Rare edges and abundant cores: range-wide variation in abundance in North American birds.

Understanding how the abundance of species varies across geographical ranges is central to ecology; however, few studies test hypotheses using detailed abundance estimates across the full ranges of species on a continental scale. Here, we use unprecedented, detailed estimates of breeding abundance for North American birds (eBird) to test two hypotheses for how abundance varies across species' ranges. We find widespread support for the rare-edge hypothesis-where the abundance of species declines near the range edge-reflecting both reduced occurrence and lower local abundance near range edges. By contrast, we find mixed support for the abundant-centre hypothesis-where the abundance of species peaks in the centre of the range and declines towards the edges-with limited support in conservative tests within species, but general support in among-species tests that control for unbalanced sampling and consider a broader definition of the range centre. Overall, results are consistent with a gradual decline in suitable conditions and increase in challenge towards the range edge that eventually limit the ability of populations to persist.

Southern Africa's Great Escarpment as an amphitheater of climate-driven diversification and a buffer against future climate change in bats.

Hosting 1460 plant and 126 vertebrate endemic species, the Great Escarpment (hereafter, Escarpment) forms a semi-circular "amphitheater" of mountains girdling southern Africa from arid west to temperate east. Since arid and temperate biota are usually studied separately, earlier studies overlooked the biogeographical importance of the Escarpment as a whole. Bats disperse more widely than other mammalian taxa, with related species and intraspecific lineages occupying both arid and temperate highlands of the Escarpment, providing an excellent model to address this knowledge gap. We investigated patterns of speciation and micro-endemism from modeled past, present, and future distributions in six clades of southern African bats from three families (Rhinolophidae, Cistugidae, and Vespertilionidae) having different crown ages (Pleistocene to Miocene) and biome affiliations (temperate to arid). We estimated mtDNA relaxed clock dates of key divergence events across the six clades in relation both to biogeographical features and patterns of phenotypic variation in crania, bacula and echolocation calls. In horseshoe bats (Rhinolophidae), both the western and eastern "arms" of the Escarpment have facilitated dispersals from the Afrotropics into southern Africa. Pleistocene and pre-Pleistocene "species pumps" and temperate refugia explained observed patterns of speciation, intraspecific divergence and, in two cases, mtDNA introgression. The Maloti-Drakensberg is a center of micro-endemism for bats, housing three newly described or undescribed species. Vicariance across biogeographic barriers gave rise to 29 micro-endemic species and intraspecific lineages whose distributions were congruent with those identified in other phytogeographic and zoogeographic studies. Although Köppen-Geiger climate models predict a widespread replacement of current temperate ecosystems in southern Africa by tropical or arid ecosystems by 2070-2100, future climate Maxent models for 13 bat species (all but one of those analyzed above) showed minimal range changes in temperate species from the eastern Escarpment by 2070, possibly due to the buffering effect of mountains to climate change.

Rapid range shifts in African Anopheles mosquitoes over the last century.

Facing a warming climate, many tropical species-including the arthropod vectors of several infectious diseases-will be displaced to higher latitudes and elevations. These shifts are frequently projected for the future, but rarely documented in the present day. Here, we use one of the most comprehensive datasets ever compiled by medical entomologists to track the observed range limits of African malaria mosquito vectors (Anopheles spp.) from 1898 to 2016. Using a simple regression approach, we estimate that these species' ranges gained an average of 6.5 m of elevation per year, and the southern limits of their ranges moved polewards 4.7 km per year. These shifts would be consistent with the local velocity of recent climate change, and might help explain the incursion of malaria transmission into new areas over the past few decades. Confirming that climate change underlies these shifts, and applying similar methods to other disease vectors, are important directions for future research.

Energetic dissociation of individual and species ranges.

