Habitat distribution models for intertidal seaweeds: responses to climatic and non‐climatic drivers

Aim Because intertidal organisms often live close to their physiological tolerance limits, they are potentially sensitive indicators of climate‐driven changes in the environment. The goals of this study were to assess the effect of climatic and non‐climatic factors on the geographical distribution of intertidal macroalgae, and to predict future distributions under different climate‐warming scenarios. Location North‐western Iberian Peninsula, southern Europe. Methods We developed distribution models for six ecologically important intertidal seaweed species. Occurrence and microhabitat data were sampled at 1‐km² resolution and analysed with climate variables measured at larger spatial scales. We used generalized linear models and applied the deviance and Bayesian information criterion to model the relationship between environmental variables and the distribution of each target species. We also used hierarchical partitioning (HP) to identify predictor variables with higher independent explanatory power. Results The distributions of Himanthalia elongata and Bifurcaria bifurcata were correlated with measures of terrestrial and marine climate, although in opposite directions. Model projections under two warming scenarios indicated the extinction of the former at a faster rate in the Cantabrian Sea (northern Spain) than in the Atlantic (west). In contrast, these models predicted an increase in the occurrence of B. bifurcata in both areas. The occurrences of Ascophyllum nodosum and Pelvetia canaliculata, species showing rather static historical distributions, were related to specific non‐climatic environmental conditions and locations, such as the location of sheltered sites. At the southernmost distributional limit, these habitats may present favourable microclimatic conditions or provide refuges from competitors or natural enemies. Model performances for Fucus vesiculosus and F. serratus were similar and poor, but several climatic variables influenced the occurrence of the latter in the HP analyses. Main conclusions The correlation between species distributions and climate was evident for two species, whereas the distributions of the others were associated with non‐climatic predictors. We hypothesize that the distribution of F. serratus responds to diverse combinations of factors in different sections of the north‐west Iberian Peninsula. Our study shows how the response of species distributions to climatic and non‐climatic variables may be complex and vary geographically. Our analyses also highlight the difficulty of making predictions based solely on variation in climatic factors measured at coarse spatial scales.     

On the Iberian endemism <Emphasis Type="Italic">Eurylophella iberica</Emphasis> Keffermuller and Da Terra 1978 (Ephemeroptera,Ephemerellidae): current and future potential distributions,and assessment of the effectiveness of the Natura 2000 network on its protection

Eurylophella iberica Keffermüller and Da Terra, 1978 is an endemic insect species of the Iberian Peninsula whose distribution has been poorly studied to date with rather old and scattered records. Here we compiled all existing distribution records and add new records from recent sampling activities. We also used this updated distributional information and environmental data (climate and geology) to estimate both current and future potential distributions in different climate change scenarios. We found that currently ca. 50% of the total Iberian region could present suitable environmental conditions for E. iberica (all the Iberian Peninsula, save the most eastern and Mediterranean areas). However, the potential distributions estimated when considering future climate change scenarios showed a marked reduction in the areas with suitable environmental conditions for the species, especially in the south. The northwest part of the Iberian Peninsula is a crucial zone for the future survival of this endemic species. We also found that most populations that occur in areas with suitable (both current and future) environmental conditions fall outside the Natura 2000 network of protected areas. Our results represent the first attempt to estimate the potential distribution of this endemic species providing important insights for its conservation.     

Geological habitat template overrides late Quaternary climate change as a determinant of range dynamics and phylogeography in some habitat‐specialist water beetles

