subfossil taxa. The response of the pelagic diatom could also be related to their broad range of nutrient tolerances. 4. The observed negative response of species richness to phosphorus enrichment, particularly in the littoral assemblages, has implications for ecosystems functioning because communities with reduced biodiversity often are less resilient to anthropogenic change.     

Dispersal ability links to cross‐scale species diversity patterns across the Eurasian Arctic tundra

Aim The role of dispersal in structuring biodiversity across spatial scales is controversial. If dispersal controls regional and local community assembly, it should also affect the degree of spatial species turnover as well as the extent to which regional communities are represented in local communities. Here we provide the first integrated assessment of relationships between dispersal ability and local‐to‐regional spatial aspects of species diversity across a large geographical area. Location Northern Eurasia. Methods Using a cross‐scale analysis covering local (0.64 m²) to continental (the Eurasian Arctic biome) scales, we compared slope parameters of the dissimilarity‐to‐distance relationship in species composition and the local‐to‐regional relationship in species richness among three plant‐like groups that differ in dispersal ability: lichens with the highest dispersal ability; mosses and moss allies with intermediate dispersal ability; and seed plants with the lowest dispersal ability. Results Diversity patterns generally differed between the three groups according to their dispersal ability, even after controlling for niche‐based processes. Increasing dispersal ability is linked to decreasing spatial species turnover and an increasing ratio of local to regional species richness. All comparisons supported our expectations, except for the slope of the local‐to‐regional relationship in species richness for mosses and moss allies which was not significantly steeper than that of seed plants. Main conclusions The negative link between dispersal ability and spatial species turnover and the corresponding positive link between dispersal ability and the ratio of local‐to‐regional species richness support the idea that dispersal affects community structure and diversity patterns across spatial scales.     

Patterns of diversity and habitat relationships in terrestrial mollusc communities of the Pukeamaru Ecological District, northeastern New Zealand

Aim The New Zealand terrestrial mollusc fauna is among the most speciose in the world, with often remarkably high richness at lowland forest sites. We sought to elucidate general explanations for patterns of richness in terrestrial mollusc communities by analysis of species coexistence and habitat relationships within a New Zealand district fauna. Location Pukeamaru Ecological District, eastern North Island, New Zealand. Methods We sampled molluscs using qualitative methods at twenty-three sites and quantitatively by frame sampling of scrubland-forest floor litter at sixteen of these sites and analysed patterns of species richness and turnover in relation to regional species pools and local habitat attributes. We then tested for nonrandom assemblage of taxa along diversity and habitat gradients. Results Ninety-four indigenous mollusc species were recorded from a district fauna estimated at 102 indigenous species: only two species were endemic. From the presumptive geological history of the district, the low endemism, and Brooks parsimony and indicator species analyses of faunal relationships, the communities were indicated to have resulted by accumulation of colonists from other New Zealand districts since the Miocene. Richness ranged from two or three indigenous species in dune habitats to fifty-nine species in a floristically rich forest. Beta diversity was high and site occupancy per species was low, indicating communities structured by successive replacement of ecological equivalents. Sites differing in vegetation had characteristic species assemblages, indicating a degree of habitat specialization. Canonical correspondence analysis indicated that canopy tree species, canopy height, floristic diversity, altitude, litter mass, and litter pH were important determinants of species assemblage in scrubland and forest. Richness was strongly associated with site floristic diversity and, for litter-dwelling species, the pH of litter substrate. High richness occurred at those sites supporting molluscs in high abundance. Shell-shape distributions were essentially Cainian unimodal, with communities dominated by snail species with subglobose to discoidal shells. Mean and variance of shell size increased with mollusc species richness and floristic diversity at sites, indicating dominance of communities by small-shelled species at early successional or floristically poor sites, and increased richness resulting from addition of larger snails into vacant niches. Shifts in shell form were associated with sympatry in several congeneric taxa. Main conclusions The underdispersion of shell shape, relative to faunas elsewhere in the world, indicates that community structure in New Zealand land snail faunas has been constrained by limited phylogenetic diversity and/or by convergence upon successful adaptations. The remarkably high richness that characterizes these communities indicates special conditions allow coexistence of numerous species. The relationship between floristic diversity at sites and the richness, diversity, and shell-size distributions of the molluscs suggests assemblages structured around niche partitioning among competing species. While there is an element of congruence between vegetation and mollusc pattern, this study indicates that assembly rules will be defined, and spatial pattern predicted, only through a better understanding of the linkage between regional species pool, organism traits, environment, and local community assemblage.     

