Mapping,Bayesian Geostatistical Analysis and Spatial Prediction of Lymphatic Filariasis Prevalence in Africa

There is increasing interest to control or eradicate the major neglected tropical diseases. Accurate modelling of the geographic distributions of parasitic infections will be crucial to this endeavour. We used 664 community level infection prevalence data collated from the published literature in conjunction with eight environmental variables, altitude and population density, and a multivariate Bayesian generalized linear spatial model that allows explicit accounting for spatial autocorrelation and incorporation of uncertainty in input data and model parameters, to construct the first spatially-explicit map describing LF prevalence distribution in Africa. We also ran the best-fit model against predictions made by the HADCM3 and CCCMA climate models for 2050 to predict the likely distributions of LF under future climate and population changes. We show that LF prevalence is strongly influenced by spatial autocorrelation between locations but is only weakly associated with environmental covariates. Infection prevalence, however, is found to be related to variations in population density. All associations with key environmental/demographic variables appear to be complex and non-linear. LF prevalence is predicted to be highly heterogenous across Africa, with high prevalences (>20%) estimated to occur primarily along coastal West and East Africa, and lowest prevalences predicted for the central part of the continent. Error maps, however, indicate a need for further surveys to overcome problems with data scarcity in the latter and other regions. Analysis of future changes in prevalence indicates that population growth rather than climate change per se will represent the dominant factor in the predicted increase/decrease and spread of LF on the continent. We indicate that these results could play an important role in aiding the development of strategies that are best able to achieve the goals of parasite elimination locally and globally in a manner that may also account for the effects of future climate change on parasitic infection.     

Climatic niche divergence and habitat suitability of eight alien invasive weeds in China under climate change

Testing climatic niche divergence and modeling habitat suitability under conditions of climate change are important for developing strategies to limit the introduction and expansion of alien invasive weeds (AIWs) and providing important ecological and evolutionary insights. We assessed climatic niches in both native and invasive ranges as well as habitat suitability under climate change for eight representative Chinese AIWs from the American continent. We used climatic variables associated with occurrence records and developed ecological niche models with Maxent. Interestingly, the climatic niches of all eight AIWs diverged significantly between the native and invasive ranges (the American continent and China). Furthermore, the AIWs showed larger climatic niche breadths in the invasive ranges than in the native ranges. Our results suggest that climatic niche shifts between native and invasive ranges occurred. Thus, the occurrence records of both native and invasive regions must be considered when modeling and predicting the spatial distributions of AIWs under current and future climate scenarios. Owing to high habitat suitability, AIWs were more likely to expand into regions of low latitude, and future climate change was predicted to result in a shift in the AIWs in Qinghai and Tibet (regions of higher altitude) as well as Heilongjiang, Jilin, Liaoning, Inner Mongolia, and Gansu (regions of higher latitude). Our results suggest that we need measures to prevent and control AIW expansion at the country‐wide level.     

Do geographic distribution,niche property and life form explain plants' vulnerability to global change?

We modelled the future distribution in 2050 of 975 endemic plant species in southern Africa distributed among seven life forms, including new methodological insights improving the accuracy and ecological realism of predictions of global changes studies by: (i) using only endemic species as a way to capture the full realized niche of species, (ii) considering the direct impact of human pressure on landscape and biodiversity jointly with climate, and (iii) taking species' migration into account. Our analysis shows important promises for predicting the impacts of climate change in conjunction with land transformation. We have shown that the endemic flora of Southern Africa on average decreases with 41% in species richness among habitats and with 39% on species distribution range for the most optimistic scenario. We also compared the patterns of species' sensitivity with global change across life forms, using ecological and geographic characteristics of species. We demonstrate here that species and life form vulnerability to global changes can be partly explained according to species' (i) geographical distribution along climatic and biogeographic gradients, like climate anomalies, (ii) niche breadth or (iii) proximity to barrier preventing migration. Our results confirm that the sensitivity of a given species to global environmental changes depends upon its geographical distribution and ecological proprieties, and makes it possible to estimate a priori its potential sensitivity to these changes.     

