Modeling a spatially restricted distribution in the Neotropics: How the size of calibration area affects the performance of five presence-only methods

We here examine species distribution models for a Neotropical anuran restricted to ombrophilous areas in the Brazilian Atlantic Forest hotspot. We extend the known occurrence for the treefrog Hypsiboas bischoffi (Anura: Hylidae) through GPS field surveys and use five modeling methods (BIOCLIM, DOMAIN, OM-GARP, SVM, and MAXENT) and selected bioclimatic and topographic variables to model the species distribution. Models were first trained using two calibration areas: the Brazilian Atlantic Forest (BAF) and the whole of South America (SA). All modeling methods showed good levels of predictive power and accuracy with mean AUC ranging from 0.77 (BIOCLIM/BAF) to 0.99 (MAXENT/SA). MAXENT and SVM were the most accurate presence-only methods among those tested here. All but the SVM models calibrated with SA predicted larger distribution areas when compared to models calibrated in BAF. OM-GARP dramatically overpredicted the species distribution for the model calibrated in SA, with a predicted area around 10⁶ km² larger than predicted by other SDMs. With increased calibration area (and environmental space), OM-GARP predictions followed changes in the environmental space associated with the increased calibration area, while MAXENT models were more consistent across calibration areas. MAXENT was the only method that retrieved consistent predictions across calibration areas, while allowing for some overprediction, a result that may be relevant for modeling the distribution of other spatially restricted organisms.     

How decisions about fitting species distribution models affect conservation outcomes

Species distribution models (SDMs) are increasingly used in conservation and land-use planning as inputs to describe biodiversity patterns. These models can be built in different ways, and decisions about data preparation, selection of predictor variables, model fitting, and evaluation all alter the resulting predictions. Commonly, the true distribution of species is unknown and independent data to verify which SDM variant to choose are lacking. Such model uncertainty is of concern to planners. We analyzed how 11 routine decisions about model complexity, predictors, bias treatment, and setting thresholds for predicted values altered conservation priority patterns across 25 species. Models were created with MaxEnt and run through Zonation to determine the priority rank of sites. Although all SDM variants performed well (area under the curve >0.7), they produced spatially different predictions for species and different conservation priority solutions. Priorities were most strongly altered by decisions to not address bias or to apply binary thresholds to predicted values; on average 40% and 35%, respectively, of all grid cells received an opposite priority ranking. Forcing high model complexity altered conservation solutions less than forcing simplicity (14% and 24% of cells with opposite rank values, respectively). Use of fewer species records to build models or choosing alternative bias treatments had intermediate effects (25% and 23%, respectively). Depending on modeling choices, priority areas overlapped as little as 10–20% with the baseline solution, affecting top and bottom priorities differently. Our results demonstrate the extent of model-based uncertainty and quantify the relative impacts of SDM building decisions. When it is uncertain what the best SDM approach and conservation plan is, solving uncertainty or considering alterative options is most important for those decisions that change plans the most.     

Impact of time series data on calibration and prediction uncertainty for a deterministic hydrogeochemical model

Model calibration is fundamental in applications of deterministic process-based models. Uncertainty in model predictions depends much on the input data and observations available for model calibration. Here we explored how model predictions (forecasts) and their uncertainties vary with the length of time series data used in calibration. As an example we used the hydrogeochemical model MAGIC and data from Birkenes, a small catchment in southern Norway, to simulate future water chemistry under a scenario of reduced acid deposition. A Bayesian approach with a Markov Chain Monte Carlo (MCMC) technique was used to calibrate the model to different lengths of observed data (4–29 years) and to estimate the prediction uncertainty each calibration. The results show that the difference between modelled and observed water chemistry (calibration goodness of fit) in general decreases with increasing length of the time series used in calibration. However, there are considerable differences for different time series of the same length. The results also show that the uncertainties in predicted future acid neutralizing capacity were lowest (i.e. the distribution peak narrowest) when using the longest time series for calibration. As for calibration success, there were considerable differences between the future distributions (prediction uncertainty) for the different calibrations.     

