Fish species richness in spring‐fed ponds: effects of habitat size versus isolation in temporally variable environments

1. Species richness in a habitat patch is determined by immigration (regional) and extinction (local) processes, and understanding their relative importance is crucial for conservation of biodiversity. In this study, we applied the Island Biogeography concept to spring ponds connected to a river in southwestern Japan to examine how immigration and extinction processes interact to determine fish species richness in temporally variable environments. 2. Fish censuses were conducted 15 times in 13 study ponds at 1–4 month intervals from August 1998 through October 2000. Effects of habitat size (pond area), isolation (distance from the river) and temporal environmental variability (water level fluctuation) on (i) species richness, (ii) immigration and extinction rates and (iii) population size and persistence of each fish species were assessed. 3. The results revealed predominant effects of distance on species richness, immigration/extinction rates and population size and persistence. Species richness decreased with increasing distance but was not related to either pond area or water level fluctuation. A negative effect of distance on immigration rate was detected, while neither pond area nor water level fluctuation had significant effects on extinction rate. Further, population size and persistence of four species increased with decreasing distance, suggesting that, in ponds close to the river, immigrants from the river reduce the probability of extinction (i.e. provide a rescue effect), contributing to the maintenance of high species richness. 4. Overall results emphasise the importance of immigration processes, rather than extinction, in shaping patterns of species richness in our system. The predominant importance of immigration was probably because of (i) high temporal variability that negates habitat‐size effects and (ii) continuous immigration that easily compensates for local extinctions. Our results suggest that consideration of regional factors (e.g. connectivity, locations of source populations and barriers to colonisation) is crucial for conservation and restoration of local habitats.     

Nitrate availability and hydrophyte species richness in shallow lakes

1. Submerged plant richness is a key element in determining the ecological quality of freshwater systems; it has often been reduced or completely lost. 2. The submerged and floating‐leaved macrophyte communities of 60 shallow lakes in Poland and the U.K. have been surveyed and species richness related to environmental factors by general linearised models. 3. Nitrogen, and more specifically winter nitrate, concentrations were most important in explaining species richness with which they were inversely correlated. Phosphorus was subsidiary. Such an inverse relationship is consistent with findings in terrestrial communities. Polish lakes, with less intensively farmed catchments, had greater richness than the U.K. lakes. 4. The richest U.K. communities were associated with winter nitrate‐N concentrations of up to about 1–2 mg L^?1 and may correspond with ‘good’ ecological quality under the terms of the European Water Framework Directive. Current concentrations in European lowlands are often much higher.     

Connectivity and cladoceran species richness in a metacommunity of shallow lakes

SUMMARY 1. The shallow ponds of the nature reserve 'De Maten' form a metacommunity, in which individual ponds are highly interconnected via a system of overflows and rivulets. This study reports on the relations between cladoceran species richness and (a) connectivity patterns and (b) local environmental variables.
2. No relation was found between local species richness and three connectivity variables or dispersal pathways.
3. Spatial configuration was related to richness, but was confounded by environmental variables for 2 of 3 years. In those 2 years, there was a significant linear relation between Secchi disc depth and species richness, suggesting an important impact of the clearwater/turbid state alternative equilibria in shallow lakes in determining cladoceran richness. Only in the year in which environmental variables were unimportant did connectivity between the ponds influence species richness.
4. These results suggest that local environmental variables related to the clearwater/turbid state alternative equilibria in shallow lakes are important in determining cladoceran species richness. Connectivity and dispersal of individuals between the different ponds only act secondarily by increasing the general species richness within a pond through dispersal from ponds with different environmental conditions.     

Dragonfly species richness on man-made ponds: effects of pond size and pond age on newly established assemblages

We studied the abundance and species richness of adult dragonflies in 11 artificial ponds which were recently established (within 2years). We found that the adult dragonfly assemblage patterns were influenced by pond size as well as pond age. The species richness was positively correlated with the pond size, which was because the distributional patterns of species were significantly nested according to pond area. The species richness was highly correlated with pond age in association with the vegetation cover within ponds. It was suggested that the species richness was enhanced by the increasing immigration rate of species which favor well-vegetated ponds.     

