Showing papers on "Species richness published in 2002"

Global Dispersal of Free-Living Microbial Eukaryote Species

Abstract: The abundance of individuals in microbial species is so large that dispersal is rarely (if ever) restricted by geographical barriers. This “ubiquitous” dispersal requires an alternative view of the scale and dynamics of biodiversity at the microbial level, wherein global species number is relatively low and local species richness is always sufficient to drive ecosystem functions.

Functional diversity (FD), species richness and community composition

Abstract: Functional diversity is an important component of biodiversity, yet in comparison to taxonomic diversity, methods of quantifying functional diversity are less well developed. Here, we propose a means for quantifying functional diversity that may be particularly useful for determining how functional diversity is related to ecosystem functioning. This measure of functional diversity “FD” is defined as the total branch length of a functional dendrogram. Various characteristics of FD make it preferable to other measures of functional diversity, such as the number of functional groups in a community. Simulating species' trait values illustrates how the relative importance of richness and composition for FD depends on the effective dimensionality of the trait space in which species separate. Fewer dimensions increase the importance of community composition and functional redundancy. More dimensions increase the importance of species richness and decreases functional redundancy. Clumping of species in trait space increases the relative importance of community composition. Five natural communities show remarkably similar relationships between FD and species richness.

Network structure and biodiversity loss in food webs: robustness increases with connectance

Abstract: Food-web structure mediates dramatic effects of biodiversity loss including secondary and ‘cascading’ extinctions. We studied these effects by simulating primary species loss in 16 food webs from terrestrial and aquatic ecosystems and measuring robustness in terms of the secondary extinctions that followed. As observed in other networks, food webs are more robust to random removal of species than to selective removal of species with the most trophic links to other species. More surprisingly, robustness increases with food-web connectance but appears independent of species richness and omnivory. In particular, food webs experience ‘rivet-like’ thresholds past which they display extreme sensitivity to removal of highly connected species. Higher connectance delays the onset of this threshold. Removing species with few trophic connections generally has little effect though there are several striking exceptions. These findings emphasize how the number of species removed affects ecosystems differently depending on the trophic functions of species removed.

The Global 200: Priority ecoregions for global conservation

Abstract: A global strategy to conserve biodiversity must aim to protect representative examples of all of the world's ecosystems, as well as those areas that contain exceptional concentrations of species and endemics. Although lacking the richness of tropical forests, deserts, tropical lakes, and subpolar seas all contain distinct species, communities, and ecological phenomena. We analyzed global patterns of biodiversity to identify a set of the Earth's terrestrial, freshwater, and marine ecoregions that harbor exceptional biodiversity and are representative of its ecosystems. We placed each of the Earth's ecoregions within a system of 30 biomes and biogeographic realms to facilitate a representation analysis. Biodiversity features were compared among ecoregions to assess their irreplaceability or distinctiveness. These features included species richness, endemic species, unusual higher taxa, unusual ecological or evolutionary phenomena, and the global rarity of habitats. This process yielded 238 ecoregions-the Global 200-comprised of 142 terrestrial, 53 freshwater, and 43 marine priority ecoregions. Effective conservation in this set of ecoregions would help conserve the most outstanding and representative habitats for biodiversity nn this planet.

Characterizing ecosystem‐level consequences of biological invasions: the role of ecosystem engineers

Abstract: Invasions by exotic species represent both threats to ecosystems as well as opportunities to learn more about them. Among the invaders that will have the largest impacts are those that directly modify ecosystems and thus have cascading effects for resident biota. Exotics can affect ecosystems by altering system-level flows, availability, or quality of nutrients, food, and physical resources (e.g. living space, water, heat or light). The invader-mediated control of physical resources, typically achieved through the modification of habitats, has received limited attention in invasion biology. This reflects a general trend in ecology, and only recently has the concept of ecosystem engineering been developed to account for the role of species that shape habitats. Plants and animals in terrestrial and aquatic systems can both create and destroy structure. When introduced into ecosystems, these exotic engineers cause physical state changes with effects that ramify throughout the system. Although the consequences of these modifications are varied and complex, insight gained from general ecological principles offers an opportunity to predict what invaders will do upon their integration into systems. Examples from the literature suggest that introduced ecosystem engineers that increase habitat complexity or heterogeneity tend to cause abundances and/or species richness to rise, while those that decrease complexity tend to have the reverse effect. In assessing such patterns, however, it is critical to also consider spatial scales and the life habits of resident organisms. In addition to providing predictive power, recognition of engineering as a major means by which invasive species affect ecosystems provides a unifying theme for invasion biology and offers a chance to consider more fully the general role of species in ecosystems.

