The distribution–abundance (density) relationship: its form and causes in a tropical mammal order, Primates

Aim Across a wide variety of organisms, taxa with high local densities (abundance) have large geographical ranges (distributions). We use primatology's detailed knowledge of its taxon to investigate the form and causes of the relationship in, unusually for macroecological analysis, a tropical taxon. Location Africa, Central and South America, Asia, Madagascar. Methods To investigate the form of the density–range relationship, we regressed local density on geographical range size, and also on female body mass, because in the Primates, density correlates strongly with mass. To investigate the biological causes of the relationship, we related (1) abundance (density × range size) and (2) residuals from the density–range regression lines to various measures of (i) resource use, (ii) reproductive rate and (iii) potential specialization. All data are from the literature. Analyses were done at the level of species (n = 140), genera (n = 60) and families/subfamilies (n = 17). We present various levels of results, including for all data, after omission of outlier data, after correction for phylogenetic dependence, and after Bonferroni correction of probabilities for multiple comparisons. Results Regarding the form of the relationship, Madagascar primates are clear outliers (high densities in small ranges). Among the remaining three realms, the relation of density to range is weak or non‐existent at the level of species and genera. However, it is strong, tight and linear at the level of families/subfamilies (r² = 0.6, F_1,10 = 19, P < 0.01). Although among primates, density is very significantly related to mass, at no taxonomic level is range size related to body mass. Consequently, removing the effects of mass makes little to no difference to density–range results. Regarding the biology of the relationship, only traits indicative of specialization are associated with abundance (meaning numbers): rare taxa are more specialized than are abundant taxa. The association is largely via range size, not density. Across families, no traits correlate significantly with the density–range relationship, nor with deviations from it, despite the strength of the relationship at this taxonomic level. Main conclusions We suggest that in macroecology, analysis at taxonomic levels deeper than that of the relatively ephemeral species can be appropriate. We argue that the several purely methodological explanations for the positive density–range size relationship in primates can be rejected. Of the various biological hypotheses, those having to do with specialization–generalization seem the only applicable ones. The fact that the relationship is entirely via range size, not via density, means that while we might have a biology of range size, we do not yet have one of the density–geographical range relationship. It is probably time to search for multivariate explanations, rather than univariate ones. However, we can for the first time, for at least primates, suggest that any association of abundance or range size with specialization is via the number of different subtaxa, not the average degree of specialization of each subtaxon. The implication for conservation is obvious.     

Neutral community dynamics, the mid-domain effect and spatial patterns in species richness

The mid‐domain effect (MDE) aims to explain spatial patterns in species richness invoking only stochasticity and geometrical constraints. In this paper, we used simulations to show that its main qualitative prediction, a hump‐shaped pattern in species richness, converges to the expectation of a spatially bounded neutral model when communities are linked by short‐distance migration. As these two models can be linked under specific situations, neutral theory may provide a mechanistic population level basis for MDE. This link also allows establishing in which situations MDE patterns are more likely to be found. Also, in this situation, MDE models could be used as a first approximation to understand the role of both stochastic (ecological drift and migration) and deterministic (adaptation to environmental conditions) processes driving the spatial structure of species richness.     

The species-area-energy relationship

Area and available energy are major determinants of species richness. Although scale dependency of the relationship between energy availability and species richness (the species-energy relationship) has been documented, the exact relationship between the species-area and the species-energy relationship has not been studied explicitly. Here we show, using two extensive data sets on avian distributions in different biogeographic regions, that there is a negative interaction between energy availability and area in their effect on species richness. The slope of the species-area relationship is lower in areas with higher levels of available energy, and the slope of the species-energy relationship is lower for larger areas. This three-dimensional species-area-energy relationship can be understood in terms of probabilistic processes affecting the proportions of sites occupied by individual species. According to this theory, high environmental energy elevates species' occupancies, which depress the slope of the species-area curve.     

Avian foraging studies: an overlooked source of distribution data for macroecological and conservation studies

Macroecological and biogeographical studies and their applicability for biological conservation vitally depend on distribution data. These are usually taken from distribution atlases and databases or maps found in identification guides. A previous study pointed to another little explored source of data — local faunistic studies. Here, I would like to draw attention to another potential source. Papers that analysed the food composition of some taxa (e.g. insectivorous birds) are an overlooked source of rich information on taxa distributions. These studies frequently include an 'Appendix' with a list of food items determined to the species level. These studies also contain data on abundances, number of samples, sampling time, and geographical location as a rule. Foraging birds naturally provide data on invertebrate distributions with good spatio-temporal coverage and reasonably large samples. Importantly, birds frequently collect rare and by entomological methods hardly detectable species (i.e. those living high, in tree canopies, in very dense vegetation, or with secretive lifestyle). Data from bird dietary studies may help to ameliorate one of the most serious problems in distribution studies — zero inflation. I briefly discuss pros and cons of this so far neglected source of biogeographical data.     

