Habitat associations of species show consistent but weak responses to climate

Different vegetation types can generate variation in microclimates at local scales, potentially buffering species from adverse climates. To determine if species could respond to such microclimates under climatic warming, we evaluated whether ectothermic species (butterflies) can exploit favourable microclimates and alter their use of different habitats in response to year-to-year variation in climate. In both relatively cold (Britain) and warm (Catalonia) regions of their geographical ranges, most species shifted into cooler, closed habitats (e.g. woodland) in hot years, and into warmer, open habitats (e.g. grassland) in cooler years. Additionally, three-quarters of species occurred in closed habitats more frequently in the warm region than in the cool region. Thus, species shift their local distributions and alter their habitat associations to exploit favourable microclimates, although the magnitude of the shift (approx. 1.3% of individuals from open to shade, per degree Celsius) is unlikely to buffer species from impacts of regional climate warming.     

Comparing species distributions modelled from occurrence data and from expert-based range maps. Implication for predicting range shifts with climate change

Species range and climate change risk are often assessed using species distribution models (SDM) that model species niche from presence points and environmental variables and project it in space and time. These presence points frequently originate from occurrence data downloaded from public biodiversity databases, but such data are known to suffer from high biases. There is thus a need to find alternative sources of information to train these models. In this regard, expert-based range maps such as those provided by the International Union for Conservation of Nature (IUCN) have the potential to be used as a source of species presence in a SDM workflow. Here, I compared the predictions of SDM built using true occurrences provided by GBIF or iNaturalist, or using pseudo-occurrences sampled from IUCN expert-based range maps, in current and future climate. I found that the agreement between both types of SDM did not depend on the spatial resolution of environmental data but instead were affected by the number of points sampled from range maps and even more by the spatial congruence between input data. A strong agreement between occurrence data and range maps resulted in very similar SDM outputs, which suggests that expert knowledge can be a valuable alternative source of data to feed SDM and assess potential range shifts when the only available occurrences are biased or fragmentary.     

Competition for S and N by associations of three annual range species in lysimeters

Summary Competition for S and N was studied on free draining lysimeters seeded to associations of three annual forages on Josephine loam (Typic Haploxerults), an important soil for revegetation to rangeland production. Forage yields increased and the botanical composition shifted toward subclover (Trifolium subterraneum L.) with added S. Sulfate uptake and plant S concentration were increased as a function of the S applied. Forage yields were related to plant S concentration and N:S ratios. Subclover and filaree (Erodium botrys (Cav.) Bertol.) obtained a larger proportion of their S from fertilizer-S than did soft chess (Bromus mollis L.). Total S uptake was largely from the applied fertilizer, 39% at the intermediate S level and 78% at high S. The replacement series design of the experiment and the use of labeled S and N permitted calculation of competition coefficients for fertilizer S, total S, soil N, and total N uptake for the species in pairwise associations. Sulfur was the factor limiting to subclover where S was not applied, and N was the limiting factor to soft chess and filaree in mixtures with subclover at high S where subclover was able to fix most of its own N. In the mixture of soft chess and filaree competition for S and N was about equally intense for both nutrients at all S levels, and filaree was dominant consistently. Comparison of two methods for estimating the fate of fertilizer S showed that differences in sulfur uptake and leachate losses over the controls provided significantly higher values for sulfur recovery than estimates based on the fate of³⁵S. Contribution of the University of California Hopland Field Station and Dept. of Agronomy and Range Science, Univ. of California, Davis, CA 95616.     

Climatic change and body size in two species of Japanese rodents

Using museum specimens, we studied temporal changes in skull size in two species of Japanese rodents, the large Japanese field mouse ( Apodemus speciosus ) and Pratt's vole ( Eothenomys smithii  =  E. kageus ) during the 20th century. We used the greatest length of the skull (GTL), zygomatic breadth (ZB), narrowest width of the skull across the interorbital region (IC) and the length of the upper cheek teeth row (M) as indicators to such changes. We found that GTL and ZB (but not IC and M) increased during the study period in mice, and that IC and M (but not GTL and ZB) increased marginally in voles. We attribute these changes to elevated ambient minimal temperatures, which increased food availability and energy savings for the mice, and required diet change in the voles.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 82 , 263–267.     

