Long-term unsustainable patterns of development rather than recent deforestation caused the emergence of Orthocoronavirinae species.

We investigated whether a set of phylogeographical tracked emergent events of Orthocoronavirinae were related to developed, urban and polluted environments worldwide. We explored coronavirus records in response to climate (rainfall parameters), population density, CO2 emission, Human Developmental Index (HDI) and deforestation. We contrasted environmental characteristics from regions with spillovers or encounters of wild Orthocoronavirinae against adjacent areas having best-preserved conditions. We used all complete sequenced CoVs genomes deposited in NCBI and GISAID databases until January 2021. Except for Deltacoronavirus, concentrated in Hong Kong and in birds, the other three genera were scattered all over the planet, beyond the original distribution of the subfamily, and found in humans, mammals, fishes and birds, wild or domestic. Spillovers and presence in wild animals were only reported in developed/densely populated places. We found significantly more occurrences reported in places with higher HDI, CO2 emission, or population density, along with more rainfall and more accentuated seasonality. Orthocoronavirinae occurred in areas with significantly higher human populations, CO2 emissions and deforestation rates than in adjacent locations. Intermediately disturbed ecosystems seemed more vulnerable for Orthocoronavirinae emergence than forested regions in frontiers of deforestation. Sadly, people experiencing poverty in an intensely consumerist society are the most vulnerable.

Large-scale Degradation of the Tocantins-Araguaia River Basin.

The Tocantins-Araguaia Basin is one of the largest river systems in South America, located entirely within Brazilian territory. In the last decades, capital-concentrating activities such as agribusiness, mining, and hydropower promoted extensive changes in land cover, hydrology, and environmental conditions. These changes are jeopardizing the basin's biodiversity and ecosystem services. Threats are escalating as poor environmental policies continue to be formulated, such as environmentally unsustainable hydropower plants, large-scale agriculture for commodity production, and aquaculture with non-native fish. If the current model persists, it will deepen the environmental crisis in the basin, compromising broad conservation goals and social development in the long term. Better policies will require thought and planning to minimize growing threats and ensure the basin's sustainability for future generations.

Searching for synthetic mechanisms on how biological traits mediate species responses to climate change / Em busca de mecanismos sintéticos sobre como os atributos biológicos mediam as respostas das espécies às mudanças climáticas

Abstract: Climate change will likely be the most significant challenge faced by species in this century, and species' ability to cope with climate change depends on their life history and ecological and evolutionary traits. Understanding how these traits mediate species' responses is beneficial for identifying more vulnerable species or prone to extinction risk. Here, we carried out a literature review describing how four traits commonly used in vulnerability assessments (i.e. clutch size, diet breadth, dispersal ability, and climatic tolerance) may determine species vulnerability. We also portray the possible mechanisms that explain how these traits govern species responses to climate change. The literature suggests different mechanisms operating for the evaluated traits. The mechanism of response to climate change differs between species inhabiting tropical and temperate regions: while species from the temperate areas may respond positively to temperature rise, tropical species may be severely affected. Since ectotherms depend on environment temperature, they are more sensitive and present different response mechanisms from endotherms.

Resumo: A mudança climática provavelmente será o maior desafio enfrentado pelas espécies neste século e a capacidade das espécies em lidar com a mudança climática depende de seus próprios atributos de história de vida, ecológicos e evolutivos. Entender como esses atributos mediam as respostas das espécies é extremamente útil para identificar espécies que são mais vulneráveis ou sujeitas ao risco de extinção. Aqui, realizamos uma revisão da literatura com foco na descrição de como quatro atributos comumente usados em avaliações de vulnerabilidade (tamanho da ninhada, amplitude da dieta, capacidade de dispersão e tolerância climática) podem realmente determinar a vulnerabilidade das espécies. Também retratamos os possíveis mecanismos que explicam como esses atributos governam as respostas das espécies à mudança climática. A literatura sugere diferentes mecanismos operando para os atributos avaliados. O mecanismo de resposta à mudança climática difere entre as espécies que habitam as regiões tropicais e temperadas: enquanto as espécies das regiões temperadas podem responder positivamente ao aumento da temperatura, as espécies tropicais podem ser severamente afetadas. Como os ectotérmicos dependem da temperatura ambiente, eles são mais sensíveis e apresentam mecanismos de resposta diferentes dos endotérmicos.

