Does climate limit species richness by limiting individual species’ ranges?

Broad-scale geographical variation in species richness is strongly correlated with climate, yet the mechanisms underlying this correlation are still unclear. We test two broad classes of hypotheses to explain this pattern. Bottom-up hypotheses propose that the environment determines individual species’ ranges. Ranges then sum up to yield species richness patterns. Top-down hypotheses propose that the environment limits the number of species that occur in a region, but not which ones. We test these two classes of hypotheses using a natural experiment: seasonal changes in environmental variables and seasonal range shifts of 625 migratory birds in the Americas. We show that richness seasonally tracks the environment. By contrast, individual species’ geographical distributions do not. Rather, species occupy different sets of environmental conditions in two seasons. Our results are inconsistent with extant bottom-up hypotheses. Instead, a top-down mechanism appears to constrain the number of species that can occur in a given region.     

Spatio‐temporal scaling of biodiversity and the species–time relationship in a stream fish assemblage

1. The increase of species richness with the area of the habitat sampled, that is the species–area relationship, and its temporal analogue, the species–time relationship (STR), are among the few general laws in ecology with strong conservation implications. However, these two scale‐dependent phenomena have rarely been considered together in biodiversity assessment, especially in freshwater systems. 2. We examined how the spatial scale of sampling influences STRs for a Central‐European stream fish assemblage (second‐order Bernecei stream, Hungary) using field survey data in two simulation‐based experiments. 3. In experiment one, we examined how increasing the number of channel units, such as riffles and pools (13 altogether), and the number of field surveys involved in the analyses (12 sampling occasions during 3 years), influence species richness. Complete nested curves were constructed to quantify how many species one observes in the community on average for a given number of sampling occasions at a given spatial scale. 4. In experiment two, we examined STRs for the Bernecei fish assemblage from a landscape perspective. Here, we evaluated a 10‐year reach level data set (2000–09) for the Bernecei stream and its recipient watercourse (third‐order Kemence stream) to complement results on experiment one and to explore the mechanisms behind the observed patterns in more detail. 5. Experiment one indicated the strong influence of the spatial scale of sampling on the accumulation of species richness, although time clearly had an additional effect. The simulation methodology advocated here helped to estimate the number of species in a diverse combination of spatial and temporal scale and, therefore, to determine how different scale combinations influence sampling sufficiency. 6. Experiment two revealed differences in STRs between the upstream (Bernecei) and downstream (Kemence) sites, with steeper curves for the downstream site. Equations of STR curves were within the range observed in other studies, predominantly from terrestrial systems. Assemblage composition data suggested that extinction–colonisation dynamics of rare, non‐resident (i.e. satellite) species influenced patterns in STRs. 7. Our results highlight that the determination of species richness can benefit from the joint consideration of spatial and temporal scales in biodiversity inventory surveys. Additionally, we reveal how our randomisation‐based methodology may help to quantify the scale dependency of diversity components (α, β, γ) in both space and time, which have critical importance in the applied context.     

Ecological niche models display nonlinear relationships with abundance and demographic performance across the latitudinal distribution of Astragalus utahensis (Fabaceae)

The potential for ecological niche models (ENMs) to accurately predict species' abundance and demographic performance throughout their geographic distributions remains a topic of substantial debate in ecology and biogeography. Few studies simultaneously examine the relationship between ENM predictions of environmental suitability and both a species' abundance and its demographic performance, particularly across its entire geographic distribution. Yet, studies of this type are essential for understanding the extent to which ENMs are a viable tool for identifying areas that may promote high abundance or performance of a species or how species might respond to future climate conditions. In this study, we used an ensemble ecological niche model to predict climatic suitability for the perennial forb Astragalus utahensis across its geographic distribution. We then examined relationships between projected climatic suitability and field‐based measures of abundance, demographic performance, and forecasted stochastic population growth (λ_s). Predicted climatic suitability showed a J‐shaped relationship with A. utahensis abundance, where low‐abundance populations were associated with low‐to‐intermediate suitability scores and abundance increased sharply in areas of high predicted climatic suitability. A similar relationship existed between climatic suitability and λ_s from the center to the northern edge of the latitudinal distribution. Patterns such as these, where density or demographic performance only increases appreciably beyond some threshold of climatic suitability, support the contention that ENM‐predicted climatic suitability does not necessarily represent a reliable predictor of abundance or performance across large geographic regions.     