Climate Change Influences on the Global Potential Distribution of Bluetongue Virus

The geographic distribution of arboviruses has received considerable attention after several dramatic emergence events around the world. Bluetongue virus (BTV) is classified among category “A” diseases notifiable to the World Organization of Animal Health (OIE), and is transmitted among ruminants by biting midges of the genus Culicoides. Here, we developed a comprehensive occurrence data set to map the current distribution, estimate the ecological niche, and explore the future potential distribution of BTV globally using ecological niche modeling and based on diverse future climate scenarios from general circulation models (GCMs) for four representative concentration pathways (RCPs). The broad ecological niche and potential geographic distribution of BTV under present-day conditions reflected the disease’s current distribution across the world in tropical, subtropical, and temperate regions. All model predictions were significantly better than random expectations. As a further evaluation of model robustness, we compared our model predictions to 331 independent records from most recent outbreaks from the Food and Agriculture Organization Emergency Prevention System for Transboundary Animal and Plant Pests and Diseases Information System (EMPRES-i); all were successfully anticipated by the BTV model. Finally, we tested ecological niche similarity among possible vectors and BTV, and could not reject hypotheses of niche similarity. Under future-climate conditions, the potential distribution of BTV was predicted to broaden, especially in central Africa, United States, and western Russia.     

洲际入侵植物生态位稳定性研究进展

人类活动引起的大规模洲际物种交换与生物入侵, 改变了当地生态系统结构与功能, 使生物多样性受到日益严重的威胁。本文通过综合分析主要国家和地区入侵植物的地理起源, 发现洲际入侵主要包括东亚—北美、东亚—南美、欧洲—南非、欧洲—北美、欧洲—东亚、北美—大洋洲等, 这些洲际入侵造成的后果往往比陆内入侵更为严重。利用物种分布模型(SDMs)预测入侵物种潜在分布范围是有效管理和提早预防生物入侵的重要依据, 但这些模型的一个关键假定是: 入侵物种的生态位在空间和时间上是保守的、稳定的。然而, 对于远离原产地种群并能快速适应新生境的洲际入侵植物来说, 生态位可能发生显著的变化。入侵种能否在入侵地保持原有的生态位, 取决于制约其生态分布的限制因素和生态过程在不同地区间是否发生变化。本文中作者总结了洲际入侵与陆内入侵的生态与进化过程的异同点, 认为这些限制物种原产地分布的因素如扩散限制、种间互作、适应性进化、生态可塑性和种群遗传特性等均可能导致入侵物种生态位的改变。建议下一步的研究应该重视: (1)对生态位属性进行多尺度的研究, 包括时间、空间、环境或系统发育等几个方面; (2)对比生态位稳定与发生偏移的物种特性, 确定什么样的入侵物种更容易改变原有的生态位; (3)进行生态位时间动态格局研究, 探讨生态位变化的倾向、历史速率和偏移程度, 以便判定生态位变化趋势。这些研究结果将会进一步提高物种分布模型的预测能力, 有助于更为准确地揭示气候变化和物种入侵对生物多样性的影响。     

Conservatism of ecological niche characteristics in North American plant species over the Pleistocene-to-Recent transition

Aim  To provide a test of the conservatism of a species' niche over the last 20,000 years by tracking the distribution of eight pollen taxa relative to climate type as they migrated across eastern North America following the Last Glacial Maximum (LGM).
Location  North America.
Methods  We drew taxon occurrence data from the North American pollen records in the Global Pollen Database, representing eight pollen types – all taxa for which ≥5 distinct geographic occurrences were available in both the present day and at the LGM (21,000 years ago ± 3000 years). These data were incorporated into ecological niche models based on present-day and LGM climatological summaries available from the Palaeoclimate Modelling Intercomparison Project to produce predicted potential geographic distributions for each species at present and at the LGM. The output for each time period was projected onto the 'other' time period, and tested using independent known occurrence information from that period.
Results  The result of our analyses was that all species tested showed general conservatism in ecological characteristics over the climate changes associated with the Pleistocene-to-Recent transition.
Main conclusions  This analysis constitutes a further demonstration of general and pervasive conservatism in ecological niche characteristics over moderate periods of time despite profound changes in climate and environmental conditions. As such, our results reinforce the application of ecological niche modelling techniques to the reconstruction of Pleistocene biodiversity distribution patterns, and to project the future potential distribution range of species in the face of global-scale climatic changes.     