Author Correction: Strategic approaches to restoring ecosystems can triple conservation gains and halve costs.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

Priority areas for conservation of and research focused on terrestrial vertebrates.

Effective conservation policies require comprehensive knowledge of biodiversity. However, knowledge shortfalls still remain, hindering possibilities to improve decision making and built such policies. During the last 2 decades, conservationists have made great efforts to allocate resources as efficiently as possible but have rarely considered the idea that if research investments are also strategically allocated, it would likely fill knowledge gaps while simultaneously improving conservation actions. Therefore, prioritizing areas where both conservation and research actions could be conducted becomes a critical endeavor that can further maximize return on investment. We used Zonation, a conservation planning tool and geographical distributions of amphibians, birds, mammals, and reptiles to suggest and compare priority areas for conservation and research of terrestrial vertebrates worldwide. We also evaluated the degree of human disturbance in both types of priority areas by describing the value of the human footprint index within such areas. The spatial concordance between priority conservation and research areas was low: 0.36% of the world's land area. In these areas, we found it would be possible to protect almost half of the currently threatened species and to gather information on nearly 42% of data-deficient (DD) species. We also found that 6199 protected areas worldwide are located in such places, although only 35% of them have strict conservation purposes. Areas of consensus between conservation and research areas represent an opportunity for simultaneously conserving and acquiring knowledge of threatened and DD species of vertebrates. Although the picture is not the most encouraging, joint conservation and research efforts are possible and should be fostered to save vertebrate species from our own ignorance and extinction.

Áreas Prioritarias para la Conservación e Investigación Enfocadas en Vertebrados Terrestres Resumen Las políticas efectivas de conservación requieren del conocimiento integral de la biodiversidad. Sin embargo, la deficiencia de conocimiento todavía obstaculiza las posibilidades de mejorar la toma de decisiones y construir dichas políticas. Durante las últimas dos décadas, los conservacionistas han realizado un gran esfuerzo por asignar los recursos de la manera más eficiente posible, pero en pocas ocasiones han considerado la idea de que, si las inversiones para la investigación también se asignan estratégicamente, probablemente llenarían los vacíos de conocimiento a la vez que mejoran las acciones de conservación. Por lo tanto, la priorización de las áreas en donde podrían realizarse acciones de conservación y de investigación se convierte en un esfuerzo importante que puede avanzar todavía más la maximización del rendimiento de la inversión. Usamos Zonation, una herramienta de planeación de la conservación, junto con las distribuciones geográficas de anfibios, aves, mamíferos y reptiles para sugerir y comparar las áreas prioritarias para la conservación e investigación de los vertebrados terrestres en todo el mundo. También evaluamos el grado de perturbación humana en ambos tipos de áreas prioritarias mediante la descripción del valor del índice de la huella ecológica humana dentro de dichas áreas. La concordancia espacial entre las áreas prioritarias para la conservación y para la investigación fue baja: 0.36% del área total de suelo mundial. Encontramos que en estas áreas sería posible proteger a casi la mitad de las especies que se encuentran actualmente amenazadas, así como recopilar información sobre casi el 42% de las especies cuya información es deficiente. También descubrimos que 6199 áreas protegidas a nivel mundial están localizadas en dichos lugares, aunque sólo el 35% de ellas tiene propósitos estrictos de conservación. Las áreas de consenso entre la conservación y la investigación representan una oportunidad para simultáneamente conservar y adquirir conocimiento sobre las especies amenazadas de vertebrados y cuya información es deficiente. Aunque el futuro no es alentador, los esfuerzos conjuntos de conservación e investigación son posibles y deberían fomentarse para salvar a las especies de vertebrados de nuestra propia ignorancia y extinción.

Meta-analyzing the likely cross-species responses to climate change.

