Regional invasion history and land use shape the prevalence of non-native species in local assemblages.

The ecological impact of non-native species arises from their establishment in local assemblages. However, the rates of non-native spread in new regions and their determinants have not been comprehensively studied. Here, we combined global databases documenting the occurrence of non-native species and residence of non-native birds, mammals, and vascular plants at regional and local scales to describe how the likelihood of non-native occurrence and their proportion in local assemblages relate with their residence time and levels of human usage in different ecosystems. Our findings reveal that local non-native occurrence generally increases with residence time. Colonization is most rapid in croplands and urban areas, while it is slower and variable in natural or semi-natural ecosystems. Notably, non-native occurrence continues to rise even 200 years after introduction, especially for birds and vascular plants, and in other land-use types rather than croplands and urban areas. The impact of residence time on non-native proportions is significant only for mammals. We conclude that the continental exchange of biotas requires considerable time for effects to manifest at the local scale across taxa and land-use types. The unpredictability of future impacts, implied by the slow spread of non-native species, strengthens the call for stronger regulations on the exchange of non-native species to reduce the long-lasting invasion debt looming on ecosystems' future.

The importance of migratory drop-off for island colonization in birds.

Seasonal migration is an underappreciated driver of animal diversification. Changes in migratory behaviour may favour the establishment of sedentary founder populations and promote speciation if there is sufficient reproductive isolation between sedentary and migratory populations. From a systematic literature review, we here quantify the role of migratory drop-off-the loss of migratory behaviour-in promoting speciation in birds on islands. We identify at least 157 independent colonization events likely initiated by migratory species that led to speciation, including 44 cases among recently extinct species. By comparing, for all islands, the proportion of island endemic species that derived from migratory drop-off with the proportion of migratory species among potential colonizers, we showed that seasonal migration has a larger effect on island endemic richness than direct dispersal. We also found that the role of migration in island colonization increases with the geographic isolation of islands. Furthermore, the success of speciation events depends in part on species biogeographic and ecological factors, here positively associated with greater range size and larger flock sizes. These results highlight the importance of shifts in migratory behaviour in the speciation process and calls for greater consideration of migratory drop-off in the biogeographic distribution of birds.

Using the IUCN Environmental Impact Classification for Alien Taxa to inform decision-making.

The Environmental Impact Classification for Alien Taxa (EICAT) is an important tool for biological invasion policy and management and has been adopted as an International Union for Conservation of Nature (IUCN) standard to measure the severity of environmental impacts caused by organisms living outside their native ranges. EICAT has already been incorporated into some national and local decision-making procedures, making it a particularly relevant resource for addressing the impact of non-native species. Recently, some of the underlying conceptual principles of EICAT, particularly those related to the use of the precautionary approach, have been challenged. Although still relatively new, guidelines for the application and interpretation of EICAT will be periodically revisited by the IUCN community, based on scientific evidence, to improve the process. Some of the criticisms recently raised are based on subjectively selected assumptions that cannot be generalized and may harm global efforts to manage biological invasions. EICAT adopts a precautionary principle by considering a species' impact history elsewhere because some taxa have traits that can make them inherently more harmful. Furthermore, non-native species are often important drivers of biodiversity loss even in the presence of other pressures. Ignoring the precautionary principle when tackling the impacts of non-native species has led to devastating consequences for human well-being, biodiversity, and ecosystems, as well as poor management outcomes, and thus to significant economic costs. EICAT is a relevant tool because it supports prioritization and management of non-native species and meeting and monitoring progress toward the Kunming-Montreal Global Biodiversity Framework (GBF) Target 6.

