Where will species go? Incorporating new advances in climate modelling into projections of species distributions

Bioclimatic models are the primary tools for simulating the impact of climate change on species distributions. Part of the uncertainty in the output of these models results from uncertainty in projections of future climates. To account for this, studies often simulate species responses to climates predicted by more than one climate model and/or emission scenario. One area of uncertainty, however, has remained unexplored: internal climate model variability. By running a single climate model multiple times, but each time perturbing the initial state of the model slightly, different but equally valid realizations of climate will be produced. In this paper, we identify how ongoing improvements in climate models can be used to provide guidance for impacts studies. In doing so we provide the first assessment of the extent to which this internal climate model variability generates uncertainty in projections of future species distributions, compared with variability between climate models. We obtained data on 13 realizations from three climate models (three from CSIRO Mark2 v3.0, four from GISS AOM, and six from MIROC v3.2) for two time periods: current (1985–1995) and future (2025–2035). Initially, we compared the simulated values for each climate variable (P, T_max, T_min, and T_mean) for the current period to observed climate data. This showed that climates simulated by realizations from the same climate model were more similar to each other than to realizations from other models. However, when projected into the future, these realizations followed different trajectories and the values of climate variables differed considerably within and among climate models. These had pronounced effects on the projected distributions of nine Australian butterfly species when modelled using the BIOCLIM component of DIVA-GIS. Our results show that internal climate model variability can lead to substantial differences in the extent to which the future distributions of species are projected to change. These can be greater than differences resulting from between-climate model variability. Further, different conclusions regarding the vulnerability of species to climate change can be reached due to internal model variability. Clearly, several climate models, each represented by multiple realizations, are required if we are to adequately capture the range of uncertainty associated with projecting species distributions in the future.     

The ability of climate envelope models to predict the effect of climate change on species distributions

Climate envelope models (CEMs) have been used to predict the distribution of species under current, past, and future climatic conditions by inferring a species' environmental requirements from localities where it is currently known to occur. CEMs can be evaluated for their ability to predict current species distributions but it is unclear whether models that are successful in predicting current distributions are equally successful in predicting distributions under different climates (i.e. different regions or time periods). We evaluated the ability of CEMs to predict species distributions under different climates by comparing their predictions with those obtained with a mechanistic model (MM). In an MM the distribution of a species is modeled based on knowledge of a species' physiology. The potential distributions of 100 plant species were modeled with an MM for current conditions, a past climate reconstruction (21 000 years before present) and a future climate projection (double preindustrial CO₂ conditions). Point localities extracted from the currently suitable area according to the MM were used to predict current, future, and past distributions with four CEMs covering a broad range of statistical approaches: Bioclim (percentile distributions), Domain (distance metric), GAM (general additive modeling), and Maxent (maximum entropy). Domain performed very poorly, strongly underestimating range sizes for past or future conditions. Maxent and GAM performed as well under current climates as under past and future climates. Bioclim slightly underestimated range sizes but the predicted ranges overlapped more with the ranges predicted with the MM than those predicted with GAM did. Ranges predicted with Maxent overlapped most with those produced with the MMs, but compared with the ranges predicted with GAM they were more variable and sometimes much too large. Our results suggest that some CEMs can indeed be used to predict species distributions under climate change, but individual modeling approaches should be validated for this purpose, and model choice could be made dependent on the purpose of a particular study.     

Latitudinal variation of life-history traits of an exotic and a native impatiens species in Europe