Biogeographic anomalies in the species richness of Chilean forests: Incorporating evolution into a climatic – historic scenario

Broad‐scale richness gradients are closely associated with temperature and water availability. However, historical and evolutionary processes have also contributed to shape current diversity patterns. In this paper we focus on the potential influences of Pleistocene glaciation and phylogenetic niche conservatism (the tendency for traits to be maintained during diversification) on the tree diversity gradient in Chile, and we quantify its primary climatic correlates. Tree species richness is greatest at mid latitudes, particularly in the Andes and Coastal ranges, and decreases abruptly to the south and north. Regression tree analysis identified annual precipitation and annual temperature as the primary probable drivers of this gradient. Ice cover during the Last Glacial Maximum was also identified as an ‘important’ variable, but the contemporary and historical predictors are strongly collinear. Geographically weighted regression indicated that the relationships between richness and environmental variables vary regionally: the relationship between tree richness and precipitation is stronger in north‐central Chile, whereas tree richness and temperature are most strongly associated in south‐central Chile. By assigning each species the age of the family to which it belongs and averaging all species in each geographical unit, we also found that species from the oldest families are distributed mainly in mid to high latitudes and species from younger families are distributed mainly at lower latitudes. This pattern is closely associated with annual precipitation. Thus, the ecological component of tree richness follows contemporary climatic gradients of both energy and water, but the aridification of the Atacama Desert was an important driver over evolutionary time. The influence of recent Pleistocene glaciation remains unresolved but it cannot be discounted.     

Global angiosperm family richness revisited: linking ecology and evolution to climate

Aim The global richness gradient of angiosperm families is correlated with current climate, and it has been claimed that historical processes are not necessary to understand patterns of plant family richness. This claim has drawn criticism, and there have been doubts about the quality of the data used to quantify the pattern. We revisit this issue using the Angiosperm Phylogeny Group (APG) III classification and revised range maps, and we incorporate an evolutionary variable, family age, to explore covariation between evolution and ecology and their links to climate via the tropical conservatism hypothesis (TCH). Location Global. Methods The richness pattern for 408 families was derived from range maps, and family ages were derived from a dated angiosperm phylogeny. Patterns were generated for all families, 143 families composed of trees, and 149 families composed of herbs. We also examined family range size patterns to test the extent to which extratropical floras are nested subsets of tropical floras. Ordinary least squares (OLS) multiple and partial regressions were used to generate climate models for richness, mean range size and mean age for each plant dataset and to evaluate the covariation between contemporary climate and clade age as correlates of family richness. Results We confirmed the strong association between contemporary climate and family richness. Age patterns predicted by TCH were also found for families comprising trees. The richness of herbaceous families, in contrast, was correlated with climate but the age pattern was not as predicted by TCH. Floras in cold and dry areas are strongly nested within richer tropical floras. Main conclusions Phylogenetic niche conservatism at the family level offers a likely explanation for the global diversity gradient of trees, but not for non‐desert herbs, probably because of the faster evolutionary rates for herbs and less constrained evolutionary responses to climate change. Thus, it appears that multiple processes account for the overall angiosperm family gradient. Our analysis also demonstrates that even very strong associations of taxon richness and climate do not preclude evolutionary processes, as has been widely argued, and that climatic and evolutionary hypotheses for richness gradients are not mutually exclusive.     