Aim We investigated the roles of lithology and climate in constraining the ranges of four co‐distributed species of Iberian saline‐habitat specialist water beetles (Ochthebius glaber, Ochthebius notabilis, Enochrus falcarius and Nebrioporus baeticus) across the late Quaternary and in shaping their geographical genetic structure. The aim was to improve our understanding of the effects of past climate changes on the biota of arid Mediterranean environments and of the relative importance of history and landscape on phylogeographical patterns. Location Iberian Peninsula, Mediterranean. Methods We combined species distribution modelling (SDM) and comparative phylogeography. We used a multi‐model inference and model‐averaging approach both for assessment of range determinants (climate and lithology) and for provision of spatially explicit estimates of the species current and Last Glacial Maximum (LGM) potential ranges. Potential LGM distributions were then contrasted with the phylogeographical and population expansion patterns as assessed using mitochondrial DNA sequence data. We also evaluated the relative importance of geographical distance, habitat resistance and historical isolation for genetic structure in a causal modelling framework. Results Lithology poses a strong constraint on the distribution of Iberian saline‐habitat specialist water beetles, with a variable, but generally moderate, additional influence by climate. The degree to which potential LGM distributions were reduced and fragmented decreased with increasing importance of lithology. These SDM‐based suitability predictions were mostly congruent with phylogeographical and population genetic patterns across the study species, with stronger geographical structure in the genetic diversity of the more temperature‐sensitive species (O. glaber and E. falcarius). Furthermore, while historical isolation was the only factor explaining genetic structure in the more temperature‐sensitive species, lithology‐controlled landscape configuration also played an important role for those species with more lithology‐determined ranges (O. notabilis and N. baeticus). Main conclusions Our data show that lithology is an important constraint on the distribution and range dynamics of endemic Iberian saline‐habitat water beetles, in interaction with climate and long‐term climate change, and overrides the latter in importance for some species. Hence, geological landscape structure and long‐term history may codetermine the overall range and the distribution of genetic lineages in endemic species with specialized edaphic requirements.     

Habitat type mediates equilibrium with climatic conditions in the distribution of Iberian diving beetles

Aim The contrasting habitat permanence over geological time‐scales of lotic and lentic habitats may impose different constraints on the dispersal ability of their macroinvertebrate populations, and ultimately on the degree of equilibrium with current climate. We aim to test for differences between species typical of either habitat type in their potential versus realized distributions as a surrogate measure of degree of climate equilibrium, both in refuges and more recently deglaciated areas. Location Western Europe. Methods We focus on 99 Iberian diving beetles (family Dytiscidae). A multidimensional envelope procedure was used to estimate their potential distributions, which were projected for different spatial scales. At the continental scale we calculated the percentage of countries with climatically suitable conditions for each species over those actually occupied (range filling). At the regional scale, we estimated realized distributions using: (1) convex hull polygons for Sweden and the Iberian Peninsula; and (2) generalized linear models for the Iberian Peninsula. Results In the Iberian Peninsula, differences in the degree of equilibrium with climatic conditions between lotic and lentic species were few, if any. However, at the continental scale we found significant differences, with lentic species closer to equilibrium than lotic species. In the recently deglaciated area (Sweden) the subset of species with ranges wide enough to encompass Iberia and Scandinavia were mostly lentic, and all were closer to climatic equilibrium without significant differences between habitat types. Main conclusions Our results show that, at continental scales, climate equilibrium is not concordant between the habitat types across western Europe. We hypothesize that: (1) the differences between refuge areas in dispersal ability are erased probably due to long‐term climatic stability, allowing enough time to reach equilibrium, and (2) the species with wide geographical ranges able to recolonize recently deglaciated areas should have the highest dispersal abilities, and are closer to climatic equilibrium.     

Exploring the distribution of Sterocorax Ortuño, 1990 (Coleoptera, Carabidae) species in the Iberian Peninsula

Aim The first aim of this paper was to evaluate the distribution of the three Sterocorax species found in the Iberian Peninsula by estimating the main environmental factors that constrain their distributions. The second aim was to explore the potential importance of competitive interactions in limiting their current distributions using predictive distribution models. Location Iberian Peninsula. Methods Species presence data were collected from records in the literature and private and public collections. Ecological niche factor analysis was performed to extract pseudo‐absences (probable absences), which, together with presence data, were modelled using generalized additive models. The models were run twice. Initially we used only environmental variables, and thereafter additional spatial variables were included in order to account for spatially structured factors not accounted for in the environmental variables. Results Highly reliable distribution models were obtained for the three species, with AUC scores (area under the receiver operating characteristics curve) higher than 0.96. The addition of spatial variables to the first model significantly improved the predicted distribution of Corax (Sterocorax) globosus and Corax (Sterocorax) insidiator, by reducing their potential distribution area. In contrast, the model of Corax (Sterocorax) galicianus was not improved by the addition of a spatial term. Main conclusions Generated pseudo‐absences, such as those used in this study, helped to avoid problems of using erroneous data (false absences) in distribution records. Pseudo‐absences greatly improved the models by only selecting absences within the area with the most unfavourable environmental conditions. The importance of spatial variables to both C. (S.) globosus and C. (S.) insidiator distributions probably relates to a number of unknown factors, such as unique historical events. The absence of established populations of C. (S.) globosus north of the Ebro Valley appears to be one such historical factor. The distribution of C. (S.) galicianus only marginally overlaps with that of C. (S.) globosus, according to our environmental factor models. As this overlap is restricted it is not likely to be a result of competitive exclusion; rather, their geographical segregation seems to be environmentally mediated. The addition of spatial variables reduced the potential habitat of C. (S.) insidiator, eliminating some environmentally optimal areas from its distribution. As no environmental barrier seems apparent in this case, competitive interaction with C. (S.) globosus is a plausible hypothesis for its absence in these optimal parts of its range.     