Understanding the responses of invasive and native populations to environmental change is crucial for reliable predictions of invasions in the face of global change. While comparisons of responses across invasive species with different life histories have been performed before, comparing functional traits of congeneric native and invasive species may help to reveal driving factors associated with invasion. Here we compared morphological functional trait patterns of an invasive species (Impatiens parviflora) with its congeneric native species (I. noli-tangere) along an approximately 1600 km European latitudinal gradient from France (49°34′N) to Norway (63°40′N). Soil nitrogen was recorded during six weeks of the growing season, and light, soil moisture, and nutrient availability were estimated for each sampled population using community weighted means of indicator values for co-occurring species. Temperature data were gathered from nearby weather stations.Both the native and invasive species are taller at higher latitudes and this response is strongest in the invasive species. Seed mass and number of seeds per capsule increase in I. noli-tangere but decrease in I. parviflora towards higher latitudes. Surprisingly, plant height in the invasive I. parviflora decreases with increasing soil nitrogen availability. The latitudinal pattern in seed mass is positively related to temperature in I. noli-tangere and negatively in I. parviflora. Leaf area of both species decreases with increasing Ellenberg indicator values for nitrogen and light but increases with increasing soil moisture. Soil nitrogen concentrations and Ellenberg indicator values for nitrogen have significant positive (I. noli-tangere) and negative (I. parviflora) effects on the number of seeds per capsule. Our results show that the native I. noli-tangere has efficient reproduction at its range edge while the invasive I. parviflora shows a marked decrease in seed size and seed number per capsule. These patterns are unrelated to the growth and obtained size of the plants: even low soil nitrogen availability in the north seemed not to limit plant growth and size. Our results suggest that the invasive I. parviflora tends to become more invasive at lower latitudes by producing heavier seeds and more seeds per capsule.     

Vulnerability of benthic habitats to the aquatic invasive species

A comparative vulnerability analysis of 16 selected benthic habitat types in the SE Baltic Sea waters and the Curonian lagoon, including Klaipeda strait, was performed using long-term monitoring datasets (1980–2003) and results of several other surveys in the lagoon and the sea. Results indicated that invasive species richness (number of alien species per habitat) in lagoon habitats was significantly higher than in the sea. Habitats formed by artificial rock and stone, sand, mud, and habitats modified by zebra mussel shell deposits appeared to be the most invaded. Highest invasive species richness occurred in habitats with high native species richness indicating that the main factors driving native species distribution (such as favourable physical conditions, habitat alterations generated by human or/and biotic activities) are also driving aquatic invaders. Physical factors distinguished to be the most important for native and invasive species distribution were salinity, depth range (expressed by the maximal and minimal depths difference within a habitat), shallowness of a habitat (expressed by a minimal depth), and availability of a hard substrate.     

Using species distributions models for designing conservation strategies of Tropical Andean biodiversity under climate change

Biodiversity in the Tropical Andes is under continuous threat from anthropogenic activities. Projected changes in climate will likely exacerbate this situation. Using species distribution models, we assess possible future changes in the diversity and climatic niche size of an unprecedented number of species for the region. We modeled a broad range of taxa (11,012 species of birds and vascular plants), including both endemic and widespread species and provide a comprehensive estimation of climate change impacts on the Andes. We find that if no dispersal is assumed, by 2050s, more than 50% of the species studied are projected to undergo reductions of at least 45% in their climatic niche, whilst 10% of species could be extinct. Even assuming unlimited dispersal, most of the Andean endemics (comprising ∼5% of our dataset) would become severely threatened (>50% climatic niche loss). While some areas appear to be climatically stable (e.g. Pichincha and Imbabura in Ecuador; and Nariño, Cauca, Valle del Cauca and Putumayo in Colombia) and hence depict little diversity loss and/or potential species gains, major negative impacts were also observed. Tropical high Andean grasslands (páramos and punas) and evergreen montane forests, two key ecosystems for the provision of environmental services in the region, are projected to experience negative changes in species richness and high rates of species turnover. Adapting to these impacts would require a landscape-network based approach to conservation, including protected areas, their buffer zones and corridors. A central aspect of such network is the implementation of an integrated landscape management approach based on sustainable management and restoration practices covering wider areas than currently contemplated.     

The checkerboard score and species distributions

Summary There has been an ongoing controversy over how to decide whether the distribution of species is random — i.e., whether it is not greatly different from what it would be if species did not interact. We recently showed (Roberts and Stone (1990)) that in the case of the Vanuatu (formerly New Hebrides) avifauna, the number of islands shared by species pairs was incompatible with a random null hypothesis. However, it was difficult to determine the causes or direction of the community's exceptionality. In this paper, the latter problem is examined further. We use Diamond's (1975) notion of checkerboard distributions (originally developed as an indicator of competition) and construct a C-score statistic which quantifies checkerboardedness. This statistic is based on the way two species might colonise a pair of islands; whenever each species colonises a different island this adds 1 to the C-score. Following Connor and Simberloff (1979) we generate a control group of random colonisation patterns (matrices), and use the C-score to determine their checkerboard characteristics. As an alternative mode of enquiry, we make slight alterations to the observed data, repeating this process many times so as to obtain another control group. In both cases, when we compare the observed data for the Vanuatu avifauna and the Antillean bat communities with that given by their respective control group, we find that these communities have significantly large checkerboard distributions, making implausible the hypothesis that their species distributions are a product of random colonisation.     