The use of energy is universal to all life forms and all levels of biological organization, potentially linking processes operating at variable scales. Individual and species ranges might be energetically constrained, yet divergent metabolic limitations at both scales can disassociate these individual and species traits. We analysed comparative energetic and range data to unravel the mechanistic basis of the dissociation between individual and species range sizes observed among mammalian species. Our results demonstrate that basal, or maintenance, metabolism negatively correlates with individual ranges, but, at the same time, it positively correlates with species ranges. High aerobic capacity, i.e. maximum metabolic rate, positively correlates with individual ranges, but it is weakly related to species range size. These antagonistic energetic constraints on both ranges could lead to a disassociation between individual and species traits and to a low covariation between home and species range sizes. We show that important organismal functions, such as basal and maximum metabolic rates, have the potential to unravel mechanisms operating at different levels of biological organization and to expose links between energy-dependent processes at different scales.

Scale-dependent effects of niche specialisation: The disconnect between individual and species ranges.

One of the most general expectations of species range dynamics is that widespread species tend to have broader niches. However, it remains unclear how this relationship is expressed at different levels of biological organisation, which involve potentially distinctive processes operating at different spatial and temporal scales. Here, we show that range sizes of terrestrial non-volant mammals at the individual and species level show contrasting relationships with two ecological niche dimensions: diet and habitat breadth. While average individual home range size appears to be mainly shaped by the interplay of diet niche breadth and body mass, species geographical range size is primarily related to habitat niche breadth but not to diet niche breadth. Our findings suggest that individual home range size is shaped by the trade-off between energetic requirements, movement capacity and trophic specialisation, whereas species geographical range size is related to the ability to persist under various environmental conditions.

Species-level drivers of mammalian ectoparasite faunas.

Traditionally, most studies have described the organization of host-parasite interaction networks by considering only few host groups at limited geographical extents. However, host-parasite relationships are merged within different taxonomic groups and factors shaping these interactions likely differ between host and parasite groups, making group-level differences important to better understand the ecological and evolutionary dynamics of these interactive communities. Here we used a dataset of 629 ectoparasite species and 251 species of terrestrial mammals, comprising 10 orders distributed across the Nearctic and Neotropical regions of Mexico to assess the species-level drivers of mammalian ectoparasite faunas. Specifically, we evaluated whether body weight, geographical range size and within-range mammal species richness (i.e. diversity field) predict mammal ectoparasite species richness (i.e. degree centrality) and their closeness centrality within the mammal-ectoparasite network. In addition, we also tested if the observed patterns differ among mammal orders and if taxonomic closely related host mammals could more likely share the same set of ectoparasites. We found that ectoparasite species richness of small mammals (mainly rodents) with large proportional range sizes was high compared to large-bodied mammals, whereas the diversity field of mammals had no predictive value (except for bats). We also observed that taxonomic proximity was a main determinant of the probability to share ectoparasite species. Specifically, the probability to share ectoparasites in congeneric species reached up to 90% and decreased exponentially as the taxonomic distance increased. Further, we also detected that some ectoparasites are generalists and capable to infect mammalian species across different orders and that rodents have a remarkable role in the network structure, being closely connected to many other taxa. Hence, because many rodent species have synanthropic habits they could act as undesired reservoirs of disease agents for humans and urban animals. Considering the reported worldwide phenomenon of the proliferation of rodents accompanying the demographic decrease or even local extinction of large-bodied mammal species, these organisms may already be an increasing health threat in many regions of the world.

Should I Stay or Should I Go: Partially Sedentary Populations Can Outperform Fully Dispersing Populations in Response to Climate-Induced Range Shifts.

Global mean temperatures have increased by 0.72 [Formula: see text]C since the 1950s, and climate warming is resulting in geographical shifts in the range limits of many species. Climate velocity is estimated to be 0.42 km/year, and if a species fails to adapt to the new climate, it must track the location of its climatically constrained niche in order to survive. Dispersal has an important role to play in enabling a population to shift is geographical range limits, but many species are partially sedentary, with only a fraction of the population dispersing each year. We ask, can partially sedentary populations keep pace with climate or will such populations be more vulnerable to extinction? Through the development of a moving-habitat integrodifference equation model, we show that, provided climate velocity is not too large, partially sedentary populations can outperform fully dispersing populations in one of two ways: (i) by persisting at climate speeds where a fully dispersing population cannot, and (ii) exhibiting higher population densities. Moreover, we find that positive density-dependent dispersal can further improve the likelihood a population can persist. Our results highlight the positive role that non-dispersers may play in mitigating the effects of overdispersal and facilitating population persistence in a warming world.

