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FIGURE 11 in Calling for a reassessment of rodent diversity in Colombia: description of a new species of Neacomys (Cricetidae: Oryzomyini) from the Magdalena Valley, with a new phylogenetic hypothesis for the genus and comments on its diversification

FIGURE 11. Selected qualitative morphological traits differentiating Trans-Andean species of Neacomys present in Colombia: A), C), E) Neacomys sp. nov. B), D), F) N. tenuipes. Note the tegmen tympani exposed above the dorsal surface of the ectotympanic, the opened ectotympanic ring, the larger orbicular apophysis of the malleus, the bifurcated paraoccipital process, the deeper folds of the upper molars, and the divided mesoflexus of M1 in the new species. ab: auditory bullae; adeect: anterodorsal edge of the ectotympanic; hyp: hypoflexus; hp: hamular process of the squamosal; M1 prc: procingulum of first upper molar; mes: mesoflexus; met: metaflexus; mt: mastoid tubercle; oa: orbicular apophysis; par: paraflexus; pgf: postglenoid foramen; pp: paraoccipital process; pro: protoflexus; ssf: subsquamosal fenestra; tt: tegmen tympani.

FIGURE 2 in Calling for a reassessment of rodent diversity in Colombia: description of a new species of Neacomys (Cricetidae: Oryzomyini) from the Magdalena Valley, with a new phylogenetic hypothesis for the genus and comments on its diversification

FIGURE 2. Map showing collection localities of the named and unamed small-bodied species of Neacomys (except N. pictus) along with salient topographic features of northern South America. Localities of N. tenuipes and Neacomys sp. nov are numbered according to Appendix I. Localities of the remaining species are detailed in Voss et al. (2001), Sánchez-Vendizú et al. (2018), and Semedo et al. (2020). Miniature in the right side of the map corresponds to the Bayesian Cytb phylogeny recovered here (FIGURE 3): species in the map and phylogeny are same colored; the numbers in some nodes correspond to the median divergence times estimated by Upham et al. (2019) within Neacomys.

FIGURE 1 in Calling for a reassessment of rodent diversity in Colombia: description of a new species of Neacomys (Cricetidae: Oryzomyini) from the Magdalena Valley, with a new phylogenetic hypothesis for the genus and comments on its diversification

FIGURE 1. Map of collecting localities of the specimens of Neacomys sp. nov. (stars) 1. Colombia, Department of Santander, Municipality of El Carmen de Chucurí, vereda La Bodega, sector Manchurrias, western slope of the Serranía de los Yariguíes, Finca Buenos Aires, in the basin of the La San Guillerma stream (Type locality); 2. Colombia, Department of Santander, Municipality of Hato, vereda Hoya Negra. Numbers also correspond to the localities listed in Appendix I. Dashed lines represent boundaries of the municipalities encompassing the two localities. Green polygon represents the area comprised by the Serranía de los Yariguíes National Natural Park.

FIGURE 9 in Calling for a reassessment of rodent diversity in Colombia: description of a new species of Neacomys (Cricetidae: Oryzomyini) from the Magdalena Valley, with a new phylogenetic hypothesis for the genus and comments on its diversification

FIGURE 9. Comparison of the dorsal and ventral views of the skulls of Trans-Andean species of Neacomys present in Colombia: A), C) Neacomys sp. nov. (UIS-MHN-M 1068, holotype). B), D) Neacomys tenuipes (UIS-MHN-M 1723). Note the nasals more expanded anteriorly, the shallower zygomatic notches, the less developed supraorbital ridges, the procingulum of M1 flattened laterally, and the incisive foramina extending almost to anterior alveolus of M1 in Neacomys sp. nov. if: incisive foramen; M1 prc: procingulum of first upper molar; nas: nasal bone; sob: supraorbital beads; zn: zygomatic notch.

opencc-zeroFeb 2021View details →

Species distribution, hybridization and connectivity in the genus Chionodraco: unveiling unknown icefish diversity in Antarctica