Post‐glacial migration lag restricts range filling of plants in the European Alps

Aim To evaluate the effect of post‐glacial migration lags on the current distribution of Alpine plants and the factors responsible for possible range‐filling differences among species. Location Austrian Alps. Methods We used species distribution models to predict environmentally suitable sites for 183 Alpine plants at a fine spatial resolution (100 × 100 m²). We overlaid these predictions with independent mapping data (3′× 5′) and calculated the extent to which species fill their potential ranges at this coarser grain based on several different approaches. Moreover, we correlated range‐filling estimates with the magnitude of improvement of distribution models when using the distance to putative glacial refugia as an additional independent variable. Finally, we compared species‐specific range‐filling estimates with traits related to dispersal capacity and competitive ability of these species as well as with characteristics of their habitats. Results Even under a conservative approach, incomplete range filling appears common, with 46% and 31% of the species studied occurring in less than 75% and 50% of their predicted suitable ranges, respectively. Proximity to glacial refugia generally accounts for a lower percentage of the deviance in species distribution data (0–20%, mean 4%) than environmental variables (9–57%, mean 27%). However, its importance correlates closely and negatively with the calculated range‐filling estimates. Range filling significantly increases with the dispersal capacity of a species' propagules and the breadth of its altitudinal niche. Calcicolous species have lower range filling than silicicolous plants and substrate generalists. Conclusions Our results suggest that the current ranges of many Alpine plants are still shaped by delayed Holocene recolonization of suitable sites. Hence, long‐term migration lags also affect plant distribution in mountainous areas, at least on regional scales. These findings question whether high mountain floras will be able to track the expected rapid, climate change driven shifts in habitat.     

Environmental responses of Pinus ponderosa and associated species in the south-western USA

Abstract Aim We addressed four objectives: (1) Determine the regional responses of species, size classes and a vegetation type to climate and parent material predictors, including their distributions in environmental space and the relative contributions of the predictors to explained variation. (2) Determine whether size classes of a species respond similarly to climate and parent material. (3) Assess the extent to which the predicted regional distribution of a vegetation type can be approximated by the distribution of its diagnostic species and vice versa. The establishment of a consistent relationship between the distribution of a vegetation type and its diagnostic species would facilitate change detection, management and conservation planning by allowing the use of one distribution to generate the other when data availability is limited. (4) Examine landscape‐scale environmental variability in predicted species and vegetation type distributions. Location South‐western USA (Arizona, New Mexico and southern Colorado). Methods Ecological response surface models were developed using a data base of 1409 vegetation plots to analyse biotic–environmental relationships of (1) Pinus ponderosa P. & C. Lawson and Abies concolor (Gord. & Glend.) Lindl. Ex Hildebr. size classes, (2) P. ponderosa, A. concolor and Quercus gambelii Nutt. combined size classes, and (3) a P. ponderosa forest type widely distributed in the south‐western USA. Results and main conclusions Pinus ponderosa and A. concolor models generally were judged to be successful. Quercus gambelii models were judged unsuccessful, which may result from the influence of variables not modelled, such as soil moisture, disturbance, biotic factors and other site limiting factors. Size classes differed in the range of environmental conditions associated with high occurrence probabilities within and between species, reflecting differences in the effects of climate variability and anthropogenic changes, such as fire suppression, on the distribution of each size class. Pinus ponderosa alliance was predicted to be distributed over a narrower range of environmental conditions than P. ponderosa species models, therefore limiting the use of this vegetation type as a surrogate for the distribution of the dominant species, and vice versa. Maps of combinations of environmental variables that produced a high probability of P. ponderosa occurrence showed that some landscapes predicted to contain the species exhibited diverse environmental conditions over short distances. The use of regional environmental relationships to characterize areas with high local environmental variability may facilitate identification of areas of potential rapid biotic change.     