Predicting species distributions from samples collected along roadsides

Predictive models of species distributions are typically developed with data collected along roads. Roadside sampling may provide a biased (nonrandom) sample; however, it is currently unknown whether roadside sampling limits the accuracy of predictions generated by species distribution models. We tested whether roadside sampling affects the accuracy of predictions generated by species distribution models by using a prospective sampling strategy designed specifically to address this issue. We built models from roadside data and validated model predictions at paired locations on unpaved roads and 200 m away from roads (off road), spatially and temporally independent from the data used for model building. We predicted species distributions of 15 bird species on the basis of point-count data from a landbird monitoring program in Montana and Idaho (U.S.A.). We used hierarchical occupancy models to account for imperfect detection. We expected predictions of species distributions derived from roadside-sampling data would be less accurate when validated with data from off-road sampling than when it was validated with data from roadside sampling and that model accuracy would be differentially affected by whether species were generalists, associated with edges, or associated with interior forest. Model performance measures (kappa, area under the curve of a receiver operating characteristic plot, and true skill statistic) did not differ between model predictions of roadside and off-road distributions of species. Furthermore, performance measures did not differ among edge, generalist, and interior species, despite a difference in vegetation structure along roadsides and off road and that 2 of the 15 species were more likely to occur along roadsides. If the range of environmental gradients is surveyed in roadside-sampling efforts, our results suggest that surveys along unpaved roads can be a valuable, unbiased source of information for species distribution models.     

Generality of Models that Predict the Distribution of Species: Conservation Activity and Reduction of Model Transferability for a Threatened Bustard

Abstract:  Predictive models can help clarify the distribution of poorly known species but should display strong transferability when applied to independent data. Nevertheless, model transferability for threatened tropical species is poorly studied. We built models predicting the incidence of the critically endangered Bengal Florican ( Houbaropsis bengalensis ) within the Tonle Sap (TLS) floodplain, Cambodia. Separate models were constructed with soil, land-use, and landscape data and species incidence sampled over the entire floodplain (12,000 km²) and from the Kompong Thom (KT) province (4000 km²). In each case, the probability of Bengal Florican presence within randomly selected 1 × 1 km squares was modeled by binary logistic regression with multimodel inference. We assessed the transferability of the KT model by comparing predictions with observed incidence elsewhere in the floodplain. In terms of standard model-validation statistics, the KT model showed good spatial transferability. Nevertheless, it overpredicted florican presence outside the KT calibration region, classifying 491 km² as suitable habitat compared with 237 km² predicted as suitable by the TLS model. This resulted from higher species incidence within the calibration region, probably owing to a program of conservation education and enforcement that has reduced persecution there. Because both research and conservation activity frequently focus on areas with higher density, such effects could be widespread, reducing transferability of predictive distribution models.     

Invasive species distribution modeling (iSDM): Are absence data and dispersal constraints needed to predict actual distributions?

Species distribution models (SDMs) based on statistical relationships between occurrence data and underlying environmental conditions are increasingly used to predict spatial patterns of biological invasions and prioritize locations for early detection and control of invasion outbreaks. However, invasive species distribution models (iSDMs) face special challenges because (i) they typically violate SDM's assumption that the organism is in equilibrium with its environment, and (ii) species absence data are often unavailable or believed to be too difficult to interpret. This often leads researchers to generate pseudo-absences for model training or utilize presence-only methods, and to confuse the distinction between predictions of potential vs. actual distribution. We examined the hypothesis that true-absence data, when accompanied by dispersal constraints, improve prediction accuracy and ecological understanding of iSDMs that aim to predict the actual distribution of biological invasions. We evaluated the impact of presence-only, true-absence and pseudo-absence data on model accuracy using an extensive dataset on the distribution of the invasive forest pathogen Phytophthora ramorum in California. Two traditional presence/absence models (generalized linear model and classification trees) and two alternative presence-only models (ecological niche factor analysis and maximum entropy) were developed based on 890 field plots of pathogen occurrence and several climatic, topographic, host vegetation and dispersal variables. The effects of all three possible types of occurrence data on model performance were evaluated with receiver operating characteristic (ROC) and omission/commission error rates. Results show that prediction of actual distribution was less accurate when we ignored true-absences and dispersal constraints. Presence-only models and models without dispersal information tended to over-predict the actual range of invasions. Models based on pseudo-absence data exhibited similar accuracies as presence-only models but produced spatially less feasible predictions. We suggest that true-absence data are a critical ingredient not only for accurate calibration but also for ecologically meaningful assessment of iSDMs that focus on predictions of actual distributions.     

