Speciation-rate dependence in species–area relationships

The general tendency for species number (S) to increase with sampled area (A) constitutes one of the most robust empirical laws of ecology, quantified by species–area relationships (SAR). In many ecosystems, SAR curves display a power-law dependence, S∝A^z. The exponent z is always less than one but shows significant variation in different ecosystems. We study the multitype voter model as one of the simplest models able to reproduce SAR similar to those observed in real ecosystems in terms of basic ecological processes such as birth, dispersal and speciation. Within the model, the species–area exponent z depends on the dimensionless speciation rate ν, even though the detailed dependence is still matter of controversy. We present extensive numerical simulations in a broad range of speciation rates from ν=10^-3 down to ν=10^-11, where the model reproduces values of the exponent observed in nature. In particular, we show that the inverse of the species–area exponent linearly depends on the logarithm of ν. Further, we compare the model outcomes with field data collected from previous studies, for which we separate the effect of the speciation rate from that of the different species lifespans. We find a good linear relationship between inverse exponents and logarithm of species lifespans. However, the slope sets bounds on the speciation rates that can hardly be justified on evolutionary basis, suggesting that additional effects should be taken into account to consistently interpret the observed exponents.     

Species–area relationships across four trophic levels – decreasing island size truncates food chains

That larger areas will typically host more diverse ecological assemblages than small ones has been regarded as one of the few fundamental ‘laws’ in ecology. Yet, area may affect not only species diversity, but also the trophic structure of the local ecological assemblage. In this context, recent theory on trophic island biogeography offers two clear‐cut predictions: that the slope of the species–area relationship should increase with trophic rank, and that food chain length (i.e. the number of trophic levels) should increase with area. These predictions have rarely been verified in terrestrial systems. To offer a stringent test of key theory, we focused on local food chains consisting of trophic specialists: plants, lepidopteran herbivores, and their primary and secondary parasitoids. For each of these four trophic levels, we surveyed species richness across a set of 20 off‐shore continental islands spanning a hundred‐fold range in size. We then tested three specific hypotheses: that species richness is affected by island size, that the slope of the species–area curve is related to trophic rank, and that such differences in slope translate into variation in food chain length with island size. Consistent with these predictions, estimates of the species–area slope steepened from plants through herbivores and primary parasitoids to secondary parasitoids. As a result of the elevated sensitivity of top consumers to island size, food chain length decreased from large to small islands. Since island size did not detectably affect the ratio between generalists and specialists among either herbivores (polyphages vs oligophages) or parasitoids (idiobionts vs koinobionts), the patterns observed seemed more reflective of changes in the overall number of nodes and levels in local food webs than of changes in their linking structure. Overall, our results support the trophic‐level hypothesis of island biogeography. Per extension, they suggest that landscape modification may imperil food web integrity and vital biotic interactions.     

Fragmentation,grazing and the species–area relationship

Habitat loss is one of the greatest threats to species persistence. Gauging the scale of this problem requires quantitative methods that can predict the number of extinctions resulting from habitat loss. For the past three decades, the species–area relationship, an empirical relationship between the number of species present in an area and the size of that area, has been this tool. However, it fails to incorporate threats to species aside from habitat loss and the heterogeneous distribution of these threats across habitats. Recent studies have improved species–area predictions by incorporating not only direct effects of area on richness, but also indirect effects of area (through area‐mediated predator abundance), on prey species richness. We extend this work to test the hypotheses that the indirect effects of the multiple threats of grazing and trampling in addition to fragmentation will amplify the effect of area on species richness and that this effect will be greatest in zones closest to the fragment edge. Further, we test for species and population level effects of fragmentation and grazing, including the non‐random pattern of species loss and the decline in population sizes. We test our hypotheses with a field study of land snail richness in fragments with and without the additional threats of grazing and trampling. Our study supports the hypotheses that fragments with multiple threats in addition to habitat loss harbour fewer species than fragments without these threats, and that this effect is non‐uniform across fragments, populations and species.     

