Showing papers in "Global Ecology and Biogeography in 2006"

Novel ecosystems: theoretical and management aspects of the new ecological world order

Abstract: We e xplore the issues relevant to those types of ecosystems containing new combinations of species that arise through human action, environmental change, and the impacts of the deliberate and inadvertent introduction of species from other regions. Novel ecosystems (also termed ‘emerging ecosystems’) result when species occur in combinations and relative abundances that have not occurred previously within a given biome. Key characteristics are novelty, in the form of new species combinations and the potential for changes in ecosystem functioning, and human agency, in that these ecosystems are the result of deliberate or inadvertent human action. As more of the Earth becomes transformed by human actions, novel ecosystems increase in importance, but are relatively little studied. Either the degradation or invasion of native or ‘wild’ ecosystems or the abandonment of intensively managed systems can result in the formation of these novel systems. Important considerations are whether these new systems are persistent and what values they may have. It is likely that it may be very difficult or costly to return such systems to their previous state, and hence consideration needs to be given to developing appropriate management goals and approaches.

Towards an integrated computational tool for spatial analysis in macroecology and biogeography

Abstract: Because most macroecological and biodiversity data are spatially autocorrelated, special tools for describing spatial structures and dealing with hypothesis testing are usually required. Unfortunately, most of these methods have not been available in a single statistical package. Consequently, using these tools is still a challenge for most ecologists and biogeographers. In this paper, we present sam (Spatial Analysis in Macroecology), a new, easy-to-use, freeware package for spatial analysis in macroecology and biogeography. Through an intuitive, fully graphical interface, this package allows the user to describe spatial patterns in variables and provides an explicit spatial framework for standard techniques of regression and correlation. Moran's I autocorrelation coefficient can be calculated based on a range of matrices describing spatial relationships, for original variables as well as for residuals of regression models, which can also include filtering components (obtained by standard trend surface analysis or by principal coordinates of neighbour matrices). sam also offers tools for correcting the number of degrees of freedom when calculating the significance of correlation coefficients. Explicit spatial modelling using several forms of autoregression and generalized least-squares models are also available. We believe this new tool will provide researchers with the basic statistical tools to resolve autocorrelation problems and, simultaneously, to explore spatial components in macroecological and biogeographical data. Although the program was designed primarily for the applications in macroecology and biogeography, most of sam's statistical tools will be useful for all kinds of surface pattern spatial analysis. The program is freely available at http://www.ecoevol.ufg.br/sam (permanent URL at http://purl.oclc.org/sam/).

On defining and quantifying biotic homogenization

Abstract: Ongoing species invasions and extinctions are changing biological diversity in different ways at different spatial scales. Biotic homogenization (or BH) refers to the process by which the genetic, taxonomic or functional similarities of regional biotas increase over time. It is a multifaceted process that encompasses species invasions, extinctions and environmental alterations, focusing on how the identities of species (or their genetic or functional attributes) change over space and time. Despite the increasing use of the term BH in conservation biology, it is often used erroneously as a synonym for patterns of species invasions, loss of native species or changes in species richness through time. This reflects the absence of an agreed-upon, cogent definition of BH. Here, we offer an operational definition for BH and review the various methodologies used to study this process. We identify the strengths and weaknesses of these approaches, and make explicit recommendations for future studies. We conclude by citing the need for researchers to: (1) consider carefully the definition of BH by recognizing the genetic, taxonomic and functional realms of this process; (2) recognize that documenting taxonomic homogenization requires tracking the identity of species (not species richness) comprising biotas through space and time; and (3) employ more rigorous methods for quantifying BH.

Not seeing the ocean for the islands: the mediating influence of matrix-based processes on forest fragmentation effects.

Abstract: The pervasive influence of island biogeography theory on forest fragmentation research has often led to a misleading conceptualization of landscapes as areas of forest/habitat and ‘non-forest/non-habitat’ and an overriding focus on processes within forest remnants at the expense of research in the human-modified matrix. The matrix, however, may be neither uniformly unsuitable as habitat nor serve as a fully‐absorbing barrier to the dispersal of forest taxa. In this paper, we present a conceptual model that addresses how forest habitat loss and fragmentation affect biodiversity through reduction of the resource base, subdivision of populations, alterations of species interactions and disturbance regimes, modifications of microclimate and increases in the presence of invasive species and human pressures on remnants. While we acknowledge the importance of changes associated with the forest remnants themselves (e.g. decreased forest area and increased isolation of forest patches), we stress that the extent, intensity and permanence of alterations to the matrix will have an overriding influence on area and isolation effects and emphasize the potential roles of the matrix as not only a barrier but also as habitat, source and conduit. Our intention is to argue for shifting the examination of forest fragmentation effects away from a patch-based perspective focused on factors such as patch area and distance metrics to a landscape mosaic perspective that recognizes the importance of gradients in habitat conditions.

