Of mice and wrens: the relation between abundance and geographic range size in British mammals and birds

We examine the relation between population size and geographic range size for British breeding birds and mammals. As for most other assemblages studied, a strong positive interspecific correlation is found in both taxa. The relation is also recovered once the phylogenetic relatedness of species has been controlled for using an evolutionary comparative method. The slope of the relation is steeper for birds than for mammals, but this is due in large part to two species of mammals that have much higher population sizes than expected from their small geographic ranges. These outlying mammal species are the only ones in Britain to be found only on small offshore islands, and so may be exhibiting density compensation effects. With them excluded, the slope of the abundance–range size relation for mammals is not significantly different to that for birds. However, the elevation of the relation is higher for mammals than for birds, indicating that mammals are approximately 30 times more abundant than birds of equivalent geographic range size. An earlier study of these assemblages showed that, for a given body mass, bats had abundances more similar to birds than to non-volant mammals, suggesting that the difference in abundance between mammals and birds might be due to constraints of flight. Our analyses show that the abundance–range size relation for bats is not different for that from other mammals, and that the anomalously low abundance of bats for their body mass may result because they have smaller than expected geographic extents for their size. Other reasons why birds and mammals might have different elevations for the relation between population size and geographic range size are discussed, together with possible reasons for why the slopes of these relations might be similar.     

Population density and geographical range size in the introduced and native passerine faunas of New Zealand

The current avifauna of New Zealand comprises species with two distinct origins: those that evolved in New Zealand or colonized naturally from neighbouring landmasses, and those that were deliberately introduced to the islands by European settlers. Elsewhere, it has been shown that for species introduced to New Zealand from Britain there is a positive interspecific correlation between the geographical range sizes attained in both countries. Since positive relationships between abundance, measured either as population size or density, and geographical range size are a near ubiquitous feature of assemblages of closely related animal species, this suggests that species’ abundances may also be so correlated between the two countries. Here, data for 12 passerine bird species introduced to New Zealand from Britain are used to compare population densities and density–range size relationships in their native and alien ranges. In addition, the density–range size relationship for 12 passerine bird species that can be considered native to New Zealand is compared to that for the introduced species. The geographical range size and the mean and maximum densities of introduced species in New Zealand were significantly positively correlated with those values for the same species in Britain. However, in no case was the relationship between mean density and range size significant. While not statistically significant, density–range size relationships for introduced species are similar in New Zealand and Britain, but those for introduced and native species in New Zealand are quite different. Implications of these patterns are discussed.     

Interspecific abundance-range size relationships: range position and phylogeny

A number of mechanisms have been proposed to explain the widely observed positive interspecific relationship between local abundance and extent of geographic distribution in animals Here, we use data on British birds to assess two of these hypotheses that the relationship results from the relative position of a study area with respect to the geographic ranges of the species which occur there, and that the relationship results from a simple difference between taxonomic groups, rather than any general tendency for more abundant species to have larger range sizes We find support for neither hypothesis Phylogenetically controlled comparative analyses reveal that the positive abundance-range size relationship is consistently found within taxa, even when abundance and range size are calculated at a variety of spatial and temporal scales Analyses both across species and within taxa show that bird species for which Britain is near to the centre of their distribution in Europe tend to have larger British range sizes and higher abundances than do species where Britain is close to the edge of their range in Europe However, these relationships do not cause that between abundance and range size, because this latter relationship persists within different range position categories Whether a species is near the centre or edge of its geographic range in Britain may affect its position on the abundance-range size relationship, but does not produce the relationship Range position in Britain does, however, seem to be related to the magnitude of temporal changes in the range sizes of British birds There is some evidence to suggest that species for which Britain is nearer to their European range centre have shown smaller changes in distribution over the period 1970–1990 than have species for which Britain is close to their European range edge     

Identifying plant functional types using floristic data bases: Ecological correlates of plant range size