Cross-validation of species distribution models: removing spatial sorting bias and calibration with a null model

Species distribution models are usually evaluated with cross-validation. In this procedure evaluation statistics are computed from model predictions for sites of presence and absence that were not used to train (fit) the model. Using data for 226 species, from six regions, and two species distribution modeling algorithms (Bioclim and MaxEnt), I show that this procedure is highly sensitive to "spatial sorting bias": the difference between the geographic distance from testing-presence to training-presence sites and the geographic distance from testing-absence (or testing-background) to training-presence sites. I propose the use of pairwise distance sampling to remove this bias, and the use of a null model that only considers the geographic distance to training sites to calibrate cross-validation results for remaining bias. Model evaluation results (AUC) were strongly inflated: the null model performed better than MaxEnt for 45% and better than Bioclim for 67% of the species. Spatial sorting bias and area under the receiver-operator curve (AUC) values increased when using partitioned presence data and random-absence data instead of independently obtained presence-absence testing data from systematic surveys. Pairwise distance sampling removed spatial sorting bias, yielding null models with an AUC close to 0.5, such that AUC was the same as null model calibrated AUC (cAUC). This adjustment strongly decreased AUC values and changed the ranking among species. Cross-validation results for different species are only comparable after removal of spatial sorting bias and/or calibration with an appropriate null model.     

Comparison of statistical and theoretical habitat models for conservation planning: the benefit of ensemble prediction

Selection of a modeling approach is an important step in the conservation planning process, but little guidance is available. We compared two statistical and three theoretical habitat modeling approaches representing those currently being used for avian conservation planning at landscape and regional scales: hierarchical spatial count (HSC), classification and regression tree (CRT), habitat suitability index (HSI), forest structure database (FS), and habitat association database (HA). We focused our comparison on models for five priority forest-breeding species in the Central Hardwoods Bird Conservation Region: Acadian Flycatcher, Cerulean Warbler, Prairie Warbler, Red-headed Woodpecker, and Worm-eating Warbler. Lacking complete knowledge on the distribution and abundance of each species with which we could illuminate differences between approaches and provide strong grounds for recommending one approach over another, we used two approaches to compare models: rank correlations among model outputs and comparison of spatial correspondence. In general, rank correlations were significantly positive among models for each species, indicating general agreement among the models. Worm-eating Warblers had the highest pairwise correlations, all of which were significant (P < 0.05). Red-headed Woodpeckers had the lowest agreement among models, suggesting greater uncertainty in the relative conservation value of areas within the region. We assessed model uncertainty by mapping the spatial congruence in priorities (i.e., top ranks) resulting from each model for each species and calculating the coefficient of variation across model ranks for each location. This allowed identification of areas more likely to be good targets of conservation effort for a species, those areas that were least likely, and those in between where uncertainty is higher and thus conservation action incorporates more risk. Based on our results, models developed independently for the same purpose (conservation planning for a particular species in a particular geography) yield different answers and thus different conservation strategies. We assert that using only one habitat model (even if validated) as the foundation of a conservation plan is risky. Using multiple models (i.e., ensemble prediction) can reduce uncertainty and increase efficacy of conservation action when models corroborate one another and increase understanding of the system when they do not.     