Climatic change is advancing the phenology of moth species in Ireland

Recent increases in global temperatures have contributed to advancing phenology of plants and animals. These increases in temperature have been shown to affect the phenological phases (phenophases) of plants and birds in Ireland, but less is known about the effect on the phenophases of Irish insects. Records of the flight periods of 59 species of Irish moths over the past 35 years (1974–2009) were obtained from a public monitoring group. Observations were analysed across the country using generalized additive models (GAMs) weighted by total yearly population numbers for each species. The results of the statistical analyses showed that 45 of the 59 species studied have a significantly earlier first sighting date now than when observations began. With this earlier emergence, 44 of the 59 species also have a significantly longer flight season over the same 35‐year period. The extent of these changes varies across the country and by species life history. In particular, species emerging in spring are advancing at a much faster rate than species emerging during the summer. Many of these changes in first sighting are negatively correlated with rising temperatures in Ireland, particularly in late spring and early summer (May and June). The variation in phenological advancement in the moth species of Ireland is extremely complex and may be influenced more by species life history than by the phenology of interacting species, such as host plants.     

Climatic change as an engine for speciation in flightless Orthoptera species inhabiting African mountains

Many East African mountains are characterized by an exceptionally high biodiversity. Here we assess the hypothesis that climatic fluctuations during the Plio-Pleistocene led to ecological fragmentation with subsequent genetic isolation and speciation in forest habitats in East Africa. Hypotheses on speciation in savannah lineages are also investigated. To do this, mitochondrial DNA sequences from a group of bush crickets consisting of both forest and savannah inhabiting taxa were analysed in relation to Plio-Pleistocene range fragmentations indicated by palaeoclimatic studies. Coalescent modelling and mismatch distributions were used to distinguish between alternative biogeographical scenarios. The results indicate two radiations: the earliest one overlaps in time with the global spread of C4 grasslands and only grassland inhabiting lineages originated in this radiation. Climatically induced retraction of forest to higher altitudes about 0.8 million years ago, promoting vicariant speciation in species inhabiting the montane zone, can explain the second radiation. Although much of the biodiversity in East Africa is presently threatened by climate change, past climatic fluctuations appear to have contributed to the species richness observed in the East African hot spots. Perceiving forests as centres of speciation reinforces the importance of conserving the remaining forest patches in the region.     

The importance of biotic interactions for modelling species distributions under climate change

Aim There is a debate as to whether biotic interactions exert a dominant role in governing species distributions at macroecological scales. The prevailing idea is that climate is the key limiting factor; thus models that use present‐day climate–species range relationships are expected to provide reasonable means to quantify the impacts of climate change on species distributions. However, there is little empirical evidence that biotic interactions would not constrain species distributions at macroecological scales. We examine this idea, for the first time, and provide tests for two null hypotheses: (H₀ 1) – biotic interactions do not exert a significant role in explaining current distributions of a particular species of butterfly (clouded Apollo, Parnassius mnemosyne) in Europe; and (H₀ 2) – biotic interactions do not exert a significant role in predictions of altered species’ ranges under climate change. Location Europe. Methods Generalized additive modelling (GAM) was used to investigate relationships between species and climate; species and host plants; and species and climate + host plants. Because models are sensitive to the variable selection strategies utilised, four alternative approaches were used: AIC (Akaike's Information Criterion), BIC (Bayesian Information Criterion), BRUTO (Adaptive Backfitting), and CROSS (Cross Selection). Results In spite of the variation in the variables selected with different methods, both hypotheses (H₀ 1 and H₀ 2) were falsified, providing support for the proposition that biotic interactions significantly affect both the explanatory and predictive power of bioclimatic envelope models at macro scales. Main conclusions Our results contradict the widely held view that the effects of biotic interactions on individual species distributions are not discernible at macroecological scales. Results are contingent on the species, type of interaction and methods considered, but they call for more stringent evidence in support of the idea that purely climate‐based modelling would be sufficient to quantify the impacts of climate change on species distributions.     