Identifying hotspots of parasite diversity from species–area relationships: host phylogeny versus host ecology

Interspecific variation in parasite species richness among host species has generated much empirical research. As in comparisons among geographical areas, controlling for variation in host body size is crucial because host size determines resource availability. Recent developments in the use of species–area relationships (SARs) to detect hotspots of biodiversity provide a powerful way to control for host body size, and to identify ‘hot’ and ‘cold hosts’ of parasite diversity, i.e. hosts with more or fewer parasites than expected from their size. Applying SAR modelling to six large datasets on parasite species richness in vertebrates, we search for hot and cold hosts and assess the effect of other ecological variables on the probability that a host species is hot/cold taking body size (and sampling effort) into account. Five non‐sigmoid SAR models were fitted to the data by optimisation; their relative likelihood was evaluated using the Bayesian information criterion, before deriving an averaged SAR function. Overall, the fit between the five SAR models and the actual data was poor; there was substantial uncertainty surrounding the fitted models, and the best model differed among the six datasets. These results show that host body size is not a strong or consistent determinant of parasite species richness across taxa. Hotspots were defined as host species lying above the upper limit of the 80% confidence interval of the averaged SAR, and coldspots as species lying below its lower limit. Our analyses revealed (1) no apparent effect of specific ecological factors (i.e. water temperature, mean depth range, latitude or population density) on the likelihood of a host species being a hot or coldspot; (2) evidence of phylogenetic clustering, i.e. hosts from certain families are more likely to be hotspots (or coldspots) than other species, independently of body size. These findings suggest that host phylogeny may sometimes outweigh specific host ecological traits as a predictor of whether or not a host species harbours more (or fewer) parasite species than expected for its size.     

Estimates of species extinctions from species–area relationships strongly depend on ecological context

Species–area (SAR) and endemics–area (EAR) relationships are amongst the most common methods used to forecast species loss resulting from habitat loss. One critical, albeit often ignored, limitation of these area‐based estimates is their disregard of the ecological context that shapes species distributions. In this study, we estimate species loss using a spatially explicit mechanistic simulation model to evaluate three important aspects of ecological context: coexistence mechanisms (e.g. species sorting, competition–colonization tradeoffs and neutral dynamics), spatial distribution of environmental conditions, and spatial pattern of habitat loss. We found that 1) area‐based estimates of extinctions are sensitive to coexistence mechanisms as well as to the pattern of environmental heterogeneity; 2) there is a strong interaction between coexistence mechanisms and the pattern of habitat loss; 3) SARs always yield higher estimates of species loss than do EARs; and 4) SARs and EARs consistently underestimate the realized species loss. Our results highlight the need to integrate ecological mechanisms in area‐estimates of species loss.     

mmSAR: an R‐package for multimodel species–area relationship inference

The species–area relationship (SAR) is one of the most fundamental tools in ecology. After almost a century of quantitative ecology, however, the quest for a “best SAR model” still remains elusive, with a substantial uncertainty about the best fitting SAR model frequently being observed. Recent research has required that this uncertainty be addressed, and a multimodel SAR framework has been devised. Here we introduce the mmSAR R‐package, which is a flexible and scalable implementation of the multimodel SAR framework for species‐area datasets, and provide some examples of its use. This R‐package provides functions for fitting SAR models, performing model selection, and the build up of multimodel SARs.     

Individual species–area relationships and spatial patterns of species diversity in a Great Basin,semi‐arid shrubland

Traditional biodiversity metrics operate at the level of a plant community but do not capture spatial variation in diversity from a ‘plant's‐eye view’ of a community. Recently‐developed statistics consider the spatial patterns of plants as well as the number and distribution of species in local plant neighborhoods to quantitatively assess multispecies spatial patterns from a ‘plant's‐eye view’. We used one such statistic, the individual species area relationship (ISAR), to assess spatial patterns of species diversity in a Great Basin (USA) semi‐arid shrubland through an analysis of a spatial dataset on shrub species and locations. In conjunction with appropriate null models, the ISAR blends species area relationships with second‐order spatial statistics to measure the expected species richness in local neighborhoods of variable size around the individuals of a focal species within a community. We found that, contrary to a previous analysis using more traditional methods, the community was well‐mixed with a typical shrub surrounded on average by 4.9 shrub neighbors of 2.1 species at a neighborhood scale of 1.0 m. We also found statistically significant fine‐scale variation in diversity patterns, such that neighborhoods of two species were more diverse than expected by a heterogeneous Poisson null model that accounted for larger‐scale habitat heterogeneity. However, this effect was caused by intraspecific aggregation of these species and was not due to positive interspecific association. Contrary to previous findings in other semi‐arid shrublands, our analysis suggests that the spatial pattern of the shrub community was not significantly structured by interspecific facilitation. This result supports growing evidence for balanced species patterns of adult plants in multispecies communities. Our approach may be used in other communities to describe complex multispecies spatial patterns, quantify species‐specific associations with diversity patterns, and to generate hypotheses regarding relationships between patterns and community‐structuring processes.     

