Expanded view of the local–regional richness relationship by incorporating functional richness and time: a large‐scale perspective

Aim We investigate the relationship between local and regional richness in marine fouling assemblages using an expanded and globally replicated approach by incorporating two dimensions of diversity (taxonomic and functional) and different successional stages. Location Global. Methods In eight different biogeographic regions (Australia, Brazil, Chile, England, Italy, Japan, Portugal and Sweden) 68 polyvinylchloride (PVC) panels (15 × 15 × 0.3 cm) were deployed for colonization. Communities colonizing panels were analysed by measuring percentage cover at each of four different successional ages: 2, 4, 6 and 8 months. Local richness was assessed as the average number of species and functional groups (FGs) per panel and regional richness was evaluated as the estimated (Jack2) asymptote of the sample‐accumulation curves for species and FG on experimental panels. Results We found that the shape of the relationship between local and regional richness depended on successional stage and the type of richness considered, i.e. taxonomic or functional richness. Hardly any relationship was detectable between local taxonomic richness and regional taxonomic richness at any successional stage. In contrast, the relation between local functional and regional functional richness shows a unimodal pattern of change during succession, passing through the stages ‘independent’, ‘unsaturated rising’, ‘saturated rising’ and once again ‘independent’. Main conclusions The relationship between local and regional richness, whether taxonomic or functional, frequently displays independence of the two scales, particularly in early and late phases of the successional process.     

Water–energy,land‐cover and heterogeneity drivers of the distribution of plant species richness in a mountain region of the European Alps

Aim To evaluate the relative importance of water–energy, land‐cover, environmental heterogeneity and spatial variables on the regional distribution of Red‐Listed and common vascular plant species richness. Location Trento Province (c. 6200 km²) on the southern border of the European Alps (Italy), subdivided regularly into 228 3′ × 5′ quadrants. Methods Data from a floristic inventory were separated into two subsets, representing Red‐Listed and common (i.e. all except Red‐Listed) plant species richness. Both subsets were separately related to water–energy, land‐cover and environmental heterogeneity variables. We simultaneously applied ordinary least squares regression with variation partitioning and hierarchical partitioning, attempting to identify the most important factors controlling species richness. We combined the analysis of environmental variables with a trend surface analysis and a spatial autocorrelation analysis. Results At the regional scale, plant species richness of both Red‐Listed and common species was primarily related to energy availability and land cover, whereas environmental heterogeneity had a lesser effect. The greatest number of species of both subsets was found in quadrants with the largest energy availability and the greatest degree of urbanization. These findings suggest that the elevation range within our study region imposes an energy‐driven control on the distribution of species richness, which resembles that of the broader latitude gradient. Overall, the two species subsets had similar trends concerning the relative importance of water–energy, land cover and environmental heterogeneity, showing a few differences regarding the selection of some predictors of secondary importance. The incorporation of spatial variables did not improve the explanatory power of the environmental models and the high original spatial autocorrelation in the response variables was reduced drastically by including the selected environmental variables. Main conclusions Water–energy and land cover showed significant pure effects in explaining plant species richness, indicating that climate and land cover should both be included as explanatory variables in modelling species richness in human‐affected landscapes. However, the high degree of shared variation between the two groups made the relative effects difficult to separate. The relatively low range of variation in the environmental heterogeneity variables within our sampling domain might have caused the low importance of this complex factor.     

The control of community composition by distance,environment and history: a regional‐scale study of the mountain grasslands of southern New Zealand

Aim To examine the roles of physical and biotic environment, distance, direction and dispersal in determining the composition of plant communities at a regional scale. Location Grassland and shrubland at 266 sites in the mountains of southern South Island, New Zealand. Methods Species abundances of vascular plants, bryophytes and lichens were measured by height‐frequency. Predictors were soils, climate, mammal grazing, physical distance and direction between plots, and geological history represented by occurrence on the same or different mountain ranges. Relationships were evaluated with quadratic multiple regression and Mantel tests. Results Spatial autocorrelation was present for both vascular and lower plants. However, distance explained only a minor part of the variation. Distance relationships were anisotropic, with the vegetation differences being greatest in the direction of the prevailing south‐westerly wind. Even using all the environmental information, much of the species composition remained unexplained. For species presence, distance alone explained 7% of the variation, environment alone 25% and both together 28%. The abundance of species was even less predictable: distance 2%, environment 11% and both 12%. With vascular species, climate‐related factors contributed the most to prediction, especially altitude. Surprisingly, soil chemical factors were more important for lower plants that are not rooted in the substrate. Species likely to disperse further showed a tendency towards weaker distance differentiation. Vegetation composition seemed unrelated to contemporary grazing, although historical grazing/burning may have been causal. There was little evidence for a relationship of species composition with biogeographic/geological history. Main conclusions Difference in community composition in these grasslands is only weakly related to distance, and only half of the distance effect can be explained by the environmental factors measured. Dispersal and geological history do not seem to explain the discrepancy. Explicability of community differences was comparable to that in some previous case studies, although weaker than that in others, but all of the previous studies examined only part of the flora, such as ‘abundant trees’ or palms – none examined even all the vascular species. The species complement of a site seems more conservative than the abundance of the species, much of which was not explicable from current information and may reflect past management history.     