Ecological Niche Models (ENMs) have different performances in predicting potential geographic distributions. Here we meta-analyzed the likely effects of climate change on the potential geographic distribution of 1,205 bird species from the Neotropical region, modeled using eight ENMs and three Atmosphere-Ocean General Circulation Models (AOGCM). We considered the variability in ENMs performance to estimate a weighted mean difference between potential geographic distributions for baseline and future climates. On average, potential future ranges were projected to be from 25.7% to 44.5% smaller than current potential ranges across species. However, we found that 0.2% to 18.3% of the total variance in range shifts occurred "within species" (i.e., owing to the use of different modeling techniques and climate models) and 81.7% to 99.8% remained between species (i.e., it could be explained by ecological correlates). Using meta-analytical techniques akin to regression, we also showed that potential range shifts are barely predicted by bird biological traits. We demonstrated that one can combine and reduce species-specific effects with high uncertainty in ENMs and also explore potential causes of climate change effect on species using meta-analytical tools. We also highlight that the search for powerful correlates of climate change-induced range shifts can be a promising line of investigation.

Strategic approaches to restoring ecosystems can triple conservation gains and halve costs.

International commitments for ecosystem restoration add up to one-quarter of the world's arable land. Fulfilling them would ease global challenges such as climate change and biodiversity decline but could displace food production and impose financial costs on farmers. Here, we present a restoration prioritization approach capable of revealing these synergies and trade-offs, incorporating ecological and economic efficiencies of scale and modelling specific policy options. Using an actual large-scale restoration target of the Atlantic Forest hotspot, we show that our approach can deliver an eightfold increase in cost-effectiveness for biodiversity conservation compared with a baseline of non-systematic restoration. A compromise solution avoids 26% of the biome's current extinction debt of 2,864 plant and animal species (an increase of 257% compared with the baseline). Moreover, this solution sequesters 1 billion tonnes of CO2-equivalent (a 105% increase) while reducing costs by US$28 billion (a 57% decrease). Seizing similar opportunities elsewhere would offer substantial contributions to some of the greatest challenges for humankind.

Climate change likely to reduce orchid bee abundance even in climatic suitable sites.

Studies have tested whether model predictions based on species' occurrence can predict the spatial pattern of population abundance. The relationship between predicted environmental suitability and population abundance varies in shape, strength and predictive power. However, little attention has been paid to the congruence in predictions of different models fed with occurrence or abundance data, in particular when comparing metrics of climate change impact. Here, we used the ecological niche modeling fit with presence-absence and abundance data of orchid bees to predict the effect of climate change on species and assembly level distribution patterns. In addition, we assessed whether predictions of presence-absence models can be used as a proxy to abundance patterns. We obtained georeferenced abundance data of orchid bees (Hymenoptera: Apidae: Euglossina) in the Brazilian Atlantic Forest. Sampling method consisted in attracting male orchid bees to baits of at least five different aromatic compounds and collecting the individuals with entomological nets or bait traps. We limited abundance data to those obtained by similar standard sampling protocol to avoid bias in abundance estimation. We used boosted regression trees to model ecological niches and project them into six climate models and two Representative Concentration Pathways. We found that models based on species occurrences worked as a proxy for changes in population abundance when the output of the models were continuous; results were very different when outputs were discretized to binary predictions. We found an overall trend of diminishing abundance in the future, but a clear retention of climatically suitable sites too. Furthermore, geographic distance to gained climatic suitable areas can be very short, although it embraces great variation. Changes in species richness and turnover would be concentrated in western and southern Atlantic Forest. Our findings offer support to the ongoing debate of suitability-abundance models and can be used to support spatial conservation prioritization schemes and species triage in Atlantic Forest.

Model uncertainties do not affect observed patterns of species richness in the Amazon.

BACKGROUND: Climate change is arguably a major threat to biodiversity conservation and there are several methods to assess its impacts on species potential distribution. Yet the extent to which different approaches on species distribution modeling affect species richness patterns at biogeographical scale is however unaddressed in literature. In this paper, we verified if the expected responses to climate change in biogeographical scale-patterns of species richness and species vulnerability to climate change-are affected by the inputs used to model and project species distribution. METHODS: We modeled the distribution of 288 vertebrate species (amphibians, birds and mammals), all endemic to the Amazon basin, using different combinations of the following inputs known to affect the outcome of species distribution models (SDMs): 1) biological data type, 2) modeling methods, 3) greenhouse gas emission scenarios and 4) climate forecasts. We calculated uncertainty with a hierarchical ANOVA in which those different inputs were considered factors. RESULTS: The greatest source of variation was the modeling method. Model performance interacted with data type and modeling method. Absolute values of variation on suitable climate area were not equal among predictions, but some biological patterns were still consistent. All models predicted losses on the area that is climatically suitable for species, especially for amphibians and primates. All models also indicated a current East-western gradient on endemic species richness, from the Andes foot downstream the Amazon river. Again, all models predicted future movements of species upwards the Andes mountains and overall species richness losses. CONCLUSIONS: From a methodological perspective, our work highlights that SDMs are a useful tool for assessing impacts of climate change on biodiversity. Uncertainty exists but biological patterns are still evident at large spatial scales. As modeling methods are the greatest source of variation, choosing the appropriate statistics according to the study objective is also essential for estimating the impacts of climate change on species distribution. Yet from a conservation perspective, we show that Amazon endemic fauna is potentially vulnerable to climate change, due to expected reductions on suitable climate area. Climate-driven faunal movements are predicted towards the Andes mountains, which might work as climate refugia for migrating species.