A comparison of methods for mapping species ranges and species richness

Abstract: Aim Maps of species richness are the basis for applied research and conservation planning as well as for theoretical research investigating patterns of richness and the processes shaping these patterns. The method used to create a richness map could influence the results of such studies, but differences between these methods have been insufficiently evaluated. We investigate how different methods of mapping species ranges can influence patterns of richness, at three spatial resolutions. Location California, USA. Methods We created richness maps by overlaying individual species range maps for terrestrial amphibians and reptiles. The methods we used to create ranges included: point-to-grid maps, obtained by overlaying point observations of species occurrences with a grid and determining presence or absence for each cell; expert-drawn maps; and maps obtained through species distribution modelling. We also used a hybrid method that incorporated data from all three methods. We assessed the correlation and similarity of the spatial patterns of richness maps created with each of these four methods at three different resolutions. Results Richness maps created with different methods were more correlated at lower spatial resolutions than at higher resolutions. At all resolutions, point-to-grid richness maps estimated the lowest species richness and those derived from species distribution models the highest. Expert-drawn maps and hybrid maps showed intermediate levels of richness but had different spatial patterns of species richness from those derived with the other methods. Main conclusions Even in relatively well-studied areas such as California, different data sources can lead to rather dissimilar maps of species richness. Evaluating the strengths and weaknesses of different methods for creating a richness map can provide guidance for selecting the approach that is most appropriate for a given application and region.

The biogeography of prediction error: why does the introduced range of the fire ant over-predict its native range?

Abstract: Aim The use of species distribution models (SDMs) to predict biological invasions is a rapidly developing area of ecology. However, most studies investigating SDMs typically ignore prediction errors and instead focus on regions where native distributions correctly predict invaded ranges. We investigated the ecological significance of prediction errors using reciprocal comparisons between the predicted invaded and native range of the red imported fire ant ( Solenopsis invicta ) (hereafter called the fire ant). We questioned whether fire ants occupy similar environments in their native and introduced range, how the environments that fire ants occupy in their introduced range changed through time relative to their native range, and where fire ant propagules are likely to have originated. Location We developed models for South America and the conterminous United States (US) of America. Methods We developed models using the Genetic Algorithm for Rule-set Prediction (GARP) and 12 environmental layers. Occurrence data from the native range in South America were used to predict the introduced range in the US and vice versa. Further, time-series data recording the invasion of fire ants in the US were used to predict the native range. Results Native range occurrences under-predicted the invasive potential of fire ants, whereas occurrence data from the US over-predicted the southern boundary of the native range. Secondly, introduced fire ants initially established in environments similar to those in their native range, but subsequently invaded harsher environments. Time-series data suggest that fire ant propagules originated near the southern limit of their native range. Conclusions Our findings suggest that fire ants from a peripheral native population established in an environment similar to their native environment, and then ultimately expanded into environments in which they are not found in their native range. We argue that reciprocal comparisons between predicted native and invaded ranges will facilitate a better understanding of the biogeography of invasive and native species and of the role of SDMs in predicting future distributions.

Could temperature and water availability drive elevational species richness patterns? A global case study for bats

Abstract: Aim A global meta-analysis was used to elucidate a mechanistic understanding of elevational species richness patterns of bats by examining both regional and local climatic factors, spatial constraints, sampling and interpolation. Based on these results, I propose the first climatic model for elevational gradients in species richness, and test it using preliminary bat data for two previously unexamined mountains. Location Global data set of bat species richness along elevational gradients from Old and New World mountains spanning 12.5 ° S to 38 ° N latitude. Methods Bat elevational studies were found through an extensive literature search. Use was made only of studies sampling ≥ 70% of the elevational gradient without significant sampling biases or strong anthropogenic disturbance. Undersampling and interpolation were explicitly examined with three levels of error analyses. The influence of spatial constraints was tested with a Monte Carlo simulation program, Mid-Domain Null. Preliminary bat species richness data sets for two test mountains were compiled from specimen records from 12 US museum collections. Results Equal support was found for decreasing species richness with elevation and mid-elevation peaks. Patterns were robust to substantial amounts of error, and did not appear to be a consequence of spatial constraints. Bat elevational richness patterns were related to local climatic gradients. Species richness was highest where both temperature and water availability were high, and declined as temperature and water availability decreased. Mid-elevational peaks occurred on mountains with dry, arid bases, and decreasing species richness occurred on mountains with wet, warm bases. A preliminary analysis of bat richness patterns on elevational gradients in western Peru (dry base) and the Olympic Mountains, WA (wet base), supported the predictions of the climate model. Main conclusions The relationship between species richness and combined temperature and water availability may be due to both direct (thermoregulatory constraints) and indirect (food resources) factors. Abundance was positively correlated with species richness, suggesting that bat species richness may also be related to productivity. The climatic model may be applicable to other taxonomic groups with similar ecological constraints, for instance certain bird, insect and amphibian clades.

Global patterns in seed size

Abstract: Aim To provide the first global quantification of the slope and shape of the latitudinal gradient in seed mass, and to determine whether global patterns in seed mass are best explained by growth form, vegetation type, seed dispersal syndrome, or net primary productivity (NPP). Location Global. Methods We collected seed mass data for 11,481 species × site combinations from around the world. We used regression to describe the latitudinal gradient in seed mass, then applied general linear models to quantify the relative explanatory power of each of the variables hypothesized to underlie the latitudinal gradient in seed size. Results There is a 320-fold decline in geometric mean seed mass between the equator and 60°. This decline is not linear. At the edge of the tropics, there is a sudden 7-fold drop in mean seed mass. The strongest correlates of the latitudinal gradient in seed mass are plant growth form, and vegetation type, followed by dispersal syndrome and NPP. A model including growth form, vegetation type, dispersal syndrome and NPP explains 51% of the variation in seed mass. Latitude explains just 0.2% of the residual variation from this model. Main conclusions This is the first demonstration of a major decrease in seed size at the edge of the tropics. This drop in seed mass is most closely correlated with changes in plant growth form and vegetation type. This suggests that the drop in seed mass might be part of a sudden change in plant strategy at the edge of the tropics.