Rarity in the tropics: biogeography and macroecology of the primates

Aim To describe rarity and elucidate its biology in a tropical mammalian order, the Primates. Location Africa, Central and South America, Asia, Madagascar. Methods A review of the literature, with some additional analyses using data from the literature. A variety of definitions of rarity are used in order to describe it and to investigate its biology by correlating the degree of rarity with a variety of biological traits indicative of resource use (e.g. size of annual home range), reproductive rate (e.g. birth interval)and specialization (e.g. number of habitat types used). Results Few primate taxa occur outside the tropics, and most taxa are rare (small geographical range size or latitudinal extent, low density or both). Latitudinal extent is narrower at lower latitudes in Africa and Asia, but the potential resultant packing of taxa appears not to explain the taxonomic diversity gradient. Whilst primate species do not show the common, positive density by range size relationship, primate genera show a significant shallow slope, and primate families/subfamilies a strongly positive slope. Rare taxa are specialized, but neither use more resources nor breed more slowly than common taxa. The correlation of rarity and specialization is via geographical range: taxa with small ranges, or small ranges for their density, are specialized, but not taxa at low density. Common taxa are generalized because they consist of more differently specialized subtaxa, not because each subtaxon is generalized. Main conclusions Most primate taxa are rare, in which case most are presumably likely to go extinct. Rare primates are specialized, but do not necessarily use more resources, nor breed more slowly. Specialization as an explanation for rarity appears to work via constriction of range size, not of density. Common primates might be common (large range size) not because subtaxa or individuals are generalized, but because they are composed of more subtaxa. A consequence could be that persistence of even common taxa will depend on conservation of several populations scattered across the taxon's geographical range.     

The island rule and a research agenda for studying ecogeographical patterns

We are currently experiencing a resurgence of interest in ecogeographical rules, which describe general trends in morphology and related traits along geographical gradients. In order to develop a more comprehensive understanding of the generality and underlying causal mechanisms for these patterns, we recommend a new, more integrated research agenda. In particular, we recommend studies that simultaneously consider different clines in morphology, geographical ranges and diversity as intricately related phenomena; all being ecological, evolutionary and biogeographical responses of organisms to selection regimes that vary non-randomly over space and time, and among species with different ecological and evolutionary histories.     

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.     

Eight (and a half) deadly sins of spatial analysis

Biogeography is spatial by nature. Over the past 20 years, the literature related to the analysis of spatially structured data has exploded, much of it focused on a perceived problem of spatial autocorrelation and ways to deal with it. However, there are a number of other issues that permeate the biogeographical and macroecological literature that have become entangled in the spatial autocorrelation web. In this piece I discuss some of the assumptions that are often made in the analysis of spatially structured data that can lead to misunderstandings about the nature of spatial data, the methods used to analyse them, and how results can be interpreted.     

More and more generalists: two decades of changes in the European avifauna

Biotic homogenization (BH) is a process whereby some species (losers) are systematically replaced by others (winners). While this process has been related to the effects of anthropogenic activities, whether and how BH is occurring across regions and the role of native species as a driver of BH has hardly been investigated. Here, we examine the trend in the community specialization index (CSI) for 234 native species of breeding birds at 10 111 sites in six European countries from 1990 to 2008. Unlike many BH studies, CSI uses abundance information to estimate the balance between generalist and specialist species in local assemblages. We show that bird communities are more and more composed of native generalist species across regions, revealing a strong, ongoing BH process. Our result suggests a rapid and non-random change in community composition at a continental scale is occurring, most likely driven by anthropogenic activities.     

Integrating ecology with biogeography using landscape characteristics: a case study of subtidal habitat across continental Australia

Aim We aimed to redress a current limitation of local ecological studies (i.e. piecemeal information on specific taxa) by integrating existing ecological knowledge with quantifiable patterns in primary habitat (i.e. composition, distribution and cover) from local to continental scales. By achieving this aim, we sought to provide a biogeographical framework for the interpretation of variation in the ecology of, and threats to, subtidal rocky landscapes. Location The subtidal rocky coast of continental Australia, with longitudinal comparisons spanning > 4000 km of southern coast (115°03′ E–153°60′ E) between latitudes of 33°05′ S and 35°36′ S, and latitudinal comparisons across 26°40′ S to 37°08′ S of eastern Australia. Methods The frequency and size of patches of major benthic habitat were quantified to indicate contemporary function (ecology) and to establish patterns that may result from contrasting regional‐scale processes (biogeography). This was achieved by quantifying the composition and patchiness of key subtidal habitats across the continent and relating them to the known ecology of subsets of locations in each region. A nested design of several spatial scales (1000s, 100s, 10–1 km) was adopted to distinguish patterns at local through to biogeographical scales. Results We show biogeography (in terms of longitude and latitude) to have a fundamental influence on the patterns of abundance and composition of subtidal habitats across regional (1000s of kilometres) to local (10s of kilometres to metres) scales. Across the continent, the most fundamental patterns related to (1) the proportion of rock covered by kelp forests, as related to particular functional groups of herbivores, and (2) the small‐scale heterogeneity (metres) that characterizes these forests. Main conclusions We interpret these results within a framework of alternative processes known to maintain habitat heterogeneity across these regions (e.g. productivity versus consumption as shapers of habitat structure). These interpretations illustrate how regional differences in ecological patterns and processes can create contradictory outcomes for the management of natural resources. We suggest that researchers and managers of natural resources alike may benefit from understanding local issues (e.g. the effects of fishing and its synergies with water pollution) in their biogeographical contexts.