Scale‐dependent effects of landscape context on three pollinator guilds

Abstract: Most ecological processes and interactions depend on scales much larger than a single habitat, and therefore it is important to link spatial patterns and ecological processes at a landscape scale. Here, we analyzed the effects of landscape context on the distribution of bees (Hymenoptera: Apoidea) at multiple spatial scales with respect to the following hypotheses: (1) Local abundance and diversity of bees increase with increasing proportion of the surrounding seminatural habitats. (2) Solitary wild bees, bumble bees, and honey bees respond to landscape context at different spatial scales. We selected 15 landscape sectors and determined the percentage of seminatural habitats and the diversity of habitat types at eight spatial scales (radius 250-3000 m) by field inspections and analyses of vegetation maps using two Geographic Information Systems. The percentage of semi- natural habitats varied between 1.4% and 28%. In the center of each landscape sector a patch of potted flowering plants (four perennial and two annual species) was placed in the same habitat type, a grassy field margin adjacent to cereal fields. In all, 865 wild bee individuals and 467 honey bees were observed and an additional 475 individuals were caught for species identification. Species richness and abundance of solitary wild bees showed a close positive correlation with the percentage of seminatural habitats at small scales up to 750 m, whereas bumble bees and honey bees did not respond to landscape context at these scales. In contrast, honey bees were correlated with landscape context at large scales. The densities of flower-visiting honey bees even increased with decreasing proportion of seminatural habitats at a radius of 3000 m. We are not aware of any empirical studies showing contrasting foraging patterns related to landscape context at different spatial scales. We conclude (1) that local landscape destruction affects solitary wild bees more than social bees, possibly changing mutualistic plant-pollinator and competitive wild bees- honey bees interactions and (2) that only analyses of multiple spatial scales may detect the importance of the landscape context for local pollinator communities.

Biodiversity as a barrier to ecological invasion

Abstract: Biological invasions are a pervasive and costly environmental problem1,2 that has been the focus of intense management and research activities over the past half century. Yet accurate predictions of community susceptibility to invasion remain elusive. The diversity resistance hypothesis, which argues that diverse communities are highly competitive and readily resist invasion3,4,5, is supported by both theory6 and experimental studies7,8,9,10,11,12,13,14 conducted at small spatial scales. However, there is also convincing evidence that the relationship between the diversity of native and invading species is positive when measured at regional scales3,11,15,16. Although this latter relationship may arise from extrinsic factors, such as resource heterogeneity, that covary with diversity of native and invading species at large scales, the mechanisms conferring greater invasion resistance to diverse communities at local scales remain unknown. Using neighbourhood analyses, a technique from plant competition studies17,18,19, we show here that species diversity in small experimental grassland plots enhances invasion resistance by increasing crowding and species richness in localized plant neighbourhoods. Both the establishment (number of invaders) and success (proportion of invaders that are large) of invading plants are reduced. These results suggest that local biodiversity represents an important line of defence against the spread of invaders.

Global Biodiversity, Biochemical Kinetics, and the Energetic-Equivalence Rule

Abstract: The latitudinal gradient of increasing biodiversity from poles to equator is one of the most prominent but least understood features of life on Earth. Here we show that species diversity can be predicted from the biochemical kinetics of metabolism. We first demonstrate that the average energy flux of populations is temperature invariant. We then derive a model that quantitatively predicts how species diversity increases with environmental temperature. Predictions are supported by data for terrestrial, freshwater, and marine taxa along latitudinal and elevational gradients. These results establish a thermodynamic basis for the regulation of species diversity and the organization of ecological communities.

Species diversity enhances ecosystem functioning through interspecific facilitation

Abstract: Facilitation between species is thought to be a key mechanism by which biodiversity affects the rates of resource use that govern the efficiency and productivity of ecosystems; however, there is no direct empirical evidence to support this hypothesis. Here we show that increasing the species diversity of a functional group of aquatic organisms induces facilitative interactions, leading to non-additive changes in resource consumption. We increased the richness and evenness of suspension-feeding caddisfly larvae (Insecta, Trichoptera) in stream mesocosms and found that the increased topographical complexity of the benthic habitat alters patterns of near-bed flow such that the feeding success of individuals is enhanced. Species diversity reduces 'current shading' (that is, the deceleration of flow from upstream to downstream neighbours), allowing diverse assemblages to capture a greater fraction of suspended resources than is caught by any species monoculture. The fundamental nature of this form of hydrodynamic facilitation suggests that it is broadly applicable to freshwater and marine habitats; in addition, it has several analogues in terrestrial ecosystems where fluxes of energy and matter can be influenced by biophysical complexity. Thus, changes in species diversity may alter the probability of positive species interactions, resulting in disproportionately large changes in the functioning of ecosystems.