The role of land cover in bioclimatic models depends on spatial resolution

Abstract: Aim We explored the importance of climate and land cover in bird species distribution models on multiple spatial scales. In particular, we tested whether the integration of land cover data improves the performance of pure bioclimatic models. Location Finland, northern Europe. Methods The data of the bird atlas survey carried out in 1986–89 using a 10 × 10 km uniform grid system in Finland were employed in the analyses. Land cover and climatic variables were compiled using the same grid system. The dependent and explanatory variables were resampled to 20-km, 40-km and 80-km resolutions. Generalized additive models (GAM) were constructed for each of the 88 land bird species studied in order to estimate the probability of occurrence as a function of (1) climate and (2) climate and land cover variables. Model accuracy was measured by a cross-validation approach using the area under the curve (AUC) of a receiver operating characteristic (ROC) plot. Results In general, the accuracies of the 88 bird–climate models were good at all studied resolutions. However, the inclusion of land cover increased the performance of 79 and 78 of the 88 bioclimatic models at 10-km and 20-km resolutions, respectively. There was no significant improvement at the 40-km resolution. In contrast to the finer resolutions, the inclusion of land cover variables decreased the modelling accuracy at 80km resolution. Main conclusions Our results suggest that the determinants of bird species distributions are hierarchically structured: climatic variables are large-scale determinants, followed by land cover at finer resolutions. The majority of the land bird species in Finland are rather clearly correlated with climate, and bioclimate envelope models can provide useful tools for identifying the relationships between these species and the environment at resolutions ranging from 10 km to 80 km. However, the notable contribution of land cover to the accuracy of bioclimatic models at 10–20-km resolutions indicates that the integration of climate and land cover information can improve our understanding and model predictions of biogeographical patterns under global change.

Scale‐related patterns in the spatial and environmental components of stream macroinvertebrate assemblage variation

Abstract: Aim We examined the relative contributions of spatial gradients and local environmental conditions to macroinvertebrate assemblages of boreal headwater streams at three hierarchical extents: bioregion, ecoregion and drainage system. We also aimed to identify the environmental variables most strongly related to assemblage structure at each study scale, and to assess how the importance of these variables is related to regional context and spatial structuring at different scales. Location Northern Finland (62–68° N, 25–32° E). Methods Variation in macroinvertebrate data was partitioned using partial canonical correspondence analysis into components explained by spatial variables (nine terms from the cubic trend surface regression), local environmental variables (15 variables) and spatially structured environmental variation. Results The strength of the relationship between assemblage structure and local environmental variables increased with decreasing spatial extent, whereas assemblage variation related to spatial variables and spatially structured environmental variation showed the opposite pattern. At the largest extents, spatial variation was related to latitudinal gradients, whereas spatial autocorrelation among neighbouring streams was the likely mechanism creating spatial structure within drainage systems. Only stream size and water acidity were consistently important in explaining assemblage structure at all study scales, while the importance of other environmental variables was more context-dependent. Main conclusions The importance of local environmental factors in explaining macroinvertebrate assemblage structure increases with decreasing spatial extent. This scale-related pattern is not caused solely by changes in study extent, however, but also by variable sample sizes at different regional extents. The importance of environmental gradients is context-dependent and few factors are likely to be universally important correlates of macroinvertebrate assemblage structure. Finally, our results suggest that bioassessment should give due attention to spatial structuring of stream assemblages, because important assemblage gradients may not only be related to local factors but also to biogeographical constraints and neighbourhood dispersal processes.

Imprints of glacial refugia in the modern genetic diversity of Pinus sylvestris

Abstract: Aim To understand the impact of glacial refugia and migration pathways on the modern genetic diversity of Pinus sylvestris Location The study was carried out throughout Europe Methods An extended set of data of pollen and macrofossil remains was used to locate the glacial refugia and reconstruct the migrating routes of P sylvestris throughout Europe A vegetation model was used to simulate the extent of the potential refugia during the last glacial period At the same time a genetic survey was carried out on this species Results The simulated distribution of P sylvestris during the last glacial period is coherent with the observed fossil data, which showed a patchy distribution of the refugia between c 40 degrees N and 50 degrees N Several migrational fronts were detected within the Iberian and the Italian peninsulas, and outside the Hungarian plain and around the Alps The modern mitochondrial DNA depicted three different haplotypes for P sylvestris Two distinct haplotypes were restricted to northern Spain and Italy, and the third haplotype dominated most of the present-day remaining distribution range of P sylvestris in Europe Main concluions During the last glacial period P sylvestris was constrained under severe climatic conditions to survive in scattered and restricted refugial areas Combining palaeoenvironmental data, vegetation modelling and the genetic data, we have shown that the long-term isolation in the glacial refugia and the migrational process during the Holocene have played a major role in shaping the modern genetic diversity of P sylvestris in Europe Document Type: Article