Aim The species of the genus Chionodraco (Notothenioidei) are the most abundant icefish on the continental shelf of the Weddell Sea. While previous studies indicated that only Chionodraco hamatus and Chionodraco myersi inhabit the Weddell Sea, the third Chionodraco species, Chionodraco rastrospinosus, was recently sampled in the area. Since C. rastrospinosus is supposed to be found only at the Antarctic Peninsula and Scotia Arc, this study aimed at confirming the species classification of C. rastrospinosus by molecular methods and identifying its putative source population. Given the documented evidence of introgression among the three species, we tested whether the newly found C. rastrospinosus shared any genetic variability with the other Chionodraco species. To explain the pattern of distribution of the Chionodraco species, we aimed at estimating the hydrodynamic connectivity between the Antarctic Peninsula and the Weddell Sea. Location Antarctic Peninsula, southern Scotia Arc and the south-eastern Weddell Sea Methods We genotyped 19 microsatellites and sequenced the mitochondrial D-loop for 560 Chionodraco individuals. We simulated the dispersal of more than 3 million drifters (Lagrangian model). Results The molecular analyses support the presence of C. rastrospinosus in the Weddell Sea and its homogeneity with C. rastrospinosus from the Antarctic Peninsula. Bayesian clustering identifies three putative hybrids among C. rastrospinosus and the other congenerics. Lagrangian simulations do not support connectivity driven by the oceanographic features of the Antarctic Peninsula and Weddell Sea via passive larval dispersal only. Main conclusions This study documents, for the first time, the presence of C. rastrospinosus in the Weddell Sea unveiling more biodiversity than previously known in this region. The sympatry of the three Chionodraco species explains the occurrence of occasional, ongoing events of hybridization in the genus. Alternative possible hypotheses need to be tested in future studies about the mechanisms maintaining the interspecific connectivity in Chionodraco spp.

opencc-zeroDec 2013View details →

Data from: Positive relationship between genetic- and species diversity on limestone outcrops in the Carpathian Mountains

We asked if the genetic diversity of Saponaria bellidifolia (a habitat specialist plant) and the species diversity of its habitat are driven by parallel landscape-level processes in an island-like system of limestone outcrops in the Carpathian Mountains. We tested the relationship of these two diversity levels at local and regional geographic scales. Local genetic and species diversity showed parallel patterns influenced by the number of plant communities. Likewise, at regional level there was strong evidence for parallel equilibrial dynamics of genotypes and species. However, a superimposed matrix effect enhanced the regional species diversity only. Genetic diversity of habitat specialist organisms and species diversity of these limestone outcrop islands on mainland are modulated by parallel landscape-level processes at different geographic scales, and mechanisms may be identified at very high spatial resolutions.

opencc-zeroSep 2019View details →

Data from: Environmental factors explain the spatial mismatches between species richness and phylogenetic diversity of terrestrial mammals

Aim: Explore the spatial variation of the relationships between species richness (SR), phylogenetic diversity (PD) and environmental factors to infer the possible mechanisms underlying patterns of diversity in different regions of the globe. Location: Global. Time period: Present day. Major taxa studied: Terrestrial mammals. Methods: We used a hexagonal grid to map SR and PD of mammals and four environmental factors (temperature, productivity, elevation and climate-change velocity since the Last Glacial Maximum). We related those variables through direct and indirect pathways using a novel combination of Path Analysis and Geographically Weighted Regression to account for spatial non-stationarity of path coefficients. Results: SR, PD and environmental factors relate differently across the geographic space, with most relationships varying in both, magnitude and direction. Species richness is associated with lower phylogenetic diversity in much of the tropics and in the Americas, which reflects the tropical origin and the recent diversification of some mammalian clades in these regions. Environmental effects on PD are predominantly mediated by their effects on SR. But once richness is controlled for, the relationships between environmental factors and PD (i.e. PDSR) highlight environmentally driven changes in species composition. Environmental-PDSR relationships suggest that the relative importance of different mechanisms driving biodiversity shifts spatially. Across most of the globe, temperature and productivity are the strongest predictors of richness, while PDSR is best predicted by temperature. Main conclusions: Richness explains most spatial variation in PD, but both dimensions of biodiversity respond differently to environmental conditions across the globe, as indicated by the spatial mismatches in the relationships between environmental factors and these two types of diversity. We show that accounting for spatial non-stationarity and environmental effects on PD while controlling for richness uncovers a more complex scenario of drivers of biodiversity than previously observed.