Irreversible impact of past land use on forest soils and biodiversity

Abstract: In western Europe, forest area has been expanding rapidly since the 19th century, mainly on former agricultural land. Previous studies show that plant diversity differs between these recent forests and ancient forests that were already forested at the time of first national cadastral surveys, around 1800. Here, we investigated the duration of such agricultural aftereffects. In northeastern France, large areas were deforested during the Roman occupation and thereafter abandoned to forest. In one such forest that was farmed during the period AD 50-250, we show that species richness and plant communities vary according to the intensity of former agriculture. These variations are linked to long- term changes of chemical and structural soil properties. Hence, we suggest that such effects of past agricultural land use on forest biodiversity may be irreversible on an historical time scale.

Geographic Range Size and Determinants of Avian Species Richness

Abstract: Geographic patterns in species richness are mainly based on wide-ranging species because their larger number of distribution records has a disproportionate contribution to the species richness counts. Here we demonstrate how this effect strongly influences our understanding of what determines species richness. Using both conventional and spatial regression models, we show that for sub-Saharan African birds, the apparent role of productivity diminishes with decreasing range size, whereas the significance of topographic heterogeneity increases. The relative importance of geometric constraints from the continental edge is moderate. Our findings highlight the failure of traditional species richness models to account for narrow-ranging species that frequently are also threatened.

Low host specificity of herbivorous insects in a tropical forest

Abstract: Two decades of research have not established whether tropical insect herbivores are dominated by specialists or generalists. This impedes our understanding of species coexistence in diverse rainforest communities. Host specificity and species richness of tropical insects are also key parameters in mapping global patterns of biodiversity. Here we analyse data for over 900 herbivorous species feeding on 51 plant species in New Guinea and show that most herbivorous species feed on several closely related plant species. Because species-rich genera are dominant in tropical floras, monophagous herbivores are probably rare in tropical forests. Furthermore, even between phylogenetically distant hosts, herbivore communities typically shared a third of their species. These results do not support the classical view that the coexistence of herbivorous species in the tropics is a consequence of finely divided plant resources; non-equilibrium models of tropical diversity should instead be considered. Low host specificity of tropical herbivores reduces global estimates of arthropod diversity from 31 million (ref. 1) to 4 6 million species. This finding agrees with estimates based on taxonomic collections, reconciling an order of magnitude discrepancy between extrapolations of global diversity based on ecological samples of tropical communities with those based on sampling regional faunas.

The ant fauna of a tropical rain forest: estimating species richness three different ways

Abstract: Species richness is an important characteristic of ecological communities, but it is difficult to quantify. We report here a thorough inventory of a tropical rain forest ant fauna and use it to evaluate species richness estimators. The study was carried out in ;1500 ha of lowland rain forest at La Selva Biological Station, Costa Rica. Diverse methods were used, including canopy fogging, Malaise traps, Berlese samples, Winkler samples, baiting, and manual search. Workers of 437 ant species were encountered. The abundance distribution was clearly lognormal, and the distribution emerged from a veil line with each doubling of sampling effort. Three richness estimates were calculated: the area under the fitted lognormal distribution, the asymptote of the Michaelis-Menten equation fit to the species accumulation curve, and the Incidence-based Coverage Estimator (ICE). The per- formance of the estimators was evaluated with sample-based rarefaction plots. The inventory was nearly complete because the species accumulation curve approached an asymptote, the richness estimates were very close to the observed species richness, and the uniques and duplicates curves were both declining. None of the richness estimators was stable in sample- based rarefaction plots, but regions of stability of estimators occurred. The explanation of rarity is one key to understanding why richness estimates fail. Fifty-one species (12% of the total) were still uniques (known from only one sample) at the end of the inventory. The rarity of 20 of these species was explained by ''edge effects'': ''methodological edge species'' (possibly abundant at the site but difficult to sample because of their microhabitat), and ''geographic edge species,'' known to be common in habitats or regions outside of La Selva. Rarity of 31 species remained unexplained. Most of the 51 rare species were known from additional collections outside of La Selva, either in other parts of Costa Rica or in other countries. Only six species were ''global uniques,'' known to date from only one sample on Earth. The study demonstrates that patterns of species occurrence early in an inventory may be inadequate to estimate species richness, but that relatively complete inventories of species-rich arthropod communities are possible if multiple sampling methods and extensive effort are applied.