The role of macroinvertebrates for conservation of freshwater systems.

Freshwater ecosystems are the most threatened ecosystems worldwide. Argentinian-protected areas have been established mainly to protect vertebrates and plants in terrestrial ecosystems. In order to create a comprehensive biodiverse conservation plan, it is crucial to integrate both aquatic and terrestrial systems and to include macroinvertebrates. Here, we address this topic by proposing priority areas of conservation including invertebrates, aquatic ecosystems, and their connectivity and land uses. LOCATION: Northwest of Argentina. We modeled the ecological niches of different taxa of macroinvertebrates such as Coleoptera, Ephemeroptera, Hemiptera, Megaloptera, Lepidoptera, Odonata, Plecoptera, Trichoptera, Acari, and Mollusca. Based on these models, we analyzed the contribution of currently established protected areas in the conservation of the aquatic biodiversity and we propose a spatial prioritization taking into account possible conflict regarding different land uses. Our analysis units were the real watersheds, to which were added longitudinal connectivity up and down the rivers. A total of 132 species were modeled in the priority area analyses. The analysis 1 showed that only an insignificant percentage of the macroinvertebrates distribution is within the protected areas in the North West of Argentina. The analyses 2 and 3 recovered similar values of protection for the macroinvertebrate species. The upper part of Bermejo, Salí-Dulce, San Francisco, and the Upper part of Juramento basins were identified as priority areas of conservation. The aquatic ecosystems need special protection and 10% or even as much as 17% of land conservation is insufficient for species of macroinvertebrates. In turn the protected areas need to combine the aquatic and terrestrial systems and need to include macroinvertebrates as a key group to sustain the biodiversity. In many cases, the land uses are in conflict with the conservation of biodiversity; however, it is possible to apply the connectivity of the watersheds and create multiple-use modules.

Global priorities for conservation across multiple dimensions of mammalian diversity.

Conservation priorities that are based on species distribution, endemism, and vulnerability may underrepresent biologically unique species as well as their functional roles and evolutionary histories. To ensure that priorities are biologically comprehensive, multiple dimensions of diversity must be considered. Further, understanding how the different dimensions relate to one another spatially is important for conservation prioritization, but the relationship remains poorly understood. Here, we use spatial conservation planning to (i) identify and compare priority regions for global mammal conservation across three key dimensions of biodiversity-taxonomic, phylogenetic, and traits-and (ii) determine the overlap of these regions with the locations of threatened species and existing protected areas. We show that priority areas for mammal conservation exhibit low overlap across the three dimensions, highlighting the need for an integrative approach for biodiversity conservation. Additionally, currently protected areas poorly represent the three dimensions of mammalian biodiversity. We identify areas of high conservation priority among and across the dimensions that should receive special attention for expanding the global protected area network. These high-priority areas, combined with areas of high priority for other taxonomic groups and with social, economic, and political considerations, provide a biological foundation for future conservation planning efforts.

Niche conservatism and the invasive potential of the wild boar.