Potential impacts of climate change on the distributions and diversity patterns of European mammals

The Intergovernmental Panel on Climate Change (IPCC) predicts an increase in global temperatures of between 1.4°C and 5.8°C during the 21st century, as a result of elevated CO₂ levels. Using bioclimatic envelope models, we evaluate the potential impact of climate change on the distributions and species richness of 120 native terrestrial non-volant European mammals under two of IPCC’s future climatic scenarios. Assuming unlimited and no migration, respectively, our model predicts that 1% or 5–9% of European mammals risk extinction, while 32–46% or 70–78% may be severely threatened (lose > 30% of their current distribution) under the two scenarios. Under the no migration assumption endemic species were predicted to be strongly negatively affected by future climatic changes, while widely distributed species would be more mildly affected. Finally, potential mammalian species richness is predicted to become dramatically reduced in the Mediterranean region but increase towards the northeast and for higher elevations. Bioclimatic envelope models do not account for non-climatic factors such as land-use, biotic interactions, human interference, dispersal or history, and our results should therefore be seen as first approximations of the potential magnitude of future climatic changes.     

Phenological stages of natural species and their use as climate indicators

 The objectives of this paper are to: (1) present 10 years of phenological data for nine natural species growing in a Mediterranean-type climate, (2) present threshold temperatures that were derived for the computation of cumulative degree-days (CDD), and (3) evaluate the sensitivity of the nine natural species to weather variability. The study was conducted at the Phenological Research Garden of Oristano, Sardinia, Italy, during the period 1986–96. The observations were made on five typical Mediterranean species and four species that are typical of higher latitudes. The mean annual pattern of phenological events and the CDD from 1 January are given for each development stage. Temperature thresholds were evaluated by comparing the standard deviation about the mean number of days in the development period for each species. A good relationship between timing of phenophase occurrence and temperature was observed for the Mediterranean species, which were little affected by variations in rainfall. Phenological development of the non-native species was affected by springtime rainfall. Accepted: 28 October 1998     

Empirical methodology for predicting changes in species range extension and richness associated with climate warming

Species richness and extent of geographical range for North American freshwater amphipods and crayfish are strongly correlated with mean annual absolute, or range of, temperature, respectively. Such spatially based relationships are shown to be useful for predicting the biological consequences arising through scenarios of temporal climate warming.     

Alien vascular plants in Iceland: Diversity,spatial patterns,temporal trends,and the impact of climate change

The present study provides first comprehensive and up-to-date results on alien plant taxa in Iceland since 1967. We evidenced the presence of 336 alien vascular plant taxa, including 277 casuals and 59 naturalised taxa, two being invasive. The distribution of the alien flora exhibits a clear spatial pattern showing hotspots of occurrence and diversity within areas of major settlement centres. Altitude above sea level and temperature-related variables proved to be the most important factors shaping alien plant distribution in Iceland. Predictive modelling evidenced that arctic areas of Iceland and the Central Highlands are under serious risk of alien plant invasion due to climate change. The results provide crucial information for alien and invasive plant management and contribute data for meta-analyses of invasion processes worldwide.     

Species abundance distributions over time

It has been known for 50 years that the time period over which data are collected affects the shape of empirical species abundance distributions. However, despite a recent resurgence of interest in characterizing and explaining these patterns the temporal component of species abundance distributions has been largely ignored. I argue that it is essential to take account of time, and not only because sampling duration can have a profound influence on the perceived shape of the distribution. Partitions of species abundance distributions based on temporal occurrence in the record will facilitate tests of both biological and neutral models and may lead to a better understanding of rarity. These temporal partitions also have interesting, but as yet barely explored, parallels with spatial ones such as the core-satellite division. Moreover, changes in abundance distributions across all three of Preston's temporal scales (sampling time, ecological time and evolutionary time) present rich opportunities for ecological research.     