Nutrient enrichment weakens the stabilizing effect of species richness

With global freshwater biodiversity declining at an even faster rate than in the most disturbed terrestrial ecosystems, understanding the effects of changing environmental conditions on relationships between biodiversity and the variability of community and population processes in aquatic ecosystems is of significant interest. Evidence is accumulating that biodiversity loss results in more variable communities; however, the mechanisms underlying this effect have been the subject of considerable debate. We manipulated species richness and nutrients in outdoor aquatic microcosms composed of naturally occurring assemblages of zooplankton and benthic invertebrates to determine how the relationship between species richness and variability might change under different nutrient conditions. Temporal variability of populations and communities decreased with increasing species richness in low nutrient microcosms. In contrast, we found no relationship between species richness and either population or community variability in nutrient enriched microcosms. Of the different mechanisms we investigated (e.g. overyielding, statistical averaging, insurance effects, and the stabilizing effect of species richness on populations) the only one that was consistent with our results was that increases in species richness led to more stable community abundances through the stabilizing effect of species richness on the component populations. While we cannot conclusively determine the mechanism(s) by which species richness stabilized populations, our results suggest that more complete resource-use in the more species-rich low nutrient communities may have dampened population fluctuations.     

Evolution of termite functional diversity: analysis and synthesis of local ecological and regional influences on local species richness

Aim To (1) describe termite functional diversity patterns across five tropical regions using local species richness sampling of standardized areas of habitat; (2) assess the relative importance of environmental factors operating at different spatial and temporal scales in influencing variation in species representation within feeding groups and functional taxonomic groups across the tropics; (3) achieve a synthesis to explain the observed patterns of convergence and divergence in termite functional diversity that draws on termite ecological and biogeographical evidence to‐date, as well as the latest evidence for the evolutionary and distributional history of tropical rain forests. Location Pantropical. Methods A pantropical termite species richness data set was obtained through sampling of eighty‐seven standardized local termite diversity transects from twenty‐nine locations across five tropical regions. Local‐scale, intermediate‐scale and large‐scale environmental data were collected for each transect. Standardized termite assemblage and environmental data were analysed at the levels of whole assemblages and feeding groups (using components of variance analysis) and at the level of functional taxonomic groups (using correspondence analysis and canonical correspondence analysis). Results Overall species richness of local assemblages showed a greater component of variation attributable to local habitat disturbance level than to region. However, an analysis accounting for species richness across termite feeding groups indicated a much larger component of variation attributable to region. Mean local assemblage body size also showed the greater overall significance of region compared with habitat type in influencing variation. Ordination of functional taxonomic group data revealed a primary gradient of variation corresponding to rank order of species richness within sites and to mean local species richness within regions. The latter was in the order: Africa > south America > south‐east Asia > Madagascar > Australia. This primary gradient of species richness decrease can be explained by a decrease in species richness of less dispersive functional taxonomic groups feeding on more humified food substrates such as soil. Hence, the transects from more depauperate sites/regions were dominated by more dispersive functional taxonomic groups feeding on less humified food substrates such as dead wood. Direct gradient analysis indicated that ‘region’ and other large‐scale factors were the most important in explaining patterns of local termite functional diversity followed by intermediate‐scale geographical and site variables and, finally, local‐scale ecological variables. Synthesis and main conclusions Within regions, centres of termite functional diversity lie in lowland equatorial closed canopy tropical forests. Soil feeding termite evolution further down food substrate humification gradients is therefore more likely to have depended on the long‐term presence of this habitat. Known ecological and energetic constraints upon contemporary soil feeders lend support for this hypothesis. We propose further that the anomalous distribution of termite soil feeder species richness is partly explained by their generally very poor dispersal abilities across oceans. Evolution, radiation and dispersal of soil feeder diversity appears to have been largely restricted to what are now the African and south American regions. The inter‐regional differences in contemporary local patterns of termite species richness revealed by the global data set point to the possibility of large differences in consequent ecosystem processes in apparently similar habitats on different continents.     

Diversity of European habitat types is correlated with geography more than climate and human pressure

Habitat richness, that is, the diversity of ecosystem types, is a complex, spatially explicit aspect of biodiversity, which is affected by bioclimatic, geographic, and anthropogenic variables. The distribution of habitat types is a key component for understanding broad‐scale biodiversity and for developing conservation strategies. We used data on the distribution of European Union (EU) habitats to answer the following questions: (i) how do bioclimatic, geographic, and anthropogenic variables affect habitat richness? (ii) Which of those factors is the most important? (iii) How do interactions among these variables influence habitat richness and which combinations produce the strongest interactions? The distribution maps of 222 terrestrial habitat types as defined by the Natura 2000 network were used to calculate habitat richness for the 10 km × 10 km EU grid map. We then investigated how environmental variables affect habitat richness, using generalized linear models, generalized additive models, and boosted regression trees. The main factors associated with habitat richness were geographic variables, with negative relationships observed for both latitude and longitude, and a positive relationship for terrain ruggedness. Bioclimatic variables played a secondary role, with habitat richness increasing slightly with annual mean temperature and overall annual precipitation. We also found an interaction between anthropogenic variables, with the combination of increased landscape fragmentation and increased population density strongly decreasing habitat richness. This is the first attempt to disentangle spatial patterns of habitat richness at the continental scale, as a key tool for protecting biodiversity. The number of European habitats is related to geography more than climate and human pressure, reflecting a major component of biogeographical patterns similar to the drivers observed at the species level. The interaction between anthropogenic variables highlights the need for coordinated, continental‐scale management plans for biodiversity conservation.     