Abstract. In an effort to identify ‘plant functional types’, the islands floras of Great Britain and Kríti (Crete, Greece) were examined separately for ecological correlates of plant range size. Plant functional types (PFTs) were defined here as categories into which plants could be grouped on the basis of attributes that predict greater or lesser sensitivity to ecological variability. Plant range size indicates commonness of a species and was assumed to be a proxy for ‘ecological flexibility’, i.e. species of larger range sizes can better withstand environmental change and differences than species of smaller range sizes. Using evolutionary comparative methods that account for the effect of taxonomic relatedness, both floras were investigated for the effects on range size of woodiness vs. non-woodiness, trees vs. shrubs, trees vs. herbs and shrubs vs. herbs. The British flora was examined additionally for the effects of wind- vs. non-wind-pollination, self vs. animal pollination and animal vs. non-animal fruit dispersal on range size. Two analyses showed significant effects on range size: for British species, trees had larger ranges than shrubs, and wind- pollinated species had larger ranges than non-wind-pollinated species. It is suggested that the lack of a similar pattern for shrubs and trees in Kríti is because the lower water availability of Kríti imbues shrubs with an ecophysiological advantage not relevant in plants of Great Britain. That trees have larger range sizes than shrubs in Great Britain is ascribed to the greater importance of competition for light when other factors are not at issue. The greater range of wind-pollinated than non-windpollinated species in Great Britain is postulated to be because both mutualists must be capable of invading new areas. This may be termed a ‘cost of mutualism’. In terms of PFTs, the results indicate that ‘life-form’ is too broad a classification category by which to differentiate relative sensitivity to environmental variability in Great Britain, in that there were significant differences in range size of trees and shrubs, but not between either of the two categories and herbs, or between woody and non-woody plants. Although pollination type may predict relative sensitivity to variation in Great Britain, dispersal type will not. Finally, differences between Great Britain and Kríti in relative range size patterns suggests that plant functional types may be specific to a region or set of conditions.     

Coincidence in the distributions of butterflies and their foodplants

The relationship between the geographic distribution of consumers and of their hosts (foodplants) is examined using the resident butterfly fauna of Britain. On average, butterfly species that feed on more widely distributed hosts are themselves more widely distributed. However, the relationship is approximately triangular and the upper constraint imposed by the range sizes of hosts is not closely followed; some species have much more restricted ranges than their hosts have. There is no relationship between the proportion of the range of the foodplant that is occupied and the size of the range of the foodplant. Monopbagous butterfly species have smaller range sizes than polyphagous species, probably as a consequence of the greater potential range sizes of the latter. Those plant species that are used as hosts by butterflies have larger range sizes than expected by chance, and individual polyphagous butterfly species tend disproportionately to be found in areas containing larger numbers of their host plant species. In sum, this study reveals a complex relationship between the distribution of butterflies and that of their resources (foodplants).     

The trait and host plant ecology of aphids and their distribution and abundance in the United Kingdom

Aim We characterized the annual populations of 170 aphid species by their log abundance, site occupancy and site continuity (i.e. the persistence of species in time) and used this information to make predictions about groups of species that displayed characteristic patterns. By doing so, we aimed to identify commonalities in functional traits (host‐alternation; mode of reproduction; life‐cycle plasticity; median body size) and host plant geographic range sizes that may indicate why some species are common and others not. Location The population dynamics of winged aphids at 27 locations in the United Kingdom were studied. Methods The annual numbers of aphids were studied using a dataset comprising over 11 million individuals across 509 site‐years. Traits and host plants were analysed using linear mixed effects models and nonlinear regression models. Results Linear mixed effects models showed that the fixed effects of host alternation and winter host plant area of occupancy were important in predicting log abundance, site occupancy and site continuity. Life‐cycle plasticity was also a significant effect, although not for log abundance. Relationships between site continuity, site occupancy and log abundance were strongly nonlinear. Site continuity always lagged site occupancy, indicating that species were less likely to retain previously occupied sites when abundances were low. Main conclusions Aphid traits are a better paradigm than taxonomic relatedness in explaining macroecological patterns. Host alternation induces an annual flux of migrants that engenders higher annual log abundances, consistent with the theory that species with high local densities tend to confer a much wider distribution than those with low densities. The abundance of aphids is monotonically related to the geographic range size of their winter host, suggestive of a strong bottom‐up effect (i.e. resource controlled). The areas of occupancy (AOO) of winter host plants constrain aphids to low abundances because a greater proportion of hosts are trees and shrubs which tend to have smaller AOO than herbs and grasses, the common summer host types.     