Assessing the effects of pseudo-absences on predictive distribution model performance

Modelling species distributions with presence data from atlases, museum collections and databases is challenging. In this paper, we compare seven procedures to generate pseudo-absence data, which in turn are used to generate GLM-logistic regressed models when reliable absence data are not available. We use pseudo-absences selected randomly or by means of presence-only methods (ENFA and MDE) to model the distribution of a threatened endemic Iberian moth species (Graellsia isabelae). The results show that the pseudo-absence selection method greatly influences the percentage of explained variability, the scores of the accuracy measures and, most importantly, the degree of constraint in the distribution estimated. As we extract pseudo-absences from environmental regions further from the optimum established by presence data, the models generated obtain better accuracy scores, and over-prediction increases. When variables other than environmental ones influence the distribution of the species (i.e., non-equilibrium state) and precise information on absences is non-existent, the random selection of pseudo-absences or their selection from environmental localities similar to those of species presence data generates the most constrained predictive distribution maps, because pseudo-absences can be located within environmentally suitable areas. This study shows that if we do not have reliable absence data, the method of pseudo-absence selection strongly conditions the obtained model, generating different model predictions in the gradient between potential and realized distributions.     

A Behaviorally Explicit Demographic Model Integrating Habitat Selection and Population Dynamics in California Sea Lions

Abstract: Although there has been a call for the integration of behavioral ecology and conservation biology, there are few tools currently available to achieve this integration. Explicitly including information about behavioral strategies in population viability analyses may enhance the ability of conservation biologists to understand and estimate patterns of extinction risk. Nevertheless, most behavioral‐based PVA approaches require detailed individual‐based data that are rarely available for imperiled species. We present a mechanistic approach that incorporates spatial and demographic consequences of behavioral strategies into population models used for conservation. We developed a stage‐structured matrix model that includes the costs and benefits of movement associated with 2 habitat‐selection strategies (philopatry and direct assessment). Using a life table for California sea lions (Zalophus californianus), we explored the sensitivity of model predictions to the inclusion of these behavioral parameters. Including behavioral information dramatically changed predicted population sizes, model dynamics, and the expected distribution of individuals among sites. Estimated population sizes projected in 100 years diverged up to 1 order of magnitude among scenarios that assumed different movement behavior. Scenarios also exhibited different model dynamics that ranged from stable equilibria to cycles or extinction. These results suggest that inclusion of behavioral data in viability models may improve estimates of extinction risk for imperiled species. Our approach provides a simple method for incorporating spatial and demographic consequences of behavioral strategies into population models and may be easily extended to other species and behaviors to understand the mechanisms of population dynamics for imperiled populations.     

Predicting tree distributions in an East African biodiversity hotspot: model selection, data bias and envelope uncertainty

The Eastern Arc Mountains (EAMs) of Tanzania and Kenya support some of the most ancient tropical rainforest on Earth. The forests are a global priority for biodiversity conservation and provide vital resources to the Tanzanian population. Here, we make a first attempt to predict the spatial distribution of 40 EAM tree species, using generalised additive models, plot data and environmental predictor maps at sub 1 km resolution. The results of three modelling experiments are presented, investigating predictions obtained by (1) two different procedures for the stepwise selection of predictors, (2) down-weighting absence data, and (3) incorporating an autocovariate term to describe fine-scale spatial aggregation. In response to recent concerns regarding the extrapolation of model predictions beyond the restricted environmental range of training data, we also demonstrate a novel graphical tool for quantifying envelope uncertainty in restricted range niche-based models (envelope uncertainty maps). We find that even for species with very few documented occurrences useful estimates of distribution can be achieved. Initiating selection with a null model is found to be useful for explanatory purposes, while beginning with a full predictor set can over-fit the data. We show that a simple multimodel average of these two best-model predictions yields a superior compromise between generality and precision (parsimony). Down-weighting absences shifts the balance of errors in favour of higher sensitivity, reducing the number of serious mistakes (i.e., falsely predicted absences); however, response functions are more complex, exacerbating uncertainty in larger models. Spatial autocovariates help describe fine-scale patterns of occurrence and significantly improve explained deviance, though if important environmental constraints are omitted then model stability and explanatory power can be compromised. We conclude that the best modelling practice is contingent both on the intentions of the analyst (explanation or prediction) and on the quality of distribution data; generalised additive models have potential to provide valuable information for conservation in the EAMs, but methods must be carefully considered, particularly if occurrence data are scarce. Full results and details of all species models are supplied in an online Appendix.     

