Mapping environments at risk under different global climate change scenarios

All global circulation models based on Intergovernmental Panel on Climate Change (IPCC) scenarios project profound changes, but there is no consensus on how to map their environmental consequences. Our multivariate representation of environmental space combines stable topographic and edaphic attributes with dynamic climatic attributes. We divide that environmental space into 500 unique domains and map their current locations and their projected locations in 2100 under contrasting emissions scenarios. The environmental domains found across half the study area today disappear under the higher emissions scenario, but persist somewhere in it under the lower emissions scenario. Locations affected least and those affected most under each scenario are mapped. This provides an explicit framework for designing conservation networks to include both areas at least risk (potential refugia) and areas at greatest risk, where novel communities may form and where sentinel ecosystems can be monitored for signs of stress.     

Projected impacts of climate change on regional capacities for global plant species richness

Climate change represents a major challenge to the maintenance of global biodiversity. To date, the direction and magnitude of net changes in the global distribution of plant diversity remain elusive. We use the empirical multi-variate relationships between contemporary water-energy dynamics and other non-climatic predictor variables to model the regional capacity for plant species richness (CSR) and its projected future changes. We find that across all analysed Intergovernmental Panel on Climate Change emission scenarios, relative changes in CSR increase with increased projected temperature rise. Between now and 2100, global average CSR is projected to remain similar to today (+0.3%) under the optimistic B1/+1.8°C scenario, but to decrease significantly (−9.4%) under the ‘business as usual’ A1FI/+4.0°C scenario. Across all modelled scenarios, the magnitude and direction of CSR change are geographically highly non-uniform. While in most temperate and arctic regions, a CSR increase is expected, the projections indicate a strong decline in most tropical and subtropical regions. Countries least responsible for past and present greenhouse gas emissions are likely to incur disproportionately large future losses in CSR, whereas industrialized countries have projected moderate increases. Independent of direction, we infer that all changes in regional CSR will probably induce on-site species turnover and thereby be a threat to native floras.     

Risk of spring frost to apple production under future climate scenarios: the role of phenological acclimation

In the context of global warming, the general trend towards earlier flowering dates of many temperate tree species is likely to result in an increased risk of damage from exposure to frost. To test this hypothesis, a phenological model of apple flowering was applied to a temperature series from two locations in an important area for apple production in Europe (Trentino, Italy). Two simulated 50-year climatic projections (A2 and B2 of the Intergovernmental Panel on Climate Change - Special Report on Emission Scenarios) from the HadCM3 general circulation model were statistically downscaled to the two sites. Hourly temperature records over a 40-year period were used as the reference for past climate. In the phenological model, the heat requirement (degree hours) for flowering was parameterized using two approaches; static (constant over time) and dynamic (climate dependent). Parameterisation took into account the trees’ adaptation to changing temperatures based on either past instrumental records or the downscaled outputs from the climatic simulations. Flowering dates for the past 40 years and simulated flowering dates for the next 50 years were used in the model. A significant trend towards earlier flowering was clearly detected in the past. This negative trend was also apparent in the simulated data. However, the significance was less apparent when the “dynamic” setting for the degree hours requirement was used in the model. The number of frost episodes and flowering dates, on an annual basis, were graphed to assess the risk of spring frost. Risk analysis confirmed a lower risk of exposure to frost at present than in the past, and probably either constant or a slightly lower risk in future, especially given that physiological processes are expected to acclimate to higher temperatures. An erratum to this article can be found at     

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.     

Allee effects under climate change

Understanding how climate change affects population dynamics is crucial for assessing future of biodiversity. Here I ask how can Allee effects, occurring when mean individual fitness is reduced in rare populations, respond to increasing temperature. Despite the role Allee effects play in ecology of invasive, threatened and harvested populations, impacts of climate change on Allee effects are practically unknown. Analysis of two population models reveals that whereas the Allee effect driven by predation generally weakens as temperature increases, the Allee effect due to need of finding mates is predicted to become stronger when warming occurs. For the former model, the metabolic theory suggests that with increasing temperature prey growth rate should increase faster than predator attack rate. Increasing temperature thus weakens the Allee effect. In the latter, gypsy moth population model, mating rate increases with warming due to enhanced female?male encounter rate and temperature‐induced modifications in female and male adult emergence distributions. However, male and female mortality rates increase, too and the net effect is strengthening of the Allee effect. These results have repercussions also for pest control, indicating that augmentation of biocontrol agents may perhaps be not as effective as using pesticides or disrupting mating.     

