Spatial and temporal variability in species richness in a temperate intertidal community

Understanding changes in estuarine benthic communities has important implications for conservation and yet it is a challenge due to the high natural variability of these systems. We addressed this challenge through the study of temporal and spatial patterns of species richness in an intertidal benthic community in New Zealand North Island. Five different locations within the estuary were monitored seasonally over 12 yr. This data set allowed the study of species–time–area relationships (STAR) and the delineation of patterns in species richness, heterogeneity and turnover in space and time. The site with the highest species richness also had the highest within‐site heterogeneity in species richness, a high number of species occurring infrequently in time, the lowest mud content and the most variable wave climate. Similarities and differences between sites were generally maintained over time, although seasonal and multi‐year patterns in species richness occurred at all sites. The STAR showed a significant negative interaction between space and time, with species accumulation rates in space and time being equivalent at 4 spatial replicates (250 m²) and 2 temporal replicates (6 months). The lowest source of variability in species turnover was within site, and the highest source was over years. This was reflected in the lack of an asymptotic relationship in the species accumulation curve despite the 12 yr of monitoring. These results contribute to the knowledge of the variability in diversity patterns in estuaries and have important implications for long‐term monitoring of natural communities and the estimation of diversity for conservation.     

Plant species richness and environmental heterogeneity in a mountain landscape: effects of variability and spatial configuration

The loss of biodiversity has become a matter of urgent concern and a better understanding of local drivers is crucial for conservation. Although environmental heterogeneity is recognized as an important determinant of biodiversity, this has rarely been tested using field data at management scale. We propose and provide evidence for the simple hypothesis that local species diversity is related to spatial environmental heterogeneity. Species partition the environment into habitats. Biodiversity is therefore expected to be influenced by two aspects of spatial heterogeneity: 1) the variability of environmental conditions, which will affect the number of types of habitat, and 2) the spatial configuration of habitats, which will affect the rates of ecological processes, such as dispersal or competition. Earlier, simulation experiments predicted that both aspects of heterogeneity will influence plant species richness at a particular site. For the first time, these predictions were tested for plant communities using field data, which we collected in a wooded pasture in the Swiss Jura mountains using a four-level hierarchical sampling design. Richness generally increased with increasing environmental variability and roughness (i.e. decreasing spatial aggregation). Effects occurred at all scales, but the nature of the effect changed with scale, suggesting a change in the underlying mechanisms, which will need to be taken into account if scaling up to larger landscapes. Although we found significant effects of environmental heterogeneity, other factors such as history could also be important determinants. If a relationship between environmental heterogeneity and species richness can be shown to be general, recently available high-resolution environmental data can be used to complement the assessment of patterns of local richness and improve the prediction of the effects of land use change based on mean site conditions or land use history.     

Environmental effects on vertebrate species richness: testing the energy, environmental stability and habitat heterogeneity hypotheses

Background

Explaining species richness patterns is a central issue in biogeography and macroecology. Several hypotheses have been proposed to explain the mechanisms driving biodiversity patterns, but the causes of species richness gradients remain unclear. In this study, we aimed to explain the impacts of energy, environmental stability, and habitat heterogeneity factors on variation of vertebrate species richness (VSR), based on the VSR pattern in China, so as to test the energy hypothesis, the environmental stability hypothesis, and the habitat heterogeneity hypothesis.

Methodology/Principal Findings

A dataset was compiled containing the distributions of 2,665 vertebrate species and eleven ecogeographic predictive variables in China. We grouped these variables into categories of energy, environmental stability, and habitat heterogeneity and transformed the data into 100×100 km quadrat systems. To test the three hypotheses, AIC-based model selection was carried out between VSR and the variables in each group and correlation analyses were conducted. There was a decreasing VSR gradient from the southeast to the northwest of China. Our results showed that energy explained 67.6% of the VSR variation, with the annual mean temperature as the main factor, which was followed by annual precipitation and NDVI. Environmental stability factors explained 69.1% of the VSR variation and both temperature annual range and precipitation seasonality had important contributions. By contrast, habitat heterogeneity variables explained only 26.3% of the VSR variation. Significantly positive correlations were detected among VSR, annual mean temperature, annual precipitation, and NDVI, whereas the relationship of VSR and temperature annual range was strongly negative. In addition, other variables showed moderate or ambiguous relations to VSR.