Predicting abundance of desert riparian birds: validation and calibration of the Effective Area Model.

Reliable prediction of the effects of landscape change on species abundance is critical to land managers who must make frequent, rapid decisions with long-term consequences. However, due to inherent temporal and spatial variability in ecological systems, previous attempts to predict species abundance in novel locations and/or time frames have been largely unsuccessful. The Effective Area Model (EAM) uses change in habitat composition and geometry coupled with response of animals to habitat edges to predict change in species abundance at a landscape scale. Our research goals were to validate EAM abundance predictions in new locations and to develop a calibration framework that enables absolute abundance predictions in novel regions or time frames. For model validation, we compared the EAM to a null model excluding edge effects in terms of accurate prediction of species abundance. The EAM outperformed the null model for 83.3% of species (N=12) for which it was possible to discern a difference when considering 50 validation sites. Likewise, the EAM outperformed the null model when considering subsets of validation sites categorized on the basis of four variables (isolation, presence of water, region, and focal habitat). Additionally, we explored a framework for producing calibrated models to decrease prediction error given inherent temporal and spatial variability in abundance. We calibrated the EAM to new locations using linear regression between observed and predicted abundance with and without additional habitat covariates. We found that model adjustments for unexplained variability in time and space, as well as variability that can be explained by incorporating additional covariates, improved EAM predictions. Calibrated EAM abundance estimates with additional site-level variables explained a significant amount of variability (P < 0.05) in observed abundance for 17 of 20 species, with R2 values >25% for 12 species, >48% for six species, and >60% for four species when considering all predictive models. The calibration framework described in this paper can be used to predict absolute abundance in sites different from those in which data were collected if the target population of sites to which one would like to statistically infer is sampled in a probabilistic way.     

Modelling species distributions with penalised logistic regressions: A comparison with maximum entropy models

An important aspect of species distribution modelling is the choice of the modelling method because a suboptimal method may have poor predictive performance. Previous comparisons have found that novel methods, such as Maxent models, outperform well-established modelling methods, such as the standard logistic regression. These comparisons used training samples with small numbers of occurrences per estimated model parameter, and this limited sample size may have caused poorer predictive performance due to overfitting. Our hypothesis is that Maxent models would outperform a standard logistic regression because Maxent models avoid overfitting by using regularisation techniques and a standard logistic regression does not. Regularisation can be applied to logistic regression models using penalised maximum likelihood estimation. This estimation procedure shrinks the regression coefficients towards zero, causing biased predictions if applied to the training sample but improving the accuracy of new predictions. We used Maxent and logistic regression (standard and penalised) to analyse presence/pseudo-absence data for 13 tree species and evaluated the predictive performance (discrimination) using presence-absence data. The penalised logistic regression outperformed standard logistic regression and equalled the performance of Maxent. The penalised logistic regression may be considered one of the best methods to develop species distribution models trained with presence/pseudo-absence data, as it is comparable to Maxent. Our results encourage further use of the penalised logistic regression for species distribution modelling, especially in those cases in which a complex model must be fitted to a sample with a limited size.     

Forecasting New Zealand Mudsnail invasion range: model comparisons using native and invaded ranges.

Evaluations of the potential distribution of invasive species can increase the efficiency of their management by focusing prevention measures. Generally, ecological models are built using occurrence data from a species' native range to predict the distribution in areas that the species may invade. However, historical and geographical constraints can limit a species' native distribution. Genetic Algorithm for Rule-set Production (GARP), an ecological niche modeling program, was used to predict the potential distribution of the invasive, freshwater New Zealand mudsnail, Potamopyrgus antipodarum, in Australia and North America. We compared the strength of the predictions made by models built with data from the snail's native range in New Zealand to models built with data from the locations invaded by the species. A time-series analysis of the Australian models demonstrated that range-of-invasion data can make better predictions about the potential distribution of invasive species than models built with native range data. Large differences among the model forecasts indicate that uncritical choice of the data set used in training the GARP models can result in misleading predictions. The models predict a large expansion in the range of P. antipodarum in both Australia and North America unless prevention measures are implemented rapidly.     