Sensitivity of predictive species distribution models to change in grain size

Predictive species distribution modelling (SDM) has become an essential tool in biodiversity conservation and management. The choice of grain size (resolution) of environmental layers used in modelling is one important factor that may affect predictions. We applied 10 distinct modelling techniques to presence-only data for 50 species in five different regions, to test whether: (1) a 10-fold coarsening of resolution affects predictive performance of SDMs, and (2) any observed effects are dependent on the type of region, modelling technique, or species considered. Results show that a 10 times change in grain size does not severely affect predictions from species distribution models. The overall trend is towards degradation of model performance, but improvement can also be observed. Changing grain size does not equally affect models across regions, techniques, and species types. The strongest effect is on regions and species types, with tree species in the data sets (regions) with highest locational accuracy being most affected. Changing grain size had little influence on the ranking of techniques: boosted regression trees remain best at both resolutions. The number of occurrences used for model training had an important effect, with larger sample sizes resulting in better models, which tended to be more sensitive to grain. Effect of grain change was only noticeable for models reaching sufficient performance and/or with initial data that have an intrinsic error smaller than the coarser grain size.     

Wing trait–inversion associations in Drosophila subobscura can be generalized within continents,but may change through time

Clinal variation is one of the most emblematic examples of the action of natural selection at a wide geographical range. In Drosophila subobscura, parallel clines in body size and inversions, but not in wing shape, were found in Europe and South and North America. Previous work has shown that a bottleneck effect might be largely responsible for differences in wing trait–inversion association between one European and one South American population. One question still unaddressed is whether the associations found before are present across other populations of the European and South American clines. Another open question is whether evolutionary dynamics in a new environment can lead to relevant changes in wing traits–inversion association. To analyse geographical variation in these associations, we characterized three recently laboratory founded D. subobscura populations from both the European and South American latitudinal clines. To address temporal variation, we also characterized the association at a later generation in the European populations. We found that wing size and shape associations can be generalized across populations of the same continent, but may change through time for wing size. The observed temporal changes are probably due to changes in the genetic content of inversions, derived from adaptation to the new, laboratory environment. Finally, we show that it is not possible to predict clinal variation from intrapopulation associations. All in all this suggests that, at least in the present, wing traits–inversion associations are not responsible for the maintenance of the latitudinal clines in wing shape and size.     

Probabilistic accounting of uncertainty in forecasts of species distributions under climate change

Forecasts of species distributions under future climates are inherently uncertain, but there have been few attempts to describe this uncertainty comprehensively in a probabilistic manner. We developed a Monte Carlo approach that accounts for uncertainty within generalized linear regression models (parameter uncertainty and residual error), uncertainty among competing models (model uncertainty), and uncertainty in future climate conditions (climate uncertainty) to produce site‐specific frequency distributions of occurrence probabilities across a species' range. We illustrated the method by forecasting suitable habitat for bull trout (Salvelinus confluentus) in the Interior Columbia River Basin, USA, under recent and projected 2040s and 2080s climate conditions. The 95% interval of total suitable habitat under recent conditions was estimated at 30.1–42.5 thousand km; this was predicted to decline to 0.5–7.9 thousand km by the 2080s. Projections for the 2080s showed that the great majority of stream segments would be unsuitable with high certainty, regardless of the climate data set or bull trout model employed. The largest contributor to uncertainty in total suitable habitat was climate uncertainty, followed by parameter uncertainty and model uncertainty. Our approach makes it possible to calculate a full distribution of possible outcomes for a species, and permits ready graphical display of uncertainty for individual locations and of total habitat.     

