Variations on a theme: sources of heterogeneity in the form of the interspecific relationship between abundance and distribution

1. A positive interspecific relationship between abundance and distribution is widely considered to be one of the most general patterns in ecology. However, the relationship appears to vary considerably across assemblages, from significant positive to significant negative correlations and all shades in between. 2. This variation has led to the suggestion that the abundance-distribution relationship has multiple forms, with the corollary that different patterns may inform about, or have different, causes. However, this variation has never been formally quantified, nor has it been determined whether the observed variation is indicative of sampling error in estimating a single effect or of real heterogeneity in such relationships. Here, we use the meta-analytical approach to assess variation in abundance-distribution relationships, and to test different hypotheses for it. 3. Analysis of 279 relationships found a mean effect size of 0.655, which was both highly significantly different from zero and indicative of a strong positive association between abundance and distribution. However, effect sizes were highly heterogeneous, supporting the contention that this relationship does indeed have multiple forms. 4. Most notably, relationships vary significantly in strength across realms, with the strongest in the marine and intertidal, intermediate relationships for terrestrial and parasitic assemblages, and the weakest relationships in freshwater systems. Effect sizes in all of the aquatic realms are homogeneous, suggesting that realm is an important source of the heterogeneity observed across all studies. We posit that this may be because the different spatial structure of the environment in each realm affects the opportunity for the dispersal of individuals between sites. 5. Some of the remaining heterogeneity in effect sizes for terrestrial assemblages could be explained by partitioning assemblages by habitat, scale, biogeographical region and taxon, but considerable heterogeneity in effect sizes for terrestrial and parasitic assemblages remained unexplained.     

Climatic niche conservatism and the evolutionary dynamics in species range boundaries: global congruence across mammals and amphibians

Aim Comparative evidence for phylogenetic niche conservatism – the tendency for lineages to retain their ancestral niches over long time scales – has so far been mixed, depending on spatial and taxonomic scale. We quantify and compare conservatism in the climatic factors defining range boundaries in extant continental mammals and amphibians in order to identify those factors that are most evolutionarily conserved, and thus hypothesized to have played a major role in determining the geographic distributions of many species. We also test whether amphibians show stronger signals of climatic niche conservatism, as expected from their greater physiological sensitivity and lower dispersal abilities. Location Global; continental land masses excluding Antarctica. Methods We used nearly complete global distributional databases to estimate the climatic niche conservatism in extant continental mammals and amphibians. We characterized the climatic niche of each species by using a suite of variables and separately investigate conservatism in each variable using both taxonomic and phylogenetic approaches. Finally, we explored the spatial, taxonomic and phylogenetic patterns in recent climatic niche evolution. Results Amphibians and mammals showed congruent patterns of conservatism in cold tolerance, with assemblages of escapee species (i.e. those escaping most from the climatic constraints of their ancestors) aggregated in the North Temperate Zone. Main conclusions The relative strength of climatic niche conservatism varies across the variables tested, but is strongest for cold tolerance in both mammals and amphibians. Despite the apparent conservatism in this variable, there is also a strong signal of recent evolutionary shifts in cold tolerance in assemblages inhabiting the North Temperate Zone. Our results thus indicate that distribution patterns of both taxa are influenced by both niche conservatism and niche evolution.     

The relationship between range size and niche breadth: a test using five species of Gammarus (Amphipoda)

Data for five closely related species of gammarid crustaceans are used to examine interspecific relationships between the breadth of fundamental tolerance or capacity and geographical range size. Gammarus duebeni is, almost without exception, the most tolerant species and that with the best physiological performance. Although there is some limited variation, the remaining species can be ranked broadly in the sequence G. zaddachi  > G. salinus  >  G. oceanicus > G. locusta . The wide tolerance and high performance of G. duebeni is associated with the occupation of a wider range of environmental 'types' than any other of the species. In terms of geographical range size, the species can be ranked from most to least widespread in the sequence G. oceanicus  > G. duebeni  >  G. zaddachi  >  G. salinus  >  G. locusta . This provides little support for Brown's hypothesis, or the argument that the more widely distributed species within a taxonomic assemblage also tend to have the widest fundamental niches. However, if marine ( G. oceanicus and G. locusta ) and estuarine ( G. duebeni , G. zaddachi , G. salinus) species are considered separately, then in each case the species with the largest geographical range is also the most tolerant/best performer. In this sense, the jack-of-all-trades is the master-of-all, rather than the master-of-none.     