Invasion triangle: an organizational framework for species invasion

Species invasion is a complex, multifactor process. To encapsulate this complexity into an intuitively appealing, simple, and straightforward manner, we present an organizational framework in the form of an invasion triangle. The invasion triangle is an adaptation of the disease triangle used by plant pathologists to help envision and evaluate interactions among a host, a pathogen, and an environment. Our modification of this framework for invasive species incorporates the major processes that result in invasion as the three sides of the triangle: (1) attributes of the potential invader; (2) biotic characteristics of a potentially invaded site; and (3) environmental conditions of the site. The invasion triangle also includes the impact of external influences on each side of the triangle, such as climate and land use change. This paper introduces the invasion triangle, discusses how accepted invasion hypotheses are integrated in this framework, describes how the invasion triangle can be used to focus research and management, and provides examples of application. The framework provided by the invasion triangle is easy to use by both researchers and managers and also applicable at any level of data intensity, from expert opinion to highly controlled experiments. The organizational framework provided by the invasion triangle is beneficial for understanding and predicting why species are invasive in specific environments, for identifying knowledge gaps, for facilitating communication, and for directing management in regard to invasive species.     

Differential effects of past climate warming on mountain and flatland species distributions: a multispecies North American mammal assessment

Aim The magnitude of predicted range shifts during climate change is likely to be different for species living in mountainous environments compared with those living in flatland environments. The southern edges of ranges in mountain species may not shift northwards during warming as populations instead migrate up available elevational gradients; overall latitudinal range appears therefore to expand. In contrast, flatland species should shift range centroids northwards but not expand or contract their latitudinal range extent. These hypotheses were tested utilizing Late Pleistocene and modern occurrence data. Location North America. Methods The location and elevation of modern and Late Pleistocene species occurrences were collected from data bases for 26 species living in mountain or flatland environments. Regressions of elevation change over latitude, and southern and northern range edges were calculated for each species for modern and fossil data sets. A combination of regressions and anova s were used to test whether flatland species shift range edges and latitudinal extents more than mountain species do. Results Flatland species had significantly larger northward shifts at southern range edges than did mountain‐dwelling species from the Late Pleistocene to the present. There was also a significant negative correlation between the amount of change in the latitude of the southern edge of the range and the amount of elevational shifting from the Late Pleistocene to the present. Although significant, only c. 25% of the variance could be explained by this relationship. In addition, there was a weak indication that overall range expansion was less in flatland‐dwelling than in mountain‐dwelling species. Main conclusions The approach used here was to examine past species’ range responses to warming that occurred after the last ice ages as a means to better predict potential future responses to continued warming. The results confirm predictions of differential southern edge and overall range shifts for species occupying mountain and flatland regions in North America. The findings may be broadly applicable in other regions, thus allowing better modelling of future range and distribution related responses.     

Modelling the potential geographic distribution of invasive ant species in New Zealand

Despite their economic and environmental impacts, there have been relatively few attempts to model the distribution of invasive ant species. In this study, the potential distribution of six invasive ant species in New Zealand are modelled using three fundamentally different methods (BIOCLIM, DOMAIN, MAXENT). Species records were obtained from museum collections in New Zealand. There was a significant relationship between the length of time an exotic species had been present in New Zealand and its geographic range. This is the first time such a time lag has been described for exotic ant species, and shows there is a considerable time lag in their spread. For example, it has taken many species several decades (40–60 years) to obtain a distribution of 17–25% of New Zealand regions. For all six species, BIOCLIM performed poorly compared to the other two modelling methods. BIOCLIM had lower AUC scores and higher omission error, suggesting BIOCLIM models under-predicted the potential distribution of each species. Omission error was significantly higher between models fitted with all 19 climate variables compared to those models with fewer climate variables for BIOCLIM, but not DOMAIN or MAXENT. Widespread species had a greater commission error. A number of regions in New Zealand are predicted to be climatically suitable for the six species modelled, particularly coastal and lowland areas of both the North and South Islands.     

