Jorge M. Lobo

Use of niche models in invasive species risk assessments

Risk maps summarizing landscape suitability of novel areas for invading species can be valuable tools for preventing species’ invasions or controlling their spread, but methods employed for development of such maps remain variable and unstandardized. We discuss several considerations in development of such models, including types of distributional information that should be used, the nature of explanatory variables that should be incorporated, and caveats regarding model testing and evaluation. We highlight that, in the case of invasive species, such distributional predictions should aim to derive the best hypothesis of the potential distribution of the species by using (1) all distributional information available, including information from both the native range and other invaded regions; (2) predictors linked as directly as is feasible to the physiological requirements of the species; and (3) modelling procedures that carefully avoid overfitting to the training data. Finally, model testing and evaluation should focus on well-predicted presences, and less on efficient prediction of absences; a k-fold regional cross-validation test is discussed.

Accounting for uncertainty when mapping species distributions: The need for maps of ignorance

Accurate mapping of species distributions is a fundamental goal of modern biogeography, both for basic and applied purposes. This is commonly done by plotting known species occurrences, expert-drawn range maps or geographical estimations derived from species distribution models. However, all three kinds of maps are implicitly subject to uncertainty, due to the quality and bias of raw distributional data, the process of map building, and the dynamic nature of species distributions themselves. Here we review the main sources of uncertainty suggesting a code of good practices in order to minimize their effects. Specifically, we claim that uncertainty should be always explicitly taken into account and we propose the creation of maps of ignorance to provide information on where the mapped distributions are reliable and where they are uncertain.

The Effectiveness of Iberian Protected Areas in Conserving Terrestrial Biodiversity

The Iberian Peninsula harbors about 50% of European plant and terrestrial vertebrate species and more than 30% of European endemic species. Despite the global recognition of its importance, the selection of protected areas has been ad hoc and the effectiveness of such choices has rarely been assessed. We compiled the most comprehensive distributional data set of Iberian terrestrial plant and vertebrate species available to date and used it to assess the degree of species representation within existing protected areas. Existing protected areas in Spain and Portugal reasonably represented the plant and animal species we considered (73-98%). Nevertheless, species of some groups (amphibians, reptiles, birds, and gymnosperms) did not accumulate in protected areas at a rate higher than expected by chance (p > 0.05). We determined that to conserve all vertebrate and plant species in the Iberian Peninsula, at least 36 additional areas are needed. Selection of additional areas for conservation would be facilitated if such areas coincided with sites of community importance (SCI) designated under the European Commission Habitats Directive. Additional areas required for full representation of the selected plant and animal species all coincide with SCI in Spain. Nevertheless, the degree of coincidence varies between 0.3% and 74.6%, and there is a possibility that important areas for conservation occur outside the SCI. Our results support the view that current SCI can be used for prioritization of areas for conservation, but a systematic reevaluation of conservation priorities in Spain and Portugal would be necessary to ensure that effective conservation of one of Europeans most important biodiversity regions is achieved.

A GLOBAL EVALUATION OF METABOLIC THEORY AS AN EXPLANATION FOR TERRESTRIAL SPECIES RICHNESS GRADIENTS

We compiled 46 broadscale data sets of species richness for a wide range of terrestrial plant, invertebrate, and ectothermic vertebrate groups in all parts of the world to test the ability of metabolic theory to account for observed diversity gradients. The theory makes two related predictions: (1) In-transformed richness is linearly associated with a linear, inverse transformation of annual temperature, and (2) the slope of the relationship is near -0.65. Of the 46 data sets, 14 had no significant relationship; of the remaining 32, nine were linear, meeting prediction 1. Model I (ordinary least squares, OLS) and model II (reduced major axis, RMA) regressions then tested the linear slopes against prediction 2. In the 23 data sets having nonlinear relationships between richness and temperature, split-line regression divided the data into linear components, and regressions were done on each component to test prediction 2 for subsets of the data. Of the 46 data sets analyzed in their entirety using OLS regression, one was consistent with metabolic theory (meeting both predictions), and one was possibly consistent. Using RMA regression, no data sets were consistent. Of 67 analyses of prediction 2 using OLS regression on all linear data sets and subsets, two were consistent with the prediction, and four were possibly consistent. Using RMA regression, one was consistent (albeit weakly), and four were possibly consistent. We also found that the relationship between richness and temperature is both taxonomically and geographically conditional, and there is no evidence for a universal response of diversity to temperature. Meta-analyses confirmed significant heterogeneity in slopes among data sets, and the combined slopes across studies were significantly lower than the range of slopes predicted by metabolic theory based on both OLS and RMA regressions. We conclude that metabolic theory, as currently formulated, is a poor predictor of observed diversity gradients in most terrestrial systems.

Ice age climate, evolutionary constraints and diversity patterns of European dung beetles

Current climate and Pleistocene climatic changes are both known to be associated with geographical patterns of diversity. We assess their associations with the European Scarabaeinae dung beetles, a group with high dispersal ability and well-known adaptations to warm environments. By assessing spatial stationarity in climate variability since the last glacial maximum (LGM), we find that current scarab richness is related to the location of their limits of thermal tolerance during the LGM. These limits mark a strong change in their current species richness-environment relationships. Furthermore, northern scarab assemblages are nested and composed of a phylogenetically clustered subset of large-range sized generalist species, whereas southern ones are diverse and variable in composition. Our results show that species responses to current climate are limited by the evolution of assemblages that occupied relatively climatically stable areas during the Pleistocene, and by post-glacial dispersal in those that were strongly affected by glaciations.

