Testing methods in species distribution modelling using virtual species: what have we learnt and what are we missing?

Species distribution models (SDMs) have become one of the major predictive tools in ecology. However, multiple methodological choices are required during the modelling process, some of which may have a large impact on forecasting results. In this context, virtual species, i.e. the use of simulations involving a fictitious species for which we have perfect knowledge of its occurrence–environment relationships and other relevant characteristics, have become increasingly popular to test SDMs. This approach provides for a simple virtual ecologist framework under which to test model properties, as well as the effects of the different methodological choices, and allows teasing out the effects of targeted factors with great certainty. This simplification is therefore very useful in setting up modelling standards and best practice principles. As a result, numerous virtual species studies have been published over the last decade. The topics covered include differences in performance between statistical models, effects of sample size, choice of threshold values, methods to generate pseudo‐absences for presence‐only data, among many others. These simulations have therefore already made a great contribution to setting best modelling practices in SDMs. Recent software developments have greatly facilitated the simulation of virtual species, with at least three different packages published to that effect. However, the simulation procedure has not been homogeneous, which introduces some subtleties in the interpretation of results, as well as differences across simulation packages. Here we 1) review the main contributions of the virtual species approach in the SDM literature; 2) compare the major virtual species simulation approaches and software packages; and 3) propose a set of recommendations for best simulation practices in future virtual species studies in the context of SDMs.     

Water relations of Fino lemon plants on two rootstocks under flooded conditions

Potted two-year-old Fino lemon plants (Citrus limon (L.) Burm. fil.) grafted on two different rootstocks: sour orange (C. aurantium) (SO), and C. macrophylla (CM) were submitted to two different treatments: non-flooded (control) and flooded for 8 days, under field conditions. Lemon/CM plants had lower plant plus soil resistance to water flow (R_(p+s)) values for both treatments. The decrease in leaf water potential (Ψ_l) and leaf turgor potential (Ψ_p) values, observed in the last part of the flooding period, in both flooded scion/rootstock combinations, can be related to the increase in the resistance to water flow. The maintenance in Ψ_l and Ψ_p values in flooded plants at values similar to those of the control plants, at the beginning of the flooding period and during the recovery period, can be ascribed to the stomatal control observed (decrease in leaf conductance (g₁) values). The later g₁ recovery in lemon/CM than in lemon/SO flooded plants, could explain the lower vegetative growth of lemon/CM plants by flooding effect. The observed g₁ response to soil flooding suggested that porometry is a reliable indicator of the altered behaviour caused by flooding in lemon plant.     

Integrating phylogenetic and functional biodiversity facets to guide conservation: a case study using anurans in a global biodiversity hotspot

The Cerrado is one of the most threatened biomes in Brazil, with little spatial representation within the Protected Area network. Recently, proposed conservation plans worldwide have advocated for the use of multiple biodiversity facets to protect unique evolutionary and functional processes. Our aim was to identify areas with high biodiversity representativeness applying this multifaceted perspective, and propose conservation plans based on the joint analysis of taxonomic, functional and phylogenetic diversity. We used a database of the Brazilian National Program for Research in Biodiversity, which employs a standard protocol for sampling tadpoles. The Cerrado database includes samples from 165 water bodies spread over 15 localities, covering most of the Central Brazilian Cerrado. We selected four morphological traits to calculate functional diversity and used a dated phylogeny available in the literature to compute phylogenetic diversity. Our approach selected five priority areas for conservation, one of which is already protected. Our results highlighted the importance of four new areas which show high values of diversity, including original lineages and traits, and urgently need conservation prioritization. Furthermore, unlike the current protected network, our approach performs significantly better than random at protecting sites with high phylogenetic and functional diversity. We therefore discuss how the multifaceted indices considered can help protect key ecosystem functions and evolutionary legacy in anuran communities of the Brazilian Cerrado.     

Evidence of environmental niche differentiation in the striped mouse (Rhabdomys sp.): inference from its current distribution in southern Africa

The aim of this study was to characterize environmental differentiation of lineages within Rhabdomys and provide hypotheses regarding potential areas of contact between them in the Southern African subregion, including the Republic of South Africa, Lesotho, and Namibia. Records of Rhabdomys taxa across the study region were compiled and georeferenced from the literature, museum records, and field expeditions. Presence records were summarized within a 10 × 10 km grid covering the study area. Environmental information regarding climate, topography, land use, and vegetation productivity was gathered at the same resolution. Multivariate statistics were used to characterize the current environmental niche and distribution of the whole genus as well as of three mitochondrial lineages known to occur in southern Africa. Distribution modeling was carried out using MAXENT in order to generate hypotheses regarding current distribution of each taxa and their potential contact zones. Results indicate that the two species within Rhabdomys appear to have differentiated across the precipitation/temperature gradient present in the region from east to west. R. dilectus occupies the wettest areas in eastern southern Africa, while R. pumilio occupies the warmer and drier regions in the west, but also penetrates in the more mesic central part of the region. We provide further evidence of environmental differentiation within two lineages of R. dilectus. Contact zones between lineages appear to occur in areas of strong environmental gradients and topographic complexity, such as the transition zones between major biomes and the escarpment area where a sharp altitudinal gradient separates coastal and plateau areas, but also within more homogeneous areas such as within grassland and savannah biomes. Our results indicate that Rhabdomys may be more specialized than previously thought when considering current knowledge regarding mitochondrial lineages. The genus appears to have differentiated along two major environmental axes in the study region, but results also suggest dispersal limitations and biological interactions having a role in limiting current distribution boundaries. Furthermore, the projection of the potential geographic distribution of the different lineages suggests several contact zones that may be interesting study fields for understanding the interplay between ecological and evolutionary processes during speciation.     