Uso de la Clasificación de Impacto Ambiental de los Taxones Exóticos de la UICN para la toma de decisiones Resumen La Clasificación de Impacto Ambiental de los Taxones Exóticos (EICAT, en inglés) es una herramienta importante para las políticas y manejo de las invasiones biológicas y ha sido adoptada como un estándar de la Unión Internacional para la Conservación de la Naturaleza (UICN) para medir la seriedad del impacto ambiental causado por los organismos que viven fuera de su extensión nativa. La EICAT ya ha sido incorporada a algunos procedimientos locales y nacionales de toma de decisiones, lo que la vuelve un recurso particularmente relevante para abordar el impacto de las especies no nativas. Algunos principios conceptuales subyacentes de la EICAT han sido cuestionados recientemente, en particular aquellos relacionados con el uso del principio de precaución. Aunque todavía son relativamente nuevas, las directrices para la aplicación e interpretación de la EICAT tendrán una revisión periódica, basada en evidencia científica, por parte de la comunidad de la UICN para mejorar el proceso. Algunas de las críticas recientes están basadas en suposiciones seleccionadas subjetivamente que no pueden generalizarse y podrían perjudicar los esfuerzos globales para manejar las invasiones biológicas. La EICAT adopta un principio de precaución cuando considera el historial de impacto de una especie en cualquier otro lugar ya que algunos taxones tienen características que podrían volverlos más dañinos. Además, las especies no nativas suelen ser factores de pérdida de bidiversidad, incluso bajo otras presiones. Cuando ignoramos el principio de precaución al abordar el impacto de las especies no nativas, hay consecuencias devastadoras para el bienestar humano, la biodiversidad y los ecosistemas, así como resultados pobres de conservación, y por lo tanto con costos económicos importantes. La EICAT es una herramienta relevante porque respalda la priorización y el manejo de las especies no nativas y ayuda con el cumplimiento y monitoreo del progreso para llegar al objetivo 6 del Marco Mundial de Biodiversidad Kunming­Montreal.

Undiscovered bird extinctions obscure the true magnitude of human-driven extinction waves.

Birds are among the best-studied animal groups, but their prehistoric diversity is poorly known due to low fossilization potential. Hence, while many human-driven bird extinctions (i.e., extinctions caused directly by human activities such as hunting, as well as indirectly through human-associated impacts such as land use change, fire, and the introduction of invasive species) have been recorded, the true number is likely much larger. Here, by combining recorded extinctions with model estimates based on the completeness of the fossil record, we suggest that at least ~1300-1500 bird species (~12% of the total) have gone extinct since the Late Pleistocene, with 55% of these extinctions undiscovered (not yet discovered or left no trace). We estimate that the Pacific accounts for 61% of total bird extinctions. Bird extinction rate varied through time with an intense episode ~1300 CE, which likely represents the largest human-driven vertebrate extinction wave ever, and a rate 80 (60-95) times the background extinction rate. Thus, humans have already driven more than one in nine bird species to extinction, with likely severe, and potentially irreversible, ecological and evolutionary consequences.

Quantifying global colonization pressures of alien vertebrates from wildlife trade.

The global trade in live wildlife elevates the risk of biological invasions by increasing colonization pressure (the number of alien species introduced to an area). Yet, our understanding of species traded as aliens remains limited. We created a comprehensive global database on live terrestrial vertebrate trade and use it to investigate the number of traded alien species, and correlates of establishment richness for aliens. We identify 7,780 species involved in this trade globally. Approximately 85.7% of these species are traded as aliens, and 12.2% of aliens establish populations. Countries with greater trading power, higher incomes, and larger human populations import more alien species. These countries, along with island nations, emerge as hotspots for establishment richness of aliens. Colonization pressure and insularity consistently promote establishment richness across countries, while socio-economic factors impact specific taxa. Governments must prioritize policies to mitigate the release or escape of traded animals and protect global biosecurity.

The impact of land use on non-native species incidence and number in local assemblages worldwide.

While the regional distribution of non-native species is increasingly well documented for some taxa, global analyses of non-native species in local assemblages are still missing. Here, we use a worldwide collection of assemblages from five taxa - ants, birds, mammals, spiders and vascular plants - to assess whether the incidence, frequency and proportions of naturalised non-native species depend on type and intensity of land use. In plants, assemblages of primary vegetation are least invaded. In the other taxa, primary vegetation is among the least invaded land-use types, but one or several other types have equally low levels of occurrence, frequency and proportions of non-native species. High land use intensity is associated with higher non-native incidence and frequency in primary vegetation, while intensity effects are inconsistent for other land-use types. These findings highlight the potential dual role of unused primary vegetation in preserving native biodiversity and in conferring resistance against biological invasions.