What drives elevational patterns of diversity? A test of geometric constraints, climate and species pool effects for pteridophytes on an elevational gradient in Costa Rica

Abstract: Aim We studied pteridophyte species richness between 100 m and 3400 m along a Neotropical elevational gradient and tested competing hypotheses for patterns of species richness. Location Elevational transects were situated at Volcan Barva in the Braulio Carrillo National Park and La Selva Biological Station (100–2800 m) and Cerro de la Muerte (2700–3400 m), both on the Atlantic slope of Costa Rica, Central America. Method We analysed species richness on 156 plots of 20 × 20 m and measured temperature and humidity at four elevations (40, 650, 1800 and 2800 m). Species richness patterns were regressed against climatic variables (temperature, humidity, precipitation and actual evapotranspiration), regional species pool, area and predicted species number of a geometric null model (the mid-domain effect, MDE). Results The species richness of the 484 recorded species showed a hump-shaped pattern with elevation with a richness peak at mid-elevations (c. 1700 m). The MDE was the single most powerful explanatory variable in linear regression models, but species richness was also associated strongly with climatic variables, especially humidity and temperature. Area and species pool were associated less strongly with observed richness patterns. Main conclusions Geometric models and climatic models exclusive of geometric constraints explained comparable amounts of the elevational variation in species richness. Discrimination between these two factor complexes is not possible based on model fits. While overall fits of geometric models were high, large- and small-ranged species were explained by geometric models to different extents. Species with narrow elevational ranges clustered at both ends of the gradient to a greater extent than predicted by the MDE null models used here. While geometric models explained much of the pattern in species richness, we cannot rule out the role of climatic factors (or vice versa) because the predicted peak in richness from geometric models, the empirical peak in richness and the overlap in favourable environmental conditions all coincide at middle elevations. Mid-elevations offer highest humidity and moderate temperatures, whereas at high elevations richness is reduced due to low temperatures, and at low elevations by reduced water availability due to high temperatures.

Geographical gradients of species richness: a test of the water-energy conjecture of Hawkins et al. (2003) using European data for five taxa

Abstract: Aims We present an analysis of grid-based species-richness data for European plants, mammals, birds, amphibians and reptiles, designed to test the proposition of Hawkins et al. (2003a) that the single best factor describing richness variation switches from the water regime to the energy regime in the mid-latitudes and that the ‘breakpoint’ is related to the physiological character of the taxa. We go on to develop subregional models showing the extent to which regional model fits vary as a function of the extent of the study system, and compare the relative performance of ‘water’, ‘energy’ and ‘water‐energy’ models of richness for southern, northern and pan-European models.

Patterns in species richness and endemism of European freshwater fish

Abstract: Aim To analyse the patterns in species richness and endemism of the native European riverine fish fauna, in the light of the Messinian salinity crisis and the Last Glacial Maximum (LGM). Location European continent. Methods After gathering native fish faunistic lists of 406 hydrographical networks, we defined large biogeographical regions with homogenous fish fauna, based on a hierarchical cluster analysis. Then we analysed and compared the patterns in species richness and endemism among these regions, as well as species‐area relationships. Results Among the 233 native species present in the data set, the Cyprinidae family was strongly dominant (> 50% of the total number of species). Seven biogeographical regions were defined: Western Peri-Mediterranea, Central Peri-Mediterranea, Eastern Peri-Mediterranea, Ponto-Caspian Europe, Northern Europe, Central Europe and Western Europe. The highest regional species richness was observed for Central Peri-Mediterranea and Ponto-Caspian Europe. The highest endemic richness was found in Central Peri-Mediterranea. Species‐area relationships were characterized by high slope values for Peri-Mediterranean Europe and low values for Central and Western Europe. Main conclusions The results were in agreement with the ‘Lago Mare’ hypothesis explaining the specificity of Peri-Mediterranean fish fauna, as well as with the history of recolonization of Central and Western Europe from Ponto-Caspian Europe following the LGM. The results also agreed with the mechanisms of speciation and extinction influencing fish diversity in hydrographical networks. We advise the use of the seven biogeographical regions for further studies, and suggest considering

Altered geographic and temporal variability in phenology in response to climate change

Abstract: Aim In response to recent climate warming, numerous studies have reported an earlier onset of spring and, to a lesser degree, a later onset of autumn, both determined from phenological observations. Here, we examine whether these reported changes have affected the synchronization of events on a regional level by examining temporal and spatial variability in phenology. In particular, we study whether years with earlier springs are associated with an altered spatial variability in phenology. Location Germany and the United Kingdom. Methods Plant phenological observations of 35 different phases (events such as flowering and leafing) collected by the German Weather Service (1951–2002) and butterfly phenological records of 29 species collected by the UK Butterfly Monitoring Scheme (1976–2003) are used. In these long-term records, we examine the temporal (year-to-year) variability and the spatial (geographic or between site) variability with particular emphasis on how they vary with time of the year and with earliness or lateness of the phase. Results Early phenological events (i.e. spring) are more variable than later events, both in time and in space, although the pattern is clearer for plants than for butterflies. Confirming previous results, we find a clear relationship between the mean date of spring and summer phases and the degree to which they have become earlier. The spatial variability of spring events is greater in warmer years that have faster plant development. However, late spring and summer events do not show a consistent relationship. Autumn events are somewhat more spatially variable in years characterized by later seasons. Main conclusions This is the first examination of spatial variability of plant and animal phenological events at a multinational scale. Earlier spring events are likely to be associated with increased spatial variability in plants, although this is unlikely to also be true for summer events. If species experience differential changes in geographic variation this may disrupt interactions among them, e.g. in food webs. On the other hand, these may offer advantages for mobile species. Further research on linked species is recommended.