Colonization and extinction of ant communities in a fragmented landscape

Abstract In this paper we tested the assumption that smaller and more isolated remnants receive fewer ant colonizers and lose more species. We also tested hypotheses to explain such a pattern. We sampled ants in Brazil for 3 years in 18 forest remnants and in 10 grasslands between them. We tested the influence of remnant area and isolation on colonization rate, as well as the effect of remnant area on extinction rate. We tested the correlation between remnant area and isolation to verify the landscape design. Colonization rate was not affected by remnant area or isolation. Extinction rate, however, was smaller in larger remnants. Remnant area and isolation were negatively correlated. We tested two hypotheses related to the decrease in ant species extinction rate with increased remnant area: (i) small remnants support smaller and more extinction‐prone populations; and (ii) small remnants are more often invaded by generalist species, which suffer higher extinction inside remnants. The density of ant populations significantly increased with area. Generalist species presented a lower colonization rate in larger remnants, contrary to the pattern observed in forest species. Generalist species suffered more extinction than expected inside remnants. The lack of response of colonization rate to remnant area can be explained by the differential colonization by generalist and forest species. The decrease of ant population density in smaller remnants could be related to loss of habitat quality or quantity. The higher colonization by generalist ant species in the smaller remnants could be related to landscape design, because smaller remnants are more similar to the matrix than larger ones. Our results have important implications for conservation strategies because small remnants seem to be more affected by secondary effects of fragmentation, losing more forest species and being invaded more often by generalist species. Studies that compare only species richness between remnants cannot detect such patterns in species composition.     

Can stochastic geographical evolution re‐create macroecological richness–environment correlations?

Aim Richness gradients are frequently correlated with environmental characteristics at broad geographic scales. In particular, richness is often associated with energy and climate, while environmental heterogeneity is rarely its best correlate. These correlations have been interpreted as evidence in favour of environmental determinants of diversity gradients, particularly energy and climate. This interpretation assumes that the expected‐by‐random correlation between richness and environment is zero, and that this is equally true for all environmental characteristics. However, these expectations might be unrealistic. We investigated to what degree basic evolutionary/biogeographical processes occurring independently of environment could lead to richness gradients that correlate with environmental characteristics by chance alone. Location Africa, Australia, Eurasia and the New World. Methods We produced artificial richness gradients based on a stochastic simulation model of geographic diversification of clades. In these simulations, species speciate, go extinct and expand or shift their distributions independently of any environmental characteristic. One thousand two hundred repetitions of this model were run, and the resulting stochastic richness gradients were regressed against real‐world environmental variables. Stochastic species–environment relationships were then compared among continents and among three environmental characteristics: energy, environmental heterogeneity and climate seasonality. Results Simulations suggested that a significant degree of correlation between richness gradients and environment is expected even when clades diversify and species distribute stochastically. These correlations vary considerably in strength; but in the best cases, environment can spuriously account for almost 80% of variation in stochastic richness. Additionally, expected‐by‐chance relationships were different among continents and environmental characteristics, producing stronger spurious relationships with energy and climate than with heterogeneity. Main conclusions We conclude that some features of empirical species–environment relationships can be reproduced just by chance when taking into account evolutionary/biogeographical processes underlying the construction of species richness gradients. Future tests of environmental effects on richness should consider structure in richness–environment correlations that can be produced by simple evolutionary null models. Research should move away from the naive non‐biological null hypotheses that are implicit in traditional statistical tests.     

Diversity and distribution of small mammals in the South American Dry Andes

The Andean mountain range has played an important role in the evolution of South American biota. However, there is little understanding of the patterns of species diversity across latitudinal and altitudinal gradients. In this paper, we examine the diversity of small mammals along the South Central Dry Andes (SCDA) within the framework of two contrasting hypotheses: (a) species richness decreases with increasing elevation and latitude; and (b) species richness peaks at altitudinal midpoints (mid‐domain). We explore the composition of the species pool, the impact of species–area relationships and the Rapoport effect (i.e. size of geographic ranges) along latitudinal and elevational gradients. First, we constructed a database of SCDA small mammals. Then, species richness patterns were analysed through generalized models, and species–area relationships were assessed by log–log regressions; the curvilinear method (c = S/Az) was use to compute richness corrected by area size. Lastly, the Rapoport effect was evaluated using the midpoint method. Our results show: (1) a richness of 67 small mammals along the SCDA, of which 36 are endemic; (2) a hump‐shaped pattern in species richness along elevation and latitudinal gradients; (3) a species–area relationship for both gradients; (4) endemic species corrected by area present a strong and positive relationship with elevation; (5) a Rapoport effect for the latitudinal ranges, but no effect across the elevational gradient; and (6) a major species turnover between 28° and 30° south latitude. This is the first study quantifying the diversity of small mammals encompassing the central Andean region. Overall, our macrogeographic analysis supports the previously postulated role of the Andes in the diversification of small mammals (i.e. in situ cladogenesis) and highlights some basic attributes (i.e. anatomy of geographic ranges; species–area relationships) when considering the consequences of climate change on biodiversity conservation of mountain ecosystems.     

