Absence Of Environmental Gradients Research Articles

Stochastic distribution of small soil eukaryotes resulting from high dispersal and drift in a local environment.

A central challenge in ecology is to understand the relative importance of processes that shape diversity patterns. Compared with aboveground biota, little is known about spatial patterns and processes in soil organisms. Here we examine the spatial structure of communities of small soil eukaryotes to elucidate the underlying stochastic and deterministic processes in the absence of environmental gradients at a local scale. Specifically, we focus on the fine-scale spatial autocorrelation of prominent taxonomic and functional groups of eukaryotic microbes. We collected 123 soil samples in a nested design at distances ranging from 0.01 to 64 m from three boreal forest sites and used 454 pyrosequencing analysis of Internal Transcribed Spacer for detecting Operational Taxonomic Units of major eukaryotic groups simultaneously. Among the main taxonomic groups, we found significant but weak spatial variability only in the communities of Fungi and Rhizaria. Within Fungi, ectomycorrhizas and pathogens exhibited stronger spatial structure compared with saprotrophs and corresponded to vegetation. For the groups with significant spatial structure, autocorrelation occurred at a very fine scale (<2 m). Both dispersal limitation and environmental selection had a weak effect on communities as reflected in negative or null deviation of communities, which was also supported by multivariate analysis, that is, environment, spatial processes and their shared effects explained on average <10% of variance. Taken together, these results indicate a random distribution of soil eukaryotes with respect to space and environment in the absence of environmental gradients at the local scale, reflecting the dominant role of drift and homogenizing dispersal.

Environmental heterogeneity predicts species richness of freshwater mollusks in sub-Saharan Africa

Species diversity and how it is structured on a continental scale is influenced by stochastic, ecological, and evolutionary driving forces, but hypotheses on determining factors have been mainly examined for terrestrial and marine organisms. The extant diversity of African freshwater mollusks is in general well assessed to facilitate conservation strategies and because of the medical importance of several taxa as intermediate hosts for tropical parasites. This historical accumulation of knowledge has, however, not resulted in substantial macroecological studies on the spatial distribution of freshwater mollusks. Here, we use continental distribution data and a recently developed method of random and cohesive allocation of species distribution ranges to test the relative importance of various factors in shaping species richness of Bivalvia and Gastropoda. We show that the mid-domain effect, that is, a hump-shaped richness gradient in a geographically bounded system despite the absence of environmental gradients, plays a minor role in determining species richness of freshwater mollusks in sub-Saharan Africa. The western branch of the East African Rift System was included as dispersal barrier in richness models, but these simulation results did not fit observed diversity patterns significantly better than models where this effect was not included, which suggests that the rift has played a more complex role in generating diversity patterns. Present-day precipitation and temperature explain richness patterns better than Eemian climatic condition. Therefore, the availability of water and energy for primary productivity during the past does not influence current species richness patterns much, and observed diversity patterns appear to be in equilibrium with contemporary climate. The availability of surface waters was the best predictor of bivalve and gastropod richness. Our data indicate that habitat diversity causes the observed species–area relationship, and hence, that environmental heterogeneity is a principal driver of freshwater mollusk richness on a continental scale.

Bat diversity in the lowland forests of the Heart of Borneo

Borneo’s rainforests are renowned for their high levels of biodiversity, yet information on the distribution and structure of this diversity is lacking, particularly for less charismatic taxonomic groups. We quantified bat diversity across ten sites within a contiguous tract of largely undisturbed rainforest in the Heart of Borneo (HoB) transboundary conservation area. Using comparative analyses of 1,362 bat captures from six sites in Brunei Darussalam, together with data from four additional sites in neighbouring territories, we show that the main differences in bat assemblage composition between sites were driven by the abundances of a few cave-roosting species. Beta diversity (distance decay) was notably low and non-significant. Bat assemblage structure in these undisturbed palaeotropical forests is therefore relatively homogenous in the absence of environmental gradients. By adding 15 bat species to the Brunei national inventory, we confirm the area of north Borneo to be species-diverse and therefore a priority for conservation efforts. However, we also highlight that coastal forest to be included in a recent extension to the HoB hosts bat assemblages with the fewest species and lowest densities. We maintain that extending the HoB in Brunei to include a more diverse portfolio of habitat types is still warranted on the grounds of maximising botanical diversity and habitat area, as long as it does not detract attention from interior forests that support higher vertebrate diversity.