Global macroecology of bird assemblages in urbanized and semi‐natural ecosystems

Aim Despite the increasing pace of urbanization, little is known about how this process affects biodiversity globally. We investigate macroecological patterns of bird assemblages in urbanized areas relative to semi‐natural ecosystems. Location World‐wide. Methods We use a database of quantitative bird surveys to compare key assemblage structure parameters for plots in urbanized and semi‐natural ecosystems controlling for spatial autocorrelation and survey methodology. We use the term ‘urbanized’ instead of ‘urban’ ecosystems as many of the plots were not located in the centre of towns but in remnant habitat patches within conurbations. Results Some macroecological relationships were conserved in urbanized landscapes. Species–area, species–abundance and species–biomass relationships did not differ significantly between urbanized and non‐urbanized environments. However, there were differences in the relationships between productivity and assemblage structure. In forests, species richness increased with productivity; in both forests and open habitats, the evenness of species abundances declined as productivity increased. Among urbanized plots, instead, both species richness and the evenness of species abundances were independent of variation in productivity. Main conclusions Remnant habitats within urbanized areas are subject to many ecological alterations, yet key macroecological patterns differ remarkably little in urbanized versus non‐urbanized plots. Our results support the need for increased conservation activities in urbanized landscapes, particularly given the additional benefits of local experiences of biodiversity for the human population. With increasing urbanization world‐wide, broad‐scale efforts are needed to understand and manage the effects of this driver of change on biodiversity.     

Development of a diverse epiphyte community in response to phosphorus fertilization

The role of terrestrial soil nutrient supply in determining the composition and productivity of epiphyte communities has been little investigated. In a montane Hawaiian rainforest, we documented dramatic increases in the abundance and species richness of canopy epiphytes in a forest that had been fertilized annually with phosphorus (P) for 15 years; there was no response in forest that had been fertilized with nitrogen (N) or other nutrients. The response of N-fixing lichens to P fertilization was particularly strong, although mosses and non-N-fixing lichens also increased in abundance and diversity. We show that enhancement of canopy P availability is the most likely factor driving the bloom in epiphytes. These results provide strong evidence that terrestrial soil fertility may structure epiphyte communities, and in particular that the abundance of N-fixing lichens – a functionally important epiphyte group – may be particularly sensitive to ecosystem P availability.     

Additive partitioning of aquatic invertebrate species diversity across multiple spatial scales

1. Additive partitioning of three measures of diversity (species richness, Shannon's diversity index H and Simpson's diversity D) was used to study the relationship between local and regional diversity of benthic macroinvertebrate communities of boreal lakes (littoral habitats) and streams (riffle habitats) across three spatial scales (sampling sites, ecoregions and biogeographic regions). 2. Alpha (α) and beta (β) diversity are defined as within‐habitat and between‐habitat diversity, respectively. According to the concept of additive partitioning, diversity can be partitioned across multiple spatial scales such that the total (γ) diversity on one spatial scale becomes within‐habitat (α) diversity at the next higher scale. Hence, the total diversity at one scale is determined by the α diversity and the between‐habitat diversity (β) at the next lower scale. Consequently, one of the advantages of additive partitioning is that it is possible to study simultaneously β diversity and the regional‐local species relationship and the scale dependence of α and β components. 3. For both lakes and streams α diversity was low for sites and ecoregions, whereas β diversity was high, indicating that among‐site factors are important in describing the variability among the lakes and streams studied here. 4. Weak, albeit significant, evidence was found for regional and local species saturation patterns. Multiple stepwise regression indicated that local processes might be more important in structuring lake‐littoral and stream‐riffle species assemblages than regional processes. From these results we conclude that environmental heterogeneity may act as an important factor contributing to species coexistence, resulting in the observed saturation patterns. 5. Our study supports the use of additive partitioning for identifying specific patterns of macroinvertebrate diversity on multiple spatial scales and the underlying processes generating these patterns. This information is needed to improve understanding of the relation between patterns and processes affecting (decreasing) trends in aquatic biodiversity.