Two Clonal Species of Phytophthora Associated to Solanaceous Crops Coexist in Central and Southern Colombia.

In Colombia, late blight is considered one of the most limiting diseases on potato and tomato production. Recently, a new Phytophthora species, P. betacei, was described infecting tree tomato crops in the south of Colombia. However, the distribution and the host range of this new emerging pathogen in the country are unknown. The main aims of this study were to determine if this novel species is confined to the south of Colombia, to assess if P. betacei represents a genetically uniform clone across Colombia and to determine if in all regions there is a clear differentiation between the two Phytophthora species. Therefore, we characterized Phytophthora isolates obtained from tree tomato and potato crops in a central region of Colombia and compared them with the strains from the south. Initially, we evaluated the genetic differentiation among Phytophthora strains obtained from tree tomato and potato crops using simple sequence repeat markers. Results showed a strong population structure between P. infestans and P. betacei. However, we did not detect any genetic differentiation within P. infestans or P. betacei populations from different regions. Furthermore, we detected significant morphological differences among the species based on growth and sporangial morphology measurements. We also showed that strains of Phytophthora spp. are predominantly of the A1 mating type and belong to EC-1 and EC-3 clonal lineages for P. infestans and P. betacei, respectively. Our results describe the expanded geographical range of the new species of P. betacei in the central region of Colombia.

What factors explain the geographical range of mammalian parasites?

Free-living species vary substantially in the extent of their spatial distributions. However, distributions of parasitic species have not been comprehensively compared in this context. We investigated which factors most influence the geographical extent of mammal parasites. Using the Global Mammal Parasite Database we analysed 17 818 individual geospatial records on 1806 parasite species (encompassing viruses, bacteria, protozoa, arthropods and helminths) that infect 396 carnivore, ungulate and primate host species. As a measure of the geographical extent of each parasite species we quantified the number and area of world ecoregions occupied by each. To evaluate the importance of variables influencing the summed area of ecoregions occupied by a parasite species, we used Bayesian network analysis of a subset ( n = 866) of the parasites in our database that had at least two host species and complete information on parasite traits. We found that parasites that covered more geographical area had a greater number of host species, higher average phylogenetic relatedness between host species and more sampling effort. Host and parasite taxonomic groups had weak and indirect effects on parasite ecoregion area; parasite transmission mode had virtually no effect. Mechanistically, a greater number of host species probably increases both the collective abundance and habitat breadth of hosts, providing more opportunities for a parasite to have an expansive range. Furthermore, even though mammals are one of the best-studied animal classes, the ecoregion area occupied by their parasites is strongly sensitive to sampling effort, implying mammal parasites are undersampled. Overall, our results support that parasite geographical extent is largely controlled by host characteristics, many of which are subsumed within host taxonomic identity.

Reconstructing geographical parthenogenesis: effects of niche differentiation and reproductive mode on Holocene range expansion of an alpine plant.

Asexual taxa often have larger ranges than their sexual progenitors, particularly in areas affected by Pleistocene glaciations. The reasons given for this 'geographical parthenogenesis' are contentious, with expansion of the ecological niche or colonisation advantages of uniparental reproduction assumed most important in case of plants. Here, we parameterized a spread model for the alpine buttercup Ranunculus kuepferi and reconstructed the joint Holocene range expansion of its sexual and apomictic cytotype across the European Alps under different simulation settings. We found that, rather than niche broadening or a higher migration rate, a shift of the apomict's niche towards colder conditions per se was crucial as it facilitated overcoming of topographical barriers, a factor likely relevant for many alpine apomicts. More generally, our simulations suggest potentially strong interacting effects of niche differentiation and reproductive modes on range formation of related sexual and asexual taxa arising from their differential sensitivity to minority cytotype disadvantage.

Extinction risk in extant marine species integrating palaeontological and biodistributional data.

