Range-size Frequency Distribution Research Articles

Diversification Rates and the Evolution of Species Range Size Frequency Distribution

The geographic range sizes frequency distribution (RFD) within clades is typically right-skewed with untransformed data, and bell-shaped or slightly left-skewed under the log-transformation. This means that most species within clades occupy diminutive ranges, whereas just a few species are truly widespread. A number of ecological and evolutionary explanations have been proposed to account for this pattern. Among the latter, much attention has been given to the issue of how extinction and speciation probabilities influence RFD. Numerous accounts now convincingly demonstrate that extinction rate decreases with range size, both in living and extinct taxa. The relationship between range size and speciation rate, though, is much less obvious, with either small or large ranged species being proposed to originate more daughter taxa. Herein, we used a large fossil database including twenty-one animal clades and more than 80,000 fossil occurrences distributed over more than 400 million years of marine metazoans (exclusive of vertebrates) evolution, to test the relationship between extinction rate, speciation rate, and range size. As expected, we found that extinction rate almost linearly decreases with range size. In contrast, speciation rate peaks at the large (but not the largest) end of the range size spectrum. This is consistent with the peripheral isolation mode of allopatric speciation being the main mechanism of species origination. The huge variation in phylogeny, fossilization potential, time of fossilization, and the overarching effect of mass extinctions suggest caution must be posed at generalizing our results, as individual clades may deviate significantly from the general pattern.

Macroecological conclusions based on IUCN expert maps: A call for caution

AbstractAimInternational Union for Conservation of Nature (IUCN) expert maps are increasingly used in macroecological research. However, they have not been produced for this purpose. Macroecological insights based exclusively on this type of data could therefore be misleading. Here we compare, for a large taxonomic group (bats) and an entire biogeographical realm (Africa), the species‐specific discrepancies between IUCN maps and species distribution models (SDMs) that approximate the complete geographical range of species. We then examine the implications for a typical macroecological analysis that explores environmental correlates of species richness.LocationContinental Africa.Time periodAround 2000.TaxaBats (Chiroptera).MethodsWe measure disagreement between IUCN expert maps and SDMs at both the species (geographical ranges) and the aggregated level (range size–frequency distributions and species richness). We further quantify the difference in absolute and relative weight assigned to three variables hypothesized to drive species richness: primary productivity, climatic seasonality and environmental heterogeneity.ResultsLocation, shape and size of individual species’ ranges, derived richness patterns and range size–frequency distributions differ substantially. SDMs predict larger and more complex geographical ranges, and species’ range sizes vary less. The spatial congruence of richness hotspots among both datasets is only 42%. These discrepancies are large enough to alter the absolute explanatory power of environmental correlates, whereas the redundancy in the variation explained increases markedly when richness is inferred using SDM‐based estimates of complete species ranges.Main conclusionsIUCN expert maps differ considerably and systematically from SDMs built to estimate complete species ranges, primarily because of their intentional greater sensitivity to geographical sampling bias. This property is desirable in a conservation context but unwanted in most macroecological analyses. We therefore caution against the use of IUCN expert maps in macroecology and recommend at least gauging the robustness of results using additional range estimates designed to approximate the complete geographical range of species.

Unequal Contribution of Widespread and Narrow-Ranged Species to Botanical Diversity Patterns

In conservation studies, solely widespread species are often used as indicators of diversity patterns, but narrow-ranged species can show different patterns. Here, we assess how well subsets of narrow-ranged, widespread or randomly selected plant species represent patterns of species richness and weighted endemism in Gabon, tropical Africa. Specifically, we assess the effect of using different definitions of widespread and narrow-ranged and of the information content of the subsets. Finally, we test if narrow-ranged species are overrepresented in species-rich areas. Based on distribution models of Gabonese plant species, we defined sequential subsets from narrow-ranged-to-widespread, widespread-to-narrow-ranged, and 100 randomly arranged species sequences using the range sizes of species in tropical Africa and within Gabon. Along these sequences, correlations between subsets and the total species richness and total weighted endemism patterns were computed. Random species subsets best represent the total species richness pattern, whereas subsets of narrow-ranged species best represent the total weighted endemism pattern. For species ordered according to their range sizes in tropical Africa, subsets of narrow-ranged species represented the total species richness pattern better than widespread species subsets did. However, the opposite was true when range sizes were truncated by the Gabonese national country borders. Correcting for the information content of the subset results in a skew of the sequential correlations, its direction depending on the range-size frequency distribution. Finally, we find a strong, positive, non-linear relation between weighted endemism and total species richness. Observed differences in the contribution of narrow-ranged, widespread and randomly selected species to species richness and weighted endemism patterns can be explained by the range-size frequency distribution and the use of different definitions of widespread or narrow-ranged. We call for a reconsideration of the use of widespread species as an indicator of diversity patterns, and advocate using the full ranges of species when assessing diversity patterns.