opencc-zeroDec 2015View details →

Data from: In the shadows: phylogenomics and coalescent species delimitation unveil cryptic diversity in a Cerrado endemic lizard (Squamata: Tropidurus)

The recognition of cryptic diversity within geographically widespread species is gradually becoming a trend in the highly speciose Neotropical biomes. The statistical methods to recognise such cryptic lineages are rapidly advancing, but have rarely been applied to genomic-scale datasets. Herein, we used phylogenomic data to investigate phylogenetic history and cryptic diversity within Tropidurus itambere, a lizard endemic to the Cerrado biodiversity hotspot. We applied a series of phylogenetic methods to reconstruct evolutionary relationships and a coalescent Bayesian species delimitation approach (BPP) to clarify species limits. The BPP results suggest that the widespread nominal taxon comprises a complex of 5 highly supported and geographically structured cryptic species. We highlight and discuss the different topological patterns recovered by concatenated and coalescent species tree methods for these closely related lineages. Finally, we suggest that the existence of cryptic lineages in the Cerrado is much more common than traditionally thought, highlighting the value of using NGS data and coalescent techniques to investigate patterns of species diversity.

opencc-zeroJun 2021View details →

eDNA metabarcoding in lakes to quantify influences of landscape features and human activity on aquatic invasive species prevalence and fish community diversity

Aim: Our goal was to use eDNA metabarcoding to characterize fish community diversity, detect aquatic invasive species (AIS), and assess how measures of community (or AIS) diversity are influenced by lake physical and environmental covariates, measures of hydrological connectivity, and human accessibility. Location: Michigan, USA. Methods: eDNA samples collected from 22 lakes were sequenced using two mitochondrial gene regions (12S and 16S rRNA). Metabarcoding data were compared to traditional fisheries survey data for a subset of lakes, and data from all 22 lakes were combined with environmental information to identify significant associations with community diversity and AIS relative abundance. Results: Occupancy modeling indicated that detection probabilities were generally higher with eDNA than traditional fisheries gear. Measures of connectivity with upstream aquatic habitats were positively associated with both AIS relative abundance and fish species diversity. We also demonstrate the use of spatial interpolation methods to map distributions of species diversity and AIS relative abundance within lakes. Conclusions: eDNA metabarcoding methods provided information on the composition and diversity of fish assemblages and the presence of AIS in freshwater lakes that varied greatly in drainage connectivity and anthropogenic development. Our case study identified associations between environmental covariates and fish diversity or AIS relative abundance across lakes. This information is of particular importance given increasing anthropogenic disturbance, invasive species spread, and associated declines in aquatic biodiversity. Incorporating eDNA metabarcoding as a supplement to traditional fisheries surveys will permit managers to identify greater numbers of taxa, including early detection of AIS, with less field effort and fish mortality. Further, eDNA methods may more accurately identify physical and biological features that correlate with diversity and abundance, and allow agencies to more effectively direct AIS management activities.

opencc-zeroDec 2012View details →

Data from: Rapid buildup of genetic diversity in founder populations of the gynodioecious plant species Origanum vulgare after semi-natural grassland restoration