Niche conservatism, i.e. the retention of a species' fundamental niche through evolutionary time, is cornerstone for biological invasion assessments. The fact that species tend to maintain their original climate niche allows predictive maps of invasion risk to anticipate potential invadable areas. Unravelling the mechanisms driving niche shifts can shed light on the management of invasive species. Here, we assessed niche shifts in one of the world's worst invasive species: the wild boar Sus scrofa. We also predicted potential invadable areas based on an ensemble of three ecological niche modelling methods, and evaluated the performance of models calibrated with native vs. pooled (native plus invaded) species records. By disentangling the drivers of change on the exotic wild boar population's niches, we found strong evidence for niche conservatism during biological invasion. Ecological niche models calibrated with both native and pooled range records predicted convergent areas. Also, observed niche shifts are mostly explained by niche unfilling, i.e. there are unoccupied areas in the exotic range where climate is analogous to the native range. Niche unfilling is expected as result of recent colonization and ongoing dispersal, and was potentially stronger for the Neotropics, where a recent wave of introductions for pig-farming and game-hunting has led to high wild boar population growth rates. The invasive potential of wild boar in the Neotropics is probably higher than in other regions, which has profound management implications if we are to prevent their invasion into species-rich areas, such as Amazonia, coupled with expansion of African swine fever and possibly great economic losses. Although the originally Eurasian-wide distribution suggests a pre-adaptation to a wide array of climates, the wild boar world-wide invasion does not exhibit evidence of niche evolution. The invasive potential of the wild boar therefore probably lies on the reproductive, dietary and morphological characteristics of this species, coupled with behavioural thermoregulation.

Assessing Mammal Exposure to Climate Change in the Brazilian Amazon.

Human-induced climate change is considered a conspicuous threat to biodiversity in the 21st century. Species' response to climate change depends on their exposition, sensitivity and ability to adapt to novel climates. Exposure to climate change is however uneven within species' range, so that some populations may be more at risk than others. Identifying the regions most exposed to climate change is therefore a first and pivotal step on determining species' vulnerability across their geographic ranges. Here, we aimed at quantifying mammal local exposure to climate change across species' ranges. We identified areas in the Brazilian Amazon where mammals will be critically exposed to non-analogue climates in the future with different variables predicted by 15 global circulation climate forecasts. We also built a null model to assess the effectiveness of the Amazon protected areas in buffering the effects of climate change on mammals, using an innovative and more realistic approach. We found that 85% of species are likely to be exposed to non-analogue climatic conditions in more than 80% of their ranges by 2070. That percentage is even higher for endemic mammals; almost all endemic species are predicted to be exposed in more than 80% of their range. Exposure patterns also varied with different climatic variables and seem to be geographically structured. Western and northern Amazon species are more likely to experience temperature anomalies while northeastern species will be more affected by rainfall abnormality. We also observed an increase in the number of critically-exposed species from 2050 to 2070. Overall, our results indicate that mammals might face high exposure to climate change and that protected areas will probably not be efficient enough to avert those impacts.

Systematic review on the conservation genetics of African savannah elephants.

BACKGROUND: In this paper we review the conservation genetics of African savannah elephants, aiming to understand the spatio-temporal research trends and their underlying factors. As such, we explore three questions associated to the conservation genetics and molecular ecology of these elephants: (1) what are the research trends concerning the conservation genetics of Loxodonta africana? (2) Do richer countries conduct more research on the genetics of African elephants? (3) Which attributes influence where scholars conduct their research? MATERIALS AND METHODS: We examined available peer-reviewed publications from 1993 to 2014 in complementary online databases, including the ISI/Web of Science (WoS), Scopus and Google Scholar (GS), and searched for publications in scientific journals as well as in the reference section of these publications. We analyzed the annual trend of publications in this field of research, including the number of authors, levels of collaboration among authors, year of publication, publishing journal and the countries from where genetic samples were collected. Additionally, we identified main research clusters, authors, and institutional collaborations, based on co-citation and co-occurrence networks. RESULTS: We found that during the study period there was a positive trend in the number of publications and a reduction in the number of authors per paper. Twenty-five countries contributed, with the majority of publications authored by researchers in the USA, Kenya and South Africa. The majority of samples were collected in Kenya, Tanzania and South Africa. Research outputs are associated with the existence of long-term conservation/research projects and research potential as measured by the literacy rate and the number of higher education institutions in a country. Five research clusters were identified, focusing on the origin and evolution of the species, methodological issues and the relatedness among elephant species. CONCLUSIONS: Research in this field should be expanded to additional countries harboring elephant populations to enable a more comprehensive understanding of the population structure and genetic differentiation of the species, and to cope with challenges associated with the conservation of the species such as illegal hunting, habitat fragmentation, species reintroduction and climate change.