Biodiversity on urban roundabouts—Hemiptera, management and the species–area relationship

The biodiversity of insects within urban areas has been relatively little studied. Given the large and ever increasing extent of urban areas, and that the insect species richness there can be high, it is important to know the factors determining that aspect of biodiversity. In this study two of these factors, namely habitat management and area, were considered. Arboreal and grassland Hemiptera, and grassland plants, were sampled on 18 roundabouts and other road enclosed sites in the town of Bracknell. Hemiptera were sampled using suction sampling and tree beating. A significant species–area relationship was found for arboreal Hemiptera, which was strongly related to habitat diversity. For both grassland plants and Hemiptera, grassland management, by mowing, had a significant effect on species richness. Despite the management grassland plants showed a significant species–area relationship. However the effect of management on Hemiptera was great enough to outweigh any area effect. As the size of open spaces is often constrained in urban areas, altering habitat management has a greater potential for enhancing biodiversity. For arboreal Hemiptera choice of trees for planting is of particular importance, while for grassland Hemiptera diversity would be increased with a reduction in the intensity of management, such a reduction in the frequency of mowing.

Zusammenfassung

Die Biodiversität der Insekten auf urbanen Flächen ist relativ wenig untersucht. Angesichts der großen und der immer größer werdenden Ausdehnung urbaner Gebiete und angesichts dessen, dass der Artenreichtum der Insekten dort groß sein kann, ist es wichtig die Faktoren zu kennen, die diesen Aspekt der Biodiversität bestimmen. In dieser Untersuchung wurden zwei dieser Faktoren, nämlich Habitatmanagement und Fläche, betrachtet. Baum- und wiesenbewohnende Hemiptera sowie Wiesenpflanzen wurden in 18 Kreisverkehren und anderen straßenumschlossenen Orten innerhalb der Stadt Bracknell gesammelt. Die Hemiptera wurden mit Saugproben und Klopfproben an den Bäumen gesammelt. Für die baumbewohnenden Hemiptera wurde eine signifikante Art-Areal-Beziehung gefunden, die in enger Beziehung zur Habitatdiversität stand. Sowohl für die Wiesenpflanzen als auch für die Hemiptera hatte das Wiesenmanagement in Form von Mahd einen signifikanten Einfluss auf den Artenreichtum. Trotz des Managements zeigten die Wiesenpflanzen eine signifikante Art-Areal-Beziehung. Die Auswirkungen des Managements auf die Hemiptera waren jedoch groß genug, um den Arealeffekt zu überwiegen. Da die Größe offener Flächen in städtischen Gebieten oft beschränkt ist, hat die Änderung des Habitatmanagements ein größeres Potenzial die Biodiversität zu erhöhen. Für baumbewohnende Hemiptera ist die Auswahl der Bäume für die Bepflanzung von besonderer Wichtigkeit, während für die wiesenbewohnenden Hemiptera die Diversität durch eine Verringerung der Managementintensität erhöht würde, wie z. B. durch die Verringerung der Mahdfrequenz.     

Species–area relationship of Neotropical waterbird assemblages in remnant wetlands: looking at the mechanisms

Aim We investigated the relative role of area, isolation, microhabitat diversity and number of individuals as explanatory factors defining the richness of waterbirds in wetland remnants. Location Freshwater marshes along the Atlantic coastal zone of South Brazil. (30°56′–30°22′S; 50°58–50°22′W; Fig. 1 ).

Figure 1 Open in figure viewer PowerPoint The study area (dashed line), in the Atlantic coastal zone of Brazil, is characterized by a high concentration of remnant wetlands used by resident and migratory waterbirds. Waterbirds were surveyed monthly along 2003 in 42 randomly selected wetland remnants.     