Tackling intraspecific genetic structure in distribution models better reflects species geographical range

Genetic diversity provides insight into heterogeneous demographic and adaptive history across organisms’ distribution ranges. For this reason, decomposing single species into genetic units may represent a powerful tool to better understand biogeographical patterns as well as improve predictions of the effects of GCC (global climate change) on biodiversity loss. Using 279 georeferenced Iberian accessions, we used classes of three intraspecific genetic units of the annual plant Arabidopsis thaliana obtained from the genetic analyses of nuclear SNPs (single nucleotide polymorphisms), chloroplast SNPs, and the vernalization requirement for flowering. We used SDM (species distribution models), including climate, vegetation, and soil data, at the whole‐species and genetic‐unit levels. We compared model outputs for present environmental conditions and with a particularly severe GCC scenario. SDM accuracy was high for genetic units with smaller distribution ranges. Kernel density plots identified the environmental variables underpinning potential distribution ranges of genetic units. Combinations of environmental variables accounted for potential distribution ranges of genetic units, which shrank dramatically with GCC at almost all levels. Only two genetic clusters increased their potential distribution ranges with GCC. The application of SDM to intraspecific genetic units provides a detailed picture on the biogeographical patterns of distinct genetic groups based on different genetic criteria. Our approach also allowed us to pinpoint the genetic changes, in terms of genetic background and physiological requirements for flowering, that Iberian A. thaliana may experience with a GCC scenario applying SDM to intraspecific genetic units.     

Forest composition in Mediterranean mountains is projected to shift along the entire elevational gradient under climate change

Aim Species distribution models have been used frequently to assess the effects of climate change on mountain biodiversity. However, the value and accuracy of these assessments have been hampered by the use of low‐resolution data for species distributions and climatic conditions. Herein we assess potential changes in the distribution and community composition of tree species in two mountainous regions of Spain under specific scenarios of climate change using data with a high spatial resolution. We also describe potential changes in species distributions and tree communities along the entire elevational gradient. Location Two mountain ranges in southern Europe: the Central Mountain Range (central west of the Iberian Peninsula), and the Iberian Mountain Range (central east). Methods We modelled current and future distributions of 15 tree species (Eurosiberian, sub‐Mediterranean and Mediterranean species) as functions of climate, lithology and availability of soil water using generalized linear models (logistic regression) and machine learning models (gradient boosting). Using multivariate ordination of a matrix of presence/absence of tree species obtained under two Intergovernmental Panel on Climate Change (IPCC) scenarios (A2 and B2) for two different periods in the future (2041–70 and 2071–2100), we assessed the predicted changes in the composition of tree communities. Results The models predicted an upward migration of communities of Mediterranean trees to higher elevations and an associated decline in communities of temperate or cold‐adapted trees during the 21st century. It was predicted that 80–99% of the area that shows a climate suitable for cold–wet‐optimum Eurosiberian coniferous and broad‐leaved species will be lost. The largest overall changes were predicted for Mediterranean species found currently at low elevations, such as Pinus halepensis, Pinus pinaster, Quercus ilex ssp. ballota and Juniperus oxycedrus, with sharp increases in their range of 350%. Main conclusions It is likely that areas with climatic conditions suitable for cold‐adapted species will decrease significantly under climate warming. Large changes in species ranges and forest communities might occur, not only at high elevations within Mediterranean mountains but also along the entire elevational gradient throughout this region, particularly at low and mid‐elevations. Mediterranean mountains might lose their key role as refugia for cold‐adapted species and thus an important part of their genetic heritage.     

Of niches and distributions: range size increases with niche breadth both globally and regionally but regional estimates poorly relate to global estimates

The relationship between species’ niche breadth (i.e. the range of environmental conditions under which a species can persist) and range size (i.e. the extent of its spatial distribution) has mostly been tested within geographically restricted areas but rarely at the global extent. Here, we not only tested the relationship between range size (derived from species’ distribution data) and niche breadth (derived from species’ distribution and co‐occurrence data) of 1255 plant species at the regional extent of the European Alps, but also at the global extent and across both spatial scales for a subset of 180 species. Using correlation analyses, linear models and variation partitioning, we found that species’ realized niche breadth estimated at the regional level is a weak predictor of species’ global niche breadth and range size. Against our expectations, distribution‐derived niche breadth was a better predictor for species’ range size than the co‐occurrence‐based estimate, which should, theoretically, account for more than the climatically determined niche dimensions. Our findings highlight that studies focusing on the niche breadth vs range size relationship must explicitly consider spatial mismatches that might have confounded and diminished previously reported relationships.     