The Co-Occurrence of Species and the Co-Diversity of Sites in Neutral Models of Biodiversity

Patterns of co-occurrence of species are widely used to assess the fit of ecological neutral models to empirical patterns. The mathematically equivalent patterns of co-diversity of sites, in contrast, have been considered only indirectly and analyses normally are focused on the spatial distribution of species richness, rather than on the patterns of species sharing. Here we use two analytical tools (range-diversity plots and rank plots) to assess the predictions of simple neutral models in relation to patterns of co-occurrence and co-diversity. Whereas a fully stochastic null model predicts zero average among species and among sites, neutral models generate systems with low levels of covariance among species and high levels of positive covariance among sites. These patterns vary with different combinations of dispersal and speciation rates, but are always linked to the shape, symmetry, and spread of the range-size and species-richness frequency distributions. Non-homogeneous patterns of diversity and distribution arise in neutral models because of the spatial arrangement of sites and their concomitant similarity, which is reflected also in the spread of the range-size frequency distribution. The nearly null covariance among species, in contrast, implies low variance in species richness of sites and very slim frequency distributions. In real world assemblages of Mexican volant and non-volant mammals, patterns of range-size and species-richness frequency distribution are similar to those generated by neutral models. However, when the comparison includes the covariance both for species (co-occurrence) and for sites (co-diversity), empirical patterns differ significantly from the predictions of neutral models. Because of the mathematical links between the covariance in the distribution of species and the variance of species-richness values and between the covariance in species sharing among sites and the variance of range-size values, a full understanding of patterns of diversity calls for the simultaneous analysis of co-occurrence and co-diversity.

Effects of diversity‐dependent colonization–extinction dynamics on the mid‐domain effect

AbstractAimWe incorporate diversity‐dependent colonization and extinction rates into process‐based models of species geographic range dynamics to explore their effects on species richness gradients, and on extent and occupancy of species ranges. In particular, we investigate whether diversity dependence promotes or inhibits the emergence of mid‐domain effects (MDEs) in homogeneous environments.LocationA theoretical one‐dimensional domain.MethodsWe formulated diversity‐independent (DI) and diversity‐dependent (DD) models that simulated colonization, local extinction and speciation within a homogenous domain. In the DD model, colonization and extinction probabilities were functions of diversity, whereas in the DI model, they were constants. For a wide range of parameter values, we examined local and regional species richness gradients and species range size frequency distributions (RSFDs).ResultsIn contrast to the DI model, for which MDEs only occurred in a very narrow parameter range, the DD model generated a MDE in regional richness (range overlap) that was robust to colonization and extinction parameters, over a broad range of speciation rates. However, neither model could produce gradients in local richness (patch occupancy). The DD model also produced more realistic RSFDs than the DI model. In the latter, all species generally either became highly pandemic or went globally extinct, depending on the balance of colonization and extinction probabilities.Main conclusionsDiversity‐dependent colonization and extinction rates can have strong effects on species richness gradients and distributions of range extent and occupancy. Models with such diversity dependence amplify MDEs in regional richness, but largely eliminate MDEs in local richness, relative to DI models. DD models also generate more realistic RSFDs. These findings suggest that diffuse species interactions can strongly influence patterns of range size and overlap, but also that environmental gradients are likely to be necessary to explain many species richness patterns in nature, which exhibit both local and regional diversity gradients.