Spatial Analysis of Twaite Shad,Alosa fallax (Lacepède, 1803), in the Southern North Sea: Application of Non‐Linear Geostatistics as a Tool to Search for Special Areas of Conservation

This study aims to evaluate the suitability of non‐linear geostatistics and indicator kriging (IK) as a tool in environmental impact assessment and nature conservation, in particular to search for potential Special Areas of Conservation (SAC) for the endangered fish species twaite shad, Alosa fallax (Lacepède , 1803) within the German Exclusive Economical Zone (EEZ) of the North Sea. To analyse the spatial distribution of this fish species, data on standardised biomass index (catch per unit effort, c.p.u.e., kg × 30 min^–1) from 1996 to 2001 were used, regarding the third and fourth quarters of each year, respectively. Thereby we assume that the spatial distribution can be described as a time‐invariant process. This assumption is supported by information on annual sampling effort, allocation of hauls and spatial distribution of the positive catches. All indicator variograms obtained for different c.p.u.e. cut‐off values displayed distinct spatial structures, clearly indicating that the indicator variables were spatially autocorrelated. Gaussian models were fitted by least‐squares methods and were evaluated with a goodness‐of‐fit statistic. Subsequently, IK was employed to estimate the probability of exceeding the c.p.u.e. cut‐off values for the twaite shad in the investigation area. These were highest in the Weser‐ and Elbe‐estuary, probably because of migrations of twaite shad to and from estuaries at the time of investigation due to spawning, while within the German EEZ of the North Sea no such areas with increased probabilities could be discerned. Thus, although available data did not allow to identify and implement any SAC in the German EEZ, the methods employed here can be regarded as a promising management tool in biological conservation issues. (© 2004 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

A new vertebrate for Europe: the discovery of a range‐restricted relict viper in the western Italian Alps

We describe Vipera walser, a new viper species from the north‐western Italian Alps. Despite an overall morphological resemblance with Vipera berus, the new species is remarkably distinct genetically from both V. berus and other vipers occurring in western Europe and shows closer affinities to species occurring only in the Caucasus. Morphologically, the new species appear to be more similar to V. berus than to its closest relatives occurring in the Caucasus, but can be readily distinguished in most cases by a combination of meristic features as confirmed by discriminant analysis. The extant population shows a very low genetic variability measured with mitochondrial markers, suggesting that the taxon has suffered a serious population reduction/bottleneck in the past. The species is extremely range‐restricted (less than 500 km²) and occurs only in two disjunct sites within the high rainfall valleys of the Alps north of Biella. This new species should be classified as globally ‘endangered’ due to its small and fragmented range, and an inferred population decline. The main near‐future threats to the species are habitat changes associated with reduced grazing, along with persecution and collecting.     

Macaronesia: a source of hidden genetic diversity for post‐glacial recolonization of western Europe in the leafy liverwort Radula lindenbergiana