Incursion and excursion of Antarctic biota: past, present and future

Abstract: Aim To investigate the major paradigms of intense isolation and little anthropogenic influence around Antarctica and to examine the timings and scales of the modification of the southern polar biota. Location Antarctica and surrounding regions. Methods First, mechanisms of and evidence for long-term isolation are reviewed. These include continental drift, the development of a surrounding deep-water channel and the Antarctic Circumpolar Current (ACC). They also include levels of endemism, richness and distinctiveness of assemblages. Secondly, evidence for past and modern opportunities for species transport are investigated. Comparative levels of alien establishments are also examined around the Southern Ocean. Discussion On a Cenozoic time-scale, it is clear that Gondwana’s fragmentation led to increasing geographical isolation of Antarctica and the initiation of the ACC, which restricted biota exchange to low levels while still permitting some movement of biota. On a shorter Quaternary time-scale, the continental ice-sheet, influenced by solar (Milankovitch) cycles, has expanded and contracted periodically, covering and exposing terrestrial and continental shelf habitats. There were probably refugia for organisms during each glacial maxima. It is also likely that new taxa were introduced into Antarctica during cycles of ice sheet and oceanic front movement. The current situation (a glacial minimum) is not ‘normal’; full interglacials represent only 10% of the last 430 ka. On short (ecological) time-scales, many natural dispersal processes (airborne, oceanic eddy, rafting and hitch-hiking on migrants) enable the passage of biota to and from Antarctica. In recent years, humans have become influential both directly by transporting organisms and indirectly by increasing survival and establishment prospects via climate change. Main conclusions Patterns of endemism and alien establishment are very different across taxa, land and sea, and north vs. south of the Polar Frontal Zone. Establishment conditions, as much as transport, are important in limiting alien establishment. Three time-scales emerge as important in the modification of Antarctica’s biota. The natural ‘interglacial’ process of reinvasion of Antarctica is being influenced strongly by humans.

Leaf flushing during the dry season: the paradox of Asian monsoon forests

Abstract: Aim Most deciduous species of dry monsoon forests in Thailand and India form new leaves 1‐2 months before the first monsoon rains, during the hottest and driest part of the year around the spring equinox. Here we identify the proximate causes of this characteristic and counterintuitive ‘spring-flushing’ of monsoon forest trees. Location Trees of 20 species were observed in semi-deciduous dry monsoon forests of northern Thailand with a 5 ‐ 6-month-long severe dry season and annual rainfall of 800 ‐1500 mm. They were growing on dry ridges (dipterocarp‐oak forest) or in moist gullies (mixed deciduous‐evergreen forest) at 680 ‐750 m altitude near Chiang Mai and in a dry lowland stand of Shorea siamensis in Uthai Thani province. Methods Two novel methods were developed to analyse temporal and spatial variation in vegetative dry-season phenology indicative of differences in root access to subsoil water reserves.

Sensitivity of African biomes to changes in the precipitation regime

Abstract: Aim Africa is identified by the Inter-governmental Panel on Climate Change (IPCC) as the least studied continent in terms of ecosystem dynamics and climate variability. The aim of this study was (1) to adapt the Lund-Postdam-Jena-GUESS (LPJ-GUESS) ecological modelling framework to Africa by providing new parameter values for tropical plant functional types (PFT), and (2) to assess the sensitivity of some African biomes to changes in precipitation regime. Location The study area was a representative transect (0-22 degrees N and 7-18 degrees E) through the transition from equatorial evergreen forests to savannas, steppes and desert northwards. The transect showed large latitudinal variation in precipitation (mean rainfall ranged from 50 to 2300 mm year(-1)). Methods New PFT parameters used to calibrate LPJ-GUESS were based on modern pollen PFTs and remote sensed leaf area index (LAI). The model was validated using independent modern pollen assemblages, LAI and through comparison with White's modern potential vegetation map. Several scenarios were developed by combining changes in total rainfall amount with variation in the length of the dry season in order to test the sensitivity of African biomes. Results Simulated vegetation compared well to observed data at local and regional scales, in terms of ecosystem functioning (LAI), and composition (pollen and White's vegetation map). The assessment of the sensitivity of biomes to changes in precipitation showed that none of the ecosystems would shift towards a new type under the range of precipitation increases suggested by the IPCC (increases from 5 to 20%). However, deciduous and semi-deciduous forests may be very sensitive to small reductions in both the amount and seasonality of precipitation. Main conclusions This version of LPJ-GUESS parameterized for Africa simulated correctly the vegetation present over a wide precipitation gradient. The biome sensitivity assessment showed that, compared with savannas and grasslands, closed canopy forests may be more sensitive to change in precipitation regime due to the synergetic effects of changed rainfall amounts and seasonality on vegetation functioning.