The influence of spatial resolution on macroecological patterns of range size variation: a case study using parrots (Aves: Psittaciformes) of the world

Aim To assess the extent to which the resolution at which geographical range sizes are measured influences macroecological patterns in this variable. Location Global. Methods Data on the geographical ranges of parrot species were digitized, and a Geographic Information System used to produce nine range size estimates for each species using different degrees of spatial resolution. The inter‐correlation of these estimates was then compared, together with their patterns of covariation with population size, body mass and migratory behaviour (across species and controlling for phylogeny), their pattern of phylogenetic correlation, and the frequency distributions of the different measures. Results Strong correlations exist among all nine range size measures across species, albeit that measures of similar spatial resolution are more strongly correlated. All measures show similar patterns of covariation with population size, body mass and migratory behaviour, and similar patterns of phylogenetic correlation. The skewness of frequency distributions increases towards zero as the resolution of the range size measure declines. Main conclusions The results of macroecological analyses are little affected by the resolution with which geographical range sizes are calculated, at least for the parrots of the world. Previously published studies based on crude measures of range size would be unlikely to have produced markedly different conclusions had they used more refined range size metrics.     

Life-history traits are correlated with geographical distribution patterns of western European forest herb species

Aim To investigate whether six plant life‐history traits that have been related to colonization ability at local scales are also related to the geographical range characteristics of 273 forest plant species. Location Continental western Europe, five countries in particular: France, Luxemburg, Belgium, the Netherlands and Germany. The region is situated between 42° and 55°N and 5°W and 15°E and has a summed total area of 971,404 km². Methods Distribution data were compiled from five national data bases and converted to a 10′ grid. Life‐history traits were taken from existing compilations of autecological information of European species. The spatial arrangement of occupied grid cells was investigated using Ripley's K. Cross‐species correlations and phylogenetically independent contrasts were used to investigate the relationships between plant life‐history traits and three range characteristics: area of occupancy, latitudinal extent and centroid latitude. Results For herbaceous species, seed dispersal mode, seed production and seed bank longevity exhibited significant associations with geographical range characteristics, including area of occupancy. Woody plant species exhibited fewer significant associations, although maximum height was positively associated with range centroid latitude within the study area. Furthermore, the ranges of species with limited dispersal ability were found to be more clustered than the ranges of species with morphological adaptations for long‐distance seed dispersal. Main conclusions For western European forest plant species, life‐history traits that are related to colonization ability at local scales are associated with variation in large‐scale geographical range characteristics. This finding implies that the distributions of some forest plant species in the study area may be limited by seed dispersal and colonization capacity rather than climate or other environmental factors.     

Relationships between local population persistence, local abundance and regional occupancy of species: distribution patterns of diatoms in boreal streams

Aims We have two aims: (1) to examine the relationship between local population persistence, local abundance and regional occupancy of stream diatoms and (2) to characterize the form of the species–occupancy frequency distribution of stream diatoms. Location Boreal streams in Finland. There were three spatial extents: (1) across ecoregions in Finland, (2) within ecoregions in Finland, and (3) within a single drainage system in southern Finland. Methods Diatoms were sampled from stones (epilithon), sediment (epipelon) and aquatic plants (epiphyton) in streams using standardized sampling methods. To assess population persistence, diatom sampling was conducted monthly at four stream sites from June to October. The relationships between local population persistence, local abundance and regional occupancy were examined using correlation analyses. Results There was a significant positive relationship between local persistence and abundance of diatoms in epilithon, epipelon and epiphyton. Furthermore, local abundance and regional occupancy showed a significant positive relationship at multiple spatial extents; that is, across ecoregions, within ecoregions and within a drainage system. The relationships between occupancy and abundance did not differ appreciably among impacted and near pristine‐reference sites. The occupancy–frequency distribution was characterized by a large number of satellite species which occurred at only a few sites, whereas core species that occurred at most sites were virtually absent. Main conclusions The positive relationship between local population persistence and abundance suggested that a high local abundance may prevent local extinction or that high persistence is facilitated by a high local cell density. High local persistence and local abundance may also positively affect the degree of regional occupancy in stream diatoms. The results further showed that anthropogenic effects were probably too weak to bias the relationship between occupancy and abundance, or that the effects have already modified the distribution patterns of stream diatoms. The small number of core species in the species–occupancy frequency distribution suggested that the regional distribution patterns of stream diatoms, or perhaps unicellular microbial organisms in general, may not be fundamentally different from those described previously for multicellular organisms, mainly in terrestrial environments, although average global range sizes may differ sharply between these two broad groups of organisms.     