Climatic and biophysical controls on conifer species distributions in mountain forests of Washington State, USA

Abstract Aim The purpose of this study was to quantify relationships between conifer species distributions and climatic and biophysical variables, in order to provide better insight into the potential for redistribution of species on the landscape in response to climatic change. Location Data are from 10,653 georeferenced sites in Washington State, USA, along a longitudinal gradient from west of the crest of the Cascade Range to the beginnings of the western slope of the Rocky Mountains, and across two physiographic provinces, the Northern Cascades, characterized by steep, rugged topography, and the Okanogan Highlands, presenting moderate slopes and broad rounded summits. Methods Tree data were drawn from the USDA Forest Service Area Ecology Program database, collected in mature, undisturbed stands. We compared simple climatic variables (annual temperature, growing‐degree days, annual and seasonal precipitation) to biophysical variables (soil, hydrologic, and solar radiation) derived from climatic variables. Climatic and biophysical variables were taken from the output of climatological and hydrological simulation models and estimated for each plot in the tree database. Generalized linear models were used, for each of fourteen tree species, at multiple spatial extents, to estimate the probability of occurrence of that species as a function of climatic and biophysical predictors. Models were validated by a combination of bootstrapping and estimating receiver operating characteristic (ROC) curves. Results For the majority of species, we were able to fit variables representing both moisture and temperature gradients, and in all but a few cases these models identified a unimodal response of species occurrence to these gradients. In some cases the ecological/environmental niche of a species had been clearly captured by the model, whereas in others a longer gradient in the predictor variable(s) would be needed. Responses of most species were consistent across three spatial scales. Main conclusions By identifying the ecological niches of multiple species, we can forecast their redistribution on the landscape in response to climatic change, evaluate the predictions of simulation models, and alert managers to particularly sensitive or vulnerable ecosystems and landscapes.     

Small-scale spatial distributions of long-finned pilot whales change over time,but foraging hot spots are consistent: Significance for marine wildlife tourism management

Data collected opportunistically aboard marine wildlife tourism vessels are an inexpensive source of spatial information on the target species. Although these data are often challenging to analyze, they can be used to monitor spatiotemporal changes in species distribution and behavior. Disruptions from whale-watching vessels to behaviors such as foraging can be particularly harmful to cetaceans, but impacts could be reduced if areas essential for these sensitive behaviors are identified. We used data collected onboard whale-watching vessels to explore space-use patterns in long-finned pilot whales (Globicephala melas) off northern Cape Breton Island, Canada, an area where tourism is essential. Encounters with pilot whales between 2011–2016 occurred twice as far offshore than during 2003–2006 and 2008, and foraging activity decreased. Despite the changes in distribution and activity budgets, we identified two hot spots of foraging activity that persisted through time. These identified foraging hot spots comprised only a small proportion (20 km² ) of the range used by whale-watching vessels. Adaptive local management (e.g., voluntary codes of conduct) focused on limiting interactions in these energetically important areas may help reduce any potential impacts from whale-watching and promote the continued viability of the whale population and the tourism industry that relies on it.     

Changes in habitat associations during range expansion: disentangling the effects of climate and residence time