Implementing plant hydraulic architecture within the LPJ Dynamic Global Vegetation Model

Abstract: Aim To implement plant hydraulic architecture within the Lund-Potsdam-Jena Dynamic Global Vegetation Model (LPJ-DGVM), and to test the model against a set of observational data. If the model can reproduce major patterns in vegetation and ecosystem processes, we consider this to be an important linkage between plant physiology and larger-scale ecosystem dynamics. Location The location is global, geographically distributed. Methods A literature review was carried out to derive model formulations and parameter values for representing the hydraulic characteristics of major global plant functional types (PFTs) in a DGVM. After implementing the corresponding formulations within the LPJ-DGVM, present-day model output was compared to observational data. Results The model reproduced observed broad-scale patterns in potential natural vegetation, but it failed to distinguish accurately between different types of grassland and savanna vegetation, possibly related to inadequate model representations of water fluxes in the soil and wildfire effects. Compared to a version of the model using an empirical formulation for calculating plant water supply without considering plant hydraulic architecture, the new formulation improved simulated patterns of vegetation in particular for dry shrublands. Global-scale simulation results for runoff and actual evapotranspiration (AET) corresponded well to available data. The model also successfully reproduced the magnitude and seasonal cycle of AET for most EUROFLUX forests, while modelled variation in NPP across a large number of sites spanning several biomes showed a strong correlation with estimates from field measurements. Main conclusions The model was generally confirmed by comparison to observational data. The novel model representation of water flow within plants makes it possible to resolve mechanistically the effects of hydraulic differences between plant functional groups on vegetation structure, water cycling, and competition. This may be an advantage when predicting ecosystem responses to nonextant climates, in particular in areas dominated by dry shrubland vegetation.

Species richness, composition and abundance of fruit‐feeding butterflies in the Brazilian Atlantic Forest: comparison between a fragmented and a continuous landscape

Abstract: Aim To evaluate frugivorous butterflies as indicators of forest disturbance in the Brazilian Atlantic Forest. Location The study area is located in the municipality of Cotia, Sao Paulo State, south-eastern Brazil (23°35′−23°50′ S, 46°45′−47°15′ W). Methods Sampling was done at four sites inside a large forest block, the Morro Grande State Reserve, and in five forest fragments in an adjacent fragmented landscape. Butterflies were sampled with portable traps, baited with a fermented mixture of banana and sugar cane juice. Sampling was carried out during the period most favourable for the capture of frugivorous butterflies in south-eastern Brazil. Results All richness-related results indicated no effect of forest fragmentation on the frugivorous butterfly guild, concurring with the suggestion of appreciable resistance of Atlantic Forest butterflies to habitat modification. However, species composition discriminated between the two landscapes, indicating that fragmentation may have effects beyond simple species richness. When species were analysed individually, clear patterns of distribution emerged, with some species that were very abundant in the fragmented landscape being practically absent in the continuous landscape, and vice versa. This pattern seems consistent even for some subfamilies. Main conclusion Our findings support the usefulness of the frugivorous butterfly guild as a biological indicator of forest disturbance effects in one of the world's most threatened ecosystems.

Are green turtles globally endangered

Abstract: Aim To examine the exploitation, recovery and current status of green turtles (Chelonia mydas) nesting at Ascension Island. Location Ascension Island (UK) (7°57′ S, 14°22′ W), South Atlantic Ocean. Methods We analysed records of the harvest of green turtles nesting at Ascension Island between 1822 and 1935, illustrating the decline in numbers over this period. Using a deterministic age-class structured model we predict the initial number of breeding females present in the population prior to the recorded harvest and compare this to our estimate of the current population based upon our recent annual surveys (1999–2004). Results Prior to 1822 we estimate the nesting population of green turtles to have been at least 19,000–22,000 individuals in order for the population to have survived the level of harvest recorded. From recent data (1999–2004), we estimate the current breeding population of green turtles at this site to be 11,000–15,000 females. Our results illustrate a dramatic recovery of the population, which is still increasing exponentially and shows no evidence of slowing, suggesting it has not reached 50% of its carrying capacity. Main conclusions We estimate that, since the 1970s, the Ascension Island population of green turtles has increased by 285% and question the recent listing of this species as endangered by the IUCN (World Conservation Union), in particular in the Atlantic Ocean, where 75% of the populations assessed by the IUCN are increasing. Indeed, we estimate the global population of this species to be in excess of 2.2 million individuals. We suggest that the IUCN's global listing process detracts attention from those populations that are truly threatened with extinction and should not, in its present form, be applied to globally distributed long-lived species such as marine turtles.

