Joaquín Hortal

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.

Climate change, humans, and the extinction of the woolly mammoth.

Woolly mammoths inhabited Eurasia and North America from late Middle Pleistocene (300 ky BP [300,000 years before present]), surviving through different climatic cycles until they vanished in the Holocene (3.6 ky BP). The debate about why the Late Quaternary extinctions occurred has centred upon environmental and human-induced effects, or a combination of both. However, testing these two hypotheses—climatic and anthropogenic—has been hampered by the difficulty of generating quantitative estimates of the relationship between the contraction of the mammoths geographical range and each of the two hypotheses. We combined climate envelope models and a population model with explicit treatment of woolly mammoth–human interactions to measure the extent to which a combination of climate changes and increased human pressures might have led to the extinction of the species in Eurasia. Climate conditions for woolly mammoths were measured across different time periods: 126 ky BP, 42 ky BP, 30 ky BP, 21 ky BP, and 6 ky BP. We show that suitable climate conditions for the mammoth reduced drastically between the Late Pleistocene and the Holocene, and 90% of its geographical range disappeared between 42 ky BP and 6 ky BP, with the remaining suitable areas in the mid-Holocene being mainly restricted to Arctic Siberia, which is where the latest records of woolly mammoths in continental Asia have been found. Results of the population models also show that the collapse of the climatic niche of the mammoth caused a significant drop in their population size, making woolly mammoths more vulnerable to the increasing hunting pressure from human populations. The coincidence of the disappearance of climatically suitable areas for woolly mammoths and the increase in anthropogenic impacts in the Holocene, the coup de grâce, likely set the place and time for the extinction of the woolly mammoth.

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.

Island Species Richness Increases with Habitat Diversity

Species richness is commonly thought to increase with habitat diversity. However, a recent theoretical model aiming to unify niche and island biogeography theories predicted a hump‐shaped relationship between richness and habitat diversity. Given the contradiction between model results and previous knowledge, we examine whether the relationship between species richness and habitat diversity is consistently monotonically increasing and under which circumstances, if at all, such relationships could be hump shaped. We review the empirical evidence about the shape of such relationships and show that species richness on islands usually increases with habitat diversity and that it never decreases. We also critically examine the assumptions of the theoretical model and modify them to incorporate a less restrictive definition of niche width. The modified assumptions lead to simulations that better capture real patterns, using either simple parameters or observed distributions of niche breadth. Further work is needed to incorporate ecological interactions and metacommunity dynamics if the aim is to merge niche and island biogeography theories in a realistic modeling framework.

Handbook of protocols for standardized measurement of terrestrial invertebrate functional traits

Summary 1. Trait-based approaches are increasingly being used to test mechanisms underlying species assemblages and biotic interactions across a wide range of organisms including terrestrial arthropods and to investigate consequences for ecosystem processes. Such an approach relies on the standardized measurement of functional traits that can be applied across taxa and regions. Currently, however, unified methods of trait measurements are lacking for terrestrial arthropods and related macroinvertebrates (terrestrial invertebrates hereafter). 2. Here, we present a comprehensive review and detailed protocol for a set of 29 traits known to be sensitive to global stressors and to affect ecosystem processes and services. We give rec- ommendations how to measure these traits under standardized conditions across various ter- restrial invertebrate taxonomic groups. 3. We provide considerations and approaches that apply to almost all traits described, such as the selection of species and individuals needed for the measurements, the importance of intraspecific trait variability, how many populations or communities to sample and over which spatial scales. 4. The approaches outlined here provide a means to improve the reliability and predictive power of functional traits to explain community assembly, species diversity patterns and ecosystem processes and services within and across taxa and trophic levels, allowing compar- ison of studies and running meta-analyses across regions and ecosystems. 5. This handbook is a crucial first step towards standardizing trait methodology across the most studied terrestrial invertebrate groups, and the protocols are aimed to balance general applicability and requirements for special cases or particular taxa. Therefore, we envision this handbook as a common platform to which researchers can further provide methodological input for additional special cases.

Challenging species distribution models : the case of Maculinea nausithous in the Iberian Peninsula

Maculinea nausithousis the most endangered Iberian butterfly, being included in the Bern and Habitat directives and in the Spanish Red Book for invertebrates. We map its potential distribution in the Iberian Peninsula, identifying potentially suitable areas where to focus future surveys. Using presence/absence data and a set of environmental variables, the distribution of M. nausithous was modelled using Generalized Additive Models. Model results were filtered using land cover data to identify the anthropogenic grasslands inhabited by the species. The model obtained indicates that some Iberian areas that currently do not support M. nausithous populations could be environmentally suitable for the species. The reasons for these absences are discussed. Indeed, field surveys identified possible sources of uncertainty at finer scale, highlighting that deficiencies in land use GIS data might constitute an important source of error, able to explain both commission and omission errors (i.e., over and underpredictions) of the model.

Information visualisation for science and policy: engaging users and avoiding bias

Visualisations and graphics are fundamental to studying complex subject matter. However, beyond acknowledging this value, scientists and science-policy programmes rarely consider how visualisations can enable discovery, create engaging and robust reporting, or support online resources. Producing accessible and unbiased visualisations from complicated, uncertain data requires expertise and knowledge from science, policy, computing, and design. However, visualisation is rarely found in our scientific training, organisations, or collaborations. As new policy programmes develop [e.g., the Intergovernmental Platform on Biodiversity and Ecosystem Services (IPBES)], we need information visualisation to permeate increasingly both the work of scientists and science policy. The alternative is increased potential for missed discoveries, miscommunications, and, at worst, creating a bias towards the research that is easiest to display.

Challenges and perspectives for species distribution modelling in the neotropics

The workshop ‘Species distribution models: applications, challenges and perspectives’ held at Belo Horizonte (Brazil), 29–30 August 2011, aimed to review the state-of-the-art in species distribution modelling (SDM) in the neotropical realm. It brought together researchers in ecology, evolution, biogeography and conservation, with different backgrounds and research interests. The application of SDM in the megadiverse neotropics—where data on species occurrences are scarce—presents several challenges, involving acknowledging the limitations imposed by data quality, including surveys as an integral part of SDM studies, and designing the analyses in accordance with the question investigated. Specific solutions were discussed, and a code of good practice in SDM studies and related field surveys was drafted.