An ED-based protocol for optimal sampling of biodiversity

While conservation planning requires good biodiversity data, our knowledge of most living groups is scarce and patchy even in well-sampled regions. Therefore, we need methodologies for rapid assessments for particular groups and regions. Maps of any biodiversity surrogate can be interpolated from even a few well-known sites, but such places are usually lacking. We therefore propose a protocol for designing field surveys to obtain good coverage of pattern variations of biodiversity in a given region. To represent biodiversity patterns comprehensively, we use a rule step site-allocation procedure, partially based on Faith and Walkers ED criterion that takes environmental and spatial variation into account, together with other criteria such as survey costs. A preliminary assessment of the adequacy of this site sampling strategy is made. Then a set of complementary sites is selected for further sampling. Using the ED criterion, during the stepwise process a p-median analysis is applied both to an environmental distance matrix and to a spatial distance matrix, to maximize the amount of variation covered by our survey planning. This rule-set allocation procedure is integrated into a continuous sampling design protocol directed to ensure we can sample all biodiversity of a region. This protocol requires the gathering of both biological and environmental information, an assessment of previously available information, the choice of sampling methods and dates, and a continuous assessment of the success of the survey being carried out. An example of the application of this protocol to the survey design of dung beetle (Coleoptera, Scarabaeoidea) diversity in the Comunidad de Madrid (Spain) is included.

Assessing the accuracy of species distribution models to predict amphibian species richness patterns

1. Evaluating the distribution of species richness where biodiversity is high but has been insufficiently sampled is not an easy task. Species distribution modelling has become a useful approach for predicting their ranges, based on the relationships between species records and environmental variables. Overlapping predictions of individual distributions could be a useful strategy for obtaining estimates of species richness and composition in a region, but these estimates should be evaluated using a proper validation process, which compares the predicted richness values and composition with accurate data from independent sources. 2. In this study, we propose a simple approach to estimate model performance for several distributional predictions generated simultaneously. This approach is particularly suitable when species distribution modelling techniques that require only presence data are used. 3. The individual distributions for the 370 known amphibian species of Mexico were predicted using maxent to model data on their known presence (66,113 presence-only records). Distributions were subsequently overlapped to obtain a prediction of species richness. Accuracy was assessed by comparing the overall species richness values predicted for the region with observed and predicted values from 118 well-surveyed sites, each with an area of c. 100 km(2), which were identified using species accumulation curves and nonparametric estimators. 4. The derived models revealed a remarkable heterogeneity of species richness across the country, provided information about species composition per site and allowed us to obtain a measure of the spatial distribution of prediction errors. Examining the magnitude and location of model inaccuracies, as well as separately assessing errors of both commission and omission, highlights the inaccuracy of the predictions of species distribution models and the need to provide measures of uncertainty along with the model results. 5. The combination of a species distribution modelling method like maxent and species richness estimators offers a useful tool for identifying when the overall pattern provided by all model predictions might be representing the geographical patterns of species richness and composition, regardless of the particular quality or accuracy of the predictions for each individual species.

Determinants of local spider (Araneidae and Thomisidae) species richness on a regional scale: climate and altitude vs. habitat structure

Abstract 1. The habitat heterogeneity hypothesis states that the more complex the habitat, the higher the species richness. The present study analyzes the effect of local factors on regional spider (Araneidae and Thomisidae) richness. The main objective is to disentangle the relative importance of habitat structure and other environmental variables.

Biogeographical and Ecological Factors Affecting the Altitudinal Variation of Mountainous Communities of Coprophagous Beetles (Coleoptera: Scarabaeoidea): a Comparative Study

Abstract The altitudinal variation in the richness and composition of communities of coprophagous beetles in a mountainous landscape in Mexico is analyzed and the results obtained are compared with those of similar studies carried out in other parts of the world. Two nonexclusive processes are proposed as responsible for the assemblage of mountain fauna: horizontal colonization by elements originating from lineages distributed at higher latitudes and vertical colonization by lineages distributed at the same latitude but at different altitudes. The current analysis supports the hypothesis that when the horizontal colonization dominates, mountain faunas show a clear altitudinal substitution between large taxa with different evolutionary histories (e.g., Scarabaeinae and Aphodiinae) and the gradient of reduction in species richness is attenuated. This occurs in mountains of different continents and depends primarily on the degree of isolation and general orientation of the mountain ranges. Conversely, when vertical colonization dominates, mountain faunas show only slight altitudinal substitution, and the reduction in richness is greater.

Historical Legacies in World Amphibian Diversity Revealed by the Turnover and Nestedness Components of Beta Diversity

Historic processes are expected to influence present diversity patterns in combination with contemporary environmental factors. We hypothesise that the joint use of beta diversity partitioning methods and a threshold-based approach may help reveal the effect of large-scale historic processes on present biodiversity. We partitioned intra-regional beta diversity into its turnover (differences in composition caused by species replacements) and nestedness-resultant (differences in species composition caused by species losses) components. We used piecewise regressions to show that, for amphibian beta diversity, two different world regions can be distinguished. Below parallel 37, beta diversity is dominated by turnover, while above parallel 37, beta diversity is dominated by nestedness. Notably, these regions are revealed when the piecewise regression method is applied to the relationship between latitude and the difference between the Last Glacial Maximum (LGM) and the present temperature but not when present energy-water factors are analysed. When this threshold effect of historic climatic change is partialled out, current energy-water variables become more relevant to the nestedness-resultant dissimilarity patterns, while mountainous areas are associated with higher spatial turnover. This result suggests that nested patterns are caused by species losses that are determined by physiological constraints, whereas turnover is associated with speciation and/or Pleistocene refugia. Thus, the new threshold-based view may help reveal the role of historic factors in shaping present amphibian beta diversity patterns.