The dynamics of food chains under climate change and nutrient enrichment

Warming has profound effects on biological rates such as metabolism, growth, feeding and death of organisms, eventually affecting their ability to survive. Using a nonlinear bioenergetic population-dynamic model that accounts for temperature and body-mass dependencies of biological rates, we analysed the individual and interactive effects of increasing temperature and nutrient enrichment on the dynamics of a three-species food chain. At low temperatures, warming counteracts the destabilizing effects of enrichment by both bottom-up (via the carrying capacity) and top-down (via biological rates) mechanisms. Together with increasing consumer body masses, warming increases the system tolerance to fertilization. Simultaneously, warming increases the risk of starvation for large species in low-fertility systems. This effect can be counteracted by increased fertilization. In combination, therefore, two main drivers of global change and biodiversity loss can have positive and negative effects on food chain stability. Our model incorporates the most recent empirical data and may thus be used as the basis for more complex forecasting models incorporating food-web structure.     

Distributional range shifts of Western Atlantic benthic Sargassum species (Fucales,Phaeophyceae) under future climate change scenarios

Climate change is altering the world’s marine biota, in particular, their geographic distribution. Sargassum species are foundation species that play critical ecological roles in tropical benthic communities, providing food, habitat heterogeneity and shelter for a wide range of marine organisms. To understand how future changes in abiotic variables could affect the distribution of Sargassum species along the Western Atlantic Ocean, we performed Ecological Niche Models (ENM) for 12 benthic Sargassum species. We projected present and future habitat suitability distributions under the RCP 4.5 and RCP 8.5 IPCC scenarios. We fit ENM and created ensembles from different algorithms. Our results predict changes in species latitudinal range (niche suitability) in the order of 0.5˚ to 8.1˚ northward, and 0˚ to 5.5˚ southward. Six species are likely to reduce their suitability area from 10% to 80%, while other six species are likely to expand their suitability area from 4% to 168%. Overall, changes in suitability area and latitudinal ranges will increase at larger latitudes for most species while suitability areas will decrease at lower latitudes for half of the species. This pattern is consistent with the expected tropicalization of temperate latitudes following global warming. Such changes can produce considerable losses in ecosystem services maintained by healthy Sargassum beds, particularly at lower latitudes. Our findings highlight the need to improve Sargassum conservation policies and management strategies to avoid the negative effects caused by losses in Sargassum forests.     

Response of the terrestrial biosphere to global climate change and human perturbation

Despite 20 years of intensive effort to understand the global carbon cycle, the budget for carbon dioxide in the atmosphere is unbalanced. To explain why atmospheric CO₂ is not increasing as rapidly as it should be, various workers have suggested that land vegetation acts as a sink for carbon dioxide. Here, I examine various possibilities and find that the evidence for a sink of sufficient magnitude on land is poor. Moreover, it is unlikely that the land vegetation will act as a sink in the postulated warmer global climates of the future. In response to rapid human population growth, destruction of natural ecosystems in the tropics remains a large net source of CO₂ for the atmosphere, which is only partially compensated by the potential for carbon storage in temperate and boreal regions. Direct and inadvertent human effects on land vegetation might increase the magnitude of regional CO₂ storage on land, but they are unlikely to play a significant role in moderating the potential rate of greenhouse warming in the future.     