Extinction risk assessments of marine invertebrate species remain scarce, which hinders effective management of marine biodiversity in the face of anthropogenic impacts. To help close this information gap, in this paper we provide a metric of relative extinction risk that combines palaeontological data, in the form of extinction rates calculated from the fossil record, with two known correlates of risk in the modern day: geographical range size and realized thermal niche. We test the performance of this metric-Palaeontological Extinction Risk In Lineages (PERIL)-using survivorship analyses of Pliocene bivalve faunas from California and New Zealand, and then use it to identify present-day hotspots of extinction vulnerability for extant shallow-marine Bivalvia. Areas of the ocean where concentrations of bivalve species with higher PERIL scores overlap with high levels of climatic or anthropogenic stressors should be considered of most immediate concern for both conservation and management.

Non-volant mammals from the Upper Paraná River Basin: a data set from a critical region for conservation in Brazil.

The data set represents the first attempt at a large-scale inventory of non-volant mammals, with potential applications to performing macroecological studies, developing conservation strategies, and undertaking population and community ecology research, but also to evaluate the ecological consequences of fragmentation and defaunation. Our objectives for compiling these data were to summarize information about inventories of non-volant mammals in the critically important area of the Upper Paraná River Basin by focusing on species richness and index of frequency of occurrence and to identify gaps in knowledge regarding non-volant mammal communities in order to guide future sampling efforts. The data set comprises studies on communities of non-volant mammals from 52 locations covering more than 1,000 km2 and comprises portion of four Brazilian states in the Upper Paraná River Basin. We listed 81 species of non-volant mammals distributed among 58 genera, 22 families, and 9 orders. Rodentia (28 species) was the richest order, followed by Carnivora (17 spp.) and Didelphimorphia (15 spp.). The richest family was Cricetidae (20 spp.), followed by Didelphidae (15 spp.), and Dasypodidae and Felidae (six spp.). Considering national conservation status, one species are considered endangered and 16 vulnerable. Considering global conservation status, 7 species are considered vulnerable, 10 are considered near threatened, and 6 are data deficient. According to the index of frequency of occurrence, Myrmecophaga tridactyla was the most frequent species, occurring at 88.64% of all sites, while 25 species were considered very restricted, occurring in just 2.56% of all sites. In general, the non-volant mammal fauna was composed of mainly very restricted (VR, 25 species) and localized species (L, 25 species), which account for 61.7% of the known species, while 38.3% are restricted (R, 8 species), common (C, 16 species), and widespread (W, 7 species). Seven marsupials and five small rodents had their distributions extended in the central-south of Brazil. All of these species are considered data deficient or threatened, which highlights the importance of these records. No copyright restrictions are associated with the use of this data set. Please cite this data paper when the data are used in publications. We also request that researchers and teachers inform us of how they are using the data.

Linking macroecology and community ecology: refining predictions of species distributions using biotic interaction networks.

Macroecological models for predicting species distributions usually only include abiotic environmental conditions as explanatory variables, despite knowledge from community ecology that all species are linked to other species through biotic interactions. This disconnect is largely due to the different spatial scales considered by the two sub-disciplines: macroecologists study patterns at large extents and coarse resolutions, while community ecologists focus on small extents and fine resolutions. A general framework for including biotic interactions in macroecological models would help bridge this divide, as it would allow for rigorous testing of the role that biotic interactions play in determining species ranges. Here, we present an approach that combines species distribution models with Bayesian networks, which enables the direct and indirect effects of biotic interactions to be modelled as propagating conditional dependencies among species' presences. We show that including biotic interactions in distribution models for species from a California grassland community results in better range predictions across the western USA. This new approach will be important for improving estimates of species distributions and their dynamics under environmental change.

Clarifying misconceptions of extinction risk assessment with the IUCN Red List.