Nest Predators and Fragmentation: a Review and Meta-Analysis

Abstract: Population declines of many avian species are often attributed to increased rates of nest predation in fragmented landscapes, yet mechanisms underlying these effects have rarely been examined. We reviewed the literature to determine the extent to which hypotheses about nest predators and fragmentation have been invoked and compared this to the number of direct tests of predators with respect to habitat edge, patch size, or landscape type. We also conducted a meta-analysis of tested predator effects to evaluate whether predator responses—numerical, functional, or species richness—to fragmentation depend on spatial scale (edge, patch, or landscape), landscape type, geographic region, or predator taxa. We found 120 papers containing hypoth- eses about nest predators and fragmentation, but only 31 with hypothesis tests. Most tests were of a single predator species or guild, whereas most cited hypotheses generalized across broader taxonomic groups. Re- sults of predator tests were variable, but some general patterns were evident. Predator effects, including in- creased abundance, activity, or species richness in edges, small patches, or certain landscapes, were more prevalent (1) in tests conducted at the landscape scale than at the local scale, (2) in agricultural landscapes than in predominantly forested landscapes, (3) in certain biogeographic regions, and (4) for avian predators than for mammalian predators. Local-scale (edge and patch) effects were most common when the land sur- rounding patches was agricultural and when tests were conducted within agricultural landscapes. The re- sponse of nest predators to fragmentation is complex, taxon-specific, and context-dependent. Conservation ef- forts for declining avian species may therefore need to be customized according to the nest-predator species primarily responsible for local nest mortality and the nature of the landscape mosaic.

Belowground ectomycorrhizal fungal community change over a nitrogen deposition gradient in Alaska

Abstract: Nitrogen availability may be a major factor structuring ectomycorrhizal fungal communities. Atmospheric nitrogen (N) deposition has been implicated in the decline of ectomycorrhizal fungal (EMF) sporocarp diversity. We previously characterized the pattern of decreased sporocarp species richness over an anthropogenic N deposition gradient in Alaska (USA). To determine whether this change in sporocarp community structure was paralleled below ground, we used molecular and morphological techniques to characterize the ectomycorrhizal community of white spruce (Picea glauca) over this gradient. We then related patterns of richness and relative abundance of taxa to various N-affected environmental parameters. Species richness of EMF declined dramatically with increasing N inputs. Over 30 taxa were identified at the low-N sites, compared with nine at the high-N sites. Low-N site dominants (Piloderma spp., Amphinema byssoides, Cortinarius spp., and various dark-mantled Tomentella spp.) disappeared completely at th...

An ecosystem engineer, the beaver, increases species richness at the landscape scale

Abstract: Ecosystem engineering - the physical modifi- cation of habitats by organisms - has been proposed as an important mechanism for maintaining high species richness at the landscape scale by increasing habitat heterogeneity. Dams built by beaver (Castor canaden- sis) dramatically alter riparian landscapes throughout much of North America. In the central Adirondacks, New York, USA, ecosystem engineering by beaver leads to the formation of extensive wetland habitat capable of supporting herbaceous plant species not found else- where in the riparian zone. We show that by increasing habitat heterogeneity, beaver increase the number of species of herbaceous plants in the riparian zone by over 33% at a scale that encompasses both beaver-modified patches and patches with no history of beaver occupa- tion. We suggest that ecosystem engineers will increase species richness at the landscape scale whenever there are species present in a landscape that are restricted to engineered habitats during at least some stages of their life cycle.

Biodiversity, invasion resistance, and marine ecosystem function: reconciling pattern and process

Abstract: A venerable generalization about community resistance to invasions is that more diverse communities are more resistant to invasion. However, results of experimental and observational studies often conflict, leading to vigorous debate about the mechanistic importance of diversity in determining invasion success in the field, as well as other eco- system properties, such as productivity and stability. In this study, we employed both field experiments and observational approaches to assess the effects of diversity on the invasion of a subtidal marine invertebrate community by three species of nonindigenous ascidians (sea squirts). In experimentally assembled communities, decreasing native diversity in- creased the survival and final percent cover of invaders, whereas the abundance of individual species had no effect on these measures of invasion success. Increasing native diversity also decreased the availability of open space, the limiting resource in this system, by buffering against fluctuations in the cover of individual species. This occurred because temporal patterns of abundance differed among species, so space was most consistently and completely occupied when more species were present. When we held diversity constant, but manipulated resource availability, we found that the settlement and recruitment of new invaders dramatically increased with increasing availability of open space. This suggests that the effect of diversity on invasion success is largely due to its effects on resource (space) availability. Apart from invasion resistance, the increased temporal stability found in more diverse communities may itself be considered an enhancement of ecosystem func- tion. In field surveys, we found a strong negative correlation between native-species richness and the number and frequency of nonnative invaders at the scale of both a single quadrat (25 3 25 cm), and an entire site (50 3 50 m). Such a pattern suggests that the means by which diversity affects invasion resistance in our experiments is important in determining the distribution of invasive species in the field. Further synthesis of mechanistic and ob- servational approaches should be encouraged, as this will increase our understanding of the conditions under which diversity does (and does not) play an important role in deter- mining the distribution of invaders in the field.