Herbivory in global climate change research: direct effects of rising temperature on insect herbivores

This review examines the direct effects of climate change on insect herbivores. Temperature is identified as the dominant abiotic factor directly affecting herbivorous insects. There is little evidence of any direct effects of CO₂ or UVB. Direct impacts of precipitation have been largely neglected in current research on climate change. Temperature directly affects development, survival, range and abundance. Species with a large geographical range will tend to be less affected. The main effect of temperature in temperate regions is to influence winter survival; at more northerly latitudes, higher temperatures extend the summer season, increasing the available thermal budget for growth and reproduction. Photoperiod is the dominant cue for the seasonal synchrony of temperate insects, but their thermal requirements may differ at different times of year. Interactions between photoperiod and temperature determine phenology; the two factors do not necessarily operate in tandem. Insect herbivores show a number of distinct life‐history strategies to exploit plants with different growth forms and strategies, which will be differentially affected by climate warming. There are still many challenges facing biologists in predicting and monitoring the impacts of climate change. Future research needs to consider insect herbivore phenotypic and genotypic flexibility, their responses to global change parameters operating in concert, and awareness that some patterns may only become apparent in the longer term.     

Phenological and elevational shifts of plants,animals and fungi under climate change in the European Alps

Mountain areas are biodiversity hotspots and provide a multitude of ecosystem services of irreplaceable socio-economic value. In the European Alps, air temperature has increased at a rate of about 0.36°C decade⁻¹ since 1970, leading to glacier retreat and significant snowpack reduction. Due to these rapid environmental changes, this mountainous region is undergoing marked changes in spring phenology and elevational distribution of animals, plants and fungi. Long-term monitoring in the European Alps offers an excellent natural laboratory to synthetize climate-related changes in spring phenology and elevational distribution for a large array of taxonomic groups. This review assesses the climatic changes that have occurred across the European Alps during recent decades, spring phenological changes and upslope shifts of plants, animals and fungi from evidence in published papers and previously unpublished data. Our review provides evidence that spring phenology has been shifting earlier during the past four decades and distribution ranges show an upwards trend for most of the taxonomic groups for which there are sufficient data. The first observed activity of reptiles and terrestrial insects (e.g. butterflies) in spring has shifted significantly earlier, at an average rate of −5.7 and −6.0 days decade⁻¹, respectively. By contrast, the first observed spring activity of semi-aquatic insects (e.g. dragonflies and damselflies) and amphibians, as well as the singing activity or laying dates of resident birds, show smaller non-significant trends ranging from −1.0 to +1.3 days decade⁻¹. Leaf-out and flowering of woody and herbaceous plants showed intermediate trends with mean values of −2.4 and −2.8 days decade⁻¹, respectively. Regarding species distribution, plants, animals and fungi (N = 2133 species) shifted the elevation of maximum abundance (optimum elevation) upslope at a similar pace (on average between +18 and +25 m decade⁻¹) but with substantial differences among taxa. For example, the optimum elevation shifted upward by +36.2 m decade⁻¹ for terrestrial insects and +32.7 m decade⁻¹ for woody plants, whereas it was estimated to range between −1.0 and +11 m decade⁻¹ for semi-aquatic insects, ferns, birds and wood-decaying fungi. The upper range limit (leading edge) of most species also shifted upslope with a rate clearly higher for animals (from +47 to +91 m decade⁻¹) than for plants (from +17 to +40 m decade⁻¹), except for semi-aquatic insects (−4.7 m decade⁻¹). Although regional land-use changes could partly explain some trends, the consistent upward shift found in almost all taxa all over the Alps is likely reflecting the strong warming and the receding of snow cover that has taken place across the European Alps over recent decades. However, with the possible exception of terrestrial insects, the upward shift of organisms seems currently too slow to track the pace of isotherm shifts induced by climate warming, estimated at about +62 to +71 m decade⁻¹ since 1970. In the light of these results, species interactions are likely to change over multiple trophic levels through phenological and spatial mismatches. This nascent research field deserves greater attention to allow us to anticipate structural and functional changes better at the ecosystem level.     