The effect of the taxon and geographic range size of host eucalypt species on the species richness of gall-forming insects

Abstract This field study was designed to test whether the taxonomic group and geographic range size of a host plant species, usually found to influence insect species richness in other parts of the world, affected the number of gall species on Australian eucalypts. We assessed the local and regional species richness of gall-forming insects on five pairs of closely related eucalypt species. One pair belonged to the subgenus Corymbia, one to Monocalyptus, and three to different sections of Symphyomyrtus. Each eucalypt pair comprised a large and a small geographic range species. Species pairs were from coastal or inland regions of eastern Australia. The total number of gall species on eucalypt species with large geographic ranges was greater than on eucalypt species with small ranges, but only after the strong effect of eucalypt taxonomic grouping was taken into account. There was no relationship between the geographic range size of eucalypt species and the size of local assemblages of gall species, but the variation in insect species composition between local sites was higher on eucalypt species with large ranges than on those with small ranges. Thus the effect of host plant range size on insect species richness was due to greater differentiation between more widespread locations, rather than to greater local species richness. This study confirms the role of the geographic range size of a host plant in the determination of insect species richness and provides evidence for the importance of the taxon of a host plant.     

The population ecology of rare species

There is no general theory of rarity, although one is sorely needed both to understand population dynamics and to determine conservation priorities. Here we suggest some of the strands that might be woven into such a theory. They include relationships between local abundance, geographic range size and body size of species, and the determinants of minimum viable population sizes. In each of these areas much can still be learnt from the classical 'compare and contrast' approach using assemblages of species from a variety of taxa. Freshwater fish have contributed relatively little to the broad ecological literature in this respect. We perform some tentative analyses for this group of species, and speculate on how they might fit into our current understanding of rarity.     

Biologic interactions determining geographic range size: a one species response to phylogenetic community structure

Range size variation in closely related species suggests different responses to biotic and abiotic heterogeneity across large geographic regions. Species turnover generates a wide spectrum of species assemblages, resulting in different competition intensities among taxa, creating restrictions as important as environmental constraints. We chose to adopt the widely used phylogenetic relatedness (NRI) measurement to define a metric that depicts competition strength (via phylogenetic similarity), which one focal species confronts in its environment. This new approach (NRI_focal) measures the potential of the community structure effect over performance of a single species. We chose two ecologically similar Peucaea sparrows, which co‐occur and have highly dissimilar range size to test whether the population response to competition intensity is different between species. We analyzed the correlation between both Peucaea species population sizes and NRI_focal using data from point counts. Results indicated that the widespread species population size was not associated with NRI_focal, whereas the population of restricted‐sized species exhibited a negative relationship with competition intensity. Consequently, a species' sensitivity to competition might be a limiting factor to range expansion, which provides new insights into geographic range analysis and community ecology.     

Genetic differentiation of starling (Sturnus vulgaris: Aves) populations in New Zealand and Great Britain

Several hundred starlings (Sturnus vulgaris) were introduced to New Zealand from Great Britain during1860–1880. Allozymic variation at 24 loci was analysed in winter populations sampled at six localities in each country. New Zealand samples had fewer alleles per locus but the same mean heterozygosity (3% per locus) and proportion of polymorphic loci as did British samples. Winter populations in Britain contain European migrants and were genetically homogeneous. Paradoxically, genetic distances among derived New Zealand populations, and between New Zealand and Great Britain were much greater, similar in magnitude to those observed among allopatric populations in other avian species. The geographical pattern of genetic variation in New Zealand suggests that reproductive isolation of populations and random drift have contributed to the development of population differentiation.     