Patterns and causes of species richness: a general simulation model for macroecology

Understanding the causes of spatial variation in species richness is a major research focus of biogeography and macroecology. Gridded environmental data and species richness maps have been used in increasingly sophisticated curve-fitting analyses, but these methods have not brought us much closer to a mechanistic understanding of the patterns. During the past two decades, macroecologists have successfully addressed technical problems posed by spatial autocorrelation, intercorrelation of predictor variables and non-linearity. However, curve-fitting approaches are problematic because most theoretical models in macroecology do not make quantitative predictions, and they do not incorporate interactions among multiple forces. As an alternative, we propose a mechanistic modelling approach. We describe computer simulation models of the stochastic origin, spread, and extinction of species' geographical ranges in an environmentally heterogeneous, gridded domain and describe progress to date regarding their implementation. The output from such a general simulation model (GSM) would, at a minimum, consist of the simulated distribution of species ranges on a map, yielding the predicted number of species in each grid cell of the domain. In contrast to curve-fitting analysis, simulation modelling explicitly incorporates the processes believed to be affecting the geographical ranges of species and generates a number of quantitative predictions that can be compared to empirical patterns. We describe three of the 'control knobs' for a GSM that specify simple rules for dispersal, evolutionary origins and environmental gradients. Binary combinations of different knob settings correspond to eight distinct simulation models, five of which are already represented in the literature of macroecology. The output from such a GSM will include the predicted species richness per grid cell, the range size frequency distribution, the simulated phylogeny and simulated geographical ranges of the component species, all of which can be compared to empirical patterns. Challenges to the development of the GSM include the measurement of goodness of fit (GOF) between observed data and model predictions, as well as the estimation, optimization and interpretation of the model parameters. The simulation approach offers new insights into the origin and maintenance of species richness patterns, and may provide a common framework for investigating the effects of contemporary climate, evolutionary history and geometric constraints on global biodiversity gradients. With further development, the GSM has the potential to provide a conceptual bridge between macroecology and historical biogeography.

Latitudinal gradients in species richness for South American Mytilidae and Ostreidae: can alternative hypotheses be evaluated by a correlative approach?

We tested to what extent mean sea surface temperature, geometric constraints in range size frequency distributions (the mid-domain effect) and geographical coastline distance to the equator are related to species richness of coastal Mytilidae and Ostreidae in the Pacific and Atlantic coasts of South America (excluding islands). The location and magnitude of the peaks in species richness, as well as the shape of the pattern, varied between oceans. Results were not biased by spatial autocorrelation, although strong multicollinearity among predictor variables was detected. However, these regional-extent regression models suggest differences in the causal factors that explain richness gradients of studied bivalves in South American coasts, most likely related to historical events such as the Southeastern Pacific Pleistocene mass extinction of bivalves. Our results reinforced the conclusion that there is no single best explanatory cause for the latitudinal gradient in species richness and showed that the correlative approach is not useful when predictor variables are strongly correlated.

Ecological Correlates and Conservation Implications of Overestimating Species Geographic Ranges

Species range maps based on extents of occurrence (EOO maps) have become the basis for many analyses in broad-scale ecology and conservation. Nevertheless, EOO maps are usually highly interpolated and overestimate small-scale occurrence, which may bias research outcomes. We evaluated geographical range overestimation and its potential ecological causes for 1158 bird species by quantifying EOO map occurrence across 4040 well-studied survey locations in Australia, North America, and southern Africa at the scale of 80-742 km2. Most species occurred in only 40-70% of the range indicated by their EOO maps. The observed proportional range overestimation affected the range-size frequency distribution, indicating that species are more range-restricted than suggested by EOO maps. The EOO maps most strongly overestimated the distribution of narrow-ranging species and ecological specialists with narrow diet and habitat breadth. These relationships support basic ecological predictions about the relationship between niche breadth and the fine-scale occurrence of species. Consequently, at-risk species were subject to particularly high proportional range overestimation, on average 62% compared with 37% of nonthreatened species. These trends affect broad-scale ecological analyses and species conservation assessments, which will benefit from a careful consideration of potential biases introduced by range overestimation.