Conclusions/Significance

The energy hypothesis and the environmental stability hypothesis were supported, whereas little support was found for the habitat heterogeneity hypothesis.     

Demographic response of plant populations to habitat fragmentation and temporal environmental variability

Many plant species currently exist in fragmented populations of different sizes, while they also experience unpredictable climatic fluctuation over time. However, we still understand little about how plant demography responds to such spatial and temporal environmental variability. We studied population dynamics of an understory perennial herb Trillium camschatcense in the Tokachi plain of Hokkaido, Japan, where a significant effect of forest fragmentation on seedling recruitment was previously reported. Four populations across a range of fragment sizes were studied for 6 years, and the data were analyzed using matrix population models. Per capita fecundity (the number of recruits per plant) varied greatly among populations, but the variation in population growth rates (λ) was mainly driven by the variation in stasis and growth rates, suggesting that the general trend of reduced fecundity in fragmented populations may not be readily translated into subsequent dynamics. Temporal variation in λ among years was more than 2 times larger than spatial variation among populations, and this result was likely attributable to the contrasting response of correlation structures among demographic rates. The among-population variation in λ was dampened by negative covariation between matrix elements possibly due to density-dependent regulation as well as an inherent constraint that some elements are not independent, whereas positive covariation between matrix elements resulted in large temporal variation in λ. Our results show that population dynamics responded differently to habitat fragmentation and temporal variability of the environment, emphasizing the need to discriminate these spatial and temporal variations in demographic models. Although no populations were projected to be declining in stochastic simulations, correlation between current habitat size and plant density implies historical λ is positively related to habitat size.     

Water and fertilizer have opposite effects on plant species richness in a mesic early successional habitat

Herbaceous plant species richness typically declines with increasing productivity, but differences in the resources underlying these gradients are often ignored. This study adds to the small number of studies examining the effects of water and mineral nutrients on biomass and richness in oldfield communities. We established 60 4 m² plots in a goldenrod-dominated oldfield to test the differential effects of water and mineral nutrients on community properties. Species richness declined with added nutrients, but increased with added water. Aboveground biomass increased only when both nutrients and water were added. Leaf area index increased with added nutrients alone, although the increase was greater when water was also added. Understory light levels decreased with added nutrients, but not with added water; however, the per-gram effect of biomass on understory light levels did not vary significantly among nutrient and water treatments. Our results suggest that water tends to enhance productivity, but does not cause the common decline in species richness that typically accompanies nitrogen additions. Rather, water increased richness through positivie effects on either germination and establishment, or increased survival. These results are consistent with either increased light limitation or increased water limitation causing loss of species from nitrogen-rich habitats.     

Distinguishing area and habitat heterogeneity effects on species richness: Birds in Victorian buloke remnants

Abstract Resolving whether area per se or habitat heterogeneity has the greater influence in controlling species richness remains a controversial yet important question. Here we show that avian species richness of same-sized transects (1 ha) is independent of the remnant area (of buloke woodland) within which a transect is positioned. We also show that avi-faunal similarity of pairs of transects randomly placed within the largest remnants (≥ 48 ha) is not consistently related to either proximity (i. e. being within the same remnant) nor to physiognomic characteristics of the transects. We believe that much of the controversy over area/habitat heterogeneity effects is probably related to scalar issues and propose a protocol by which some resolution of the question might be reached. The protocol involves ‘zoom’ sampling in which successively larger transect sizes are used, and measures of faunal richness and habitat heterogeneity are made at these different grains of resolution. One of our intentions is to stimulate discussion on how heterogeneity might be measured when grains increase from typical transect sizes (ca 1 ha) up to much larger grains (ca 128 ha).     

Stability and species richness in complex communities

Using both numerical simulations and analytical methods, we investigate how the stability of ecological communities depends on the number of species they contain. To investigate complex communities, we construct communities from modular subcommunities that can have arbitrary community structure; e.g. subcommunities could consist of pairs of predator and prey species, trios of prey, specialist predator and generalist predator, or any collection of interacting species. By building entire communities from subcommunities, we can change the number of species in the community without changing community structure. We further suppose that species sharing the same ecological role in different subcommunities act additively on the per capita population growth rates of other species. Under these assumptions, the inter-actions between species from different subcommunities have no effect on community-level stability, measured by the variability in the combined densities of species sharing the same ecological role in different subcommunities. Furthermore, increasing species richness (i.e. the number of subcommunities comprising the community) increases community-level stability only when it introduces species that respond differently to environmental fluctuations. Therefore, our results support the insurance hypothesis that species richness increases community-level stability by insuring that some species in a community are tolerant of different environmental fluctuations.     