Comparing classical community models: theoretical consequences for patterns of diversity.

Abstract: Mechanisms proposed to explain the maintenance of species diversity within ecological communities of sessile organisms include niche differentiation mediated by competitive trade-offs, frequency dependence resulting from species-specific pests, re- cruitment limitation due to local dispersal, and a speciation- extinction dynamic equilibrium mediated by stochasticity (drift). While each of these processes, and more, have been shown to act in particular communities, much remains to be learned about their relative importance in shaping community-level patterns. We used a spatially-explicit, individual-based model to assess the effects of each of these processes on species richness, relative abundance, and spatial patterns such as the species-area curve. Our model com- munities had an order-of-magnitude more individuals than any previous such study, and we also developed a finite-size scaling analysis to infer the large-scale properties of these systems in order to establish the generality of our conclusions across system sizes. As expected, each mechanism can promote diversity. We found some qualitative differences in community patterns across com- munities in which different combinations of these mechanisms operate. Species-area curves follow a power law with short-range dispersal and a logarithmic law with global dispersal. Relative- abundance distributions are more even for systems with compet- itive differences and trade-offs than for those in which all species are competitively equivalent, and they are most even when fre- quency dependence (even if weak) is present. Overall, however, communities in which different processes operated showed sur- prisingly similar patterns, which suggests that the form of com- munity-level patterns cannot in general be used to distinguish among mechanisms maintaining diversity there. Nevertheless, pa- rameterization of models such as these from field data on the

Phylogenetic Approaches for Describing and Comparing the Diversity of Microbial Communities

Abstract: Diversity is the hard currency of ecologists. Various statistics have been developed for summarizing the diversity of an ecological community. A commonly adopted summary statistic is the Shannon-Weiner index: H p ilnpi, where pi is the frequency of the ith species. In addition, species richness (the number of different species) often is reported, and recent work emphasizes the importance of accurate estimates of species richness when ecological communities and processes that affect the composition of communities and the function of ecosystems are described (5). The significance of diversity is often inferred by comparing communities characterized from different environments. Typically, such comparisons rely on standard measures of overlap, including the percentage of species shared by two communities or similarity indices. One of the indices used is Sorensen’s index: S S 12/[0.5(S1 S2)], where S 12 is the number of species common to both sites and Si is the number of species found at site i. A limitation of traditional statistics for describing and comparing diversity is that species (or operational taxonomic units [OTUs]) are defined inconsistently. For instance, Kroes et al. (6) defined an OTU as a 16S ribosomal DNA (rDNA) sequence group in which sequences differed by less than 1%. By contrast, the definition of McCaig et al. (11) included sequences that were less than 3% different, and other studies have used 5% as the magic number. The lack of consensus limits the comparative utility of statistics based solely on identification of species (or OTUs). A second, and perhaps more important, limitation of the standard statistics of diversity is that OTUs are counted equivalently even though some may be highly divergent and phylogenetically unique, whereas others may be part of a closely related group of species and are therefore phylogenetically redundant (4). The contrast can be illustrated by comparing two hypothetical communities in which the numbers of species, the richness profiles of species, and the rarefaction profiles are identical but which differ in the magnitude of phylogenetic diversity (i.e., the degree of divergence among the sampled sequences). Standard ecological statistics of diversity would miss the genetic difference between the two communities, and ecologists would most likely consider the two communities equally diverse when, in fact, one community harbors more genetic diversity (or disparity) than the other. Because genetic variation and phenotypic variance often are positively correlated in populations of animals (12), plants (7), and microbes (15), descriptions of microbial communities based on DNA data should include information about diversity and disparity. This is especially important in light of studies demonstrating an association between ecosystem function and community diversity (14, 28). In this review I introduce various statistics borrowed from population genetics and systematics for describing and comparing the diversity evident from samples of gene sequences. I briefly introduce the statistics and methodological underpinnings of tests for differences between communities, and I use the methods to analyze well-described microbial communities. I show that information gained from analysis of DNA sequences provides the basis for statistical analysis of communities in ways that advance inferences about the processes that may govern the compositions and functions of microbial communities. Furthermore, the advocated analytical approaches make it possible to accomplish broad comparisons of ecological communities. The methods of analysis explored in this paper are meant to be complementary to other methods, such as the robust estimation of richness advocated by Hughes et al. (5) and approaches for estimating functional properties of bacteria from phylogenetic inference (16).