The distributions of many species are not at equilibrium with their environment. This includes spreading non-native species and species undergoing range shifts in response to climate change. The habitat associations of these species may change during range expansion as less favourable climatic conditions at expanding range margins constrain species to use only the most favourable habitats, violating the species distribution model assumption of stationarity. Alternatively, changes in habitat associations could result from density-dependent habitat selection; at range margins, population densities are initially low so species can exhibit density-independent selection of the most favourable habitats, while in the range core, where population densities are higher, species spread into less favourable habitat. We investigate if the habitat preferences of the non-native common waxbill Estrilda astrild changed as they spread in three directions (north, east and south-east) in the Iberian Peninsula. There are different degrees of climatic suitability and colonization speed across range expansion axes, allowing us to separate the effects of climate from residence time. In contrast to previous studies we find a stronger effect of residence time than climate in influencing the prevalence of common waxbills. As well as a strong additive effect of residence time, there were some changes in habitat associations, which were consistent with density-dependent habitat selection. The combination of broader habitat associations and higher prevalence in areas that have been colonised for longer means that species distribution models constructed early in the invasion process are likely to underestimate species’ potential distribution.     

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.     

Cross‐biome transplants of plant litter show decomposition models extend to a broader climatic range but lose predictability at the decadal time scale

We analyzed results from 10‐year long field incubations of foliar and fine root litter from the Long‐term Intersite Decomposition Experiment Team (LIDET) study. We tested whether a variety of climate and litter quality variables could be used to develop regression models of decomposition parameters across wide ranges in litter quality and climate and whether these models changed over short to long time periods. Six genera of foliar and three genera of root litters were studied with a 10‐fold range in the ratio of acid unhydrolyzable fraction (AUF, or ‘lignin’) to N. Litter was incubated at 27 field sites across numerous terrestrial biomes including arctic and alpine tundra, temperate and tropical forests, grasslands and warm deserts. We used three separate mathematical models of first‐order (exponential) decomposition, emphasizing either the first year or the entire decade. One model included the proportion of relatively stable material as an asymptote. For short‐term (first‐year) decomposition, nonlinear regressions of exponential or power function form were obtained with r² values of 0.82 and 0.64 for foliar and fine‐root litter, respectively, across all biomes included. AUF and AUF : N ratio were the most explanative litter quality variables, while the combined temperature‐moisture terms AET (actual evapotranspiration) and CDI (climatic decomposition index) were best for climatic effects. Regressions contained some systematic bias for grasslands and arctic and boreal sites, but not for humid tropical forests or temperate deciduous and coniferous forests. The ability of the regression approach to fit climate‐driven decomposition models of the 10‐year field results was dramatically reduced from the ability to capture drivers of short‐term decomposition. Future work will require conceptual and methodological improvements to investigate processes controlling decadal‐scale litter decomposition, including the formation of a relatively stable fraction and its subsequent decomposition.     

Evidence for evolutionary change associated with the recent range expansion of the British butterfly, Aricia agestis, in response to climate change

Poleward range expansions are widespread responses to recent climate change and are crucial for the future persistence of many species. However, evolutionary change in traits such as colonization history and habitat preference may also be necessary to track environmental change across a fragmented landscape. Understanding the likelihood and speed of such adaptive change is important in determining the rate of species extinction with ongoing climate change. We conducted an amplified fragment length polymorphism (AFLP)‐based genome scan across the recently expanded UK range of the Brown Argus butterfly, Aricia agestis, and used outlier‐based (DFDIST and BayeScan) and association‐based (Isolation‐By‐Adaptation) statistical approaches to identify signatures of evolutionary change associated with range expansion and habitat use. We present evidence for (i) limited effects of range expansion on population genetic structure and (ii) strong signatures of selection at approximately 5% AFLP loci associated with both the poleward range expansion of A. agestis and differences in habitat use across long‐established and recently colonized sites. Patterns of allele frequency variation at these candidate loci suggest that adaptation to new habitats at the range margin has involved selection on genetic variation in habitat use found across the long‐established part of the range. Our results suggest that evolutionary change is likely to affect species’ responses to climate change and that genetic variation in ecological traits across species’ distributions should be maximized to facilitate range shifts across a fragmented landscape, particularly in species that show strong associations with particular habitats.     