Moderate grazing promotes the root biomass in Kobresia meadow on the northern Qinghai–Tibet Plateau

Grazing is an important modulator of both plant productivity and biodiversity in grassland community, yet how to determine a suitable grazing intensity in alpine grassland is still controversy. Here, we explore the effects of different grazing intensities on plant biomass and species composition, both at community level and functional group level, and examines the productivity–species richness relationship under four grazing patterns: no grazing (CK), light grazing (LG), moderate grazing, (MG) and heavy grazing (HG), attempt to determine a suitable grazing intensity in alpine grassland. The results were as follows. The total aboveground biomass (AGB) reduced with increasing grazing intensity, and the response of plant functional groups was different. AGB of both sedges and legumes increased from MG to HG, while the AGB of forbs reduced sharply and the grass AGB remained steady. There was a significant positive relationship between productivity and species richness both at community level and functional group level. In contrast, the belowground biomass (BGB) showed a unimodal relationship from CK to HG, peaking in MG (8,297.72 ± 621.29 g/m²). Interestingly, the grassland community tends to allocate more root biomass to the upper soil layer under increasing grazing intensities. Our results suggesting that moderate levels of disturbance may be the optimal grassland management strategy for alpine meadow in terms of root production.     

Allometric ecological distributions in a local community of Hymenoptera

The present paper describes basic ecological distributions in a community of beech forest Hymenoptera. It shows that the species diversity–body weight and the density–body weight distributions give rise to a new distribution that relates total community biomass to species diversity. For Hymenoptera this distribution follows a power function with a slope of 1.3. Combining this relation with the species–area and the individuals–area relations resulted in two other distributions that relate community biomass to area and individual numbers. It appeared that population densities decrease when computed over larger areas. The biomass–species diversity relation offers a new and simple way to estimate total community biomass from samples. The possible implications of this distribution to the productivity–diversity debate are discussed.     

sars: an R package for fitting,evaluating and comparing species–area relationship models

The species–area relationship (SAR) constitutes one of the most general ecological patterns globally. A number of different SAR models have been proposed. Recent work has shown that no single model universally provides the best fit to empirical SAR datasets: multiple models may be of practical and theoretical interest. However, there are no software packages available that a) allow users to fit the full range of published SAR models, or b) provide functions to undertake a range of additional SAR‐related analyses. To address these needs, we have developed the R package ‘sars’ that provides a wide variety of SAR‐related functionality. The package provides functions to: a) fit 20 SAR models using non‐linear and linear regression, b) calculate multi‐model averaged curves using various information criteria, and c) generate confidence intervals using bootstrapping. Plotting functions allow users to depict and scrutinize the fits of individual models and multi‐model averaged curves. The package also provides additional SAR functionality, including functions to fit, plot and evaluate the random placement model using a species–sites abundance matrix, and to fit the general dynamic model of oceanic island biogeography. The ‘sars’ R package will aid future SAR research by providing a comprehensive set of simple to use tools that enable in‐depth exploration of SARs and SAR‐related patterns. The package has been designed to allow other researchers to add new functions and models in the future and thus the package represents a resource for future SAR work that can be built on and expanded by workers in the field.     

Why some parasites are widespread and abundant while others are local and rare?