Parameterizing,evaluating and comparing metapopulation models with data from individual-based simulations

Due to the lack of sufficient data and appropriate ecological information parameterizing predictive population dynamical models usually is a difficult task. The approach proposed in this study is meant to overcome this problem by using detailed individual-based simulations to generate artificial data. With short-term data samples, the models to be investigated can be parameterized and their predictions be compared. The flexibility of individual-based simulations as experimental tools also facilitates the evaluation and comparison of different (aggregated) model types. The presented approach is a step towards unifying models of different complexity. As an example we applied it to two metapopulation models of insect species in a highly fragmented landscape: the well-known incidence function model with a patch-based representation of space and a grid-based analogue. The models are tested with respect to their data requirement and recommendations for a better data sampling are derived.     

Use of Coarse‐Resolution Models of Species’ Distributions to Guide Local Conservation Inferences

Abstract: Distribution models are used increasingly for species conservation assessments over extensive areas, but the spatial resolution of the modeled data and, consequently, of the predictions generated directly from these models are usually too coarse for local conservation applications. Comprehensive distribution data at finer spatial resolution, however, require a level of sampling that is impractical for most species and regions. Models can be downscaled to predict distribution at finer resolutions, but this increases uncertainty because the predictive ability of models is not necessarily consistent beyond their original scale. We analyzed the performance of downscaled, previously published models of environmental favorability (a generalized linear modeling technique) for a restricted endemic insectivore, the Iberian desman (Galemys pyrenaicus), and a more widespread carnivore, the Eurasian otter (Lutra lutra), in the Iberian Peninsula. The models, built from presence–absence data at 10 × 10 km resolution, were extrapolated to a resolution 100 times finer (1 × 1 km). We compared downscaled predictions of environmental quality for the two species with published data on local observations and on important conservation sites proposed by experts. Predictions were significantly related to observed presence or absence of species and to expert selection of sampling sites and important conservation sites. Our results suggest the potential usefulness of downscaled projections of environmental quality as a proxy for expensive and time‐consuming field studies when the field studies are not feasible. This method may be valid for other similar species if coarse‐resolution distribution data are available to define high‐quality areas at a scale that is practical for the application of concrete conservation measures.     

Incorporating low-resolution historic species location data decreases performance of distribution models

Developing robust species distribution models is important as model outputs are increasingly being incorporated into conservation policy and management decisions. A largely overlooked component of model assessment and refinement is whether to include historic species occurrence data in distribution models to increase the data sample size. Data of different temporal provenance often differ in spatial accuracy and precision. We test the effect of inclusion of historic coarse-resolution occurrence data on distribution model outputs for 187 species of birds in Australian tropical savannas. Models using only recent (after 1990), fine-resolution data had significantly higher model performance scores measured with area under the receiver operating characteristic curve (AUC) than models incorporating both fine- and coarse-resolution data. The drop in AUC score is positively correlated with the total area predicted to be suitable for the species (R² = 0.163-0.187, depending on the environmental predictors in the model), as coarser data generally leads to greater predicted areas. The remaining unexplained variation is likely to be due to the covariate errors resulting from resolution mismatch between species records and environmental predictors. We conclude that decisions regarding data use in species distribution models must be conscious of the variation in predictions that mixed-scale datasets might cause.     