Consequences of the genetic threshold model for observing partial migration under climate change scenarios

Migration is a widespread phenomenon across the animal kingdom as a response to seasonality in environmental conditions. Partially migratory populations are populations that consist of both migratory and residential individuals. Such populations are very common, yet their stability has long been debated. The inheritance of migratory activity is currently best described by the threshold model of quantitative genetics. The inclusion of such a genetic threshold model for migratory behavior leads to a stable zone in time and space of partially migratory populations under a wide range of demographic parameter values, when assuming stable environmental conditions and unlimited genetic diversity. Migratory species are expected to be particularly sensitive to global warming, as arrival at the breeding grounds might be increasingly mistimed as a result of the uncoupling of long‐used cues and actual environmental conditions, with decreasing reproduction as a consequence. Here, we investigate the consequences for migratory behavior and the stability of partially migratory populations under five climate change scenarios and the assumption of a genetic threshold value for migratory behavior in an individual‐based model. The results show a spatially and temporally stable zone of partially migratory populations after different lengths of time in all scenarios. In the scenarios in which the species expands its range from a particular set of starting populations, the genetic diversity and location at initialization determine the species’ colonization speed across the zone of partial migration and therefore across the entire landscape. Abruptly changing environmental conditions after model initialization never caused a qualitative change in phenotype distributions, or complete extinction. This suggests that climate change‐induced shifts in species’ ranges as well as changes in survival probabilities and reproductive success can be met with flexibility in migratory behavior at the species level, which will reduce the risk of extinction.     

Potential distribution patterns of scorpions in north‐eastern Brazil under scenarios of future climate change

Current predictions about the responses of species to climate change strongly rely on projecting altered environmental conditions on their distributions. In this study, we investigated the effects of future climate change scenarios on the potential distribution of 10 species of scorpions in north‐eastern Brazil in the context of their degree of specialisation to closed (Atlantic and Amazon Forests) and open (Caatinga and Cerrado) habitats. Scorpion species were classified as habitat specialists or generalists according to the IndVal index, and present and future species distribution models were prepared using minimum volume ellipsoids. According to IndVal, four species were classified as closed‐forest specialists (Ananteris mauryi, Tityus brazilae, Tityus pusillus and Tityus neglectus), four as open‐forest specialists (Jaguajir agamemnon, Jaguajir rochae, Physoctonus debilis and Bothriurus rochai), and two as generalists (Tityus stigmurus and Bothriurus asper). All species presented a drastic reduction in potential distribution, ranging from 44% to 72%, when compared with their current distribution. In addition, we found a reduction in scorpion species richness under future climate change scenarios. This finding has implications for scorpion conservation. Further, the results show that climate change may impact the composition of scorpion assemblages in north‐eastern Brazil, revealing important implications for human–scorpion interactions.     

Potential effects of future climate change on global reptile distributions and diversity

Aim

Until recently, complete information on global reptile distributions has not been widely available. Here, we provide the first comprehensive climate impact assessment for reptiles on a global scale.

Location

Global, excluding Antarctica.

Time period

1995, 2050 and 2080.

Major taxa studied

Reptiles.

Methods

We modelled the distribution of 6296 reptile species and assessed potential global and realm-specific changes in species richness, the change in global species richness across climate space, and species-specific changes in range extent, overlap and position under future climate change. To assess the future climatic impact on 3768 range-restricted species, which could not be modelled, we compared the future change in climatic conditions between both modelled and non-modelled species.

Results

Reptile richness was projected to decline significantly over time, globally but also for most zoogeographical realms, with the greatest decreases in Brazil, Australia and South Africa. Species richness was highest in warm and moist regions, with these regions being projected to shift further towards climate extremes in the future. Range extents were projected to decline considerably in the future, with a low overlap between current and future ranges. Shifts in range centroids differed among realms and taxa, with a dominant global poleward shift. Non-modelled species were significantly stronger affected by projected climatic changes than modelled species.

Main conclusions

With ongoing future climate change, reptile richness is likely to decrease significantly across most parts of the world. This effect, in addition to considerable impacts on species range extent, overlap and position, was visible across lizards, snakes and turtles alike. Together with other anthropogenic impacts, such as habitat loss and harvesting of species, this is a cause for concern. Given the historical lack of global reptile distributions, this calls for a re-assessment of global reptile conservation efforts, with a specific focus on anticipated future climate change.     