Effects of habitat fragmentation and isolation on species richness: evidence from biogeographic patterns

Summary Habitat subdivision by geography or human activity may be an important determinant of regional species richness. Cumulative species-area relationships for vertebrates, land plants, and insects on island archipelagoes show that collections of small islands generally harbor more species than comparable areas composed of one or a few large islands. The effect of the degree of habitat subdivision in increasing species richness appears to increase with the distance from potential sources of colonists. Mountaintop biotas show no clear differences between species richness on large alpine areas and collections of smaller peaks. National park faunas generally have more species in collections of small parks than in the larger parks. In all cases where a consistent effect of subdivision is observed, the more subdivided collection of islands or isolates contains more species. To the degree that these data provide guidance for establishing nature reserves, they suggest that increasing the numbers of reserves may be an important component of conservation strategies.     

Trophic structure, species richness and biodiversity in Danish lakes: changes along a phosphorus gradient

1. Using data from 71, mainly shallow (an average mean depth of 3 m), Danish lakes with contrasting total phosphorus concentrations (summer mean 0.02–1.0 mg P L?l), we describe how species richness, biodiversity and trophic structure change along a total phosphorus (TP) gradient divided into five TP classes (class 1–5: <0.05, 0.05–0.1, 0.1–0.2, 0.2–0.4,> 0.4 mg P L?1).
2. With increasing TP, a significant decline was observed in the species richness of zooplankton and submerged macrophytes, while for fish, phytoplankton and floating‐leaved macrophytes, species richness was unimodally related to TP, all peaking at 0.1–0.4 mg P L?1. The Shannon–Wiener and the Hurlbert probability of inter‐specific encounter (PIE) diversity indices showed significant unimodal relationships to TP for zooplankton, phytoplankton and fish. Mean depth also contributed positively to the relationship for rotifers, phytoplankton and fish.
3. At low nutrient concentrations, piscivorous fish (particularly perch, Perca fluviatilis) were abundant and the biomass ratio of piscivores to plankti‐benthivorous cyprinids was high and the density of cyprinids low. Concurrently, the zooplankton was dominated by large‐bodied forms and the biomass ratio of zooplankton to phytoplankton and the calculated grazing pressure on phytoplankton were high. Phytoplankton biomass was low and submerged macrophyte abundance high.
4. With increasing TP, a major shift occurred in trophic structure. Catches of cyprinids in multiple mesh size gill nets increased 10‐fold from class 1 to class 5 and the weight ratio of piscivores to planktivores decreased from 0.6 in class 1 to 0.10–0.15 in classes 3–5. In addition, the mean body weight of dominant cyprinids (roach, Rutilus rutilus, and bream, Abramis brama) decreased two–threefold. Simultaneously, small cladocerans gradually became more important, and among copepods, a shift occurred from calanoid to cyclopoids. Mean body weight of cladocerans decreased from 5.1 μg in class 1 to 1.5 μg in class 5, and the biomass ratio of zooplankton to phytoplankton from 0.46 in class 1 to 0.08–0.15 in classes 3–5. Conversely, phytoplankton biomass and chlorophyll a increased 15‐fold from class 1 to 5 and submerged macrophytes disappeared from most lakes.
5. The suggestion that fish have a significant structuring role in eutrophic lakes is supported by data from three lakes in which major changes in the abundance of planktivorous fish occurred following fish kill or fish manipulation. In these lakes, studied for 8 years, a reduction in planktivores resulted in a major increase in cladoceran mean size and in the biomass ratio of zooplankton to phytoplankton, while chlorophyll a declined substantially. In comparison, no significant changes were observed in 33 ‘control’ lakes studied during the same period.     

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).     