Field sampling is biased against small-ranged species of high conservation value: a case study on the sphingid moths of East Africa

The range size of species co-occurring in local assemblages is a pivotal variable in assessments of a site’s conservation value. Assemblages featuring many small-ranged species are given more priority than assemblages consisting mainly of wide-ranging species. However, the assembly of relevant information can be challenging and local range size distributions of tropical invertebrates are rarely available for conservation planning. We present such data for sphingid moths in East Africa, a highly diverse region of high conservation value. We compare geographic range size distributions based on field samples with predictions from modelled range map data. Using this system as a case study, we provide evidence for a systematic sampling bias when inferring average local range sizes from field data. Unseen species (i.e., species present but missed in local sampling) are often those with small ranges (hence, of high conservation value). Using an elevational gradient, we illustrate how this bias can lead to false, counterintuitive assessments of environmental effects on local range size distributions. Furthermore, with particular reference to sphingid moths in the study region, we show that current protected areas appear unrelated to the spatial distribution of species richness or average geographic range sizes at a local scale. We discuss the need to treat field sampled data with caution and in concert with other data sources such as probabilistic models.     

Abundance-range size relationships in British birds: is unexplained variation a product of life history?

Positive interspecific relationships between local abundance and geographic range size are a common feature of animal assemblages However, range size typically explains only a moderate proportion of the variation m abundance, begging the question of whether species of differing life history deviate from the underlying relationship in any systematic fashion Using data for the avifauna of Britain, and applying a comparative method to control for the effects of phylogenetic association, we demonstrate that this does not appear to be the case Only adult survivorship, age at independence and incubation period explain significant variation in abundance once range size is controlled for statistically, and then only a few percent There are two probable reasons why this result is contrary to general expectation First, although many life history variables are expected to show simple correlations with abundance or range size It is not obvious how these might relate to variation about the abundance–range size relationship Second, intuitive ideas about the form such variation might take may be seriously confounded by phylogenetic non–independence     

Reserve mass and dispersal investment in relation to geographic range of plant species: phylogenetically independent contrasts

Recent studies have reported conflicting evidence about correlations between seed size and plant species geographic range sizes. Using phylogenetically independent contrasts (PICs) within genera, we found no consistent differences in reserve mass between species with similar dispersal morphology and «wide>> versus «narrow>> geographic ranges. There was also no tendency within genera for broad ranged species to be those that allocate a larger percentage of the resources invested in each diaspora to dispersal structures. PICs were also constructed between species having a tenfold difference in seed size. In these PICs, the larger seeded species often occupied a greater number of regions than species with smaller seed sizes. This result was generated primarily through the comparison of species from different genera, families or higher level taxa which differed not only in seed mass but also in dispersal modes and growth forms. Finally, comparing species within Acacia and Eucalyptus having similar seed size but different dispersal modes, we found that bird dispersal (in Acacia ) and possession of a wing for wind dispersal (in Eucalyptus ) was associated with wider geographic range compared to lower-investment dispersal modes. Taken together, these comparisons indicate that seed size is not itself important as a factor influencing breadth of geographic range. Dispersal mode and growth form may have an influence, however, and seed size differences may be associated with contrasts in dispersal mode or growth form.     

On the heritability of geographic range sizes

Within taxonomic groups, most species are restricted in their geographic range sizes, with only a few being widespread. The possibility that species-level selection on range sizes contributes to the characteristic form of such species-range size distributions has previously been raised. This would require that closely related species have similar range sizes, an indication of "heritability" of range sizes at the species level. Support for this view came from a positive correlation between the range sizes of closely related pairs of fossil mollusc species. We extend this analysis by considering the relationship between the geographic range sizes of 103 pairs of contemporary avian sister species. Range sizes in these sister species show no evidence of being more similar to each other than expected by chance. A reassessment of the mollusc data also suggests that the high correlation was probably overestimated because of the skewed nature of range size data. The fact that sister species tend to have similar life histories and ecologies suggests that any relationship between range sizes and biology is likely to be complicated and will be influenced by historical factors, such as mode of speciation and postspeciation range size transformations.     