Multiple environmental determinants of regional species richness and effects of geographic range size

Understanding patterns of species richness at broad geographic extents remains one of the most challenging yet necessary scientific goals of our time. Many hypotheses have been proposed to account for spatial variation in species richness; among them, environmental determinants have played a central role. In this study, we use data on regional bat species richness in the New World to study redundancy and complementarity of three environmental hypotheses: energy, heterogeneity and seasonality. We accomplish this by partitioning variation in species richness among components associated with unique and combined effects of variables from each hypotheses, and by partitioning the overall richness gradient into gradients of species with varying breadths of geographic distribution. These three environmental hypotheses explain most variation in the species richness gradient of all bats, but do not account for all positive spatial autocorrelation at short distances. Although environmental predictors are highly redundant, energy and seasonality explain different and complementary fractions of variation in species richness of all bats. On the other hand, heterogeneity variables contribute little to explain this gradient. However, results change dramatically when richness is estimated for groups of species with different sizes of geographic distribution. First, the amount of variation explained by environment decreases with a decrease in range size; this suggests that richness gradients of small‐ranged species can not be explained as easily as those of broadly distributed species, as has been implied by analyses that do not consider differences in range size among species. Second, the relative contribution of environmental predictors to explained variation also changes with change in range size. Seasonality and energy are good predictors of species with broad distributions, but they loose almost all explanatory power for richness of species with small ranges. In contrast, heterogeneity, which is a relatively poor predictor of richness of species with large ranges, becomes the main predictor of richness gradients of species with restricted distributions. This suggests that range size is a different dimension on which heterogeneity and other environmental characteristics are complementary to each other. Our results suggest that determinants of species richness gradients might be complex, or at least more complex than many studies have previously suggested.     

Stochastic relations between species richness and the variability of species composition

Ecological communities change over time and space, and ecologists have long suggested that biodiversity might influence the rate of change. Here we cast new light on this question by demonstrating statistical covariation between species richness and the variability of species abundances and identities within a community (compositional variability). We provide a new analytical framework for several previously published measures of ecosystem functioning and compositional variability. We derive three related variances, each of which measures compositional variability due solely to stochastic sampling processes. Our analyses show that whether relations between species richness and compositional variability are positive, negative or zero, depends on two factors. Not only does the particular variance used affect the relation, but, more importantly, the underlying determinants of species richness can strongly affect the stochastic relations between species richness and compositional variability. This analysis makes clear for the first time how species richness should correlate with important measures of community variability, even in the absence of systematic processes.     

Species richness, temporal variability and resistance of biomass production in a Mediterranean grassland

We studied the temporal variability and resistance to perturbation of the biomass production of grassland communities from an experimental diversity gradient (the Portuguese BIODEPTH project site). With increasing species richness relative temporal variability (CV) of plant populations increased but that of communities decreased, supporting the insurance hypothesis and related theory. Species‐rich communities were more productive than species‐poor communities in all three years although a natural climatic perturbation in the third year (frequent frost and low precipitation) caused an overall decrease in biomass production. Resistance to this perturbation was constant across the experimental species richness gradient in relative terms, supporting a similar response from the Swiss BIODEPTH experiment. The positive biomass response was generated by different combinations of the complementarity and selection effects in different years. Complementarity effects were positive across mixtures on average in all three years and positively related to diversity in one season. The complementarity effect declined following perturbation in line with total biomass but, counter to predictions, in relative terms overyielding was maintained in all years. Selection effects were positively related to diversity in one year and negative overall in the other two years. The response to perturbation varied among species and for the same species growing in monoculture and mixture, but following the frost communities were more strongly dominated by species with lower monoculture biomass and the selection effect was more negative. In total, our results support previous findings of a positive relationship between diversity and productivity and between diversity and the temporal stability of production, but of no effect of diversity on the resistance to perturbation. We demonstrate for the first time that the relative strength of overyielding remained constant during an exceptional natural environmental perturbation.     