The identification of species at risk of extinction is a central goal of conservation. As the use of data compiled for IUCN Red List assessments expands, a number of misconceptions regarding the purpose, application and use of the IUCN Red List categories and criteria have arisen. We outline five such classes of misconception; the most consequential drive proposals for adapted versions of the criteria, rendering assessments among species incomparable. A key challenge for the future will be to recognize the point where understanding has developed so markedly that it is time for the next generation of the Red List criteria. We do not believe we are there yet but, recognizing the need for scrutiny and continued development of Red Listing, conclude by suggesting areas where additional research could be valuable in improving the understanding of extinction risk among species.

Three explanations for biodiversity hotspots: small range size, geographical overlap and time for species accumulation. An Australian case study.

To understand the generation and maintenance of biodiversity hotspots, we tested three major hypotheses: rates of diversification, ecological limits to diversity, and time for species accumulation. Using dated molecular phylogenies, measures of species' range size and geographical clade overlap, niche modelling, and lineages-through-time plots of Australian Fabaceae, we compared the southwest Australia Floristic Region (SWAFR; a global biodiversity hotspot) with a latitudinally equivalent non-hotspot, southeast Australia (SEA). Ranges of species (real and simulated) were smaller in the SWAFR than in SEA. Geographical overlap of clades was significantly greater for Daviesia in the SWAFR than in SEA, but the inverse for Bossiaea. Lineage diversification rates over the past 10 Myr did not differ between the SWAFR and SEA in either genus. Interaction of multiple factors probably explains the differences in measured diversity between the two regions. Steeper climatic gradients in the SWAFR probably explain the smaller geographical ranges of both genera there. Greater geographical overlap of clades in the SWAFR, combined with a longer time in the region, can explain why Daviesia is far more species-rich there than in SEA. Our results indicate that the time for speciation and ecological limits hypotheses, in concert, can explain the differences in biodiversity.

Ecological traits influence the phylogenetic structure of bird species co-occurrences worldwide.

The extent to which species' ecological and phylogenetic relatedness shape their co-occurrence patterns at large spatial scales remains poorly understood. By quantifying phylogenetic assemblage structure within geographic ranges of >8000 bird species, we show that global co-occurrence patterns are linked - after accounting for regional effects - to key ecological traits reflecting diet, mobility, body size and climatic preference. We found that co-occurrences of carnivorous, migratory and cold-climate species are phylogenetically clustered, whereas nectarivores, herbivores, frugivores and invertebrate eaters tend to be more phylogenetically overdispersed. Preference for open or forested habitats appeared to be independent from the level of phylogenetic clustering. Our results advocate for an extension of the tropical niche conservatism hypothesis to incorporate ecological and life-history traits beyond the climatic niche. They further offer a novel species-oriented perspective on how biogeographic and evolutionary legacies interact with ecological traits to shape global patterns of species coexistence in birds.

Variation and evolution of sex ratios at the northern range limit of a sexually polymorphic plant.

Gender strategies involve three fundamental sex phenotypes - female, male and hermaphrodite. Their frequencies in populations typically define plant sexual systems. Patterns of sex-ratio variation in a geographical context can provide insight into transitions among sexual systems, because environmental gradients differentially influence sex phenotype fitness. Here, we investigate sex-ratio variation in 116 populations of Sagittaria latifolia at the northern range limit in eastern N. America and evaluate mechanisms responsible for the patterns observed. We detected continuous variation in sex phenotype frequencies from monoecy through subdioecy to dioecy. There was a decline in the frequency and flower production of females in northerly populations, whereas hermaphrodite frequencies increased at the range limit, and in small populations. Tests of a model of sex-ratio evolution, using empirical estimates of fitness components, indicated that the relative female and male contribution of males and hermaphrodites to fitness is closer to equilibrium expectations than female frequencies. Plasticity in sex expression and clonality likely contribute to deviations from equilibrium expectations.

Geographical differentiation in floral traits across the distribution range of the Patagonian oil-secreting Calceolaria polyrhiza: do pollinators matter?