Geographical potential of Argentine ants (Linepithema humile Mayr) in the face of global climate change

Determining the spread and potential geographical distribution of invasive species is integral to making invasion biology a predictive science. We assembled a dataset of over 1000 occurrences of the Argentine ant (Linepithema humile), one of the world's worst invasive alien species. Native to central South America, Argentine ants are now found in many Mediterranean and subtropical climates around the world. We used this dataset to assess the species' potential geographical and ecological distribution, and to examine changes in its distributional potential associated with global climate change, using techniques for ecological niche modelling. Models developed were highly predictive of the species' overall range, including both the native distributional area and invaded areas worldwide. Despite its already widespread occurrence, L. humile has potential for further spread, with tropical coastal Africa and southeast Asia apparently vulnerable to invasion. Projecting ecological niche models onto four general circulation model scenarios of future (2050s) climates provided scenarios of the species' potential for distributional expansion with warming climates: generally, the species was predicted to retract its range in tropical regions, but to expand at higher latitude areas.     

Mapping suitability for Buruli ulcer at fine spatial scales across Africa: A modelling study

Buruli ulcer (BU) is a disabling and stigmatising neglected tropical disease (NTD). Its distribution and burden are unknown because of underdiagnosis and underreporting. It is caused by Mycobacterium ulcerans, an environmental pathogen whose environmental niche and transmission routes are not fully understood. The main control strategy is active surveillance to promote early treatment and thus limit morbidity, but these activities are mostly restricted to well-known endemic areas. A better understanding of environmental suitability for the bacterium and disease could inform targeted surveillance, and advance understanding of the ecology and burden of BU. We used previously compiled point-level datasets of BU and M. ulcerans occurrence, evidence for BU occurrence within national and sub-national areas, and a suite of relevant environmental covariates in a distribution modelling framework. We fitted relationships between BU and M. ulcerans occurrence and environmental predictors by applying regression and machine learning based algorithms, combined in an ensemble model to characterise the optimal ecological niche for the disease and bacterium across Africa at a resolution of 5km x 5km. Proximity to waterbodies was the strongest predictor of suitability for BU, followed potential evapotranspiration. The strongest predictors of suitability for M. ulcerans were deforestation and potential evapotranspiration. We identified patchy foci of suitability throughout West and Central Africa, including areas with no previous evidence of the disease. Predicted suitability for M. ulcerans was wider but overlapping with that of BU. The estimated population living in areas predicted suitable for the bacterium and disease was 46.1 million.These maps could be used to inform burden estimations and case searches which would generate a more complete understanding of the spatial distribution of BU in Africa, and may guide control programmes to identify cases beyond the well-known endemic areas.     

Conclusions about niche expansion in introduced Impatiens walleriana populations depend on method of analysis

Determining the degree to which climate niches are conserved across plant species' native and introduced ranges is valuable to developing successful strategies to limit the introduction and spread of invasive plants, and also has important ecological and evolutionary implications. Here, we test whether climate niches differ between native and introduced populations of Impatiens walleriana, globally one of the most popular horticultural species. We use approaches based on both raw climate data associated with occurrence points and ecological niche models (ENMs) developed with Maxent. We include comparisons of climate niche breadth in both geographic and environmental spaces, taking into account differences in available habitats between the distributional areas. We find significant differences in climate envelopes between native and introduced populations when comparing raw climate variables, with introduced populations appearing to expand into wetter and cooler climates. However, analyses controlling for differences in available habitat in each region do not indicate expansion of climate niches. We therefore cannot reject the hypothesis that observed differences in climate envelopes reflect only the limited environments available within the species' native range in East Africa. Our results suggest that models built from only native range occurrence data will not provide an accurate prediction of the potential for invasiveness if applied to areas containing a greater range of environmental combinations, and that tests of niche expansion may overestimate shifts in climate niches if they do not control carefully for environmental differences between distributional areas.     