Challenging Wallacean and Linnean shortfalls: knowledge gradients and conservation planning in a biodiversity hotspot

ABSTRACTKnowledge about biodiversity remains inadequate because most species living on Earth were still not formally described (the Linnean shortfall) and because geographical distributions of most species are poorly understood and usually contain many gaps (the Wallacean shortfall). In this paper, we developed models to infer the size and placement of geographical ranges of hypothetical non‐described species, based on the range size frequency distribution of anurans recently described in the Cerrado Biome, on the level of knowledge (number of inventories) and on surrogates for habitat suitability. The rationale for these models is as follow: (1) the range size frequency distribution of these species should be similar to the range‐restricted species, which have been most recently described in the Cerrado Biome; (2) the probability of new discoveries will increase in areas with low biodiversity knowledge, mainly in suitable areas, and (3) assuming range continuity, new species should occupy adjacent cells only if the level of knowledge is low enough to allow the existence of undiscovered species. We ran a model based on the number of inventories only, and two models combining effects of number of inventories and two different estimates of habitat suitability, for a total of 100 replicates each. Finally, we performed a complementary analysis using simulated annealing to solve the set‐covering problem for each simulation (i.e. finding the smallest number of cells so that all species are represented at least once), using extents of occurrence of 160 species (131 real anuran species plus 29 new simulated species). The revised reserve system that included information about unknown or poorly sampled taxa significantly shifted northwards, when compared to a system based on currently known species. This main result can be explained by the paucity of biodiversity data in this part of the biome, associated with its relatively high habitat suitability. As a precautionary measure, weighted by the inferred distribution data, the prioritization of a system of reserves in the north part of the biome appears to be defensible.

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.

Mean latitudinal range sizes of bird assemblages in six Neotropical forest chronosequences

AbstractAim The geographical range size frequency distributions of animal and plant assemblages are among the most important factors affecting large‐scale patterns of diversity. Nonetheless, the relationship between habitat type and the range size distributions of species forming assemblages remains poorly understood. We examined how the mean latitudinal range sizes of species in Neotropical bird species assemblages shift during forest clearance and subsequent regeneration. We tested the hypothesis that bird species assemblages in early successional habitats tend to have larger latitudinal ranges than those in more mature forests.Location We considered breeding bird chronosequence data from six Neotropical forests.Results Breeding bird assemblages were found to have the species with the largest average latitudinal range sizes in cleared areas, intermediate in young secondary forests and smallest in old secondary and mature forests. Similar differences were also found when we compared congeners differing in their successional preferences. Sizes of regional ranges (within the Neotropics) did not, however, differ consistently among successional stages. The larger latitudinal (but not regional) ranges of early successional species was as a result in part of the tendency of early successional species to have ranges that extend beyond the Neotropical forest biome.Conclusions Our analysis of chronosequences suggests that as early successional habitats mature, a consistent shift from large‐ranged species towards more small ranged species occurs. Even relatively old secondary forests have bird species with larger average ranges than mature forests. As a consequence, conservation of secondary forests alone will miss many of the species most at risk of extinction and most unlikely to be conserved in other locations or biomes.

Factors shaping the range‐size frequency distribution of the endemic fish fauna of the Tropical Eastern Pacific

AbstractAim To assess the effect of habitat fragmentation and isolation in determining the range‐size frequency distribution (RFD) of the shorefish fauna endemic to a discrete biogeographical region.Location The Tropical Eastern Pacific (TEP).Methods Habitat isolation represents the separation between oceanic islands and the continental shore of the TEP and habitat fragmentation the degree of spatial continuity of habitats (i.e. reefs, soft bottom, nearshore waters) along the continental coast of the TEP. The effects of habitat isolation and fragmentation were quantified by comparing the RFDs of (1) the species found on oceanic islands vs. the continental shore, and (2) species on the continental shore that use different habitat types.Results The RFD of the entire TEP fauna was bimodal, with peaks at both small‐ and large‐range ends of the spectrum. The small‐range peak was due almost entirely to island species and the large‐range peak due mainly to species found in both the continental shore and oceanic islands. RFDs varied among species using different habitats on the continental shore: reef‐fishes had a right‐skewed RFD, soft‐bottom species a flat RFD, and coastal‐pelagic fishes a left‐skewed RFD.Main conclusions Variation in dispersal capabilities associated with habitat isolation and fragmentation in the TEP appears to be the main mechanism contributing to differences among RFD structure, although variation in tolerances arising from the dynamic regional environment may contribute to some patterns. Because diversity patterns are strongly affected by RFD structure, it is now evident that the insular and continental components of a fauna should be treated separately when analysing such patterns. Furthermore, contrasts in RFD structure among species using different habitats demonstrate that a full understanding of the causes of diversity patterns requires analyses of complete regional faunas in relation to regional geography.