Rethinking patch size and isolation effects: the habitat amount hypothesis

I challenge (1) the assumption that habitat patches are natural units of measurement for species richness, and (2) the assumption of distinct effects of habitat patch size and isolation on species richness. I propose a simpler view of the relationship between habitat distribution and species richness, the ‘habitat amount hypothesis’, and I suggest ways of testing it. The habitat amount hypothesis posits that, for habitat patches in a matrix of non‐habitat, the patch size effect and the patch isolation effect are driven mainly by a single underlying process, the sample area effect. The hypothesis predicts that species richness in equal‐sized sample sites should increase with the total amount of habitat in the ‘local landscape’ of the sample site, where the local landscape is the area within an appropriate distance of the sample site. It also predicts that species richness in a sample site is independent of the area of the particular patch in which the sample site is located (its ‘local patch’), except insofar as the area of that patch contributes to the amount of habitat in the local landscape of the sample site. The habitat amount hypothesis replaces two predictor variables, patch size and isolation, with a single predictor variable, habitat amount, when species richness is analysed for equal‐sized sample sites rather than for unequal‐sized habitat patches. Studies to test the hypothesis should ensure that ‘habitat’ is correctly defined, and the spatial extent of the local landscape is appropriate, for the species group under consideration. If supported, the habitat amount hypothesis would mean that to predict the relationship between habitat distribution and species richness: (1) distinguishing between patch‐scale and landscape‐scale habitat effects is unnecessary; (2) distinguishing between patch size effects and patch isolation effects is unnecessary; (3) considering habitat configuration independent of habitat amount is unnecessary; and (4) delineating discrete habitat patches is unnecessary.     

Island theory, matrix effects and species richness patterns in habitat fragments

Island biogeography theory, created initially to study diversity patterns on islands, is often applied to habitat fragments. A key but largely untested assumption of this application of theory is that landscape matrix species composition is non‐overlapping with that of the islands. We tested this assumption in successional old field patches in a closely mowed matrix, and because our patches are appropriately viewed as sets of contiguous habitat units we studied patterns of species richness per unit area. Previous studies at our site did not find that diversity patterns on patch ‘islands’ conformed to predictions of island biogeography theory. Our results indicate that when matrix species are removed from the patch samples, diversity patterns conform better to theory. We suggest that classical island theory remains an appropriate tool to study diversity patterns in fragmented habitats, but that allowances should be made for spill‐over colonization of ‘islands’ from the ‘sea’.     

Mixed effects of habitat fragmentation on species richness and community structure in a microarthropod microecosystem

Abstract.  1. Theory is unclear about the optimal degree of isolation of habitat fragments where the aim is to maximise species richness. In a field-based microecosystem of Collembola and predatory and non-predatory mites, moss patches of the same total area were fragmented to varying degrees. The habitat was left for several months to allow the communities to approach a new state of equilibrium.
2. The species richness (in particular of predatory mites) of a given area of habitat was greater when it was part of a large mainland area than part of an island, in agreement with theory.
3. Conversely, species richness and abundance were largely unaffected by fragmentation of a fixed area of island habitat. In this case, it is suggested here that the advantages of several small patches (e.g. reduced impact of environmental stochasticity, wider range of habitats overall) were equally balanced by the advantages of a single large patch (e.g. reduced effect of demographic stochasticity, wider range of habitats within a single patch, reduced edge effect), or that both effects were small.
4. The shapes of rank–abundance curves were similar among the levels of fragmentation of a fixed area of island habitat, implying that fragmentation had little impact on community structure. Conversely, the species composition of non-predatory mites varied weakly, but significantly, by fragmentation.     

Predicting distributions of species richness and species size in regional floras: Applying the species pool hypothesis to the habitat templet model

The species pool hypothesis is applied here to the interpretation of ‘hump-shaped’ (unimodal) species richness patterns along gradients of both habitat fertility and disturbance level (the habitat templet). A ‘left-wall’ effect analogous to that proposed for the evolution of organismal complexity predicts a right-skewed unimodal distribution of historical habitat commonness on both gradients. According to the species pool hypothesis, therefore, the distribution of opportunity for net species accumulation (speciation minus extinction) should also have a corresponding unimodal central tendency on both habitat gradients. Two assumptions of this hypothesis are illustrated with particular reference to highly fertile, relatively undisturbed habitats: (i) such habitats have been relatively uncommon in space and time, thus providing relatively little historical opportunity for the origination of species with the traits necessary for effective competitive ability under these habitat conditions; and (ii) species that have evolved adaptation to these habitats are relatively large, thus imposing fundamental ‘packing’ limitations on the number of species that can ‘fit’ within such habitats. Based on these assumptions, the species pool hypothesis defines two associated predictions that are both supported by available data: (a) resident species richness will be relatively low in highly fertile, relatively undisturbed contemporary habitats; and (b) species sizes within regional floras should display as a right-skewed unimodal (log-normal) distribution. The latter is supported here by an analysis of data for 2,715 species in the vascular flora of northeastern North America.     