Abundance, species richness and energy availability in the North American avifauna

Aim To determine how species richness, abundance, biomass, energy use and mean number of individuals per species scale with environmental energy availability in wintering and breeding avian assemblages, and to contrast assemblages of (i) common and rare species and (ii) breeding residents and migrants. To assess whether such patterns are compatible with the ‘more individuals hypothesis’ (MIH) that high‐energy areas are species‐rich because they support larger populations that are buffered against extinction. Location The North American continent (latitudinal range 23.4 °?48.1 °N; longitudinal range 124.2°?68.7° W). Methods Avian species richness, abundance, biomass and energy use were calculated for 295 Resident Bird Count plots. Environmental energy availability was measured using ambient temperature and the Normalized Difference Vegetation Index (NDVI), a close correlate of plant productivity. Analyses took plot area into account, and were conducted (with and without taking habitat type into account) using general linear models and spatial mixed models. Results Positive species–energy relationships were exhibited by both wintering and breeding assemblages, but were stronger in the former. The structure of winter assemblages responded more strongly to temperature than NDVI, while breeding assemblages tended to respond more strongly to NDVI. Breeding residents responded to annual measures of energy availability while breeding migrants and the winter assemblage responded more strongly to seasonal measures. In the winter assemblage, rare and common species exhibited species–energy relationships of a similar strength, but common breeding species exhibited a much stronger relationship than rare breeding species. In both breeding and wintering assemblages, abundance, biomass and energy use increased with energy availability and species richness. Energy availability was a poor predictor of the mean number of individuals per species. Main conclusions The nature of the species–energy relationship varies seasonally and with the manner in which energy availability is measured. Our data suggest that residents are less able to respond to seasonal fluxes in resource availability than long‐distance migrants. Increasing species richness and energy availability is associated with increasing numbers of individuals, biomass and energy use. While these observations are compatible with the MIH our data provide only equivocal support for this hypothesis, as the rarest species do not exhibit the strongest species–energy relationships.     

Winter range compression of migrants in Central America

Where there is seasonal disparity among opportunities, the season with those in shortest supply is most likely to limit populations. Among migrant birds that travel between different breeding and winter ranges, any of breeding, migratory or winter conditions could exclusively constitute such population‐limiting factors. In both the New and Old Worlds, landmass is disproportionately concentrated in temperate latitudes. In the Americas, most passerine bird species that breed in the USA and Canada spend the winter further south, commonly in parts of the tropics where landmass is significantly less. Using a sample of 89 migratory species (eight passerine families) that breed in eastern North America, I considered patterns of geographic breeding range size, winter range size and winter distribution. Winter range size is usually smaller than breeding range size (84 of 89 species), often substantially so (minimum 8%, mean 52%). Wintering latitude explains significant variation in both breeding range size and winter range size, as well as in winter range size relative to breeding range size. In particular, all three measures vary latitudinally in patterns similar to latitudinal variation in landmass. These patterns collectively suggest that the reduction in landmass in the latitudes of Central America and the Caribbean is a limiting factor for migrant bird populations, adding to other research concluding that winter conditions sometimes prevail over breeding conditions in the limitation of populations. Hectare for hectare, habitat destruction in the tropics is likely to have the greater impact on the welfare of passerine populations breeding in North America.     