Range size in mid-domain models of species diversity

Geographical patterns of species diversity have been examined using mid-domain null models, in which the ranges of individual species are simulated by randomly arranging them on a bounded one- or two-dimensional continent. These models have shown that structured patterns in the geographical distribution of biodiversity can arise even under a fully stochastic procedure. In particular, mid-domain models have demonstrated that the random generation of ranges of different sizes and locations can produce a gradient of species diversity similar to the one found in real assemblages, with a peak at the middle of a continent. A less explored feature of mid-domain models is the pattern of range-size frequency distribution. Numerical simulations have provided some insights about the geographic pattern of average range size, but no exploration of the shape of range-size frequency distributions has been carried out. Here I present analytical and numerical models that generate explicit predictions for patterns of range size under the assumptions of mid-domain models of species diversity. Some generalizations include: (1) Mid-domain models predict no geographic gradient of average range size; the mean range size of species occurring at any point on a continent is constant (0.5 of the extent of the continent in the one-dimensional model, 0.25 of the area of the continent in the two-dimensional case); (2) Variance in range size is lowest at the middle of a continent and highest near the corners of a square-shaped continent; (3) The range-size frequency distribution is highly right-skewed at any point of a continent, but the skewness is highest near the corners. Despite their alleged weaknesses, mid-domain models are adequate null models against which real-world patterns can be contrasted.

The mid-domain effect and species richness patterns:what have we learned so far?

If species' ranges are randomly shuffled within a bounded geographical domain free of environmental gradients, ranges overlap increasingly toward the center of the domain, creating a "mid-domain" peak of species richness. This "mid-domain effect" (MDE) has been controversial both in concept and in application. Empirical studies assess the degree to which the evolutionary, ecological, and historical processes that undeniably act on individual species and clades produce geographical patterns that resemble those produced by MDE models. MDE models that resample empirical range size frequency distributions (RSFDs) balance the risk of underestimating and overestimating the role of MDE, whereas theoretical RSFDs are generally biased toward underestimating MDE. We discuss the inclusion of nonendemic species in MDE models, rationales for setting domain limits, and the validity of one- and two-dimensional MDE models. MDE models, though null models, are not null hypotheses to be simplistically rejected or accepted. They are a means of estimating the expected effect of geometric constraints within the context of multiple causality. We call for assessment of MDE on an equal statistical footing with other candidate explanations for richness gradients. Although some critics have categorically dismissed MDE, an overview of the 21 MDE studies published to date reveals a substantial signature of MDE in natural patterns and justifies continued work.

Occupancy frequency distributions: patterns, artefacts and mechanisms.

Numerous hypotheses have been proposed to explain the shape of occupancy frequency distributions (distributions of the numbers of species occupying different numbers of areas). Artefactual effects include sampling characteristics, whereas biological mechanisms include organismal, niche-based and meta-population models. To date, there has been little testing of these models. In addition, although empirically derived occupancy distributions encompass an array of taxa and spatial scales, comparisons between them are often not possible because of differences in sampling protocol and method of construction. In this paper, the effects of sampling protocol (grain, sample number, extent, sampling coverage and intensity) on the shape of occupancy distributions are examined, and approaches for minimising artefactual effects recommended. Evidence for proposed biological determinants of the shape of occupancy distributions is then examined. Good support exists for some mechanisms (habitat and environmental heterogeneity), little for others (dispersal ability), while some hypotheses remain untested (landscape productivity, position in geographic range, range size frequency distributions), or are unlikely to be useful explanations for the shape of occupancy distributions 'species specificity and adaptation to habitat, extinction-colonization dynamics). The presence of a core (class containing species with the highest occupancy) mode in occupancy distributions is most likely to be associated with larger sample units, and small homogenous sampling areas positioned well within and towards the range centers of a sufficient proportion of the species in the assemblage. Satellite (class with species with the lowest occupancy) modes are associated with sampling large, heterogeneous areas that incorporate a large proportion of the assemblage range. However, satellite modes commonly also occur in the presence of a core mode, and rare species effects are likely to contribute to the presence of a satellite mode at most sampling scales. In most proposed hypotheses, spatial scale is an important determinant of the shape of the observed occupancy distribution. Because the attributes of the mechanisms associated with these hypotheses change with spatial scale, their predictions for the shape of occupancy distributions also change. To understand occupancy distributions and the mechanisms underlying them, a synthesis of pattern documentation and model testing across scales is thus needed. The development of null models, comparisons of occupancy distributions across spatial scales and taxa, documentation of the movement of individual species between occupancy classes with changes in spatial scale, as well as further testing of biological mechanisms are all necessary for an improved understanding of the distribution of species and assemblages within their geographic ranges.