On the worrying fate of Data Deficient amphibians.

The 'Data Deficient' (DD) category of the IUCN Red List assembles species that cannot be placed in another category due to insufficient information. This process generates uncertainty about whether these species are safe or actually in danger. Here, we give a global overview on the current situation of DD amphibian species (almost a quarter of living amphibians) considering land-use change through habitat modification, the degree of protection of each species and the socio-political context of each country harboring DD species. We found that DD amphibians have, on average, 81% of their ranges totally outside protected areas. Worryingly, more than half of DD species have less than 1% of their distribution represented in protected areas. Furthermore, the percentage of overlap between species' range and human-modified landscapes is high, at approximately 58%. Many countries harboring a large number of DD species show a worrying socio-political trend illustrated by substantial, recent incremental increases in the Human Development Index and lower incremental increases in the establishment of protected areas. Most of these are African countries, which are located mainly in the central and southern regions of the continent. Other countries with similar socio-political trends are in southeastern Asia, Central America, and in the northern region of South America. This situation is concerning, but it also creates a huge opportunity for considering DD amphibians in future conservation assessments, planning, and policy at different levels of government administration.

The reduced effectiveness of protected areas under climate change threatens Atlantic forest tiger moths.

Climate change leads to species' range shifts, which may end up reducing the effectiveness of protected areas. These deleterious changes in biodiversity may become amplified if they include functionally important species, such as herbivores or pollinators. We evaluated how effective protected areas in the Brazilian Atlantic Forest are in maintaining the diversity of tiger moths (Arctiinae) under climate change. Specifically, we assessed whether protected areas will gain or lose species under climate change and mapped their locations in the Atlantic Forest, in order to assess potential spatial patterns of protected areas that will gain or lose species richness. Comparisons were completed using modeled species occurrence data based on the current and projected climate in 2080. We also built a null model for random allocation of protected areas to identify where reductions in species richness will be more severe than expected. We employed several modern techniques for modeling species' distributions and summarized results using ensembles of models. Our models indicate areas of high species richness in the central and southern regions of the Atlantic Forest both for now and the future. However, we estimate that in 2080 these regions should become climatically unsuitable, decreasing the species' distribution area. Around 4% of species were predicted to become extinct, some of them being endemic to the biome. Estimates of species turnover from current to future climate tended to be high, but these findings are dependent on modeling methods. Our most important results show that only a few protected areas in the southern region of the biome would gain species. Protected areas in semideciduous forests in the western region of the biome would lose more species than expected by the null model employed. Hence, current protected areas are worse off, than just randomly selected areas, at protecting species in the future.

Multifaceted diversity-area relationships reveal global hotspots of mammalian species, trait and lineage diversity.

AIM: To define biome-scale hotspots of phylogenetic and functional mammalian biodiversity (PD and FD, respectively) and compare them to 'classical' hotspots based on species richness (SR) only. LOCATION: Global. METHODS: SR, PD & FD were computed for 782 terrestrial ecoregions using distribution ranges of 4616 mammalian species. We used a set of comprehensive diversity indices unified by a recent framework that incorporates the species relative coverage in each ecoregion. We build large-scale multifaceted diversity-area relationships to rank ecoregions according to their levels of biodiversity while accounting for the effect of area on each diversity facet. Finally we defined hotspots as the top-ranked ecoregions. RESULTS: While ignoring species relative coverage led to a relative good congruence between biome top ranked SR, PD and FD hotspots, ecoregions harboring a rich and abundantly represented evolutionary history and functional diversity did not match with top ranked ecoregions defined by species richness. More importantly PD and FD hotspots showed important spatial mismatches. We also found that FD and PD generally reached their maximum values faster than species richness as a function of area. MAIN CONCLUSIONS: The fact that PD/FD reach faster their maximal value than SR may suggest that the two former facets might be less vulnerable to habitat loss than the latter. While this point is expected, it is the first time that it is quantified at global scale and should have important consequences in conservation. Incorporating species relative coverage into the delineation of multifaceted hotspots of diversity lead to weak congruence between SR, PD and FD hotspots. This means that maximizing species number may fail at preserving those nodes (in the phylogenetic or functional tree) that are relatively abundant in the ecoregion. As a consequence it may be of prime importance to adopt a multifaceted biodiversity perspective to inform conservation strategies at global scale.