Species Invasions Exceed Extinctions on Islands Worldwide: A Comparative Study of Plants and Birds

Abstract: Species richness is decreasing at a global scale. At subglobal scales, that is, within any defined area less extensive than the globe, species richness will increase when the number of nonnative species becoming naturalized is greater than the number of native species becoming extinct. Determining whether this has occurred is usually difficult because detailed records of species extinctions and naturalizations are rare; these records often exist, however, for oceanic islands. Here we show that species richness on oceanic islands has remained relatively unchanged for land birds, with the number of naturalizations being roughly equal to the number of extinctions, and has increased dramatically for vascular plants, with the number of naturalizations greatly exceeding the number of extinctions. In fact, for plants, the net number of species on islands has approximately doubled. We show further that these patterns are robust to differences in the history of human occupation of these islands and to th...

Benthic indicators to use in Ecological Quality classification of Mediterranean soft bottom marine ecosystems, including a new Biotic Index

Abstract: A general scheme for approaching the objective of Ecological Quality Status (EcoQ) classification of zoobenthic marine ecosystems is presented. A system based on soft bottom benthic indicator species and related habitat types is suggested to be used for testing the typological definition of a given water body in the Mediterranean. Benthic indices including the Shannon-Wiener diversity index and the species richness are re-evaluated for use in classification. Ranges of values and of ecological quality categories are given for the diversity and species richness in different habitat types. A new biotic index (BENTIX) is proposed based on the relative percentages of three ecological groups of species grouped according to their sensitivity or tolerance to disturbance factors and weighted proportionately to obtain a formula rendering a five step numerical scale of ecological quality classification. Its advantage against former biotic indices lies in the fact that it reduces the number of the ecological groups involved which makes it simpler and easier in its use. The Bentix index proposed is tested and validated with data from Greek and western Mediterranean ecosystems and examples are presented. Indicator species associated with specific habitat types and pollution indicator species, scored according to their degree of tolerance to pollution, are listed in a table. The Bentix index is compared and evaluated against the indices of diversity and species richness for use in classification. The advantages of the BENTIX index as a classification tool for ECoQ include independence from habitat type, sample size and taxonomic effort, high discriminative power and simplicity in its use which make it a robust, simple and effective tool for application in the Mediterranean Sea.

Geographic patterns in plant/pollinator mutualistic networks

Abstract: Recent reviews of plant-pollinator mutualistic networks showed that gen- eralization is a common pattern in this type of interaction. Here we examine the ecological correlates of generalization patterns in plant-pollinator networks, especially how interaction patterns covary with latitude, elevation, and insularity. We review the few published anal- yses of whole networks and include unpublished material, analyzing 29 complete plant- pollinator networks that encompass arctic, alpine, temperate, Mediterranean, and subtrop- ical-tropical areas. The number of interactions observed (I) was a linear function of network size (M) the maximum number of interactions: ln I 5 0.575 1 0.61 ln M; R 2 5 0.946. The connectance (C), the fraction of observed interactions relative to the total possible, decreased exponentially with species richness, the sum of animal and plant species in each community (A 1 P): C 5 13.83 exp(20.003(A 1 P)). After controlling for species richness, the residual connectance was significantly lower in highland (.1500 m elevation) than in lowland networks and differed marginally among biogeographic regions, with both alpine and trop- ical networks showing a trend for lower residual connectance. The two Mediterranean networks showed the highest residual connectance. After correcting for variation in network size, plant species were shown to be more generalized at higher latitude and lowland habitats, but showed increased specialization on islands. Oceanic island networks showed an im- poverishment of potential animal pollinators (lower ratio of animal to plant species, A : P, compared to mainland networks) associated with this trend of increased specialization. Plants, but not their flower-visiting animals, supported the often-repeated statements about higher specificity in the tropics than at higher latitudes. The pattern of interaction build- up as diversity increases in pollination networks does not differ appreciably from other

Assessing the vulnerability of species richness to anthropogenic climate change in a biodiversity hotspot

Abstract: Aim To compare theoretical approaches towards estimating risks of plant species loss to anthropogenic climate change impacts in a biodiversity hotspot, and to develop a practical method to detect signs of climate change impacts on natural populations. Location The Fynbos biome of South Africa, within the Cape Floristic Kingdom. Methods Bioclimatic modelling was used to identify environmental limits for vegetation at both biome and species scale. For the biome as a whole, and for 330 species of the endemic family Proteaceae, tolerance limits were determined for five temperature and water availability-related parameters assumed critical for plant survival. Climate scenarios for 2050 generated by the general circulation models HadCM2 and CSM were interpolated for the region. Geographic Information Systems-based methods were used to map current and future modelled ranges of the biome and 330 selected species. In the biome-based approach, predictions of biome areal loss were overlayed with species richness data for the family Proteaceae to estimate extinction risk. In the species-based approach, predictions of range dislocation (no overlap between current range and future projected range) were used as an indicator of extinction risk. A method of identifying local populations imminently threatened by climate change-induced mortality is also described. Results A loss of Fynbos biome area of between 51% and 65% is projected by 2050 (depending on the climate scenario used), and roughly 10% of the endemic Proteaceae have ranges restricted to the area lost. Species range projections suggest that a third could suffer complete range dislocation by 2050, and only 5% could retain more than two thirds of their range. Projected changes to individual species ranges could be sufficient to detect climate change impacts within ten years. Main conclusions The biome-level approach appears to underestimate the risk of species diversity loss from climate change impacts in the Fynbos Biome because many narrow range endemics suffer range dislocation throughout the biome, and not only in areas identified as biome contractions. We suggest that targeted vulnerable species could be monitored both for early warning signs of climate change and as empirical tests of predictions.