TESTING GENERALITIES IN THE SHAPE OF PATCH OCCUPANCY FREQUENCY DISTRIBUTIONS

Bimodal patch occupancy frequency distributions regularly appear in studies of a variety of taxa at a variety of spatial scales. Metapopulation models associated with the core–satellite hypothesis and a strong rescue effect predict the occurrence of this bimodality. However, there have been numerous other explanations, both artifactual and biological, of bimodality in patch occupancy frequency distributions. One approach to a better understanding of the phenomenon is to conduct studies on a variety of taxa and spatial scales. Here we examine patterns of patch occupancy frequency distributions for plants, dung beetles, and birds at four spatial scales in sand forest and mixed woodland in Tembe Elephant Park and Sileza Nature Reserve in South Africa. We then examine the predictions of each alternative mechanism proposed to explain bimodality with respect to the patterns found in this study. The incidence of bimodality was high. Little evidence was found for any of the artifactual or nonmetapopulation-based explanations of bimodality. Although there were no between-taxon differences in the shape of occupancy distributions, there was a clear trend of decreasing frequency of bimodality with an increase in the size of the area examined. The general shape of range-size frequency distributions in combination with an area effect is suggested as a mechanism underlying this pattern. Metapopulation dynamics associated with a strong rescue effect could also not be excluded as a possible mechanism playing a role in generating the bimodality found at the patch scale.

Testing Generalities in the Shape of Patch Occupancy Frequency Distributions

Bimodal patch occupancy frequency distributions regularly appear in studies of a variety of taxa at a variety of spatial scales. Metapopulation models associated with the core–satellite hypothesis and a strong rescue effect predict the occurrence of this bimodality. However, there have been numerous other explanations, both artifactual and biological, of bimodality in patch occupancy frequency distributions. One approach to a better understanding of the phenomenon is to conduct studies on a variety of taxa and spatial scales. Here we examine patterns of patch occupancy frequency distributions for plants, dung beetles, and birds at four spatial scales in sand forest and mixed woodland in Tembe Elephant Park and Sileza Nature Reserve in South Africa. We then examine the predictions of each alternative mechanism proposed to explain bimodality with respect to the patterns found in this study. The incidence of bimodality was high. Little evidence was found for any of the artifactual or nonmetapopulation-based explanations of bimodality. Although there were no between-taxon differences in the shape of occupancy distributions, there was a clear trend of decreasing frequency of bimodality with an increase in the size of the area examined. The general shape of range-size frequency distributions in combination with an area effect is suggested as a mechanism underlying this pattern. Metapopulation dynamics associated with a strong rescue effect could also not be excluded as a possible mechanism playing a role in generating the bimodality found at the patch scale.

Flight areas of British butterflies: assessing species status and decline

Geographical range size is a key ecological variable, but the consequences of measuring range size in different ways are poorly understood. We use high-resolution population data from British butterflies to demonstrate that conventional distribution maps, widely used by conservation biologists, grossly overestimate the areas occupied by species and grossly underestimate decline. The approximate flight areas occupied by 20 out of 45 colonial British species were estimated to cover a median of only 1.44% of the land surface within occupied regions. Common species were found to be declining faster than conventional distribution maps suggest: common and rare species had no significant difference in their population-level rates of extinction. This, combined with the log-normal form of the range-size frequency distribution, implies that species-level extinction rates may accelerate in the medium to long term. Population-level conservation is a matter of great urgency for all species, not just for the rarest.