The relationship between dispersal ability and geographic range size

There are a variety of proposed evolutionary and ecological explanations for why some species have more extensive geographical ranges than others. One of the most common explanations is variation in species' dispersal ability. However, the purported relationship between dispersal distance and range size has been subjected to few theoretical investigations, and empirical tests reach conflicting conclusions. We attempt to reconcile the equivocal results of previous studies by reviewing and synthesizing quantitative dispersal data, examining the relationship between average dispersal ability and range size for different spatial scales, regions and taxonomic groups. We use extensive data from marine taxa whose average dispersal varies by seven orders of magnitude. Our results suggest dispersal is not a general determinant of range size, but can play an important role in some circumstances. We also review the mechanistic theories proposed to explain a positive relationship between range size and dispersal and explore their underlying rationales and supporting or refuting evidence. Despite numerous studies assuming a priori that dispersal influences range size, this is the first comprehensive conceptual evaluation of these ideas. Overall, our results indicate that although dispersal can be an important process moderating species' distributions, increased attention should be paid to other processes responsible for range size variation.     

Relationships between geographical range size, body size, local abundance, and habitat breadth in North American suckers and sunfishes

Aim I examine the relationship between geographical range size and three variables (body size, an index of habitat breadth, and an index of local abundance) within a phylogenetic framework in North American species of suckers and sunfishes. Location North America Methods Regressions after independent contrasts of geographical range size, body size, habitat breadth, and local abundance. Results Species with large range sizes tend to be larger-bodied, be more locally abundant, and have higher habitat breadths. Character reconstructions support the prediction that variables associated with rarity (small geographical range size, low local abundance, low niche breadth, and large body size) evolve in unison, although large body size was associated with the opposite traits in these taxa. Gaston & Blackburn (1996a) suggested using visual identification of the lower boundary of the geographical range-body size relationship to identify extinction-prone species; this resulted in thirteen species that are potentially extinction-prone. Main conclusions Similar evolutionary mechanisms appear to operate on body size and other variables related to rarity, even in distantly related taxa.     

Niche breadth, competitive strength and range size of tree species: a trade-off based framework to understand species distribution

Understanding the mechanisms causing latitudinal gradients in species richness and species range size is a central issue in ecology, particularly in the current context of global climate change. Different hypotheses have been put forward to explain these patterns, emphasizing climatic variability, energy availability and competition. Here we show, using a comparative analysis controlling for phylogeny on 234 temperate/boreal tree species, that these hypotheses can be included into a single framework in an attempt to explain latitudinal gradients in species range size. We find that species tend to have larger ranges when (i) closer to the poles, (ii) successionally seral, (iii) having small and light seeds, and (iv) having short generations. The patterns can simply be explained by energy constraints associated with different life-history strategies. Overall, these findings shed a new light on our understanding of species distribution and biodiversity patterns, bringing new insights into underlying large-scale evolutionary processes.     

净多样化速率和进化时间对虎耳草目科间物种多样性差异的影响

不同生物类群包含的物种数目常存在巨大差异,这是生态学和生物学研究中普遍观察到的现象。然而,这一现象产生的原因仍然是未解之谜。从宏观进化的角度,进化时间假说和多样化速率假说是两个比较流行的假说。进化时间假说认为类群的演化时间越长,积累的物种丰富度越高;而多样化速率假说认为类群的净多样化速率越快,则其物种丰富度越高。为验证这两个假说,该文以一棵包含1 539个物种化石定年的虎耳草目系统发育树为基础,通过宏观进化分析获取了虎耳草目内15个科的物种形成和灭绝速率,并计算了每个科的平均多样化速率。结果表明：(1)虎耳草目的物种多样化速率有着增加的趋势,并且多样化速率的增加主要出现在温带和高山类群,如茶藨子科、景天科和芍药科等。(2)采用系统发育广义最小二乘模型(PGLS)和线性回归模型(LM)结果表明,虎耳草目15个科的物种丰富度与科的分化时间和科内物种的最近共同祖先年龄都没有显著相关关系,而与净多样化速率显著正相关(R² =0.380,P<0.05)。该研究支持了多样化速率假说,认为不同科的净多样化速率的差异是导致虎耳草目科间物种数目差异的主要原因之一。全球气候变冷...     