The influence of different disturbance frequencies on the species richness,diversity and equitability of phytoplankton in shallow lakes

The relationships between the species richness, diversity and equitability of phytoplankton is discussed in the context of Connell's (1978, Science 199: 1304–1310) Intermediate Disturbance Hypothesis (IDH). The records of 759 vertical phytoplankton samples, which were obtained from four shallow central European lakes (Balaton, Neusiedlersee, and two small artificial ponds) at daily to weekly intervals were analysed.

1. The Shannon-Weaver function was used to measure diversity of the recorded species compositionof the phytoplankton. It is shown on fictitious data that compositional diversity is sensitive to the numberof coequilibrating species provided that the suspected interrelationship between diversity and ‘complexity’is amenable to the application of this method.
2. The disturbance scale that was developed on the basis of the field records fits well to Reynolds'(1988, Verh. int. Ver. Limnol. 23: 683–691) derivation: < 3 days qualifies as high frequency, approximatel3–8 days as intermediate frequency and > 8–9 days as low frequency of disturbance for phytoplankton.
3. Arithmetical means of the compositional diversity of phytoplankton under different frequencies ofdisturbance support the hypothesis that maximal diversity appears at intermediate frequencies.
4. There are different reasons for decrease in diversity at higher and lower frequencies. Inequitabilitydiminishes diversity at low disturbance; while species number decreases at high frequencies.
5. The case of Neusiedlersee calls attention to the fact that it is difficult, if at all possible, to differentiatebetween the indices under continuous stress and high frequency of disturbance in lakes in temperateregions. Similar species number-equitability pattern are induced by both and it is also presumablethat high frequency disturbance can itself effect a serious stress.
6. The striking effects that regular major periodic events (e.g. significant changes in the grazing pressureat the onset of the clear-water phase, autumnal cooling) in the plankton have on its species diversity areevident. Thus, the relative importance of intermediate frequency disturbances has its own seasonality:it is increasingly important in periods (partly in the spring, but mostly in the summer-autumn equilibriumphases), in which competition among phytoplankton species is increasing. This observation suggestsa way by which the stochasticity-based IDH can be incorporated into rather more deterministicexplanations (e.g. PEG-model; Sommer et al., 1986. Archiv für Hydrobiologie 106: 433–471) of planktonsuccession.
7. The most controversial issue and, therefore, the main difficulty, with IDH is that it not onlymaintains species richness in an ecosystem but it also supposes its presence. The lack of either earlyor late successional species in a given community can inactivate the mechanism. From the point of viewof the diversity-species richness relationship, the persistence of disturbance at given frequencies is ofgreater importance than the temporal alterations themselves in the evolutionary ecology of the phytoplankton.
8. For characteristically unperturbed phytoplankton communities (no case was studied here), equilibriumconcepts (niche diversification, etc.) should be more strongly applicable to their diversity andspecies richness.

     

The Habitat Amount Hypothesis implies negative effects of habitat fragmentation on species richness

The habitat amount hypothesis (HAH) predicts that species richness in a habitat site increases with the amount of habitat in the ‘local landscape’ defined by an appropriate distance around the site, with no distinct effects of the size of the habitat patch in which the site is located. It has been stated that a consequence of the HAH, if supported, would be that it is unnecessary to consider habitat configuration to predict or manage biodiversity patterns, and that conservation strategies should focus on habitat amount regardless of fragmentation. Here, I assume that the HAH holds and apply the HAH predictions to all habitat sites over entire landscapes that have the same amount of habitat but differ in habitat configuration. By doing so, I show that the HAH actually implies clearly negative effects of habitat fragmentation, and of other spatial configuration changes, on species richness in all or many of the habitat sites in the landscape, and that these habitat configuration effects are distinct from those of habitat amount in the landscape. I further show that, contrary to current interpretations, the HAH is compatible with a steeper slope of the species–area relationship for fragmented than for continuous habitat, and with higher species richness for a single large patch than for several small patches with the same total area (SLOSS). This suggests the need to revise the ways in which the HAH has been interpreted and can be actually tested. The misinterpretation of the HAH has arisen from confounding and overlooking the differences in the spatial scales involved: the individual habitat site at which the HAH gives predictions, the local landscape around an individual site and the landscapes or regions (with multiple habitat sites and different local landscapes) that need to be analysed and managed. The HAH has been erroneously viewed as negating or diminishing the relevance of fragmentation effects, while it actually supports the importance of habitat configuration for biodiversity. I conclude that, even in the cases where the HAH holds, habitat fragmentation and configuration are important for understanding and managing species distributions in the landscape.     