Identification of current ecosystem functional types in the Iberian Peninsula

Abstract: Aim To e xamine the geographical patterns of the interception of photosynthetically active radiation by vegetation and to describe its spatial heterogeneity through the definition of ecosystem functional types (EFTs) based on the annual dynamics of the Normalized Difference Vegetation Index (NDVI), a spectral index related to carbon gains. Location The Iberian Peninsula. Methods EFTs were derived from three attributes of the NDVI obtained from NOAA/AVHRR sensors: the annual integral (NDVI-I), as a surrogate of primary production, an integrative indicator of ecosystem functioning; and the intra-annual relative range (RREL) and month of maximum NDVI (MMAX), which represent key features of seasonality. Results NDVI-I decreased south-eastwards. The highest values were observed in the Eurosiberian Region and in the highest Mediterranean ranges. Low values occurred in inner plains, river basins and in the southeast. The Eurosiberian Region and Mediterranean mountains presented the lowest RREL, while Eurosiberian peaks, river basins, inner-agricultural plains, wetlands and the southeastern part of Iberia presented the highest. Eurosiberian ecosystems showed a summer maximum of NDVI, as did high mountains, wetlands and irrigated areas in the Mediterranean Region. Mediterranean mountains had autumn‐early-winter maxima, while semi-arid zones, river basins and continental plains had spring maxima. Based on the behaviour in the functional traits, 49 EFTs were defined. Main conclusions The classification, based on only the NDVI dynamics, represents the spatial heterogeneity in ecosystem functioning by means of the interception of radiation by vegetation in the Iberian Peninsula. The patterns of the NDVI attributes may be used as a reference in evaluating the impacts of environmental changes. Iberia had a high spatial variability: except for biophysically impossible combinations (high NDVI-I and high seasonality), almost any pattern of seasonal dynamics of radiation interception was represented in the Peninsula. The approach used to define EFTs opens the possibility of monitoring and comparing ecosystem functioning through time.

A global model of island biogeography

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

Understorey plant species richness and composition in metropolitan forest archipelagos: effects of forest size, adjacent land use and distance to the edge

Abstract: Aim To address the relative role of adjacent land use, distance to forest edge, forest size and their interactions on understorey plant species richness and composition in perimetropolitan forests. Location The metropolitan area of Barcelona, north-eastern Spain.

Abundance, species richness and energy availability in the North American avifauna

Abstract: Aim To determine how species richness, abundance, biomass, energy use and mean number of individuals per species scale with environmental energy availability in wintering and breeding avian assemblages, and to contrast assemblages of (i) common and rare species and (ii) breeding residents and migrants. To assess whether such patterns are compatible with the ‘more individuals hypothesis’ (MIH) that high-energy areas are species-rich because they support larger populations that are buffered against extinction. Location The North American continent (latitudinal range 23.4 °−48.1 °N; longitudinal range 124.2°−68.7° W). Methods Avian species richness, abundance, biomass and energy use were calculated for 295 Resident Bird Count plots. Environmental energy availability was measured using ambient temperature and the Normalized Difference Vegetation Index (NDVI), a close correlate of plant productivity. Analyses took plot area into account, and were conducted (with and without taking habitat type into account) using general linear models and spatial mixed models. Results Positive species–energy relationships were exhibited by both wintering and breeding assemblages, but were stronger in the former. The structure of winter assemblages responded more strongly to temperature than NDVI, while breeding assemblages tended to respond more strongly to NDVI. Breeding residents responded to annual measures of energy availability while breeding migrants and the winter assemblage responded more strongly to seasonal measures. In the winter assemblage, rare and common species exhibited species–energy relationships of a similar strength, but common breeding species exhibited a much stronger relationship than rare breeding species. In both breeding and wintering assemblages, abundance, biomass and energy use increased with energy availability and species richness. Energy availability was a poor predictor of the mean number of individuals per species. Main conclusions The nature of the species–energy relationship varies seasonally and with the manner in which energy availability is measured. Our data suggest that residents are less able to respond to seasonal fluxes in resource availability than long-distance migrants. Increasing species richness and energy availability is associated with increasing numbers of individuals, biomass and energy use. While these observations are compatible with the MIH our data provide only equivocal support for this hypothesis, as the rarest species do not exhibit the strongest species–energy relationships.

Treeline dynamics in relation to climatic variability in the central Tianshan Mountains, northwestern China

Abstract: Aim Climate variability may be an important mediating agent of ecosystem dynamics in cold, arid regions such as the central Tianshan Mountains, northwestern China. Tree-ring chronologies and the age structure of a Schrenk spruce ( Picea schrenkiana ) forest were developed to examine treeline dynamics in recent decades in relation to climatic variability. Of particular interest was whether tree-ring growth and population recruitment patterns responded similarly to climate warming. Location The study was conducted in eight stands that ranged from 2500 m to 2750 m a.s.l. near the treeline in the Tianchi Nature Reserve (43 ° 45 ′− 43 ° 59 ′ N, 88 ° 00 ′− 88 ° 20 ′ E) in the central Xinjiang Uygur Autonomous Region, northwestern China. Methods Tree-ring cores were collected and used to develop tree-ring chronologies. The age of sampled trees was determined from basal cores sampled as close as possible to the ground. Population age structure and recruitment information were obtained using an age‐d.b.h. (diameter at breast height) regression from the sampled cores and the d.b.h. measured on all trees in the plots. Ring-width chronologies and tree age structure were both used to investigate the relationship between treeline dynamics and climate change. Results Comparisons with the climatic records showed that both the radial growth of trees and tree recruitment were influenced positively by temperature and precipitation in the cold high treeline zone, but the patterns of their responses differed. The annual variation in tree rings could be explained largely by the average monthly minimum temperatures during February and August of the current year and by the monthly precipitation of the previous August and January, which had a significant and positive effect on tree radial growth. P. schrenkiana recruitment was influenced mainly by consecutive years of high minimum summer temperatures and high precipitation during spring. Over the last several decades, the treeline did not show an obvious upward shift and new recruitment was rare. Some trees had established at the treeline at least 200 years ago. Recruitment increased until the early 20th century (1910s) but then decreased with poor recruitment over the past several decades (1950‐2000). Main conclusions There were strong associations between climatic change and ring-width patterns, and with recruitments in Schrenk spruce. Average minimum temperatures in February and August, and total precipitation in the previous August and January, had a positive effect on tree-ring width, and several consecutive years of high minimum summer temperature and spring precipitation was a main factor favouring the establishment of P. schrenkiana following germination within the treeline ecotone. Both dendroclimatology and recruitment analysis were useful and compatible to understand and reconstruct treeline dynamics in the central Tianshan Mountains.