Dominant species constrain effects of species diversity on temporal variability in biomass production of tallgrass prairie

Species diversity is thought to stabilize functioning of plant communities. An alternative view is that stability depends more on dynamics of dominant species than on diversity. We compared inter-annual variability (inverse of stability) of aboveground biomass in paired restored and remnant tallgrass prairies at two locations in central Texas, USA. Data from these two locations were used to test the hypothesis that greater richness and evenness in remnant than restored prairies would reduce variability in aboveground biomass in response to natural variation in rainfall. Restored prairies were chosen to be similar to paired remnant prairies in characteristics other than species diversity that affect temporal variability in biomass. Variability was measured as the coefficient of variation among years (square root of variance/mean; CV), where variance in community biomass equals the sum of variances of individual plant species plus the summed covariances between species pairs. Species diversity over five years was greater by a factor of 2 or more in remnant than restored prairies because richness and evenness were greater in remnant than restored prairies. Still, the CV of community biomass during spring and CV of annual biomass production did not differ consistently between prairie types. Neither the sum of species covariances nor total community biomass differed between prairies. Biomass varied relatively little in restored compared to remnant prairies because biomass of the dominant species in restored prairies (the grass Schizachyrium scoparium ) varied less than did biomass of other dominant and sub-dominant species. In these grasslands, biomass response to natural variation in precipitation depended as much on characteristics of a dominant grass as on differences in diversity.     

Small-scale variability in a mosaic tropical rainforest influences habitat use of long-tailed macaques

Pristine habitats have generally been considered to be the most important ecological resource for wildlife conservation, but due to forest degradation caused by human activities, mosaics of secondary forests have become increasingly prominent. We studied three forest types in a mosaic tropical forest consisting of short secondary forest (SS), tall secondary forest (TS) and freshwater swamp forest (SF). These forests differed in stand structure and floristic composition, as well as phenological productivity of fruits, flowers and young leaves. We examined habitat use of long-tailed macaques (Macaca fascicularis) in relation to indices of phenological activity. The macaques used the SS for feeding/foraging more than the TS and the SF. This was because the SS had higher productivity of fruit, which is a preferred food resource for macaques. Stem densities of young leaves in the SS and the TS also influenced habitat use, as they provided more clumped resources. Use of SF was limited, but these forests provided more species-rich resources. Our results showed that M. fascicularis responded to small-scale variability in phenological activity between forest types found in a heterogeneous mosaic forest, with young secondary regrowth forests likely providing the most important food resources. Mosaic landscapes may be important as they can buffer the effects of temporal food resource variability in any given forest type. In our increasingly human-altered landscapes, a better understanding of the role of secondary forest mosaics is crucial to the conservation and management of wildlife habitats and the animals they support.     

Mobility-dependent effects on species richness in fragmented landscapes

In fragmented landscapes, mobility is an important trait for population persistence but the predictions on the relationship between habitat fragmentation and extinction risk are contradictory. Here, we test the effects of the two main aspects of fragmentation, patch area and isolation, on the species richness of groups of butterflies associated with semi-natural grasslands, differing in mobility. Total species richness increased with increasing patch area and with decreasing isolation, but the strength of these effects differed between mobility classes. The effect of patch area was strongest for the sedentary species, while the effect of isolation was only statistically significant for the mobile species. We interpret these results as evidence for a predominant influence of local processes on sedentary species, and an increasing influence of regional compared to local processes with increasing mobility. When groups of species respond differently to habitat loss and fragmentation this affects community composition, with potential implications for ecosystem processes. Similar effects can be expected for other traits than mobility, and this should be an important question for future studies.     

Enhancement of habitat heterogeneity and species richness on rocky shores inundated by sand

Summary Many rocky shores are subject to periodic inundation by sand, which is often thought to reduce species richness by eliminating organisms intolerant of sand scour or sand smothering. However, regular disturbance (e.g. inundation) should promote richness by preventing the development of low diversity climax communities. A study of faunal richness on 10 regularly inundated shores showed that inundation does promote richness, but by increasing habitat heterogeneity. Some species are excluded from parts of the shore by sand, but because of the patchiness of sand deposits they are rarely excluded from the entire shore. Other species are found only on rocks associated with sand, while typically sandy shore animals occur in the sand deposits themselves. Total richness (281 species) was greater than for local noninundated shores and sandy beaches combined.     