Predicting the influence of wolf-provided carrion on scavenger community dynamics under climate change scenarios

Climate change poses an immediate threat to the persistence and distribution of many species, yet our ability to forecast changes in species composition is hindered by poor understanding of the extent to which higher trophic‐level interactions may buffer or exacerbate the adverse effects of warming. We incorporated species‐specific consumption data from 240 wolf‐killed elk carcasses from Yellowstone National Park into stochastic simulation models to link trends in the El Niño Southern Oscillation (ENSO) to food procurement by a guild of scavengers as a function of gray wolf reintroduction. We find that a shift in ENSO towards the El Niño (warming) phase of the cycle coincident with increasing global temperatures reduces carrion for scavengers, particularly those with strong seasonal patterns in resource use such as grizzly bears. Wolves alleviate these warming‐induced food shortages by rendering control over this crucial resource to biotic rather than abiotic factors. Ecosystems with intact top predators are likely to exhibit stronger biotic regulation and should be more resistant to climate change than ecosystems lacking them.     

Bird migration times, climate change, and changing population sizes

Past studies of bird migration times have shown great variation in migratory responses to climate change. We used 33 years of bird capture data (1970–2002) from Manomet, Massachusetts to examine variation in spring migration times for 32 species of North American passerines. We found that changes in first arrival dates – the unit of observation used in most studies of bird migration times – often differ dramatically from changes in the mean arrival date of the migration cohort as a whole. In our study, the earliest recorded springtime arrival date for each species occurred 0.20 days later each decade. In contrast, the mean arrival dates for birds of each species occurred 0.78 days earlier each decade. The difference in the two trends was largely explained by declining migration cohort sizes, a factor not examined in many previous studies. We found that changes in migration cohort or population sizes may account for a substantial amount of the variation in previously documented changes in migration times. After controlling for changes in migration cohort size, we found that climate variables, migration distance, and date of migration explained portions of the variation in migratory changes over time. In particular, short-distance migrants appeared to respond to changes in temperature, while mid-distance migrants responded particularly strongly to changes in the Southern Oscillation Index. The migration times of long-distance migrants tended not to change over time. Our findings suggest that previously reported changes in migration times may need to be reinterpreted to incorporate changes in migration cohort sizes.     

Global assessment of coral bleaching and required rates of adaptation under climate change

Elevated ocean temperatures can cause coral bleaching, the loss of colour from reef‐building corals because of a breakdown of the symbiosis with the dinoflagellate Symbiodinium. Recent studies have warned that global climate change could increase the frequency of coral bleaching and threaten the long‐term viability of coral reefs. These assertions are based on projecting the coarse output from atmosphere–ocean general circulation models (GCMs) to the local conditions around representative coral reefs. Here, we conduct the first comprehensive global assessment of coral bleaching under climate change by adapting the NOAA Coral Reef Watch bleaching prediction method to the output of a low‐ and high‐climate sensitivity GCM. First, we develop and test algorithms for predicting mass coral bleaching with GCM‐resolution sea surface temperatures for thousands of coral reefs, using a global coral reef map and 1985–2002 bleaching prediction data. We then use the algorithms to determine the frequency of coral bleaching and required thermal adaptation by corals and their endosymbionts under two different emissions scenarios. The results indicate that bleaching could become an annual or biannual event for the vast majority of the world's coral reefs in the next 30–50 years without an increase in thermal tolerance of 0.2–1.0°C per decade. The geographic variability in required thermal adaptation found in each model and emissions scenario suggests that coral reefs in some regions, like Micronesia and western Polynesia, may be particularly vulnerable to climate change. Advances in modelling and monitoring will refine the forecast for individual reefs, but this assessment concludes that the global prognosis is unlikely to change without an accelerated effort to stabilize atmospheric greenhouse gas concentrations.     