The geographic distribution of mammal body size in Europe

Abstract: Aims To describe the pattern of mean body size of native mammals in Europe, and to investigate its relationships with environmental predictors related to four hypotheses: (1) dispersal; (2) heat conservation; (3) heat dissipation; and (4) resource availability. Location Continental western Europe and Great Britain. Methods We used range maps to estimate the mean body size (average log mass) of mammals in 386 cells of 12,100 km(2) each. Environmental conditions in each cell were quantified using nine historical, climatic and primary production variables. We attempted to tease apart the effects of these variables using correlation, multiple regression and spatial autocorrelation analyses. Results In the part of the continent covered by ice during the Pleistocene, body mass decreases southwards, and annual average temperature explains 73% of the variance in body size, consistent with the heat-conservation hypothesis. However, in warmer, non-glaciated areas the best predictor is an estimate of seasonality in plant production, but it explains only 18% of the variance. Carnivores, omnivores and herbivores show similar relationships, but the pattern for herbivores is substantially weaker than for the other groups. Main conclusions Overall, the relationship between mean body size and temperature is non-linear, being strong in cold environments but virtually disappearing above a temperature threshold.

The influence of matrix and edges on species richness patterns of ground beetles (Coleoptera: Carabidae) in habitat islands

Abstract: Aim The aim of this study was to analyse whether, and how, the inclusion of habitat specialists and edge-preferring species modifies the species–area relationship predictions of the island biogeography theory for an insect group (ground beetles, Coloptera: Carabidae) living in natural fragments. Species–habitat island area relationships applied to terrestrial habitat islands can be distorted by the indiscriminate inclusion of all species occurring in the fragments. Matrices surrounding terrestrial habitat fragments can provide colonists that do not necessarily distinguish the fragment from the matrix and can survive and reproduce there. Edge-preferring species can further distort the expected relationship, as smaller fragments have larger edge:core ratios. Location Nineteen forest fragments were studied in the Bereg Plain, Hungary, and SW Ukraine. This area contains natural forest patches, mainly of oak and hornbeam, and supports a mountain entomofauna. Methods Ground beetles (Carabidae) present in the 19 forest patches were categorized into generalists, forest specialists and edge-preferring species. We analysed the relationship between species richness and fragment area using species richness in the different categories. Results The assemblages contained a high share of generalist species (species that occur also in the surrounding matrix). Forest patch size and the number of generalist species showed a marginally significant negative relationship, indicating that generalist species were more important in smaller patches. Forest specialist species richness was correlated positively with patch area. Edge-preferring species were shown to influence the species–area relationship: the number of edge-preferring species increased with the edge:area ratio. Main conclusions Both generalist and edge-preferring species can considerably distort the species–area relationship. Island biogeography theory can be applied to habitat islands only if the habitat islands are defined correctly from the viewpoint of the target species.

Beyond Rapoport's rule: evaluating range size patterns of New World birds in a two‐dimensional framework

Abstract: Aim To evaluate Rapoport's rule for New World birds in two-dimensional geographical space. We specifically test for a topography × climate interaction that predicts little difference in range sizes between lowlands and mountains in cold climates, whereas in the tropics, montane species have narrow ranges and lowland species have broad ranges. Location The western hemisphere. Methods We used digitized range maps of breeding birds to generate mean range sizes in grids of 27.5 × 27.5 km and 110 × 110 km across North and South America. We examined the geographical pattern with respect to range in elevation, mean temperature in the coldest month, their interaction, biome size and continental width, using model II analysis of variance, multiple regression and simple correlation. Results In northern latitudes species have broad ranges in both mountainous and flat areas. However, range sizes in the mountains and lowlands diverge southwards, with the most extreme differences in the tropics. Further, there are minimal differences in range sizes across latitudes in lowlands. The smallest mean ranges occur in the tropical Andes. Mean range sizes in north-central Canada, Central America and Argentina/Chile are also small, reflecting the narrowing of the continents in these areas. The best regression model explained 51% of the variation in mean range size. Main conclusions The two-dimensional range size pattern indicates that neither winter temperature nor annual variability in temperature strongly influences the distribution of range sizes directly; rather, climate influences bird range sizes indirectly via effects on habitat size. Also, macroclimate interacts with topographic relief across latitudes, generating sharp mesoscale habitat gradients in tropical mountains but not in high latitude mountains or in lowlands at any latitude. Birds respond to these habitat gradients, resulting in ‘latitudinal’ range size gradients in topographically complex landscapes but not in simple landscapes.