Aim Bryophytes exhibit apparently low rates of endemism in Macaronesia and differ from angiosperms in their diversity patterns by the widespread occurrence of endemics within and among archipelagos. This paper investigates the phylogeography of the leafy liverwort Radula lindenbergiana to determine: (1) whether or not morphologically cryptic diversification has occurred in Macaronesia, and (2) the relationships between Macaronesian and continental populations. Location Macaronesia, Europe, Africa. Methods Eighty‐four samples were collected across the species’ distribution range and sequenced at four chloroplast DNA (cpDNA) loci (atpB–rbcL, trnG, trnL and rps4). Phylogenetic reconstructions and Bayesian ancestral area reconstructions were used in combination with population genetics statistics (H, N_ST, F_ST) to describe the pattern of present genetic diversity in R. lindenbergiana and infer its biogeographic history. Results Patterns of genetic diversity in R. lindenbergiana exhibit a striking westwards gradient, wherein haplotype (0.90) and nucleotide (0.0038 ± 0.0019) diversity peak in Macaronesia, with a substantial endemic component. We found 20.9% of the genetic variance between biogeographic regions, and most pairwise F_ST comparisons between regions are significantly different from zero. The global N_ST (0.78) is significantly higher than the global F_ST (0.20), providing evidence for the presence of phylogeographic signal in the data. Ancestral area reconstructions suggest that the haplotypes currently found in western Europe share a Macaronesian common ancestor. Main conclusions The haplotype diversification exhibited by R. lindenbergiana in Macaronesia is comparable to that reported for many angiosperm groups at the species level. The apparent lack of radiation among Macaronesian bryophytes may thus reflect the reduced morphology of bryophytes in comparison with angiosperms. The high diversity found among Macaronesian haplotypes, especially in Madeira and the Canary Islands, and the significant N_ST/F_ST ratio between Macaronesia and all the other biogeographic regions (an indication that mutation rate exceeds dispersal rates) suggest that Macaronesian archipelagos could have served as a refugium during the Quaternary glaciations. Many haplotypes currently found in Europe share a Macaronesian common ancestor, and this further suggests that Macaronesia might have played a key role in the back‐colonization of the continent.     

Determinants of fine‐scale homogenization and differentiation of native freshwater fish faunas in a Mediterranean Basin: implications for conservation

Aim Increasing threats to freshwater biodiversity are rapidly changing the distinctiveness of regional species pools and local assemblages. Biotic homogenization/differentiation processes are threatening the integrity and persistence of native biodiversity patterns at a range of spatial scales and pose a challenge for effective conservation planning. Here, we evaluate the extent and determinants of fine‐scale alteration in native freshwater fish assemblages among stream reaches throughout a large river basin and consider the implications of these changes for the long‐term conservation of native fishes. Location Guadiana River basin (South‐Western Iberian Peninsula). Methods We quantified the magnitude of change in compositional similarity between observed and reference assemblages and its potential effect on natural patterns of compositional distinctiveness. Reference assemblages were defined as the native species expected to occur naturally (in absence of anthropogenic alterations) and were reconstructed using a multivariate adaptive regression splines predictive model. We also evaluated the role of habitat degradation and introduced species as determinants of biotic homogenization/differentiation. Results We found a significant trend towards homogenization for native fish assemblages. Changes in native fish distributions led to the loss of distinctiveness patterns along natural environmental gradients. Introduced species were the most important factor explaining the homogenization process. Homogenization of native assemblages was stronger in areas close to reservoirs and in lowland reaches where introduced species were more abundant. Main conclusions The implementation of efficient conservation for the maintenance of native fish diversity is seriously threatened by the homogenization processes. The identification of priority areas for conservation is hindered by the fact that the most diverse communities are vanishing, which would require the selection of broader areas to adequately protect all the species. Given the principal role that introduced species play in the homogenization process and their relation with reservoirs, special attention must be paid to mitigating or preventing these threats.     

Effects of vegetation state on biodiversity and nitrogen retention in created wetlands: a test of the biodiversity–ecosystem functioning hypothesis

1. Nitrogen retention in wetlands provides an example of an ecosystem function that is desired by human society, and is a rationale for the creation of wetlands to decrease nitrogen fluxes from nitrate‐loaded river catchments to coastal waters. 2. Here, we tested the impact of different vegetation states on species diversity and nitrogen retention during 4 years in surface‐flow wetlands receiving nitrate‐rich water. Tall emergent vegetation or submerged vegetation was introduced to six experimental wetlands each and six wetlands were left as unplanted controls for free development of vegetation. This resulted in three vegetation states dominated by emergent vegetation, by a mixture of submerged vegetation and filamentous green algae or by filamentous green algae. 3. Species diversity (species richness and Shannon diversity) of plants was initially lowest in free development wetlands, but during the study became lower in the emergent vegetation wetlands than in the other wetlands. Diversity of macroinvertebrates was initially lower in the submerged vegetation wetlands than in the other wetlands, but this difference disappeared during the study. Nitrogen retention was consistently higher in emergent vegetation wetlands than in the other wetlands throughout the study. 4. We conclude that plant diversity in wetlands dominated by tall emergent vegetation gradually became lower than in other wetlands, due to dominant species competitively excluding other plants. However, these wetlands were more efficient at removing nitrogen than those dominated by filamentous algae or submerged macrophytes. 5. Management of wetlands often aims to decrease the dominance of tall emergent vegetation for the benefit of plant species diversity and habitat heterogeneity. Our results demonstrate a biodiversity benefit, but also show that this strategy may decrease the ability of wetlands to remove nitrogen. In this case, there is no support for the hypothesis that biodiversity enhances ecosystem function.     