Environmental correlates of plant and invertebrate species richness in ponds

Ponds (lentic water bodies <2 ha) constitute a considerable biodiversity resource. Understanding the environmental factors that underlie this diversity is important in protecting and managing the habitat. We surveyed 425 ponds for biological and physical characteristics with 78 of those also surveyed for chemical characteristics. A total of 277 invertebrate species and 265 plant species were found. Species richness varied between 2 and 99 (mean 27.2 ± 0.6 SE) for invertebrates and 1 and 58 (mean 20.8 ± 0.4 SE) for plants. Generalised additive models were used to investigate variables that correlate with the species richness of plants and invertebrates, with additional models to investigate insect, Coleoptera, Odonata, Hemiptera, Trichoptera and Mollusca species richness. Models performed reasonably well for invertebrates in general (R ² = 30.3%) but varied between lower-order invertebrate taxa (12.7–34.7%). Ponds with lower levels of shading and no history of drying contained higher numbers of species of plants and all invertebrate groups. Aquatic plant coverage positively correlated with species richness in all invertebrate groups apart from Trichoptera and the presence of fish was associated with high invertebrate species richness in all groups apart from Coleoptera. The addition of chemistry variables suggested non-linear relationships between oxygen demand and phosphate concentration and higher-order richness. We demonstrate that the composition of biological communities varies along with their species richness and that less diverse ponds are more variable compared to more diverse ponds. Variables positively correlated with richness of one taxon may be negatively correlated with that of another, making comprehensive management recommendations difficult. Promoting a high landscape-level pond biodiversity will involve the management of a high diversity of pond types within that landscape.     

Opposite effects of environmental variability and species richness on temporal turnover of species in a complex habitat mosaic

Species are continuously lost and added to a local community. Dynamics of this process in a complex habitat mosaic (multiple habitats in a landscape), particularly of its rates (species turnover) are of primary concern for biodiversity conservation. Various studies suggest that species traits such as habitat specialization should affect species turnover. In communities where habitat specialization is a function of abiotic constraints, habitat specialists should respond faster to changing environment than generalists. We thus predicted a higher temporal turnover for specialists than for generalists in the presence of environmental variability (EV). In addition, we predicted that temporal turnover should decrease with increasing species richness of the communities they live in. We tested these predictions in a model system of 49 natural rock pools inhabited by 70 invertebrate species for which long-term (9 years) environmental and population dynamics data are available. We computed standard deviation of salinity measurements to represent EV for each pool. We further obtained the number of combined colonization and extinction events weighted by the number of years a species was recorded as a temporal turnover for each species in individual pools. We found that EV induced greater temporal turnover, however, the turnover depended on the species habitat traits (habitat specialization)—it has been higher in specialists but that relationship between EV and temporal turnover dissolved with increasing niche breadth (generalists). We further found that for some species, temporal turnover decreased with higher species richness and for other species, temporal turnover increased with higher species richness. The effect of species richness on temporal turnover was unrelated to species traits. This study suggests that whenever habitat is complex and heterogeneous and species pool diversified, local community dynamics becomes a composite of differential responses.     

Effects of habitat area, isolation, and landscape diversity on plant species richness of calcareous grasslands

Calcareous grasslands harbour a high biodiversity, but are highly fragmented and endangered in central Europe. We tested the relative importance of habitat area, habitat isolation, and landscape diversity for species richness of vascular plants. Plants were recorded on 31 calcareous grasslands in the vicinity of the city of Göttingen (Germany) and were divided into habitat specialist and generalist species. We expected that habitat specialists were more affected by area and isolation, and habitat generalists more by landscape diversity. In multiple regression analysis, the species richness of habitat specialists (n = 66 species) and habitat generalists (n = 242) increased with habitat area, while habitat isolation or landscape diversity did not have significant effects. Contrary to predictions, habitat specialists were not more affected by reduced habitat area than generalists. This may have been caused by delayed extinction of long-living plant specialists in small grasslands. Additionally, non-specialists may profit more from high habitat heterogeneity in large grasslands compared to habitat specialists. Although habitat isolation and landscape diversity revealed no significant effect on local plant diversity, only an average of 54% of habitat specialists of the total species pool were found within one study site. In conclusion, habitat area was important for plant species conservation, but regional variation between habitats contributed also an important 46% of total species richness.     