Rethinking edge effects: the unaccounted role of geometric constraints

Edge effects strongly affect the abundance and distribution of organisms across landscapes, with wide‐ranging implications in ecology and conservation biology. The extensive literature on the subject has traditionally considered that edge effects result from the active avoidance or preference of organisms for certain portions of the habitat patch, assuming that abundance is uniform across a patch when environmental conditions are uniform. We demonstrate that this assumption is incorrect due to the so‐far ignored ‘geometric edge effect’ (GEE). In the absence of environmental gradients, abundance of any organism living in a bounded habitat patch will tend to be lower in areas located near the edges compared to areas in the centre of the patch, simply because the areas in the centre receive individuals from all directions, whereas areas near the edge do not receive individuals from outside the patch. This geometric effect was already known for species richness at large geographic scales, the mid‐domain effect, but its importance in the literature of edge effects remained neglected so far. Using simulations, we show that the GEE tends to reduce population abundance and community richness near the edges of bounded habitat patches, and that apparently neutral or negative responses to the edge may occur even when habitat quality is higher near the edges. A published study that detected significant edge effects is reanalyzed, demonstrating that interpreting observed abundance patterns without taking the GEE into account – as traditionally done in the vast literature on edge effects – could provide misleading conclusions. The incorporation of the GEE into sampling and analytical protocols of future studies could advance substantially our ability to understand and predict edge effects in heterogeneous landscapes.

The Mid‐Domain Effect and Diversity Gradients: Is There Anything to Learn?

The mid-domain effect (MDE) has been proposed as a null model for diversity gradients and an explanation for observed patterns. Here we respond to a recent defense of the concept, explaining that it cannot represent a viable model in either real or null worlds. First, the MDE misrepresents the nature of species ranges. There is also an internal logical inconsistency underlying the MDE because the range size frequency distribution, necessary to generate a hump-shaped pattern under randomization, cannot exist in the absence of environmental gradients and is generated by the ecological and historical processes that the MDE claims to exclude.

The Mid‐Domain Effect Revisited

We revisit the proposition that boundary constraints on species' ranges cause species richness gradients (the mid-domain effect [MDE] hypothesis). In the absence of environmental gradients, species should not retain their observed range sizes as assumed by MDE models. Debate remains regarding the definition of domain limits, valid predictions for testing the models, and their statistical assessment. Empirical support for the MDE is varied but often weak, suggesting that geometric constraints on species' ranges do not provide a general explanation for richness gradients. Criticism of MDE model assumptions does not, however, imply opposition to the use of null models in ecology.

The community context of species’ borders: ecological and evolutionary perspectives

Species distributional limits may coincide with hard dispersal barriers or physiological thresholds along environmental gradients, but they may also be influenced by species interactions. We explore a number of models of interspecific interactions that lead to (sometimes abrupt) distribution limits in the presence and absence of environmental gradients. We find that gradients in competitive ability can lead to spatial segregation of competitors into distinct ranges, but that spatial movement tends to broaden the region of sympatry between the two species, and that Allee effects tend to sharpen these boundaries. We generalize these simple models to include metapopulation dynamics and other types of interactions including predator–prey and host–parasite interactions. We derive conditions for range limits in each case. We also consider models that include coevolution and gene flow and find that character displacement along environmental gradients can lead to stable parapatric distributions. We conclude that it is essential to consider coevolved species interactions as a potential mechanism limiting species distributions, particularly when barriers to dispersal are weak and environmental gradients are gradual.

