Projecting future expansion of invasive species: Comparing and improving methodologies for species distribution modeling

Modeling the distributions of species, especially of invasive species in non-native ranges, involves multiple challenges. Here, we developed some novel approaches to species distribution modeling aimed at reducing the influences of such challenges and improving the realism of projections. We estimated species-environment relationships with four modeling methods run with multiple scenarios of (1) sources of occurrences and geographically isolated background ranges for absences, (2) approaches to drawing background (absence) points, and (3) alternate sets of predictor variables. We further tested various quantitative metrics of model evaluation against biological insight. Model projections were very sensitive to the choice of training dataset. Model accuracy was much improved by using a global dataset for model training, rather than restricting data input to the species’ native range. AUC score was a poor metric for model evaluation and, if used alone, was not a useful criterion for assessing model performance. Projections away from the sampled space (i.e. into areas of potential future invasion) were very different depending on the modeling methods used, raising questions about the reliability of ensemble projections. Generalized linear models gave very unrealistic projections far away from the training region. Models that efficiently fit the dominant pattern, but exclude highly local patterns in the dataset and capture interactions as they appear in data (e.g. boosted regression trees), improved generalization of the models. Biological knowledge of the species and its distribution was important in refining choices about the best set of projections. A post-hoc test conducted on a new Partenium dataset from Nepal validated excellent predictive performance of our “best” model. We showed that vast stretches of currently uninvaded geographic areas on multiple continents harbor highly suitable habitats for Parthenium hysterophorus L. (Asteraceae; parthenium). However, discrepancies between model predictions and parthenium invasion in Australia indicate successful management for this globally significant weed. This article is protected by copyright. All rights reserved.

Projecting future expansion of invasive species: Comparing and improving methodologies for species distribution modeling

Modeling the distributions of species, especially of invasive species in non-native ranges, involves multiple challenges. Here, we developed some novel approaches to species distribution modeling aimed at reducing the influences of such challenges and improving the realism of projections. We estimated species-environment relationships with four modeling methods run with multiple scenarios of (1) sources of occurrences and geographically isolated background ranges for absences, (2) approaches to drawing background (absence) points, and (3) alternate sets of predictor variables. We further tested various quantitative metrics of model evaluation against biological insight. Model projections were very sensitive to the choice of training dataset. Model accuracy was much improved by using a global dataset for model training, rather than restricting data input to the species’ native range. AUC score was a poor metric for model evaluation and, if used alone, was not a useful criterion for assessing model performance. Projections away from the sampled space (i.e. into areas of potential future invasion) were very different depending on the modeling methods used, raising questions about the reliability of ensemble projections. Generalized linear models gave very unrealistic projections far away from the training region. Models that efficiently fit the dominant pattern, but exclude highly local patterns in the dataset and capture interactions as they appear in data (e.g. boosted regression trees), improved generalization of the models. Biological knowledge of the species and its distribution was important in refining choices about the best set of projections. A post-hoc test conducted on a new Partenium dataset from Nepal validated excellent predictive performance of our “best” model. We showed that vast stretches of currently uninvaded geographic areas on multiple continents harbor highly suitable habitats for Parthenium hysterophorus L. (Asteraceae; parthenium). However, discrepancies between model predictions and parthenium invasion in Australia indicate successful management for this globally significant weed. This article is protected by copyright. All rights reserved.

Thesis abstract. Applying species distribution modeling for the conservation of Iberian protected invertebrates

This article outlines the approaches to modeling the distribution of threatened invertebrates using data from atlases, museums and databases. Species Distribution Models (SDMs) are useful for estimating species’ ranges, identifying suitable habitats, and identifying the primary factors affecting species’ distributions. The study tackles the strategies used to obtain SDMs without reliable absence data while exploring their applications for conservation. I examine the conservation status of Copris species and Graellsia isabelae by delimiting their populations and exploring the effectiveness of protected areas. I show that the method of pseudo‐absence selection strongly determines the model obtained, generating different model predictions along the gradient between potential and realized distributions. After assessing the effects of species’ traits and data characteristics on accuracy, I found that species are modeled more accurately when sample sizes are larger, no matter the technique used.