BACKGROUND AND AIMS: The underlying evolutionary processes of pollinator-driven floral diversification are still poorly understood. According to the Grant-Stebbins model speciation begins with adaptive local differentiation in the response to spatial heterogeneity in pollinators. Although this crucial process links the micro- and macroevolution of floral adaptation, it has received little attention. In this study geographical phenotypic variation was investigated in Patagonian Calceolaria polyrhiza and its pollinators, two oil-collecting bee species that differ in body size and geographical distribution. METHODS: Patterns of phenotypic variation were examined together with their relationships with pollinators and abiotic factors. Six floral and seven vegetative traits were measured in 45 populations distributed across the entire species range. Climatic and edaphic parameters were determined for 25 selected sites, 2-16 bees per site of the most frequent pollinator species were captured, and a critical flower-bee mechanical fitting trait involved in effective pollination was measured. Geographical patterns of phenotypic and environmental variation were examined using uni- and multivariate analyses. Decoupled geographical variation between corolla area and floral traits related to the mechanical fit of pollinators was explored using a Mantel test. KEY RESULTS: The body length of pollinators and the floral traits related to mechanical fit were strongly correlated with each other. Geographical variation of the mechanical-fit-related traits was decoupled from variation in corolla size; the latter had a geographical pattern consistent with that of the vegetative traits and was mainly affected by climatic gradients. CONCLUSIONS: The results are consistent with pollinators playing a key role in shaping floral phenotype at a geographical scale and promoting the differentiation of two floral ecotypes. The relationship between the critical floral-fit-related trait and bee length remained significant even in models that included various environmental variables and an allometric predictor (corolla area). The abiotic environment also has an important role, mainly affecting floral size. Decoupled geographical variation between floral mechanical-fit-related traits and floral size would represent a strategy to maintain plant-pollinator phenotypic matching in this environmentally heterogeneous area.

Global patterns of freshwater species diversity, threat and endemism.

AIM: Global-scale studies are required to identify broad-scale patterns in the distributions of species, to evaluate the processes that determine diversity and to determine how similar or different these patterns and processes are among different groups of freshwater species. Broad-scale patterns of spatial variation in species distribution are central to many fundamental questions in macroecology and conservation biology. We aimed to evaluate how congruent three commonly used metrics of diversity were among taxa for six groups of freshwater species. LOCATION: Global. METHODS: We compiled geographical range data on 7083 freshwater species of mammals, amphibians, reptiles, fishes, crabs and crayfish to evaluate how species richness, richness of threatened species and endemism are distributed across freshwater ecosystems. We evaluated how congruent these measures of diversity were among taxa at a global level for a grid cell size of just under 1°. RESULTS: We showed that although the risk of extinction faced by freshwater decapods is quite similar to that of freshwater vertebrates, there is a distinct lack of spatial congruence in geographical range between different taxonomic groups at this spatial scale, and a lack of congruence among three commonly used metrics of biodiversity. The risk of extinction for freshwater species was consistently higher than for their terrestrial counterparts. MAIN CONCLUSIONS: We demonstrate that broad-scale patterns of species richness, threatened-species richness and endemism lack congruence among the six freshwater taxonomic groups examined. Invertebrate species are seldom taken into account in conservation planning. Our study suggests that both the metric of biodiversity and the identity of the taxa on which conservation decisions are based require careful consideration. As geographical range information becomes available for further sets of species, further testing will be warranted into the extent to which geographical variation in the richness of these six freshwater groups reflects broader patterns of biodiversity in fresh water.

Risk of local extinction of Odonata freshwater habitat generalists and specialists.

Understanding the risk of a local extinction in a single population relative to the habitat requirements of a species is important in both theoretical and applied ecology. Local extinction risk depends on several factors, such as habitat requirements, range size of species, and habitat quality. We studied the local extinctions among 31 dragonfly and damselfly species from 1930 to 1975 and from 1995 to 2003 in Central Finland. We tested whether habitat specialists had a higher local extinction rate than generalist species. Approximately 30% of the local dragonfly and damselfly populations were extirpated during the 2 study periods. The size of the geographical range of the species was negatively related to extinction rate of the local populations. In contrast to our prediction, the specialist species had lower local extinction rates than the generalist species, probably because generalist species occurred in both low- and high-quality habitat. Our results are consistent with source-sink theory.