Climatic stability in the Brazilian Cerrado: implications for biogeographical connections of South American savannas,species richness and conservation in a biodiversity hotspot

Aim To investigate the historical distribution of the Cerrado across Quaternary climatic fluctuations and to generate historical stability maps to test: (1) whether the ‘historical climate’ stability hypothesis explains squamate reptile richness in the Cerrado; and (2) the hypothesis of Pleistocene connections between savannas located north and south of Amazonia. Location The Cerrado, a savanna biome and a global biodiversity hotspot distributed mainly in central Brazil. Methods We generated occurrence datasets from 1000 presence points randomly selected from the entire distribution of the Cerrado, as determined by two spatial definitions. We modelled the potential Cerrado distribution by implementing a maximum‐entropy machine‐learning algorithm across four time projections: current, mid‐Holocene (6 ka), Last Glacial Maximum (LGM, 21 ka) and Last Interglacial (LIG, 120 ka). We generated historical stability maps (refugial areas) by overlapping presence/absence projections of all scenarios, and checked consistencies with qualitative comparisons with available fossil pollen records. We built a spatially explicit simultaneous autoregressive model to explore the relationship between current climate, climatic stability, and squamate species richness. Results Models predicted the LGM and LIG as the periods of narrowest and widest Cerrado distributions, respectively, and were largely corroborated by palynological evidence. We found evidence for two savanna corridors (eastern coastal during the LIG, and Andean during the LGM) and predicted a large refugial area in the north‐eastern Cerrado (Serra Geral de Goiás refugium). Variables related to climatic stability predicted squamate richness better than present climatic variables did. Main conclusions Our results indicate that Bolivian savannas should be included within the Cerrado range and that the Cerrado’s biogeographical counterparts are not Chaco and Caatinga but rather the disjunct savannas of the Guyana shield plateaus. Climatic stability is a good predictor of Cerrado squamate richness, and our stability maps could be used in future studies to test diversity patterns and genetic signatures of different taxonomic groups and as a higher‐order landscape biodiversity surrogate for conservation planning.     

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.     

Environmental and spatial controls of biotic assemblages in a discrete semi‐terrestrial habitat: comparison of organisms with different dispersal abilities sampled in the same plots

Aim The development of metacommunity theory inspired a series of studies exploring the importance of environmental and spatial effects on the composition of biotic assemblages. However, the comparison of different groups of organisms has been hampered by differences in sampling design, spatial scales or the environmental variables involved. Our aim was to test how dispersal ability affects metacommunity structure and associated species distributions by sampling different species groups in the same plots to avoid these problems. Location Western Carpathian Mountains (Europe). Methods In 191 fens we sampled the composition of diatom, bryophyte, vascular plant and mollusc assemblages, water chemistry, and macroclimatic data. We then generated spatial variables covering all relevant spatial scales using analysis of principal coordinates of neighbour matrices (PCNM). We applied the adjusted variation partitioning algorithm to quantify the effects of environment and space. Results Pure effects of water chemistry and space were highly significant for all groups of organisms. Spatial effects were stronger for groups with larger propagules (vascular plants, molluscs) than for those with smaller propagules (diatoms, bryophytes). Assemblages of macroscopic bryophytes were structured slightly less by geography and much more by environment than were those of microscopic diatoms. Vascular plant and mollusc assemblages turned out to be more spatially structured (as compared to diatom and bryophyte assemblages), with small differences between the two groups. Coarse‐scale spatial effects dominated in the bryophyte metacommunity, while in the other groups, including diatoms, finer‐scale effects were also important. Main conclusions Given that our analyses are based on a standardized sampling and analytical framework, our findings provide strong support for the hypothesis that both environmental and spatial variables structure metacommunities of organisms with very different dispersal abilities, including microscopic diatoms. In addition, we show for the first time that the strengths of these effects and their scale dependence may be predicted using important trait differences between organisms, for example differences in propagule size.