Strengthening the link between climate, hydrological and species distribution modeling to assess the impacts of climate change on freshwater biodiversity

To understand the resilience of aquatic ecosystems to environmental change, it is important to determine how multiple, related environmental factors, such as near-surface air temperature and river flow, will change during the next century. This study develops a novel methodology that combines statistical downscaling and fish species distribution modeling, to enhance the understanding of how global climate changes (modeled by global climate models at coarse-resolution) may affect local riverine fish diversity. The novelty of this work is the downscaling framework developed to provide suitable future projections of fish habitat descriptors, focusing particularly on the hydrology which has been rarely considered in previous studies. The proposed modeling framework was developed and tested in a major European system, the Adour-Garonne river basin (SW France, 116,000 km(2)), which covers distinct hydrological and thermal regions from the Pyrenees to the Atlantic coast. The simulations suggest that, by 2100, the mean annual stream flow is projected to decrease by approximately 15% and temperature to increase by approximately 1.2 °C, on average. As consequence, the majority of cool- and warm-water fish species is projected to expand their geographical range within the basin while the few cold-water species will experience a reduction in their distribution. The limitations and potential benefits of the proposed modeling approach are discussed. Copyright © 2012 Elsevier B.V. All rights reserved.

Species Distribution Modeling for Conservation Purposes

Species distribution models (SDMs) can be useful for different conservation purposes. We discuss the importance of fitting spatial scale and using current records and relevant predictors aiming conservation. We choose jaguar (Panthera onca) as a target species and Brazil and Atlantic Forest biome as study areas. We tested two different extents (continent and biome) and resolutions (similar to 4 Km and similar to 1 Km) in Maxent with 186 records and 11 predictors (bioclimatic, elevation, land-use and landscape structure). All models presented satisfactory AUC values (>0.70) and low omission errors (<23%). SDMs were scale-sensitive as the use of reduced extent implied in significant gains to model performance generating more constrained and real predictive distribution maps. Continental-scale models performed poorly in predicting potential current jaguar distribution, but they reached the historic distribution. Specificity increased significantly from coarse to finer-scale models due to the reduction of overprediction. The variability of environmental space (E-space) differed for most of climatic variables between continental and biome-scale and the representation of the E-space by predictors differed significantly (t = 2.42, g.I. = 9, P < 0.05). Refining spatial scale, incorporating landscape variables and improving the quality of biological data are essential for improving model prediction for conservation purposes.

Amphibian Hotspots and Conservation Priorities in Eastern Cuba Identified by Species Distribution Modeling

The high rate of amphibian endemism and the severe habitat modification in the Caribbean islands make them an ideal place to test if the current protected areas network might protect this group. In this study, we model distribution and map species richness of the 40 amphibian species from eastern Cuba with the objectives of identify hotspots, detect gaps in species representation in protected areas, and select additional areas to fill these gaps. We used two modeling methods, Maxent and Habitat Suitability Models, to reach a consensus distribution map for each species, then calculate species richness by combining specific models and finally performed gap analyses for species and hotspots. Our results showed that the models were robust enough to predict species distributions and that most of the amphibian hotspots were represented in reserves, but 50 percent of the species were incompletely covered and Eleutherodactylus rivularis was totally uncovered by the protected areas. We identified 1441 additional km2 (9.9% of the study area) that could be added to the current protected areas, allowing the representation of every species and all hotspots. Our results are relevant for the conservation planning in other Caribbean islands, since studies like this could contribute to fill the gaps in the existing protected areas and to design a future network. Both cases would benefit from modeling amphibian species distribution using available data, even if they are incomplete, rather than relying only in the protection of known or suspected hotspots.

Three species distribution models for the marine diatom Fragilariopsis kerguelensis of the Southern Ocean, links to NetCDF files

Fragilariopsis kerguelensis, a dominant diatom species throughout the Antarctic Circumpolar Current, is coined to be one of the main drivers of the biological silicate pump. Here, we study the distribution of this important species and expected consequences of climate change upon it, using correlative species distribution modeling and publicly available presence-only data. As experience with SDM is scarce for marine phytoplankton, this also serves as a pilot study for this organism group. We used the maximum entropy method to calculate distribution models for the diatom F. kerguelensis based on yearly and monthly environmental data (sea surface temperature, salinity, nitrate and silicate concentrations). Observation data were harvested from GBIF and the Global Diatom Database, and for further analyses also from the Hustedt Diatom Collection (BRM). The models were projected on current yearly and seasonal environmental data to study current distribution and its seasonality. Furthermore, we projected the seasonal model on future environmental data obtained from climate models for the year 2100. Projected on current yearly averaged environmental data, all models showed similar distribution patterns for F. kerguelensis. The monthly model showed seasonality, for example, a shift of the southern distribution boundary toward the north in the winter. Projections on future scenarios resulted in a moderately to negligibly shrinking distribution area and a change in seasonality. We found a substantial bias in the publicly available observation datasets, which could be reduced by additional observation records we obtained from the Hustedt Diatom Collection. Present-day distribution patterns inferred from the models coincided well with background knowledge and previous reports about F. kerguelensis distribution, showing that maximum entropy-based distribution models are suitable to map distribution patterns for oceanic planktonic organisms. Our scenario projections indicate moderate effects of climate change upon the biogeography of F. kerguelensis.