Impact of the quality of climate models for modelling species occurrences in countries with poor climatic documentation: a case study from Bolivia

The quality of climate models has largely been overlooked as a possible source of uncertainty that may affect the outcomes of species distribution models, especially in the tropics, where comparatively few climatic stations are available. We compared the geographical discrepancies and potential conservation implications of using two different climate models (Saga and Worldclim) in combination with the species modelling approach Maxent in Bolivia. We estimated ranges of selected bird and fern species biogeographically restricted to either humid montane forest of the northern Bolivian Andes or seasonal dry tropical forests (in the Andes and southern lowlands). Saga and Worldclim predicted roughly similar climate patterns of temperature that were significantly correlated. Precipitation layers of both climate models were also roughly similar, but showed important differences. Species ranges estimated with Worldclim and Saga likewise produced different results. Ranges of species endemic to humid montane forests estimated with Saga had higher AUC (Area under the curve) values than those estimated with Worldclim, which for example predicted the occurrence of humid montane forest bird species near Lake Titicaca, an area that is clearly unsuitable for these species. Likewise, Worldclim overpredicted the occurrence of fern and bird species in the lowlands of the Chapare region and well south of the Andean Elbow, where more seasonal biomes occur. By contrast, Saga predictions were coherent with the known distribution of humid montane forests in the northern Bolivian Andes. Estimated ranges of species endemic to seasonal dry tropical forests predicted with Saga and Worldclim were not statistically different in most cases. However, detailed comparisons revealed that Saga was able to distinguish fragments of seasonal dry tropical forests in rain-shadow valleys of the northern Bolivian Andes, whereas Worldclim was not. These differences highlight the neglected influence of climate layers on modelling results and the importance of using the most accurate climate data available when modelling species distributions.     

Contrasting ecological information content in whaling archives with modern cetacean surveys for conservation planning and identification of historical distribution changes

Many species are restricted to a marginal or suboptimal fraction of their historical range due to anthropogenic impacts, making it hard to interpret their ecological preferences from modern-day data alone. However, inferring past ecological states is limited by the availability of robust data and biases in historical archives, posing a challenge for policy makers . To highlight how historical records can be used to understand the ecological requirements of threatened species and inform conservation, we investigated sperm whale (Physeter macrocephalus) distribution in the Western Indian Ocean. We assessed differences in information content and habitat suitability predictions based on whale occurrence data from Yankee whaling logs (1792–1912) and from modern cetacean surveys (1995–2020). We built maximum entropy habitat suitability models containing static (bathymetry-derived) variables to compare models comprising historical-only and modern-only data. Using both historical and modern habitat suitability predictions  we assessed marine protected area (MPA) placement by contrasting suitability in- and outside MPAs. The historical model predicted high habitat suitability in shelf and coastal regions near continents and islands, whereas the modern model predicted a less coastal distribution with high habitat suitability more restricted to areas of steep topography. The proportion of high habitat suitability inside versus outside MPAs was higher when applying the historical predictions than the modern predictions, suggesting that different marine spatial planning optimums can be reached from either data sources. Moreover, differences in relative habitat suitability predictions between eras were consistent with the historical depletion of sperm whales from coastal regions, which were easily accessed and targeted by whalers, resulting in a modern distribution limited more to steep continental margins and remote oceanic ridges. The use of historical data can provide important new insights and, through cautious interpretation, inform conservation planning and policy, for example, by identifying refugee species and regions of anticipated population recovery.     

Hard or soft environmental systems?

Recent trends in lake and stream water quality modeling indicate a conflict between the search for improved accuracy through increasing model size and complexity, and the search for applicability through simplification of already existing models. Much of this conflict turns on the fact that that which can be simulated in principle issimply not matched by that which can be observed and verified in practice. This paper is concerned with that conflict. Its aim is to introduce and clarify some of the arguments surrounding two issues of key importance in resolving the conflict: uncertainty in the mathematical relationships hypothesized for a particular model (calibration and model structure identification); and uncertainty associated with the predictions obtained from the model (prediction error analysis). These are issues concerning the reliability of models and model-based forecasts. The paper argues, in particular, that there is an intimate relationship between prediction and model calibration. This relationship is especially important in accounting for uncertainty in the development and use of models. Using this argument it is possible to state a dilemma which captures some limiting features of both large and small models.