A Panax-centric view of invasive species

Plant-centric sampling provides a novel approach to quantifying the potential impact of invasive species on native plant species. The aim of this study was to determine the level of exposure of individuals and populations of Panax quinquefolius to invasive plant species using this approach in thirty natural ginseng populations. A high level of invasion was found with 63–70% of ginseng populations containing at least one invasive species. Approximately one-third of all individuals were found in close proximity to invasive plants. The most prevalent invasive species were Rosa multiflora and Berberis thunbergii. The exposure to invasives of plants in different size classes varied among populations. Invasive species presence increased with greater ginseng population sizes and presence of harvest. The abundance of invasives plants within forest interiors near this valuable medicinal herb suggests that the economic and ecological costs of competitive interactions with native species could be high.     

Scale, time, space, and predictability: species distributions of preimaginal black flies (Diptera: Simuliidae)

This study examines species-specific distributions of preimaginal black flies (Diptera: Simuliidae) over two large spatial scales (within and across ecoregions) and two seasons (spring and summer). The study area in the western two-thirds of South Carolina, United States, was divided into three ecoregions (Mountains, Piedmont, Sandhills). The mean correct classification of species distributions among stream sites was 81.3%. Predictability of species distributions varied with spatial scale, location, and time. Species occurrence was not independent of ecoregion; distributions of 12 of the 13 species examined at this scale were predictable on the basis of changes in water column (temperature, percentage dissolved oxygen) and channel characters (velocity, streambed-particle size). However, with the exception of the Mountains during the summer, predicting species distributions within ecoregions was far less successful than predicting distributions across these regions; predictability was particularly poor in the Sandhills. Seasonal effects on predictability were most pronounced in the Mountains. As stream sites became more homogeneous, predictive capability declined. Species-level identifications, aided by cytogenetic techniques, were of paramount importance in recognizing the association between species distributions and stream conditions, emphasizing that considerable ecological information is lost when species are not considered. This study emphasizes the critical role of taxonomic resolution, linked with spatial and temporal scale of observation, in elucidating patterns of species assemblages. It also demonstrates that predictability of species distributions at a fixed scale can vary with geographic location and time. Received: 2 May 1997 / Accepted: 22 September 1997     

Optimal invasive species management under multiple uncertainties

The management programs for invasive species have been proposed and implemented in many regions of the world. However, practitioners and scientists have not reached a consensus on how to control them yet. One reason is the presence of various uncertainties associated with the management. To give some guidance on this issue, we characterize the optimal strategy by developing a dynamic model of invasive species management under uncertainties. In particular, focusing on (i) growth uncertainty and (ii) measurement uncertainty, we identify how these uncertainties affect optimal strategies and value functions. Our results suggest that a rise in growth uncertainty causes the optimal strategy to involve more restrained removals and the corresponding value function to shift up. Furthermore, we also find that a rise in measurement uncertainty affects optimal policies in a highly complex manner, but their corresponding value functions generally shift down as measurement uncertainty rises. Overall, a rise in growth uncertainty can be beneficial, while a rise in measurement uncertainty brings about an adverse effect, which implies the potential gain of precisely identifying the current stock size of invasive species.     

Seed banks of invasive Australian Acacia species in South Africa: Role in invasiveness and options for management

Despite impressive efforts at clearing stands of invasive Australian Acacia species in South Africa, insufficient attention has been given to understanding the role of seed banks in the invasiveness and long-term persistence of populations. We review information on seeds of these species, considering seed production, seed rain, and the dynamics of seeds in three layers: leaf litter, and upper and lower seed banks in the soil. Many factors affect the accumulation and susceptibility to destruction of seed banks and thus the opportunities for intervention to reduce seed numbers for each of these components. Reduction of seed banks is crucial for the overall success of the multi-million dollar management initiatives against these species. Classical biological control of buds, flower and young pods has reduced the seed production of many Australian acacias in South Africa. Fire can be applied to reduce seed numbers in the leaf litter and upper seed bank in some cases, although there are serious problems associated with high fire intensities in dense acacia stands. Other options, e.g. soil inversion and solarisation, exist to exercise limited reduction of seed numbers in some situations. There is little prospect of meaningful reduction of seed numbers in the lower seed bank. Preventing the accumulation of seed banks by limiting seed production through biological control is by far the most effective means, and in almost all cases the only practical means, of reducing seed numbers. This must be an integral part of management strategies. Several invasive Australian acacias are already under effective biological control, and further work to identify additional potential agents for all the currently invasive species and potentially invasive alien species is the top priority for improving the efficiency of management programmes.