Importance of estimating matrix quality for modeling species distribution in complex tropical landscapes: a test with Atlantic forest small mammals

Information to guide decision making is especially urgent in human dominated landscapes in the tropics, where urban and agricultural frontiers are still expanding in an unplanned manner. Nevertheless, most studies that have investigated the influence of landscape structure on species distribution have not considered the heterogeneity of altered habitats of the matrix, which is usually high in human dominated landscapes. Using the distribution of small mammals in forest remnants and in the four main altered habitats in an Atlantic forest landscape, we investigated 1) how explanatory power of models describing species distribution in forest remnants varies between landscape structure variables that do or do not incorporate matrix quality and 2) the importance of spatial scale for analyzing the influence of landscape structure. We used standardized sampling in remnants and altered habitats to generate two indices of habitat quality, corresponding to the abundance and to the occurrence of small mammals. For each remnant, we calculated habitat quantity and connectivity in different spatial scales, considering or not the quality of surrounding habitats. The incorporation of matrix quality increased model explanatory power across all spatial scales for half the species that occurred in the matrix, but only when taking into account the distance between habitat patches (connectivity). These connectivity models were also less affected by spatial scale than habitat quantity models. The few consistent responses to the variation in spatial scales indicate that despite their small size, small mammals perceive landscape features at large spatial scales. Matrix quality index corresponding to species occurrence presented a better or similar performance compared to that of species abundance. Results indicate the importance of the matrix for the dynamics of fragmented landscapes and suggest that relatively simple indices can improve our understanding of species distribution, and could be applied in modeling, monitoring and managing complex tropical landscapes.

Predictive distribution modeling for rare Himalayan medicinal plant Berberis aristata DC

Predictive distribution modelling of Berberis aristata DC, a rare threatened plant with high medicinal values has been done with an aim to understand its potential distribution zones in Indian Himalayan region. Bioclimatic and topographic variables were used to develop the distribution model with the help of three different algorithms viz. GeneticAlgorithm for Rule-set Production (GARP), Bioclim and Maximum entroys(MaxEnt). Maximum entropy has predicted wider potential distribution (10.36%) compared to GARP (4.63%) and Bioclim (2.44%). Validation confirms that these outputs are comparable to the present distribution pattern of the B. atistata. This exercise highlights that this species favours Western Himalaya. However, GARP and MaxEnt's prediction of Eastern Himalayan states (i.e. Arunachal Pradesh, Nagaland and Manipur) are also identified as potential occurrence places require further exploration.

Otter (Lutra lutra) distribution modeling at two resolution scales suited to conservation planning in the Iberian Peninsula

We used the results of the Spanish Otter Survey of 1994–1996, a Geographic Information System and stepwise multiple logistic regression to model otter presence/absence data in the continental Spanish UTM 10 10-km squares. Geographic situation, indicators of human activity such as highways and major urban centers, and environmental variables related with productivity, water availability, altitude, and environmental energy were included in a logistic model that correctly classified about 73% of otter presences and absences. We extrapolated the model to the adjacent territory of Portugal, and increased the model’s spatial resolution by extrapolating it to 1 1-km squares in the whole Iberian Peninsula. The model turned out to be rather flexible, predicting, for instance, the species to be very restricted to the courses of rivers in some areas, and more widespread in others. This allowed us to determine areas where otter populations may be more vulnerable to habitat changes or harmful human interventions. # 2003 Elsevier Ltd. All rights reserved.

Spatial scale effects of sampling on the interpolation of species distribution models in the southwestern Amazon.

Knowledge of the geographical distribution of timber tree species in the Amazon is still scarce. This is especially true at the local level, thereby limiting natural resource management actions. Forest inventories are key sources of information on the occurrence of such species. However, areas with approved forest management plans are mostly located near access roads and the main industrial centers. The present study aimed to assess the spatial scale effects of forest inventories used as sources of occurrence data in the interpolation of potential species distribution models. The occurrence data of a group of six forest tree species were divided into four geographical areas during the modeling process. Several sampling schemes were then tested applying the maximum entropy algorithm, using the following predictor variables: elevation, slope, exposure, normalized difference vegetation index (NDVI) and height above the nearest drainage (HAND). The results revealed that using occurrence data from only one geographical area with unique environmental characteristics increased both model overfitting to input data and omission error rates. The use of a diagonal systematic sampling scheme and lower threshold values led to improved model performance. Forest inventories may be used to predict areas with a high probability of species occurrence, provided they are located in forest management plan regions representative of the environmental range of the model projection area.