In most landscapes the success of habitat restoration is largely dependent on spontaneous colonization of plant species. This colonization process, and the outcome of restoration practices, can only be considered successful if the genetic makeup of founding populations is not eroded through founder effects and subsequent genetic drift. Here we used 10 microsatellite markers to investigate the genetic effects of recent colonization of the long-lived gynodioecious species Origanum vulgare in restored semi-natural grassland patches. We compared the genetic diversity and differentiation of fourteen recent populations with that of thirteen old, putative source populations, and we evaluated the effects of spatial configuration of the populations on colonization patterns. We did not observe decreased genetic diversity in recent populations, or inflated genetic differentiation among them. Nevertheless, a significantly higher inbreeding coefficient was observed in recent populations, although this was not associated with negative fitness effects. Overall population genetic differentiation was low (FST = 0.040). Individuals of restored populations were assigned to on average 6.1 different source populations (likely following the 'migrant pool' model). Gene flow was, however, affected by the spatial configuration of the grasslands, with gene flow into the recent populations mainly originating from nearby source populations. This study demonstrates how spontaneous colonization after habitat restoration can lead to viable populations in a relatively short time, overcoming pronounced founder effects, when several source populations are nearby. Restored populations can therefore rapidly act as stepping stones and sources of genetic diversity, likely increasing overall metapopulation viability of the study species.

Data from: Tropical tree species diversity in a mountain system in southern Mexico: local and regional patterns and determinant factors

Mechanisms explaining patterns of biodiversity along elevation gradients in tropical mountain systems remain controversial. We use a set of climatic, topographic and soil variables encompassing regional, landscape and local-level spatial scales to explain the spatial variation of tree species diversity in the Sierra Madre of Chiapas, Mexico. We sampled 128 circular plots (0.1-ha each) in four elevational bands along four elevation gradients or transects encompassing 100-2200 m. A total of 12,533 trees belonging to 444 species were recorded. Diversity patterns along the elevation gradient and the explanatory power of independent variables were dependent on spatial scale (regional vs transect) and functional group (total vs late-successional or pioneer species). Diversity of all species and late-successional species (1 - proportion of pioneer species) showed a constant pattern at the regional and transect scales, with low predictive power of climatic variables and/or elevation. A linear decrease of either number or proportion of pioneer species diversity was observed with increasing elevation, which was correlated with temperature, rainfall, and human disturbance trends. Total species diversity showed an increase with rainfall of the warmest quarter, indicating a regional-level limiting effect of seasonality (drought duration). Yet the explanatory power of climatic and topographic variables was higher at the individual transect level than at the regional scale, suggesting the parallel but differential influence of evolutionary and geological history factors on diversification not so far studied to explain elevation patterns of species diversity in tropical mountain systems.

opencc-zeroDec 2015View details →

Data from: Effects of species diversity on fine root productivity increase with stand development and associated mechanisms in a boreal forest

There is a growing interest in understanding the relationship between diversity and below-ground productivity due to the critical contribution of below-ground systems to overall terrestrial productivity. Yet, the temporal (seasonal and developmental) changes in diversity effects on below-ground productivity and their underlying mechanisms remain unclear. We hypothesized that (i) diversity effects on fine root productivity increase with stand development, and (ii) increased diversity effects associated with stand development result from augmented horizontal soil space utilization, increased forest floor depth for rooting, enhanced effects in nutrient-poor soil layers and/or foraging towards high nutrient availability. We investigated the effects of tree species diversity on fine root productivity by sampling 18 stands dominated by single species and their mixtures in post-fire boreal forests of two stand ages (8 and 34 years following stand-replacing fire). Species evenness was significantly higher in species mixtures than in single-species-dominated stands at both age classes, while species richness did not differ across stand types and age classes. We found that the annual fine root production was higher in mixtures than the mean of single-species-dominated stands in both stand ages, with a significantly higher magnitude of effects in the 34-year-old than 8-year-old stands. Mixtures had higher horizontal soil volume filling than single-species-dominated stands with a more pronounced increase in the 34-year-old than 8-year-old stands. Compared with the 8-year-old stands, the 34-year-old stands had increased forest floor depth and greater overyielding with soil depth, and their fine root productivity was more responsive to the vertical variation in soil phosphorus concentrations among soil layers. Synthesis. Our results provide evidence for increasing positive diversity effects on fine root productivity with stand development in heterogeneous natural forests. Moreover, our results indicate that the increased positive diversity effects with stand development was the result of multiple mechanisms, including higher horizontal soil volume filling, a thicker forest floor layer for rooting, a higher magnitude of complementarity in nutrient-poor deep soil layers and stronger nutrient foraging towards soil layers with high nutrient concentrations in older than younger stands.