Risk map for the range expansion of Thrips palmi in Korea under climate change: Combining species distribution models with land-use change

Climate change and land-use change are the most powerful drivers for the invasion of alien species. To understand the integrated effects of these two drivers on pest invasion risk in the future, this study assessed how they impact the invasion risk of Thrips palmi Karny, which is the most serious invasive species in the Korean peninsula. The potential distribution of T. palmi was projected with a MaxEnt model for current and future climate change scenarios (RCP 4.5 and 8.5) based on occurrence records. The potential distribution extends to the north over time, except the eastern high mountainous area, for both RCPs in 2075. The MaxEnt outputs were filtered with agricultural area using data from three land-use change scenarios derived from the Shared Socio-economic Pathways (SSPs), because T. palmi populations can only be sustained in agricultural areas. The potential risk of T. palmi, based on the potential distribution probability in the future agricultural area, increased over time under all RCPs-SSPs combinations. The total area of T. palmi occurrence increased under RCPs-SSP1 and -SSP2 but decreased under RCPs-SSP3, due to agricultural areas being converted to urban areas. In conclusion, based on future climate change scenarios, T. palmi could be distributed throughout the Korean peninsula in the future. The invasion risk in agricultural areas will increase substantially; thus, intensive control measures for T. palmi are required in the future. Our research suggests that using both climate change and land-use change in pest risk mapping study can provide informative data for management strategy.     

Potential changes in the distributions of latitudinally restricted Australian butterfly species in response to climate change

This study assessed potential changes in the distributions of Australian butterfly species in response to global warming. The bioclimatic program, BIOCLIM, was used to determine the current climatic ranges of 77 butterfly species restricted to Australia. We found that the majority of these species had fairly wide climatic ranges in comparison to other taxa, with only 8% of butterfly species having a mean annual temperature range spanning less than 3 °C. The potential changes in the distributions of 24 butterfly species under four climate change scenarios for 2050 were also modelled using BIOCLIM. Results suggested that even species with currently wide climatic ranges may still be vulnerable to climate change; under a very conservative climate change scenario (with a temperature increase of 0.8–1.4 °C by 2050) 88% of species distributions decreased, and 54% of species distributions decreased by at least 20%. Under an extreme scenario (temperature increase of 2.1–3.9 °C by 2050) 92% of species distributions decreased, and 83% of species distributions decreased by at least 50%. Furthermore, the proportion of the current range that was contained within the predicted range decreased from an average of 63% under a very conservative scenario to less than 22% under the most extreme scenario. By assessing the climatic ranges that species are currently exposed to, the extent of potential changes in distributions in response to climate change and details of their life histories, we identified species whose characteristics may make them particularly vulnerable to climate change in the future.     

长白山云冷杉针阔混交林主要树种空间分布及其关联性

应用点格局分析方法 O-ring函数,分析了长白山云冷杉针阔混交林主要树种以及各生长阶段的空间分布格局和空间关联性。结果表明:(1)林分整体、主要建群种冷杉、云杉和红松的径级分布呈倒"J"型分布,属于增长型种群。(2)林分整体、冷杉、色木槭在0—10 m小尺度上均呈聚集分布,其他尺度上以随机分布为主,云杉、红松和椴树在整个研究尺度上以随机分布为主。(3)冷杉和色木槭在较低龄级阶段小尺度上呈明显的聚集分布,较大尺度上呈随机分布或者均匀分布,冷杉聚集程度随着龄级增大而降低。(4)在冷杉、云杉、红松、椴树和色木槭5个树种组成的10个种对中,冷杉与红松、云杉与红松在小尺度0—10 m范围上以正相关为主;色木槭与冷杉、红松、椴树在小尺度0—5 m范围上主要以正相关为主;云杉和冷杉、椴树、色木槭在整个研究尺度上均以不相关为主。椴树和冷杉、色木槭在10—25 m尺度范围上以负相关为主。(5)幼龄层冷杉与乔木层冷杉在小尺度上呈显著负相关,与乔木层其他树种主要表现为不相关。