Global genetic diversity and geographical and host-species distribution of beak and feather disease virus isolates

Psittacine beak and feather disease (PBFD) has a broad host range and is widespread in wild and captive psittacine populations in Asia, Africa, the Americas, Europe and Australasia. Beak and feather disease circovirus (BFDV) is the causative agent. BFDV has an ~2 kb single stranded circular DNA genome encoding just two proteins (Rep and CP). In this study we provide support for demarcation of BFDV strains by phylogenetic analysis of 65 complete genomes from databases and 22 new BFDV sequences isolated from infected psittacines in South Africa. We propose 94% genome-wide sequence identity as a strain demarcation threshold, with isolates sharing > 94% identity belonging to the same strain, and strain subtypes sharing> 98% identity. Currently, BFDV diversity falls within 14 strains, with five highly divergent isolates from budgerigars probably representing a new species of circovirus with three strains (budgerigar circovirus; BCV-A, -B and -C). The geographical distribution of BFDV and BCV strains is strongly linked to the international trade in exotic birds; strains with more than one host are generally located in the same geographical area. Lastly, we examined BFDV and BCV sequences for evidence of recombination, and determined that recombination had occurred in most BFDV and BCV strains. We established that there were two globally significant recombination hotspots in the viral genome: the first is along the entire intergenic region and the second is in the C-terminal portion of the CP ORF. The implications of our results for the taxonomy and classification of circoviruses are discussed. © 2011 SGM.

Uncertain species distribution: From interpolation to extrapolation

Species distribution models (SDMs) are considered to exemplify Pattern rather than Process based models of a species' response to its environment. Hence when used to map species distribution, the purpose of SDMs can be viewed as interpolation, since species response is measured at a few sites in the study region, and the aim is to interpolate species response at intermediate sites. Increasingly, however, SDMs are also being used to also extrapolate species-environment relationships beyond the limits of the study region as represented by the training data. Regardless of whether SDMs are to be used for interpolation or extrapolation, the debate over how to implement SDMs focusses on evaluating the quality of the SDM, both ecologically and mathematically. This paper proposes a framework that includes useful tools previously employed to address uncertainty in habitat modelling. Together with existing frameworks for addressing uncertainty more generally when modelling, we then outline how these existing tools help inform development of a broader framework for addressing uncertainty, specifically when building habitat models. As discussed earlier we focus on extrapolation rather than interpolation, where the emphasis on predictive performance is diluted by the concerns for robustness and ecological relevance. We are cognisant of the dangers of excessively propagating uncertainty. Thus, although the framework provides a smorgasbord of approaches, it is intended that the exact menu selected for a particular application, is small in size and targets the most important sources of uncertainty. We conclude with some guidance on a strategic approach to identifying these important sources of uncertainty. Whilst various aspects of uncertainty in SDMs have previously been addressed, either as the main aim of a study or as a necessary element of constructing SDMs, this is the first paper to provide a more holistic view.

Eliciting and encoding expert knowledge on variable selection into classical or Bayesian species distribution models

The quality of species distribution models (SDMs) relies to a large degree on the quality of the input data, from bioclimatic indices to environmental and habitat descriptors (Austin, 2002). Recent reviews of SDM techniques, have sought to optimize predictive performance e.g. Elith et al., 2006. In general SDMs employ one of three approaches to variable selection. The simplest approach relies on the expert to select the variables, as in environmental niche models Nix, 1986 or a generalized linear model without variable selection (Miller and Franklin, 2002). A second approach explicitly incorporates variable selection into model fitting, which allows examination of particular combinations of variables. Examples include generalized linear or additive models with variable selection (Hastie et al. 2002); or classification trees with complexity or model based pruning (Breiman et al., 1984, Zeileis, 2008). A third approach uses model averaging, to summarize the overall contribution of a variable, without considering particular combinations. Examples include neural networks, boosted or bagged regression trees and Maximum Entropy as compared in Elith et al. 2006. Typically, users of SDMs will either consider a small number of variable sets, via the first approach, or else supply all of the candidate variables (often numbering more than a hundred) to the second or third approaches. Bayesian SDMs exist, with several methods for eliciting and encoding priors on model parameters (see review in Low Choy et al. 2010). However few methods have been published for informative variable selection; one example is Bayesian trees (O’Leary 2008). Here we report an elicitation protocol that helps makes explicit a priori expert judgements on the quality of candidate variables. This protocol can be flexibly applied to any of the three approaches to variable selection, described above, Bayesian or otherwise. We demonstrate how this information can be obtained then used to guide variable selection in classical or machine learning SDMs, or to define priors within Bayesian SDMs.