Predicting the potential global distribution of Ageratina adenophora under current and future climate change scenarios

AimInvasive alien species (IAS) threaten ecosystems and humans worldwide, and future climate change may accelerate the expansion of IAS. Predicting the suitable areas of IAS can prevent their further expansion. Ageratina adenophora is an invasive weed over 30 countries in tropical and subtropical regions. However, the potential suitable areas of A. adenophora remain unclear along with its response to climate change. This study explored and mapped the current and future potential suitable areas of Ageratina adenophora.LocationGlobal.TaxaAsteraceae A. adenophora (Spreng.) R.M.King & H.Rob. Commonly known as Crofton weed.MethodsBased on A. adenophora occurrence data and climate data, we predicted its suitable areas of this weed under current and future (four RCPs in 2050 and 2070) by MaxEnt model. We used ArcGIS 10.4 to explore the potential suitable area distribution characteristics of this weed and the “ecospat” package in R to analyze its altitudinal distribution changes.ResultsThe area under the curve (AUC) value (>0.9) and true skill statistics (TSS) value (>0.8) indicated excelled model performance. Among environment factors, mean temperature of coldest quarter contributed most to the model. Globally, the suitable areas for A. adenophora invasion decreased under climate change scenarios, although regional increases were observed, including in six biodiversity hotspot regions. The potential suitable areas of A. adenophora under climate change would expand in regions with higher elevation (3,000–3,500 m).Main conclusionsMean temperature of coldest quarter was the most important variable influencing the potential suitable area of A. Adenophora. Under the background of a warming climate, the potential suitable area of A. adenophora will shrink globally but increase in six biodiversity hotspot regions. The potential suitable area of A. adenophora would expand at higher elevation (3,000–3,500 m) under climate change. Mountain ecosystems are of special concern as they are rich in biodiversity and sensitive to climate change, and increasing human activities provide more opportunities for IAS invasion.     

Global diversity patterns of larger benthic foraminifera under future climate change

Global warming threatens the viability of tropical coral reefs and associated marine calcifiers, including symbiont-bearing larger benthic foraminifera (LBF). The impacts of current climate change on LBF are debated because they were particularly diverse and abundant during past warm periods. Studies on the responses of selected LBF species to changing environmental conditions reveal varying results. Based on a comprehensive review of the scientific literature on LBF species occurrences, we applied species distribution modeling using Maxent to estimate present-day and future species richness patterns on a global scale for the time periods 2040–2050 and 2090–2100. For our future projections, we focus on Representative Concentration Pathway 6.0 from the Intergovernmental Panel on Climate Change, which projects mean surface temperature changes of +2.2°C by the year 2100. Our results suggest that species richness in the Central Indo-Pacific is two to three times higher than in the Bahamian ecoregion, which we have identified as the present-day center of LBF diversity in the Atlantic. Our future predictions project a dramatic temperature-driven decline in low-latitude species richness and an increasing widening bimodal latitudinal pattern of species diversity. While the central Indo-Pacific, now the stronghold of LBF diversity, is expected to be most pushed outside of the currently realized niches of most species, refugia may be largely preserved in the Atlantic. LBF species will face large-scale non-analogous climatic conditions compared to currently realized climate space in the near future, as reflected in the extensive areas of extrapolation, particularly in the Indo-Pacific. Our study supports hypotheses that species richness and biogeographic patterns of LBF will fundamentally change under future climate conditions, possibly initiating a faunal turnover by the late 21st century.     

A review of indicators of climate change for use in Ireland

Impact indicators are systems/organisms, the vitality of which alters in response to changes in environmental condition. The indicators assessed in this review fall within the impact category of the driver-pressure-state-impact-response (DPSIR) framework. Instrumental records have shown unequivocal changes in climatic conditions over the past 30 years at a global level but impact indicators allow these changes to be monitored at a finer resolution. Our main aim was to review sets of indicators of climate change currently used in various countries and to make recommendations for their use in the Irish environment. We review a preliminary set of climate change impact indicators in five sectors: agriculture; plant and animal distribution patterns; phenology; palaeoecology and human health. Currently, the most effective impact indicators of climate change have proved to be phenological observations of tree developmental stages. The strongest factor limiting the use of indicators is the lack of long-term data sets from which a climatic signal can be extracted.