How to describe species richness patterns for bryophyte conservation?

A large amount of data for inconspicuous taxa is stored in natural history collections; however, this information is often neglected for biodiversity patterns studies. Here, we evaluate the performance of direct interpolation of museum collections data, equivalent to the traditional approach used in bryophyte conservation planning, and stacked species distribution models (S‐SDMs) to produce reliable reconstructions of species richness patterns, given that differences between these methods have been insufficiently evaluated for inconspicuous taxa. Our objective was to contrast if species distribution models produce better inferences of diversity richness than simply selecting areas with the higher species numbers. As model species, we selected Iberian species of the genus Grimmia (Bryophyta), and we used four well‐collected areas to compare and validate the following models: 1) four Maxent richness models, each generated without the data from one of the four areas, and a reference model created using all of the data and 2) four richness models obtained through direct spatial interpolation, each generated without the data from one area, and a reference model created with all of the data. The correlations between the partial and reference Maxent models were higher in all cases (0.45 to 0.99), whereas the correlations between the spatial interpolation models were negative and weak (−0.3 to −0.06). Our results demonstrate for the first time that S‐SDMs offer a useful tool for identifying detailed richness patterns for inconspicuous taxa such as bryophytes and improving incomplete distributions by assessing the potential richness of under‐surveyed areas, filling major gaps in the available data. In addition, the proposed strategy would enhance the value of the vast number of specimens housed in biological collections.     

Large-scale environmental correlates of forest tree distributions in Catalonia (NE Spain)

Aim To explore the environmental correlates of tree species distributions in Catalonia according to the chorological status of the species. Location The study area is the region of Catalonia, in north‐eastern Spain. Methods We used presence‐absence data for 24 species, sampled in random plots distributed throughout forests of Catalonia. A climate model for the Catalonia region provided environmental variables. We used classification tree analysis to explore the environmental correlates of the realized niches of tree species. The predictive accuracy of the models was assessed using the ROC curve approach. Potential distribution maps of tree species were generated for the whole Catalonia region. Results Models were ranked from low to high accuracy for the 24 species. Differences in accuracy among species were related to the chorological status of species. Zonal species, or species at the core of their range (Mediterranean and Sub‐Mediterranean species), were generally well predicted, while extrazonal species, or species at the edge of their range, were predicted only moderately well. Mediterranean species distributions showed good correlations with extreme temperatures and annual precipitation. Main conclusions The above trends confirmed the difficulty of identifying the realized niche of species at the edges of their ranges. In contrast, Mediterranean and Sub‐Mediterranean species, which were at the core of their range, were well‐predicted, confirming the importance of extremes of temperature and annual precipitation as effective surrogates for variables having more direct physiological roles in limiting the ability of plants to survive and grow. Maps of potential tree distributions allowed us to define suitable habitats and to highlight areas where species have been planted outside their natural distribution.     

Co‐occurrence patterns of trees along macro‐climatic gradients and their potential influence on the present and future distribution of Fagus sylvatica L.

Aim During recent and future climate change, shifts in large‐scale species ranges are expected due to the hypothesized major role of climatic factors in regulating species distributions. The stress‐gradient hypothesis suggests that biotic interactions may act as major constraints on species distributions under more favourable growing conditions, while climatic constraints may dominate under unfavourable conditions. We tested this hypothesis for one focal tree species having three major competitors using broad‐scale environmental data. We evaluated the variation of species co‐occurrence patterns in climate space and estimated the influence of these patterns on the distribution of the focal species for current and projected future climates. Location Europe. Methods We used ICP Forest Level 1 data as well as climatic, topographic and edaphic variables. First, correlations between the relative abundance of European beech (Fagus sylvatica) and three major competitor species (Picea abies, Pinus sylvestris and Quercus robur) were analysed in environmental space, and then projected to geographic space. Second, a sensitivity analysis was performed using generalized additive models (GAM) to evaluate where and how much the predicted F. sylvatica distribution varied under current and future climates if potential competitor species were included or excluded. We evaluated if these areas coincide with current species co‐occurrence patterns. Results Correlation analyses supported the stress‐gradient hypothesis: towards favourable growing conditions of F. sylvatica, its abundance was strongly linked to the abundance of its competitors, while this link weakened towards unfavourable growing conditions, with stronger correlations in the south and at low elevations than in the north and at high elevations. The sensitivity analysis showed a potential spatial segregation of species with changing climate and a pronounced shift of zones where co‐occurrence patterns may play a major role. Main conclusions Our results demonstrate the importance of species co‐occurrence patterns for calibrating improved species distribution models for use in projections of climate effects. The correlation approach is able to localize European areas where inclusion of biotic predictors is effective. The climate‐induced spatial segregation of the major tree species could have ecological and economic consequences.     