The relationship between local and regional diatom richness is mediated by the local and regional environment

Aim In this continental study, species richness at local (LSR) and regional (RSR) scales was correlated and examined as a function of stream (local) and watershed (regional) environment in an effort to elucidate what factors control diatom biodiversity across scales. Location Conterminous United States. Methods Data on diatom richness, stream conditions and watershed properties were generated by the US Geological Survey. In the present investigation, RSR was estimated as the total diatom richness in a hydrologic study unit and, together with stream and watershed characteristics, was included in stepwise multiple regressions of LSR. The unique and shared contributions of RSR, stream and watershed environment to the explained variance in LSR were determined by variance partitioning. RSR was regressed against stream and basin features averaged per study unit. Results LSR responded most strongly to variability in stream manganese concentration and RSR. Other predictors included stream discharge and iron concentration, soil organic matter content and fertilization, and proportions of open water, barren land and forest in the watershed. Variance partitioning revealed that RSR had the lowest independent contribution to explained variance in LSR. Multiple regressions identified average stream iron concentration as the most important predictor of RSR. Main conclusions Local micronutrient concentration was the major predictor of LSR, followed by RSR. Since average micronutrient supply in the region was the chief determinant of RSR, it is proposed that micronutrients had both a direct effect on LSR and an indirect effect through RSR. The same argument is extended to watershed features with an impact on stream trophic status, because of their substantial contributions to the explained variance in both LSR and RSR. Considering that the major proportion of LSR variance explained by RSR originated from the covariance of RSR with stream and watershed properties, it is concluded that the LSR–RSR relationship was mediated by the local and regional environment.     

Local-regional relationships and the geographical distribution of species

Aim Local‐regional (LR) species diversity plots were conceived to assess the contribution of regional and local processes in shaping the patterns of biological diversity, but have been used also to explore the scaling of diversity in terms of its alpha, beta, and gamma components. Here we explore the idea that patterns in the geographical ranges of species over a continent can determine the shape of small region to large region (SRLR) plots, which are equivalent to LR plots when comparing the diversity of sites at two regional scales. Location To test that idea, we analysed the diversity patterns at two regional scales for the mammals of North America, defined as the mainland from Alaska and Canada to Panama. Method We developed a theoretical model relating average range size of species over a large‐scale region with its average regional point species diversity (RPD). Then, we generated a null model of expected SRLR plots based on theoretical predictions. Species diversities at two scales were modelled using linear and saturation functions for Type I and Type II SRLR relationships, respectively. We applied the models to the case of North American mammals by examining the regional diversity and the RPD for 21 large‐scale quadrats (with area equal to 160,000 km²), arranged along a latitudinal gradient. Results Our model showed that continental and large‐scale regional patterns of distribution of species can generate both types of SRLR relationship, and that these patterns can be reflected in LR plots without invoking any kind of local processes. We found that North American nonvolant mammals follow a Type I SRLR relationship, whereas bats follow a Type II pattern. This difference was linked to patterns in which species of the two mammalian groups distribute in geographical space. Conclusion Traditional LR plots and the new SRLR plots are useful tools in exploring the scaling of species diversity and in showing the relationship between distribution and diversity. Their usefulness in comparing the relative role of local and regional processes is, however, very limited.     

The anatomy of the interspecific abundance–range size relationship for the British avifauna: I. Spatial patterns

Data from the British Trust for Ornithology Common Birds Census and two atlases of breeding birds were used to examine the form of the interspecific abundance–range size relationship for the British avifauna. The relationship is positive for both farmland and woodland habitats and over two different periods, with some evidence of curvilinearity, using either proportion of occupied sites or numbers of occupied 10 × 10 km squares as measures of range size, and mean density at occupied sites as a measure of abundance. A log-linear plot gives the highest correlation. The relationship is stronger if based on maximum local densities than if based on average densities, but there is no relationship using minimum local densities. Relationships based on abundances at individual sites are uniformly positive for all sites, although the relationships for many sites also show evidence of curvilinearity, especially when range size is measured as the proportion of occupied sites. Species show significant concordance in their rank abundances across sites. We discuss some implications of these results.