Geographic patterns in plant–pollinator mutualistic networks

Abstract: Recent reviews of plant-pollinator mutualistic networks showed that gen- eralization is a common pattern in this type of interaction. Here we examine the ecological correlates of generalization patterns in plant-pollinator networks, especially how interaction patterns covary with latitude, elevation, and insularity. We review the few published anal- yses of whole networks and include unpublished material, analyzing 29 complete plant- pollinator networks that encompass arctic, alpine, temperate, Mediterranean, and subtrop- ical-tropical areas. The number of interactions observed (I) was a linear function of network size (M) the maximum number of interactions: ln I 5 0.575 1 0.61 ln M; R 2 5 0.946. The connectance (C), the fraction of observed interactions relative to the total possible, decreased exponentially with species richness, the sum of animal and plant species in each community (A 1 P): C 5 13.83 exp(20.003(A 1 P)). After controlling for species richness, the residual connectance was significantly lower in highland (.1500 m elevation) than in lowland networks and differed marginally among biogeographic regions, with both alpine and trop- ical networks showing a trend for lower residual connectance. The two Mediterranean networks showed the highest residual connectance. After correcting for variation in network size, plant species were shown to be more generalized at higher latitude and lowland habitats, but showed increased specialization on islands. Oceanic island networks showed an im- poverishment of potential animal pollinators (lower ratio of animal to plant species, A : P, compared to mainland networks) associated with this trend of increased specialization. Plants, but not their flower-visiting animals, supported the often-repeated statements about higher specificity in the tropics than at higher latitudes. The pattern of interaction build- up as diversity increases in pollination networks does not differ appreciably from other

Plant Diversity of the Cape Region of Southern Africa

Abstract: Comprising a land area of ca. 90,000 km 2 , less than one twentieth (5%) the land area of the southern African subcontinent, the Cape Floristic Region (CFR) is, for its size, one of the world's richest areas of plant species diversity. A new synoptic flora for the Region has made possible an accurate reassessment of the flora, which has an estimated 9030 vascular plant species (68.7% endemic), of which 8920 species are flowering plants (69.5% endemic). The number of species packed into so small an area is remarkable for the temperate zone and compares favorably with species richness for areas of similar size in the wet tropics. The Cape region consists of a mosaic of sandstone and shale substrata with local areas of limestone. It has a highly dissected, rugged topography, and a diversity of climates with rainfall mostly falling in the winter months and varying from 2000 mm locally to less than 100 mm. Ecological gradients are steep as a result of abrupt differences in soil, altitude, aspect, and precipitation. These factors combine to form an unusually large number of local habitats for plants. Sandstone-derived soils have characteristically low nutrient status, and many plants present on such soils have low seed dispersal capabilities, a factor promoting localized distributions. An unusual family composition includes Iridaceae, Aizoaceae, Ericaceae, Scrophulariaceae, Proteaceae, Restionaceae, Rutaceae, and Orchidaceae among the 10 largest families in the flora, following Asteraceae and Fabaceae, as the most speciose families. Disproportionate radiation has resulted in over 59.2% species falling in the 10 largest families and 77.4% in the largest 20 families. Twelve genera have more than 100 species and the 20 largest genera contribute some 31% of the total species. Species richness of the Cape flora is hypothesized to be the result of geographic and parapatric radiation in an area with a mosaic of different habitats due to local soil, climate, and altitudinal differences that combine to produce steep ecological gradients. Also contributing to the diversity has been a relatively stable geological history since the end of the Miocene that saw the establishment of a semi-arid and extreme seasonal climate at the southwestern part of southern Africa.