Distribution of cryptic blue oat mite species in Australia: current and future climate conditions

• 1 Invertebrate pests, such as blue oat mites Penthaleus spp., cause significant economic damage to agricultural crops in Australia. Climate is a major driver of invertebrate species distributions and climate change is expected to shift pest assemblages and pest prevalence across Australia. At this stage, little is known of how individual species will respond to climate change.
• 2 We have mapped the current distribution for each of the three pest Penthaleus spp. in Australia and built ecological niche models for each species using the correlative modelling software, maxent . Predictor variables useful for describing the climate space of each species were determined and the models were projected into a range of future climate change scenarios to assess how climate change may alter species‐specific distribution patterns in Australia.
• 3 The distributions of the three cryptic Penthaleus spp. are best described with different sets of climatic variables. Suitable climate space for all species decreases under the climate change scenarios investigated in the present study. The models also indicate that the assemblage of Penthaleus spp. is likely to change across Australia, particularly in Western Australia, South Australia and Victoria.
• 4 These results show the distributions of the three Penthaleus spp. are correlated with different climatic variables, and that regional control of mite pests is likely to change in the future. A further understanding of ecological and physiological processes that may influence the distribution and pest status of mites is required.

     

Habitat availability and gene flow influence diverging local population trajectories under scenarios of climate change: a place‐based approach

Ecological niche theory holds that species distributions are shaped by a large and complex suite of interacting factors. Species distribution models (SDMs) are increasingly used to describe species’ niches and predict the effects of future environmental change, including climate change. Currently, SDMs often fail to capture the complexity of species’ niches, resulting in predictions that are generally limited to climate‐occupancy interactions. Here, we explore the potential impact of climate change on the American pika using a replicated place‐based approach that incorporates climate, gene flow, habitat configuration, and microhabitat complexity into SDMs. Using contemporary presence–absence data from occupancy surveys, genetic data to infer connectivity between habitat patches, and 21 environmental niche variables, we built separate SDMs for pika populations inhabiting eight US National Park Service units representing the habitat and climatic breadth of the species across the western United States. We then predicted occurrence probability under current (1981–2010) and three future time periods (out to 2100). Occurrence probabilities and the relative importance of predictor variables varied widely among study areas, revealing important local‐scale differences in the realized niche of the American pika. This variation resulted in diverse and – in some cases – highly divergent future potential occupancy patterns for pikas, ranging from complete extirpation in some study areas to stable occupancy patterns in others. Habitat composition and connectivity, which are rarely incorporated in SDM projections, were influential in predicting pika occupancy in all study areas and frequently outranked climate variables. Our findings illustrate the importance of a place‐based approach to species distribution modeling that includes fine‐scale factors when assessing current and future climate impacts on species’ distributions, especially when predictions are intended to manage and conserve species of concern within individual protected areas.     

Genome-wide genetic variation coupled with demographic and ecological niche modeling of the dusky-footed woodrat (Neotoma fuscipes) reveal patterns of deep divergence and widespread Holocene expansion across northern California

Understanding how species have responded to past climate change may help refine projections of how species and biotic communities will respond to future change. Here, we integrate estimates of genome-wide genetic variation with demographic and niche modeling to investigate the historical biogeography of an important ecological engineer: the dusky-footed woodrat, Neotoma fuscipes. We use RADseq to generate a genome-wide dataset for 71 individuals from across the geographic distribution of the species in California. We estimate population structure using several model-based methods and infer the demographic history of regional populations using a site frequency spectrum-based approach. Additionally, we use ecological niche modeling to infer current and past (Last Glacial Maximum) environmental suitability across the species’ distribution. Finally, we estimate the directionality and possible spatial origins of regional population expansions. Our analyses indicate this species is subdivided into three regionally distinct populations, with the deepest divergence occurring ~1.7 million years ago across the modern-day San Francisco-Bay Delta region; a common biogeographic barrier for the flora and fauna of California. Our models of environmental suitability through time coincide with our estimates of population expansion, with relative long-term stability in the southern portion of the range, and more recent expansion into the northern end of the range. Our study illustrates how the integration of genome-wide data with spatial and demographic modeling can reveal the timing and spatial extent of historic events that determine patterns of biotic diversity and may help predict biotic response to future change.Subject terms: Evolutionary genetics, Molecular evolution     

Future Risks of Pest Species under Changing Climatic Conditions

Most agricultural pests are poikilothermic species expected to respond to climate change. Currently, they are a tremendous burden because of the high losses they inflict on crops and livestock. Smallholder farmers in developing countries of Africa are likely to suffer more under these changes than farmers in the developed world because more severe climatic changes are projected in these areas. African countries further have a lower ability to cope with impacts of climate change through the lack of suitable adapted management strategies and financial constraints. In this study we are predicting current and future habitat suitability under changing climatic conditions for Tuta absoluta, Ceratitis cosyra, and Bactrocera invadens, three important insect pests that are common across some parts of Africa and responsible for immense agricultural losses. We use presence records from different sources and bioclimatic variables to predict their habitat suitability using the maximum entropy modelling approach. We find that habitat suitability for B. invadens, C. cosyra and T. absoluta is partially increasing across the continent, especially in those areas already overlapping with or close to most suitable sites under current climate conditions. Assuming a habitat suitability at three different threshold levels we assessed where each species is likely to be present under future climatic conditions and if this is likely to have an impact on productive agricultural areas. Our results can be used by African policy makers, extensionists and farmers for agricultural adaptation measures to cope with the impacts of climate change.     