Effects of community structure on the species–area relationship in China's forests

The species–area relationship (SAR) is the oldest and most frequently documented law in ecology. In a community, the SAR is regulated by the abiotic environment and biotic interactions and depends on the individual–spatial distribution of species (ISD) and the species–abundance distribution (SAD). In this study, we explored the effects of aggregation of ISDs and unevenness of SADs on SARs in forests of China by comparing the empirical and simulated SARs of 32 nested plots distributed along an extensive latitudinal gradient. Both aggregation and unevenness affected the shape of SARs significantly: ISDs accounted for 12.6 ± 4.0% of the incremental increase in species richness with area, and SADs accounted for 18.7 ± 3.8 and 23.5 ± 3.9% under the broken‐stick model and even abundance model, respectively. Effects of both aggregation and unevenness decreased as temperature increased, suggesting that individuals of a species were spatially more aggregated than random, and the individuals among species were more discrepant from the null distribution (broken‐stick model and even abundance model in this study), in the cold than in the warm areas. Taken together, our results demonstrate that ISDs and SADs within communities can shape SARs, but these effects vary along latitudinal gradients, and are likely mediated by temperature.     

Ecological thresholds: an assessment of methods to identify abrupt changes in species–habitat relationships

Habitat thresholds are usually defined as “points of abrupt change” in the species–habitat relationships. Habitat thresholds can be a key tool for understanding species requirements, and provide an objective definition of conservation targets, by identifying when habitat loss leads to a rapid loss of species, and the minimum amount of habitat necessary for species persistence. However, a large variety of statistical methods have been used to analyse them. In this context, we reviewed these methods and, using simulated data sets, we tested the main models to compare their performance on the identification of thresholds. We show that researchers use very different analytical tools, corresponding to different operational definitions of habitat thresholds, which can considerably affect their detection. Piecewise regression and generalized additive models allow both the distinction between linear and nonlinear dynamics, and the correct identification of break point position. In contrast, other methods such as logistic regression fail because they may incorrectly detect thresholds in gradual patterns, or they may over or underestimate the threshold position. In conservation or habitat modelling, it is important to focus efforts efficiently and the inappropriate choice of statistical methods may have detrimental consequences.     

Plant communities affect the species–area relationship on Carex sempervirens tussocks

The species–area relationship (SAR) has been extensively studied in a wide range of plant communities, but very few studies have directly addressed how plant communities affect the SAR and what are the underlying mechanisms. Many graminoids form distinct tussocks where many other plant species grow, but no study has investigated whether the SAR holds true for the vegetation on tussocks. In four plant communities on an abandoned subalpine pasture in the Swiss National Park, we made releves on 600 tussocks of Carex sempervirens and measured tussock basal area and other tussock traits. In all four communities, species richness on C. sempervirens tussocks was strongly positively related to tussock basal area (R²0.74), while other tussock traits explained very little (R²<0.04). Slope and intercept of the SAR on C. sempervirens tussocks differed significantly among the four communities. This was because plant communities affected richness in smaller tussocks (basal diameter <10 cm) but not that in large tussocks (basal diameter10 cm). We conclude that the SAR holds true for vegetation on C. sempervirens tussocks and changes with plant communities. Changes in the SAR on C. sempervirens tussocks are very likely because smaller tussocks are less independent of the plant communities than the larger ones, regarding disturbance or nutrients.     

The species–energy theory: a role for energy variability

Species–energy theory posits that energy availability regulates population sizes, extinction rates and ultimately species richness. This theory has focused mostly on total energy as a measure of energy availability. However, because energy variation can also influence population sizes and extinction rates, species–energy theory should arguably consider simultaneously both total energy and its variation. Using data on species richness of land birds and mammals, we compared the fit of three species–energy models including total energy, energy variation or both combined. We show that the combination of total energy and energy variation has greater predictive power than any of them considered separately. We also evaluate three crucial assumptions of this modified species–energy theory and show that they are supported by available data. These results illuminate the current debate on climate change, given that both average conditions and variability of climatic conditions are likely to change in the future.     