Interspecific Competition, Environmental Gradients, Gene Flow, and the Coevolution of Species' Borders

Darwin viewed species range limits as chiefly determined by an interplay between the abiotic environment and interspecific interactions. Haldane argued that species' ranges could be set intraspecifically when gene flow from a species' populous center overwhelms local adaptation at the periphery. Recently, Kirkpatrick and Barton have modeled Haldane's process with a quantitative genetic model that combines density-dependent local population growth with dispersal and gene flow across a linear environmental gradient in optimum phenotype. To address Darwin's ideas, we have extended the Kirkpatrick and Barton model to include interspecific competition and the frequency-dependent selection that it generates, as well as stabilizing selection on a quantitative character. Our model includes local population growth, movements over space, natural selection, and gene flow. It simultaneously addresses the evolution of character displacement and species borders. It reproduces the Kirkpatrick and Barton single-species result that limited ranges can be produced with sufficiently steep environmental gradients and strong dispersal. Further, in the absence of environmental gradients or barriers to dispersal, interspecific competition will not limit species ranges at evolutionary equilibrium. However, interspecific competition can interact with environmental gradients and gene flow to generate limited ranges with much less extreme gradient and dispersal parameters than in the single-species case. Species display character displacement in sympatry, yet the reduction in competition that results from this displacement does not necessarily allow the two species to become sympatric everywhere. When species meet, competition reduces population densities in the region of overlap, which, in turn, intensifies the asymmetry in gene flow from center to margin. This reduces the ability of each species to adapt to local physical conditions at their range limits. If environmental gradients are monotonic but not linear, the transition zone between species at coevolutionary equilibrium occurs where the environmental gradient is steepest. If productivity gradients are also introduced into the model, then patterns similar to Rapoport's rule emerge. Interacting species respond to climate change, as it affects the optimal phenotype over space, by a combination of range shifts and local evolution in mean phenotype, while solitary species respond solely by range shifts. Finally, we compare empirical estimates for intrinsic growth rates and diffusion coefficients for several species to those needed by the single-species model to produce a stable limited range. These empirical values are generally insufficient to produce limited ranges in the model suggesting a role for interspecific interactions.

A Metapopulation Model of Species Boundaries

The majority, if not all, species have a limited geographic range bounded by a distribution edge. Violent ecotones such as sea coasts clearly produce edges for many species; however such ecotones, while sufficient for the formation of an edge, are not always necessary. We demonstrate this by simulation in discrete time of a spatially structured finite size metapopulation subjected to a spatial gradient in per-unit-time population extinction probability together with spatially structured dispersal and recolonisation. We find that relatively sharp edges separating a homeland or main geographical range from an outland or zone of relatively sparse and ephemeral colonisation can form in gradual environmental gradients. The form and placing of the edge is an emergent property of the metapopulation dynamics. The sharpness of the edge declines with increasing dispersal distance, and is dependent on the relative scales of dispersal distance and gradient length. The space over which the edge develops is short relative to the potential species range. The edge is robust against changes in both the shape of the environmental gradient and to a lesser extent to alterations in the kind of dispersal operating. Persistence times in the absence of environmental gradients are virtually independent of the shape of the dispersal function describing migration. The common finding of bell shaped population density distributions across geographic ranges may occur without the strict necessity of a niche mediated response to a spatially autocorrelated environment.

Movement and zonation of the intertidal anemone Actinia tenebrosa Farqu. (Cnidaria: Anthozoa) under experimental conditons

Actinia tenebrosa Farqu, is viviparous and the juvenile anemones are ejected from the gastrovascular cavity ready for immediate attachment to the first solid substrate they contact. It is several days after the initial attachment before the juveniles have the potential to move, and then it is at a rate approaching that of the adults. This may be at least 2.5 cm/hr for 8 hr and often more. When these animals are in the preferred zone in the intertidal region movement is much less. The movement of Actinia may be resolved into three categories: firstly, movement in the vertical plane, where prolonged submergence initiates negative geotaxis and prolonged emergence initiates positive geotaxis; secondly, slight inherent activity which continues even in the absence of environmental gradients; thirdly, directed lateral movements in response to stimuli other than submergence, emergence, or gravity. In the tidal tank, both adult and juvenile Actinia could be induced to form a zone in the intertidal region which corresponded to that formed in the natural environment. The distributions formed by adults and juveniles under identical conditions were significantly different although the range of the distributions was very similar. The high mortality of juveniles observed in the laboratory is attributed to the effects of dehydration during emergence, and it is suggested that at this stage of development Actinia is very susceptible to comparatively low relative humidities. The response of juvenile Actinia to a humidity gradient is examined. The zonation of Actinia tenebrosa is discussed with reference to coelenterate behaviour patterns and to the zonation of intertidal molluscs.