Conservation of phylogeographic lineages under climate change

Aim We address the unexplored question of whether the lack of information on intra‐specific diversity inherent in species‐level niche modelling might bias evaluation of the conservation requirements of species and phylogeographic lineages under changing climates. We test for directional biases that might arise due to these methodological differences in ways of assessing risks from climate change. Location The African continent. Methods We identified from peer‐reviewed studies that used both nuclear and plastid markers the distribution of deep phylogeographic divisions within nine species of African mammals and their phylogeographic lineages. We fitted ecological niche models to describe currently suitable, occupied climates and to project the shift of suitable climate to two future time slices. We applied gap analysis to reveal potential changes in the protection of phylogeographic diversity owing to climatic shifts. Results We found that, within species, most phylogeographic lineages differ in the climates they experience and have substantial geographic separation. Models that do not distinguish these subspecific units often fail to identify potential risks of climate change to lineages. Modelled potential effects of climate change on the geographic extent of suitable climate vary in both direction and magnitude. Predictions of the persistence of suitable climate in current protected areas for the resident lineages differ on average by factor of 2 between species and lineage models. Main conclusions Our study develops an original synthetic approach by combining niche modelling, projected climate change, phylogeographic information and gap analysis. We clearly identify the potential benefits of using the new approach to evaluate risks to the conservation of intra‐specific genetic diversity that are posed by climate change. Our results suggest that prudent conservation strategies need to incorporate potential differences in climate tolerance among lineages when planning conservation measures for species confronted with environmental change.     

Contrasting climate niches among co‐occurring subdominant forbs of the sagebrush steppe

Aim

Abiotic conditions are key components that determine the distribution of species. However, co‐occurring species can respond differently to the same factors, and determining which climate components are most predictive of geographic distributions is important for understanding community response to climate change. Here, we estimate and compare climate niches of ten subdominant, herbaceous forb species common in sagebrush steppe systems, asking how niches differ among co‐occurring species and whether more closely related species exhibit higher niche overlap.

Location

Western United States.

Methods

We used herbarium records and ecological niche modelling to estimate area of occupancy, niche breadth and overlap, and describe characteristics of suitable climate. We compared mean values and variability in summer precipitation and minimum temperatures at occurrence locations among species, plant families, and growth forms, and related estimated phylogenetic distances to niche overlap.

Results

Species varied in the size and spatial distribution of suitable climate and in niche breadth. Species also differed in the variables contributing to their suitable climate and in mean values, spatial variation and interannual variation in highly predictive climate variables. Only two of ten species shared comparable climate niches. We found family‐level differences associated with variation in summer precipitation and minimum temperatures, as well as in mean minimum temperatures. Growth forms differed in their association with variability in summer precipitation and minimum temperatures. We found no relationship between phylogenetic distance and niche overlap among our species.

Main conclusions

We identified contrasting climate niches for ten Great Basin understorey forbs, including differences in both mean values and climate variability. These estimates can guide species selection for restoration by identifying species with a high tolerance for climate variability and large climatic niches. They can also help conservationists to understand which species may be least tolerant of climate variability, and potentially most vulnerable to climate change.