The effects of edge, fragment size and degree of isolation on avian species richness in highly fragmented forest in West Africa

Almost nothing is known of the effects of forest fragmentation on bird diversity within the heavily degraded and fragmented forest remnants in West Africa. We examined the effects of edge, fragment size and isolation on bird species richness in southwestern Nigeria where forest fragmentation is pronounced. In total, 122 km of line transects were used to survey birds and vegetation within 45 forest patches between January 2000 and March 2002: 197 species were recorded. Avian species number and total counts in forest patches were unrelated to fragment area (within the observed range of 14–445 ha), but were negatively influenced by degree of isolation and increasing distance from the edge. As the total area of forested land within 15 km of a patch fell from 4 to 0%, so 21% of species were lost. In total, six and zero species (of 154 recorded more than once) were consistently recorded in the larger and smaller forest fragments, respectively, and four and two bird species were consistently recorded in unisolated and isolated forest fragments, respectively, suggesting that the addition of ‘edge’ species did not compensate for loss of species sensitive to fragmentation. Diversity index was not affected by either fragment area or degree of isolation, but decreased with distance from the edge. When individual species counts were considered, 68% of species (n = 62) showed no significant effect of distance to edge. Of those 20 species which showed an effect, 12 were less common close to the edge. Most species (65%) did not respond significantly to increasing isolation but of those 22 species that did, 20 were less common in more isolated fragments. Ninety‐seven per cent of species showed no significant response to area. As avian diversity and species composition, but not species number, were apparently insensitive to forest fragmentation, our findings suggest that fragmentation reduces the probability of occurrence of a wide range of West African bird species, rather than a subset of fragmentation‐sensitive species. The greater apparent sensitivity of present‐day West African forest bird communities to fragmentation rather than patch size might reflect previous extinctions of area‐sensitive species. Minimizing further forest fragmentation might be the most effective means of conserving avian diversity in current West African landscapes where most remaining forest patches are small (i.e. < 500 ha).     

Forest patch size and isolation as drivers of bird species richness in Maputaland,Mozambique

Aim To examine the response of forest‐dependent and generalist bird assemblages and feeding guilds to patch characteristics of forest fragments in the context of island biogeography and metapopulation theories in an area that supports high levels of species diversity and endemism. Location Dune, riverine, sand and swamp forest fragments in southern Mozambique’s Maputaland. Methods We recorded 20 forest‐dependent and 69 generalist bird species at 220 survey points in 30 forest fragments that ranged in size from 5 to 7432 ha and that were 100 to 5100 m apart. We used linear regressions to relate the number of species to forest fragment size, isolation, perimeter, fractal dimension, shape and core area for frugivores, insectivores, granivores, nectarivores, carnivores and omnivores separately for forest‐dependent and generalist bird assemblages. We tested for modality in occupancy frequency distributions of species in forest fragments and used the binary matrix temperature calculator to determine whether the assemblages had a nested structure separately for forest‐dependent and generalist bird species. Results The number of forest‐dependent and generalist bird species categorized into the feeding guilds varied independently from forest patch parameter values. Occupancy models showed a random distribution for forest‐dependent birds, and generalists had a unimodal distribution. Both the forest‐dependent and generalist bird assemblages had a non‐nested structure. The presence of eight rare forest‐dependent and four endemic bird species in 22 of the 30 fragments contributed to the non‐nested structure of these bird assemblages. Main conclusions Fragmented forests did not induce the expected responses of the forest bird assemblages in that the number of species was not explained by patch characteristics, nor could we find evidence for metapopulation dynamics. Non‐nestedness may be caused by the few rare and endemic species occupying forest fragments with a wide range of patch characteristics. Knowledge of the response of species to forest fragmentation can benefit conservation planning catering for rare and endemic species. Our study suggests that all forest fragments in the region may have to be included in a conservation network to complement existing landscape‐based plans to protect the region.     

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.     

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.