Species Richness and Altitude: A Comparison between Null Models and Interpolated Plant Species Richness along the Himalayan Altitudinal Gradient, Nepal

Abstract: We compare different null models for species richness patterns in the Nepalese Himalayas, the largest altitudinal gradient in the world. Species richness is estimated by interpolation of presences between the extreme recorded altitudinal ranges. The number of species in 100-m altitudinal bands increases steeply with altitude until 1,500 m above sea level. Between 1,500 and 2,500 m, little change in the number of species is observed, but above this altitude, a decrease in species richness is evident. We simulate different null models to investigate the effect of hard boundaries and an assumed linear relationship between species richness and altitude. We also stimulate the effect of interpolation when incomplete sampling is assumed. Some modifications on earlier simulations are presented. We demonstrate that all three factors in combination may explain the observed pattern in species richness. Estimating species richness by interpolating species presence between maximum and minimum altitudes creates an artificially steep decrease in species richness toward the ends of the gradient. The addition of hard boundaries and an underlying linear trend in species richness is needed to simulate the observed broad pattern in species richness along altitude in the Nepalese Himalayas.

Faunal and environmental change in the late Miocene Siwaliks of northern Pakistan

Abstract: The Siwalik formations of northern Pakistan consist of deposits of ancient rivers that existed throughout the early Miocene through the late Pliocene. The formations are highly fossil- iferous with a diverse array of terrestrial and freshwater vertebrates, which in combination with exceptional lateral exposure and good chronostratigraphic control allows a more detailed and tem- porally resolved study of the sediments and faunas than is typical in terrestrial deposits. Conse- quently the Siwaliks provide an opportunity to document temporal differences in species richness, turnover, and ecological structure in a terrestrial setting, and to investigate how such differences are related to changes in the fluvial system, vegetation, and climate. Here we focus on the interval between 10.7 and 5.7 Ma, a time of significant local tectonic and global climatic change. It is also the interval with the best temporal calibration of Siwalik faunas and most comprehensive data on species occurrences. A methodological focus of this paper is on controlling sampling biases that confound biological and ecological signals. Such biases include uneven sampling through time, differential preservation of larger animals and more durable skeletal elements, errors in age-dating imposed by uncertainties in correlation and paleomagnetic timescale calibrations, and uneven tax- onomic treatment across groups. We attempt to control for them primarily by using a relative-abun- dance model to estimate limits for the first and last appearances from the occurrence data. This model also incorporates uncertainties in age estimates. Because of sampling limitations inherent in the terrestrial fossil record, our 100-Kyr temporal resolution may approach the finest possible level of resolution for studies of vertebrate faunal changes over periods of millions of years. Approximately 40,000 specimens from surface and screenwash collections made at 555 localities form the basis of our study. Sixty percent of the localities have maximum and minimum age esti- mates differing by 100 Kyr or less, 82% by 200 Kyr or less. The fossils represent 115 mammalian species or lineages of ten orders: Insectivora, Scandentia, Primates, Tubulidentata, Proboscidea, Pholidota, Lagomorpha, Perissodactyla, Artiodactyla, and Rodentia. Important taxa omitted from this study include Carnivora, Elephantoidea, and Rhinocerotidae. Because different collecting methods were used for large and small species, they are treated separately in analyses. Small spe- cies include insectivores, tree shrews, rodents, lagomorphs, and small primates. They generally weigh less than 5 kg. The sediments of the study interval were deposited by coexisting fluvial systems, with the larger emergent Nagri system being displaced between 10.1 and 9.0 Ma by an interfan Dhok Pathan sys- tem. In comparison to Nagri floodplains, Dhok Pathan floodplains were less well drained, with smaller rivers having more seasonally variable flow and more frequent avulsions. Paleosol se- quences indicate reorganization of topography and drainage accompanying a transition to a more seasonal climate. A few paleosols may have formed under waterlogged, grassy woodlands, but most formed under drier conditions and more closed vegetation. The oxygen isotopic record also indicates significant change in the patterns of precipitation be- ginning at 9.2 Ma, in what may have been a shift to a drier and more seasonal climate. The carbon isotope record demonstrates that after 8.1 Ma significant amounts of C 4 grasses began to appear and that by 6.8 Ma floodplain habitats included extensive C4 grasslands. Plant communities with predominantly C3 plants were greatly diminished after 7.0 Ma, and those with predominantly C4 plants, which would have been open woodlands or grassy woodlands, appeared as early as 7.4 Ma. Inferred first and last appearances show a constant, low level of faunal turnover throughout the interval 10.7-5.7-Ma, with three short periods of elevated turnover at 10.3, 7.8, and 7.3-7.0 Ma. The three pulses account for nearly 44% of all turnover. Throughout the late Miocene, species richness declined steadily, and diversity and richness indices together with data on body size imply that community ecological structure changed abruptly just after 10 Ma, and then again at 7.8 Ma. Be- tween 10 and 7.8 Ma the large-mammal assemblages were strongly dominated by equids, with more balanced faunas before and after. The pattern of appearance and disappearance is selective with respect to inferred habits of the animals. Species appearing after 9.0 Ma are grazers or typical of more open habitats, whereas many species that disappear can be linked to more closed vege- tation. We presume exceptions to this pattern were animals of the mixed C