opencc-zeroDec 2012View details →

Data from: Multilocus phylogeny and Bayesian estimates of species boundaries reveal hidden evolutionary relationships and cryptic diversity in Southeast Asian monitor lizards

Recent conceptual, technological, and methodological advances in phylogenetics have enabled increasingly robust statistical species delimitation in studies of biodiversity. As the variety of evidence purporting species diversity has increased, so too have the kinds of tools and inferential power of methods for delimiting species. Here we showcase an organismal system for a data-rich, comparative molecular approach to evaluating strategies of species delimitation among monitor lizards of the genus Varanus. The water monitors (Varanus salvator Complex), a widespread group distributed throughout Southeast Asia and southern India, have been the subject of numerous taxonomic treatments, which have drawn recent attention due to the possibility of undocumented species diversity. To date, studies of this group have relied on purportedly diagnostic morphological characters, with no attention given to the genetic underpinnings of species diversity. Using a 5-gene dataset, we estimated phylogeny and used multilocus genetic networks, analysis of population structure, and a Bayesian coalescent approach to infer species boundaries. Our results contradict previous systematic hypotheses, reveal surprising relationships between island and mainland lineages, and uncover novel, cryptic evolutionary lineages (i.e. new putative species). Our study contributes to a growing body of literature suggesting that, used in concert with other sources of data (e.g., morphology, ecology, biogeography), multilocus genetic data can be highly informative to systematists and biodiversity specialists when attempting to estimate species diversity and identify conservation priorities. We recommend holding in abeyance taxonomic decisions until multiple, converging lines of evidence are available to best inform taxonomists, evolutionary biologists, and conservationists.

opencc-zeroDec 2015View details →

Data from: Cryptic diversity in a fig wasp community – morphologically differentiated species are sympatric but cryptic species are allopatric

A key debate in ecology centres on the relative importance of niche and neutral processes in determining patterns of community assembly with particular focus on whether ecologically similar species with similar functional traits are able to coexist. Meanwhile, molecular studies are increasingly revealing morphologically indistinguishable cryptic species with presumably similar ecological roles. Determining the geographic distribution of such cryptic species provides opportunities to contrast predictions of niche versus neutral models. Discovery of sympatric cryptic species increases alpha diversity and supports neutral models, while documentation of allopatric/parapatric cryptic species increases beta diversity and supports niche models. We tested these predictions using morphological and molecular data, coupled with environmental niche modelling analyses, of a fig wasp community along its 2700 km latitudinal range. Molecular methods increased previous species diversity estimates from eight to eleven species, revealing morphologically cryptic species in each of the four wasp genera studied. Congeneric species pairs that were differentiated by a key morphological functional trait (ovipositor length) coexisted sympatrically over large areas. In contrast, morphologically similar species, with similar ovipositor lengths, typically showed parapatric ranges with very little overlap. Despite parapatric ranges, environmental niche models of cryptic congeneric pairs indicate large regions of potential sympatry, suggesting that competitive processes are important in determining the distributions of ecologically similar species. Niche processes appear to structure this insect community and cryptic diversity may typically contribute mostly to beta rather than alpha diversity.

opencc-zeroDec 2012View details →

Data from: Patterns of selection and allele diversity of class I and class II major histocompatibility loci across the species range of sockeye salmon (Oncorhynchus nerka)

The major histocompatibility complex (MHC), an important component of the vertebrate immune system, provides an important suite of genes to examine the role of genetic diversity at non-neutral loci for population persistence. We contrasted patterns of diversity at the two classical MHC loci in sockeye salmon (Oncorhynchus nerka), MHC class I (UBA) and MHC class II (DAB), and neutral microsatellite loci across 70 populations spanning the species range from Washington State to Japan. There was no correlation in allelic richness or heterozygosity between MHC loci or between MHC loci and microsatellites. The two unlinked MHC loci may be responding to different selective pressures; the distribution of FST values for the two loci was uncorrelated, and evidence for both balancing and directional selection on alleles and lineages of DAB and UBA was observed in populations throughout the species range but rarely on both loci within a population. These results suggest that fluctuating selection has resulted in the divergence of MHC loci in contemporary populations.