A phylogenetic perspective on the evolution of Mediterranean teleost fishes

The Mediterranean Sea is a highly diverse, highly studied, and highly impacted biogeographic region, yet no phylogenetic reconstruction of fish diversity in this area has been published to date. Here, we infer the timing and geographic origins of Mediterranean teleost species diversity using nucleotide sequences collected from GenBank. We assembled a DNA supermatrix composed of four mitochondrial genes (12S ribosomal DNA, 16S ribosomal DNA, cytochrome c oxidase subunit I and cytochrome b) and two nuclear genes (rhodopsin and recombination activating gene I), including 62% of Mediterranean teleost species plus 9 outgroups. Maximum likelihood and Bayesian phylogenetic and dating analyses were calibrated using 20 fossil constraints. An additional 124 species were grafted onto the chronogram according to their taxonomic affinity, checking for the effects of taxonomic coverage in subsequent diversification analyses. We then interpreted the time-line of teleost diversification in light of Mediterranean historical biogeography, distinguishing non-endemic natives, endemics and exotic species. Results show that the major Mediterranean orders are of Cretaceous origin, specifically ~100-80 Mya, and most Perciformes families originated 80-50 Mya. Two important clade origin events were detected. The first at 100-80 Mya, affected native and exotic species, and reflects a global diversification period at a time when the Mediterranean Sea did not yet exist. The second occurred during the last 50 Mya, and is noticeable among endemic and native species, but not among exotic species. This period corresponds to isolation of the Mediterranean from Indo-Pacific waters before the Messinian salinity crisis. The Mediterranean fish fauna illustrates well the assembly of regional faunas through origination and immigration, where dispersal and isolation have shaped the emergence of a biodiversity hotspot.     

From diversity indices to community assembly processes: a test with simulated data

Ecological theory suggests that spatial distribution of biodiversity is strongly driven by community assembly processes. Thus the study of diversity patterns combined with null model testing has become increasingly common to infer assembly processes from observed distributions of diversity indices. However, results in both empirical and simulation studies are inconsistent. The aim of our study is to determine with simulated data which facets of biodiversity, if any, may unravel the processes driving its spatial patterns, and to provide practical considerations about the combination of diversity indices that would produce significant and congruent signals when using null models. The study is based on simulated species’ assemblages that emerge under various landscape structures in a spatially explicit individual‐based model with contrasting, predefined assembly processes. We focus on four assembly processes (species‐sorting, mass effect, neutral dynamics and competition colonization trade‐off) and investigate the emerging species’ distributions with varied diversity indices (alpha, beta and gamma) measured at different spatial scales and for different diversity facets (taxonomic, functional and phylogenetic). We find that 1) the four assembly processes result in distinct spatial distributions of species under any landscape structure, 2) a broad range of diversity indices allows distinguishing between communities driven by different assembly processes, 3) null models provide congruent results only for a small fraction of diversity indices and 4) only a combination of these diversity indices allows identifying the correct assembly processes. Our study supports the inference of assembly processes from patterns of diversity only when different types of indices are combined. It highlights the need to combine phylogenetic, functional and taxonomic diversity indices at multiple spatial scales to effectively infer underlying assembly processes from diversity patterns by illustrating how combination of different indices might help disentangling the complex question of coexistence.     

Conservation of temperate forest birds in Chile: implications from the study of an isolated forest relict

We characterize the bird assemblage of an isolated relict temperaterainforest (Santa Inés) in terms of its structure, composition anddynamics. The relict character and long-term isolation of this forest remnant,immersed in a matrix of semiarid scrub, provide a natural experiment to assessthe consequences of long-term isolation and forest area reduction for temperatebird species. Bird surveys were conducted in a forest fragment and thesurrounding scrub matrix between April 1999 (austral autumn) and October 2000(austral spring) on a seasonal basis. Within the forest fragment wedistinguished two microhabitat types: creeks and areas far from creeks (i.e.slopes). A total of 36 species were recorded in the study site, of which 21were inside the relict forest. Highest bird abundance and richness wererecorded during winter and spring, and these were always higher in creek plotsrather than slope plots. Comparisons between this assemblage and thosedistributed in the continuous temperate forest (located more than 900km southwards) indicate that this bird assemblage shares moresimilarities, with regard to bird species composition, to southern temperateforest localities and to other relict forests, than to nearer scrub habitats.However, there are eight species, endemic to temperate forests in southern SouthAmerica, missing from our system. In this regard, our analyses indicate thatthese eight endemic species should be of great conservation concern and willlikely be the ones that will go extinct first if the fragmentation of thetemperate forest continues.