Ecological and historical filters constraining spatial caddisfly distribution in Mediterranean rivers

1. Contemporary species distributions are determined by a mixture of ecological and historical filters acting on several spatial and temporal scales. Mediterranean climate areas are one of the world's biodiversity hotspots with a high level of endemicity, which is linked to complex ecological and historical factors. 2. This paper explores the ecological and historical factors constraining the distribution of caddisfly species on a large regional scale. A total of 69 taxa were collected from 140 sampling sites in 10 Iberian Mediterranean river basins. Approximately 74% of taxa can be considered rare, with the southern basins (the Baetic–Riffian region) having greater endemicity. The greatest richness, involving a mixture of northern and southern species, was found in the transitional area between the Baetic–Riffian region and the Hesperic Massif. 3. The historical processes occurring during the Tertiary (i.e. the junction of the Eurasian and African plates) explained 3.1% of species distribution, whereas ecological factors accounted for 20.7%. Only 0.3% was explained by the interaction of history and ecology. A set of multi‐scale ecological variables (i.e. basin, reach and bedform characteristics) defined five river types with specific caddisfly assemblages. The commonest caddisfly species accounted for the regional distribution pattern, while rare taxa contributed to the explanation of subtle patterns not shown by common species. 4. Despite the importance of historical factors for biogeography and the large scale used in our study, ecological variables better explained caddisfly distribution. This may be explained by the length of time since the historical process we are considering, the high dispersion and colonisation capacity of many caddisfly species, and the strong environmental gradient in the area. Because of the historical and environmental complexity of Mediterranean areas, rare taxa should be included in ecological studies so that the singularity of these ecosystems is not missed.     

Integrating individual habitat choices and regional distribution of a biodiversity indicator and top predator

Aim Habitat selection studies have mainly focused on behavioural choices of individuals or on the habitat‐related regional distribution of a population, with little integration of the two approaches. This is despite the fact that traditional biogeography theory sees the geographical distribution of a species as the collective outcome of the adaptive habitat choices of individuals. Here, we integrate individual habitat choices with regional distribution through a bottom‐up Geographical Information System (GIS)‐based approach, by using a 9‐year data set on a large avian predator, the eagle owl (Bubo bubo L.). We further examine the potential population level and biodiversity consequences of this approach. Location The study was conducted in the Trento Region (central‐eastern Italian Alps) and in six other areas of the nearby Lombardia Region in the central Alps. Methods We used stepwise logistic regression to build a habitat suitability model discriminating between eagle owl territories and an equal number of random locations. The model was applied to the whole Trento region by means of a GIS so as to predict suitable habitat patches. The predicted regional distribution (presence–absence in 10‐km grid quadrats) was then compared with the observed one. Furthermore, we compared estimates of biodiversity in quadrats with and without eagle owls, so as to test whether the presence of this top predator may signal macro‐areas of high biodiversity. Results The logistic habitat suitability model showed that, compared with a random distribution, eagle owls selected low‐elevation breeding sites with high availability of prey‐rich habitats in their surroundings. Breeding performance increased with the availability of prey‐rich habitats, confirming the adaptiveness of the detected habitat choices. We applied the habitat suitability model to the 6200 km² study region by means of a GIS and found a close fit between the observed and predicted regional distribution. Furthermore, population abundance was positively related to the availability of habitat defined as suitable by the above analyses. Finally, high biodiversity levels were associated with owl presence and with the amount of suitable owl habitat, demonstrating that modelling habitat suitability of a properly chosen indicator species may provide key conservation information at the wider ecosystem level. Main conclusions Our bottom‐up modelling approach may increase the conservation‐value of habitat selection models, by (1) predicting local and regional distribution, (2) estimating regional population size, (3) stimulating further hypothesis testing, (4) forecasting the population effects of future habitat loss and degradation and (5) aiding in the identification and prioritization of high‐biodiversity areas.     