Range size and environmental calcium requirements of British freshwater gastropods

Calcium is an essential requirement for the successful growth and development of gastropod molluscs. Data for British freshwater gastropods were used to examine the relationship between environmental calcium requirements and British and European range sizes. At both spatial scales calciphile species, which require a high level of environmental calcium, had significantly smaller range sizes than species able to exploit a wide range of environmental calcium levels. However, at least in Britain, range size may also be influenced by the availability of suitable habitat. British and European range sizes were significantly correlated. This study provides evidence for niche‐based explanations of range size variation, and suggests that both niche breadth and niche availability are important in determining range size.     

Body Size and Geographic Range Do Not Explain Long Term Variation in Fish Populations: A Bayesian Phylogenetic Approach to Testing Assembly Processes in Stream Fish Assemblages

We combine evolutionary biology and community ecology to test whether two species traits, body size and geographic range, explain long term variation in local scale freshwater stream fish assemblages. Body size and geographic range are expected to influence several aspects of fish ecology, via relationships with niche breadth, dispersal, and abundance. These traits are expected to scale inversely with niche breadth or current abundance, and to scale directly with dispersal potential. However, their utility to explain long term temporal patterns in local scale abundance is not known. Comparative methods employing an existing molecular phylogeny were used to incorporate evolutionary relatedness in a test for covariation of body size and geographic range with long term (1983 – 2010) local scale population variation of fishes in West Fork White River (Indiana, USA). The Bayesian model incorporating phylogenetic uncertainty and correlated predictors indicated that neither body size nor geographic range explained significant variation in population fluctuations over a 28 year period. Phylogenetic signal data indicated that body size and geographic range were less similar among taxa than expected if trait evolution followed a purely random walk. We interpret this as evidence that local scale population variation may be influenced less by species-level traits such as body size or geographic range, and instead may be influenced more strongly by a taxon’s local scale habitat and biotic assemblages.     

Patterns in the geographic ranges of the world's woodpeckers

We used data on the world's woodpeckers to test for patterns in the geographic distributions of a single group of closely related species. The frequency distribution of woodpecker geographic range sizes is approximately lognormal. Most variation in range sizes is explained by differences between species within genera; that is, range size seems to be an evolutionarily labile trait. The largest woodpecker ranges are found in Eurasia, both when absolute differences are compared and when range size is measured as a proportion of estimated available habitat. Notably, there is a negative relationship between the mean range sizes attained by species in a genus or tribe in South America and the mean ranges attained by species in the same tribe or genus in North America. Large-bodied species tend to be more widely distributed and to live at higher latitudes, but both tendencies disappear if the taxonomic relatedness of species is controlled for. Species living at high latitudes also tend to be more widely distributed. This relationship seems largely due to the effect of North American woodpeckers, which show it even when the taxonomic relatedness of species is controlled. Small continents generally have more woodpecker species than do large ones. Woodpecker geographic range sizes are smaller the more woodpecker species inhabit an area. Species show less overlap in their geographic ranges with species of similar than with species of dissimilar body size. The implications of these results for our understanding of patterns in geographic range sizes are discussed.     

Range size heritability in Carnivora is driven by geographic constraints

Range size heritability refers to an intriguing pattern where closely related species occupy geographic ranges of similar extent. Its existence may indicate selection on traits emergent only at the species level, with interesting consequences for evolutionary processes. We explore whether range size heritability may be attributable to the fact that range size is largely driven by the size of geographic domains (i.e., continents, biomes, areas given by species' climatic tolerance) that tend to be similar in phylogenetically related species. Using a well-resolved phylogeny of Carnivora, we show that range sizes are indeed constrained by geographic domains and that the phylogenetic signal in range sizes diminishes if the domain sizes are accounted for. Moreover, more detailed delimitation of species' geographic domain leads to a weaker signal in range size heritability, indicating the importance of definition of the null model against which the pattern is tested. Our findings do not reject the hypothesis of range size heritability but rather unravel its underlying mechanisms. Additional analyses imply that evolutionary conservatism in niche breadth delimits the species' geographic domain, which in turn shapes the species' range size. Range size heritability patterns thus emerge as a consequence of this interplay between evolutionary and geographic constraints.