A meta‐analysis of species–abundance distributions

The species–abundance distribution (SAD) describes the abundances of all species within a community. Many different models have been proposed to describe observed SADs. Best known are the logseries, the lognormal, and a variety of niche division models. They are most often visualized using either species richness – log abundance class (Preston) plots or abundance – species rank order (Whittaker) plots. Because many of the models predict very similar shapes, model distinction and testing become problematic. However, the variety of models can be classified into three basic types: one that predicts a double S‐shape in Whittaker plots and a unimodal distribution in Preston plots (the lognormal type), a second that lacks the mode in Preston plots (the logseries type), and a third that predicts power functions in both plotting types (the power law type). Despite the interest of ecologists in SADs no formal meta‐analysis of models and plotting types has been undertaken so far. Here we use a compilation of 558 species–abundance distributions from 306 published papers to infer the frequency of the three SAD shapes in dependence of environmental variables and type of plotting. Our results highlight the importance of distinguishing between fully censused and incompletely sampled communities in the study of SADs. We show that completely censused terrestrial or freshwater animal communities tend to follow lognormal type SADs more often than logseries or power law types irrespective of species richness, spatial scale, and geographic position. However, marine communities tend to follow the logseries type, while plant communities tend to follow the power law. In incomplete sets the power law fitted best in Whittaker plots, and the logseries in Preston plots. Finally our study favors the use of Whittaker over Preston plots.     

Species–topography association in a species-rich subtropical forest of China

Habitat specialization has been considered as a primary factor in determining the distribution of species. In this study, we investigated species–habitat associations while controlling for spatial neighbourhood effects in a large-scale (20 ha) stem-mapping plot in a species-rich subtropical forest of China. Habitat specialization was measured by topographic variation and its effects on species distributions were modelled at three different spatial scales (10×10, 20×20 and 25×25 m²) using log-linear regression models and randomization tests. Our results showed: (1) 83% of the species were related to at least one or more topographic variables. Among them, 66%, 60%, 65% and 70% were closely dependent on slope, aspect, elevation and convexity, respectively. (2) Topographic variables have much stronger non-linear (quadratic and cubic) effects on species distributions than linear effects. (3) The effects of topographic heterogeneity on the distribution of shrubs species are smaller than on the distribution of canopy species, and smaller effects were also found in less abundant species. (4) There was a strong neighbourhood effect on species distribution: In 85% of the species, abundance in a focal quadrat was significantly correlated with abundance in the neighbour quadrats. We conclude that habitat specialization plays an important role in maintaining the diversity of this species-rich subtropical forest.     

Lichens on dead wood: species‐substrate relationships in the epiphytic lichen floras of the Pacific Northwest and Fennoscandia

Dead wood is an important habitat feature for lichens in forest ecosystems, but little is known about how many and which lichens are dependent on dead wood. We reviewed substrate use by epiphytic lichens in the combined floras of Fennoscandia and the Pacific Northwest of North America based on literature and herbarium data and analyzed substrate affinity relative to life form, reproductive mode and major phylogenetic group within the floras. A total of 550 (43%) of the 1271 epiphytic species in the combined floras use wood, and 132 species (10%) are obligately associated with dead wood in one or both regions. Obligate and facultative wood‐dwelling guilds in the two floras were strongly similar in terms of internal guild structure in each region, but differ somewhat in species composition, while the bark‐dwelling guild differs strongly in both. Most obligate dead wood users are sexually reproducing crustose lichens. The largest numbers of species are associated with forest structural features such as logs and snags that have been greatly reduced by forest practices. Conservation of lichens inhabiting wood requires greater attention to crustose lichen species and the development of conservation strategies that look beyond numbers and volumes of dead wood and consider biologically meaningful dead wood structure types.     

Gel–water relationships: Dielectric dispersion

At low frequencies, the dielectric constant of gels of agar, carboxymethyl cellulose, gelatin, mid maize starch is much higher than that of water. It decreases continuously as the frequency increases, tending to level off at about 10⁸ cps. The dielectric constant is lower the higher the concentration of polymer; it increases in the order: carboxymethyl cellulose, agar, starch, and gelatin. Results are most reasonably explained by assuming a more solid structure of water the lower the dielectric constant.