     

Divergent lineages and conserved niches: using ecological niche modeling to examine the evolutionary patterns of the Nile monitor (<Emphasis Type="Italic">Varanus niloticus</Emphasis>)

Ecological niche modeling is a useful tool that can support phylogeographic analyses, offering insight into the evolutionary processes that have generated present-day patterns of biodiversity. Findings of ecological divergence across evolutionary lineages can be utilized to bolster inferences of parapatric or sympatric modes of speciation, and provide support for species-level classifications. Conversely, conserved ecological niches across evolutionary timescales are thought to have facilitated allopatric speciation. Here, we examined the climatic niche of three genetic lineages of the Nile monitor (Varanus niloticus) to better understand the processes involved in generating patterns of genetic variation, and to potentially clarify their taxonomic status. We built ecological niche models using genetically confirmed occurrence points from the three evolutionary lineages of V. niloticus, occupying the western, northern, and southern regions of Africa. Pairwise comparisons of climatic niche overlap provided evidence in support of niche conservatism across all V. niloticus lineages. These findings are consistent with an allopatric mode of differentiation. Furthermore, climatic niche conservatism could have played a role in isolating V. niloticus populations during historic climate oscillations, generating the observed genetic patterns across Africa.     

Niche-based modelling as a tool for predicting the risk of alien plant invasions at a global scale

Predicting the probability of successful establishment of plant species by matching climatic variables has considerable potential for incorporation in early warning systems for the management of biological invasions. We select South Africa as a model source area of invasions worldwide because it is an important exporter of plant species to other parts of the world because of the huge international demand for indigenous flora from this biodiversity hotspot. We first mapped the five ecoregions that occur both in South Africa and other parts of the world, but the very coarse definition of the ecoregions led to unreliable results in terms of predicting invasible areas. We then determined the bioclimatic features of South Africa's major terrestrial biomes and projected the potential distribution of analogous areas throughout the world. This approach is much more powerful, but depends strongly on how particular biomes are defined in donor countries. Finally, we developed bioclimatic niche models for 96 plant taxa (species and subspecies) endemic to South Africa and invasive elsewhere, and projected these globally after successfully evaluating model projections specifically for three well‐known invasive species (Carpobrotus edulis, Senecio glastifolius, Vellereophyton dealbatum) in different target areas. Cumulative probabilities of climatic suitability show that high‐risk regions are spatially limited globally but that these closely match hotspots of plant biodiversity. These probabilities are significantly correlated with the number of recorded invasive species from South Africa in natural areas, emphasizing the pivotal role of climate in defining invasion potential. Accounting for potential transfer vectors (trade and tourism) significantly adds to the explanatory power of climate suitability as an index of invasibility. The close match that we found between the climatic component of the ecological habitat suitability and the current pattern of occurrence of South Africa alien species in other parts of the world is encouraging. If species' distribution data in the donor country are available, climatic niche modelling offers a powerful tool for efficient and unbiased first‐step screening. Given that eradication of an established invasive species is extremely difficult and expensive, areas identified as potential new sites should be monitored and quarantine measures should be adopted.     

Does local habitat fragmentation affect large‐scale distributions? The case of a specialist grassland bird

Aim

Although the negative effects of habitat fragmentation have been widely documented at the landscape scale, much less is known about its impacts on species distributions at the biogeographical scale. We hypothesize that fragmentation influences the large‐scale distribution of area‐ and edge‐sensitive species by limiting their occurrence in regions with fragmented habitats , despite otherwise favourable environmental conditions. We test this hypothesis by assessing the interplay of climate and landscape factors influencing the distribution of the calandra lark, a grassland specialist that is highly sensitive to habitat fragmentation.

Location

Iberia Peninsula, Europe.

Methods

Ecological niche modelling was used to investigate the relative influence of climate/topography, landscape fragmentation and spatial structure on calandra lark distribution. Modelling assumed explicitly a hierarchically structured effect among explanatory variables, with climate/topography operating at broader spatial scales than landscape variables. An eigenvector‐based spatial filtering approach was used to cancel bias introduced by spatial autocorrelation. The information theoretic approach was used in model selection, and variation partitioning was used to isolate the unique and shared effects of sets of explanatory variables.

Results

Climate and topography were the most influential variables shaping the distribution of calandra lark, but incorporating landscape metrics contributed significantly to model improvement. The probability of calandra lark occurrence increased with total habitat area and declined with the number of patches and edge density. Variation partitioning showed a strong overlap between variation explained by climate/topography and landscape variables. After accounting for spatial structure in species distribution, the explanatory power of environmental variables remained largely unchanged.

Main conclusions

We have shown here that landscape fragmentation can influence species distributions at the biogeographical scale. Incorporating fragmentation metrics into large‐scale ecological niche models may contribute for a better understanding of mechanism driving species distributions and for improving predictive modelling of range shifts associated with land use and climate changes.