Survival in king penguins Aptenodytes patagonicus: temporal and sex-specific effects of environmental variability

We investigated annual adult survival rates of king penguins Aptenodytes patagonicus breeding at South Georgia during 6 years in relation to age/breeding experience, sex, and food availability. During the first 3 years of the study, when food availability was good, survival was 97.7% for experienced breeders, which confirmed the very high survival rates observed in penguins in general. In these years survival did not differ between the sexes, presumably because parental investment is shared equally between the sexes, and the sexual dimorphism is small in king penguins. Survival was lower for young, first-time breeders (83.0%). In experienced birds the annual survival rate decreased to 68-82% following a catastrophic year when food availability was extremely low. We address the question how parents balance their current investment in offspring against their chances to reproduce in the future. We argue that the high mortality rate among breeding individuals after the year of food stress provides support for previous suggestions that the response to increased costs in seabirds might be complex to predict and does not always follow intuitive expectations according to general life-history theory. We also found that females survived significantly less well than males following the bad year. We explain this result as follows: the male-biased sex ratio (56:44) that we observed in our study colony clearly does not result from lower female survival during normal conditions. An already existing skewed sex ratio forces males to delay the onset of breeding because of a lack of breeding partners. This in turn causes breeding females to be, on average, younger and less experienced than males and to have lower survival following a year of food shortage. In this study survival was linked with food availability and we suggest that this was connected to climatic/oceanographic features, such as the position of the Antarctic Polar Front Zone. We could, however, not verify this by anomalies in sea surface temperature data.     

Determinants of species richness, endemism and turnover in European longhorn beetles

This study assessed the diversity patterns of a large family of beetles, Cerambycidae, in Europe and tested the following hypotheses: 1) richness gradients of this hyperdiverse taxon are driven by water and energy variables; 2) endemism is explained by the same factors, but variation between areas also reflects post‐glacial re‐colonization processes; and 3) faunal composition is determined by the same climatic variables and, therefore, beta diversity (species turnover) is related to richness gradients. Species richness, endemism and beta diversity were modelled using inventories of 37 European territories, built from a database containing the distributions of 609 species. Area, spatial position, and nine topographical and climatic variables were used as predictors in regression and constrained analysis of principal coordinates modelling. Species richness was mostly explained by a temperature gradient, which produced a south‐to‐north decreasing richness gradient. Endemism followed the same pattern, but was also determined by longitudinal variation, peaking in the southwestern and southeastern corners of the continent. Faunal turnover was explained by an important purely spatial pattern and a spatially structured environmental gradient. Thus, contrary to other groups, cerambycid richness was mostly explained by environmental energy, but not by water availability. Endemism was concentrated in the Iberian and Greek peninsulas, but not in Italy. Thus, the latter area may have been the major source of post‐glacial re‐colonization for European longhorn beetles or, otherwise, a poor refuge during glaciations. Turnover patterns were independent of the richness gradient, because northern faunas are nested in southern ones. Turnover, in contrast to richness, was driven by both the independent effects of climate and geographic constraints that might reflect dispersal limitation or stochastic colonization events, suggesting that richness gradients are more environmentally deterministic phenomena than turnover patterns.     

Lizard species richness patterns in China and its environmental associations

Many ecological hypotheses have been widely used to explain species richness variation across the globe. We investigated lizard species richness patterns in China, and identified areas of high species richness. Furthermore, we tested hypotheses concerning the relationships between lizard richness and environmental variables. A large data including 30,902 records of point locality data for 151 lizard species occurring in China were retrieved from Herpetology museums of CIB/CAS and other museums through HerpNET, and published sources, and then predicted distributions maps were generated using ecological niche modeling. We overlaid all species prediction maps into a composite map to describe species richness patterns. A multiple regression analysis using eigenvector-based spatial filtering (SEVM) was performed to examine the best environmental predictors of species richness. Richness peaked mainly in southern China located in the Oriental realm. Our best multiple regression models explained a total of 80.1% variance of lizard richness (r² = 0.801; F = 203.47; P < 0.001). Among related factors in shaping species richness distribution, the best environmental predictors of species richness were: frost-day frequency, elevation, vegetation, and wet-day frequency. Based on models selection, our results revealed that underlying mechanisms related to different ecological hypotheses might work together and best explain lizard richness in China. We are in an initial step to develop a large data set on species richness, and provide the necessary conservation implications from habitat loss. Additional studies that test species richness at different geographical scale are required to better understand the factors that may influence the species richness distribution in East Asia.