Abundances and distributions of species are usually associated. This implies that as a species declines in abundance so does the number of sites it occupies. Conversely, when there is an increase in a species' range size, it is usually followed by an increase in population size (Gaston et al. 2000 ). This ecological phenomenon, also known as the abundance–occupancy relationship (AOR), is well documented in several species of animals and plants (Gaston et al. 2000 ) but has been little investigated in parasites. In this issue of Molecular Ecology, Drovetski et al. ( 2014 ) investigated the AOR in avian haemosporidians (vector‐borne blood parasites) using data from four well‐sampled bird communities. In support of the AOR, the research group found that the abundance of parasite cytochrome b lineages (a commonly used proxy for species identification within this group of parasites) was positively linked with the abundance of susceptible avian host species and that the most abundant haemospordian lineages were those with larger ranges. Drovetski et al. ( 2014 ) also found evidence for both hypotheses proposed to explain the AOR in parasites: the trade‐off hypothesis (TOH) and the niche‐breadth hypothesis (NBH). Interestingly, the main predictor of the AOR was the number of susceptible hosts (i.e. number of infected birds) and not the number of host species the parasites were able to exploit.     

Island biogeography is not a single‐variable discipline: the small island effect debate

In some island systems, an ‘anomalous’ feature of species richness on smaller islands, in comparison with larger ones, has been observed. This has been described as the small island effect (SIE). The precise meaning of the term remains unresolved, as does the explanation for the phenomenon and even whether it exists. Dengler (2010 ; Diversity Distrib, 16 , 256–266.) addresses a number of conceptual and methodological issues concerning the nature and the detection of the SIE but fails to settle conclusively most of the issues he raises. We contend that his approach is theoretically flawed, especially in its treatment of habitat diversity. We offer a few suggestions of what is needed to advance understanding of the SIE.     

The theory of the nested species–area relationship: geometric foundations of biodiversity scaling

The relationship between sampled area and the number of species within that area, the species–area relationship (SAR), is a major biodiversity pattern and one of a few law‐like regularities in ecology. While the SAR for isolated units (islands or continents) is assumed to result from the dynamics of species colonization, speciation and extinction, the SAR for contiguous areas in which smaller plots are nested within larger sample areas can be attributed to spatial patterns in the distribution of individuals. The nested SAR is typically triphasic in logarithmic space, so that it increases steeply at smaller scales, decelerates at intermediate scales and increases steeply again at continental scales. I will review current theory for this pattern, showing that all three phases of the SAR can be derived from simple geometric considerations. The increase of species richness with area in logarithmic space is generally determined by overall species rarity, so that the rarer the species are on average, the higher is the local slope z. Rarity is scale‐dependent: species occupy only a minor proportion of area at broad spatial scales, leading to upward accelerating shape of the SAR at continental scales. Similarly, species are represented by only a few individuals at fine spatial scales, leading to high SAR slope also at small areas. Geometric considerations reveal links of the SAR to other macroecological patterns, namely patterns of β‐diversity, the species–abundance distribution, and the relationship between energy availability (or productivity) and species richness. Knowledge of the regularities concerning nested SARs may be used for standardizing unequal areas, upscaling species richness and estimating species loss due to area loss, but all these applications have their limits, which also follow from the geometric considerations.     

Does the niche breadth or trade‐off hypothesis explain the abundance–occupancy relationship in avian Haemosporidia?

Two hypotheses have been proposed to explain the abundance–occupancy relationship (AOR) in parasites. The niche breadth hypothesis suggests that host generalists are more abundant and efficient at colonizing different host communities than specialists. The trade‐off hypothesis argues that host specialists achieve high density across their hosts' ranges, whereas generalists incur the high cost of adaptation to diverse immuno‐defence systems. We tested these hypotheses using 386 haemosporidian cytochrome‐b lineages (1894 sequences) recovered from 2318 birds of 103 species sampled in NW Africa, NW Iberia, W Greater Caucasus and Transcaucasia. The number of regions occupied by lineages was associated with their frequency suggesting the presence of AOR in avian Haemosporidia. However, neither hypothesis provided a better explanation for the AOR. Although the host generalist Plasmodium SGS1 was over three times more abundant than other widespread lineages, both host specialists and generalists were successful in colonizing all study regions and achieved high overall prevalence.