Combining physiological threshold knowledge to species distribution models is key to improving forecasts of the future niche for macroalgae

Species distribution models (SDM) are a useful tool for predicting species range shifts in response to global warming. However, they do not explore the mechanisms underlying biological processes, making it difficult to predict shifts outside the environmental gradient where the model was trained. In this study, we combine correlative SDMs and knowledge on physiological limits to provide more robust predictions. The thermal thresholds obtained in growth and survival experiments were used as proxies of the fundamental niches of two foundational marine macrophytes. The geographic projections of these species’ distributions obtained using these thresholds and existing SDMs were similar in areas where the species are either absent‐rare or frequent and where their potential and realized niches match, reaching consensus predictions. The cold‐temperate foundational seaweed Himanthalia elongata was predicted to become extinct at its southern limit in northern Spain in response to global warming, whereas the occupancy of southern‐lusitanic Bifurcaria bifurcata was expected to increase. Combined approaches such as this one may also highlight geographic areas where models disagree potentially due to biotic factors. Physiological thresholds alone tended to over‐predict species prevalence, as they cannot identify absences in climatic conditions within the species’ range of physiological tolerance or at the optima. Although SDMs tended to have higher sensitivity than threshold models, they may include regressions that do not reflect causal mechanisms, constraining their predictive power. We present a simple example of how combining correlative and mechanistic knowledge provides a rapid way to gain insight into a species’ niche resulting in consistent predictions and highlighting potential sources of uncertainty in forecasted responses to climate change.     

Squares of different sizes: effect of geographical projection on model parameter estimates in species distribution modeling

In species distribution analyses, environmental predictors and distribution data for large spatial extents are often available in long‐lat format, such as degree raster grids. Long‐lat projections suffer from unequal cell sizes, as a degree of longitude decreases in length from approximately 110 km at the equator to 0 km at the poles. Here we investigate whether long‐lat and equal‐area projections yield similar model parameter estimates, or result in a consistent bias. We analyzed the environmental effects on the distribution of 12 ungulate species with a northern distribution, as models for these species should display the strongest effect of projectional distortion. Additionally we choose four species with entirely continental distributions to investigate the effect of incomplete cell coverage at the coast. We expected that including model weights proportional to the actual cell area should compensate for the observed bias in model coefficients, and similarly that using land coverage of a cell should decrease bias in species with coastal distribution. As anticipated, model coefficients were different between long‐lat and equal‐area projections. Having progressively smaller and a higher number of cells with increasing latitude influenced the importance of parameters in models, increased the sample size for the northernmost parts of species ranges, and reduced the subcell variability of those areas. However, this bias could be largely removed by weighting long‐lat cells by the area they cover, and marginally by correcting for land coverage. Overall we found little effect of using long‐lat rather than equal‐area projections in our analysis. The fitted relationship between environmental parameters and occurrence probability differed only very little between the two projection types. We still recommend using equal‐area projections to avoid possible bias. More importantly, our results suggest that the cell area and the proportion of a cell covered by land should be used as a weight when analyzing distribution of terrestrial species.     

Geographical distributions of spiny pocket mice in South America: insights from predictive models

Aim Predictive models of species’ distributions use occurrence records and environmental data to produce a model of the species’ requirements and a map of its potential distribution. To determine regions of suitable environmental conditions and assess biogeographical questions regarding their ranges, we modelled the potential geographical distributions of two spiny pocket mice (Rodentia: Heteromyidae) in north‐western South America. Location North‐western South America. Methods We used the Genetic Algorithm for Rule‐Set Prediction (GARP), environmental data from GIS maps and georeferenced collection localities from a recent systematic review of Heteromys australis and H. anomalus to produce the models. Results GARP models indicate the potential presence of H. australis throughout mesic montane regions of north‐western South America, as well as in some lowland regions of moderately high precipitation. In contrast, H. anomalus is predicted to occur primarily in drier areas of the Caribbean coast and rain‐shadowed valleys of the Andes. Conclusions The models support the disjunct status of the population of H. australis in the Cordillera de Mérida, but predict a continuous distribution between known populations of H. anomalus in the upper Magdalena Valley and the Caribbean coast. Regions of suitable environmental conditions exist disjunct from known distributional areas for both species, suggesting possible historical restrictions to their ranges. This technique holds wide application to other study systems.