Data from: Genetic diversity is largely unpredictable but scales with museum occurrences in a species-rich clade of Australian lizards

Genetic diversity is a fundamental characteristic of species and is affected by many factors, including mutation rate, population size, life history and demography. To better understand the processes that influence levels of genetic diversity across taxa, we collected genome-wide restriction-associated DNA data from more than 500 individuals spanning 76 nominal species of Australian scincid lizards in the genus Ctenotus. To avoid potential biases associated with variation in taxonomic practice across the group, we used coalescent-based species delimitation to delineate 83 species-level lineages within the genus for downstream analyses. We then used these genetic data to infer levels of within-population genetic diversity. Using a phylogenetically informed approach, we tested whether variation in genetic diversity could be explained by population size, environmental heterogeneity or historical demography. We find that the strongest predictor of genetic diversity is a novel proxy for census population size: the number of vouchered occurrences in museum databases. However, museum occurrences only explain a limited proportion of the variance in genetic diversity, suggesting that genetic diversity might be difficult to predict at shallower phylogenetic scales.

Data from: Evolution of sociality in spiders leads to depleted genomic diversity at both population and species level

Across several animal taxa, the evolution of sociality involves a suite of characteristics, a 'social syndrome', that includes cooperative breeding, reproductive skew, primary female biased sex-ratio, and the transition from outcrossing to inbreeding mating system, factors that are expected to reduce effective population size (Ne). This social syndrome may be favoured by short-term benefits but come with long-term costs, because the reduction in Ne amplifies loss of genetic diversity by genetic drift, ultimately restricting the potential of populations to respond to environmental change. To investigate the consequences of this social life form on genetic diversity, we used a comparative RAD-sequencing approach to estimate genome-wide diversity in spider species that differ in level of sociality, reproductive skew, and mating system. We analysed multiple populations of three independent sister-species pairs of social inbreeding and subsocial outcrossing Stegodyphus spiders, and a subsocial outgroup. Heterozygosity and within population diversity were 6-10 fold lower in social compared to subsocial species, and demographic modelling revealed a tenfold reduction in Ne of social populations. Species-wide genetic diversity depends on population divergence and the viability of genetic lineages. Population genomic patterns were consistent with high lineage turnover, which homogenizes the genetic structure that builds up between inbreeding populations, ultimately depleting genetic diversity at the species level. Indeed, species-wide genetic diversity of social species was 5-8 times lower than that of subsocial species. The repeated evolution of species with this social syndrome is associated with severe loss of genome-wide diversity, likely to limit their evolutionary potential.

Data from: Mitochondrial genomes of Australian chicken Eimeria support the presence of ten species with low genetic diversity among strains

Modern molecular approaches have vastly improved diagnostic capabilities for differentiating among species of chicken infecting Eimeria. Consolidating information from multiple genetic markers, adding additional poultry Eimeria species and increasing the size of available data-sets is improving the resolving power of the DNA, and consequently our understanding of the genus. This study adds information from 25 complete mitochondrial DNA genomes from Australian chicken Eimeria isolates representing all 10 species known to occur in Australia, including OTU-X, −Y and −Z. The resulting phylogeny provides a comprehensive view of species relatedness highlighting where the OTUs align with respect to others members of the genus. All three OTUs fall within the Eimeria clade that contains only chicken-infecting species with close affinities to E. maxima, E. brunetti and E. mitis. Mitochondrial genetic diversity was low among Australian isolates likely reflecting their recent introduction to the country post-European settlement. The lack of observed genetic diversity is a promising outcome as it suggests that the currently used live vaccines should continue to offer widespread protection against Eimeria outbreaks in all states and territories. Flocks were frequently found to host multiple strains of the same species, a factor that should be considered when studying disease epidemiology in the field.

Data from: 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.