Drivers of alien species composition in bird markets across the world.

The global pet trade is a major pathway for the introduction of invasive alien species. The composition of species selected for transport is driven by market demands, which may be influenced by a combination of both historical and cultural factors. We compared Eastern (Taiwan) and Western (Australia and the Iberian Peninsula) bird markets to explore factors associated with the species composition and geographic origin of the birds for sale. We used a bespoke randomization test to compare species composition, geographic origins, and species overlap at different taxonomic levels among bird markets across countries. Alien species identified in the study accounted for more than 10% of the world's bird species. Parrots and songbirds were the most common alien bird taxa traded across all markets. In both Iberian and Australian markets, there was a strong bias toward parrots, waxbills, gamebirds, and finches. In Taiwan, species traded more than expected were parrots, waxbills, starlings, and leafbirds. Neotropical species were the most traded group in the three markets. Afrotropical species were also traded more than expected in Iberian and Australian markets, while the Taiwanese traded more alien species from neighboring Asian regions. The bird trade focuses on the same few bird groups worldwide. The composition and origin of species preferred in the Western markets may be influenced by colonial histories, cultural similarity, and strict regulations on wildlife importation, while species preferences in Eastern markets are strongly influenced by regional culture and proximity. Propagule pressure is a dominant factor influencing the success of biological invasions; it is important to recognize differences in the composition of bird markets among regions because they can translate into different invasion risks, among other factors.

Loss of functional diversity through anthropogenic extinctions of island birds is not offset by biotic invasions.

Human impacts reshape ecological communities through the extinction and introduction of species. The combined impact of these factors depends on whether non-native species fill the functional roles of extinct species, thus buffering the loss of functional diversity. This question has been difficult to address, because comprehensive information about past extinctions and their traits is generally lacking. We combine detailed information about extinct, extant, and established alien birds to quantify historical changes in functional diversity across nine oceanic archipelagos. We found that alien species often equal or exceed the number of anthropogenic extinctions yet apparently perform a narrower set of functional roles as current island assemblages have undergone a substantial and ubiquitous net loss in functional diversity and increased functional similarity among assemblages. Our results reveal that the introduction of alien species has not prevented anthropogenic extinctions from reducing and homogenizing the functional diversity of native bird assemblages on oceanic archipelagos.

DAMA: the global Distribution of Alien Mammals database.

We developed the DAMA (Distribution of Alien Mammals) database, a comprehensive source reporting the global distribution of the 230 species of mammals that have established self-sustaining and free-ranging populations outside their native range due to direct or indirect human action. Every alien range is accompanied by information on its invasion stage, pathway, method of introduction, and date of introduction. We collected information from 827 different sources (scientific literature, books, risk assessments, reports, online biodiversity databases and websites), and used it to draw alien range maps for these species following the IUCN mapping framework. DAMA comprises 2,726 range polygons, covering 199 countries, 2,190 level 1 administrative areas, and 11 zoogeographic realms for the period 21500 BC-AD 2017. The most represented orders among introduced mammal species are Rodentia (n = 58, 25.22%), Cetartiodactyla (n = 49 species, 21.30%), Carnivora (n = 30 species, 13.04%), Diprotodontia (n = 28, 12.17%), and Primates (n = 26, 11.30%). Mammal species have been frequently introduced for hunting (n = 100), pet trade (n = 57), conservation (n = 51), and fauna improvement (n = 42). The majority of range polygons are placed on islands (n = 2,196, 80.56%), encompass populations that have moved beyond establishment and into the invasion stage (n = 1,655, 60.71%), and originated from 1500 AD to the present (n = 1,496, 54.88%). Despite inheriting literature biases towards more studied regions (e.g., developed countries), DAMA is the most up-to-date picture of alien mammal global distribution and can be used to investigate their invasion ecology across different biogeographical regions. There are no copyright or proprietary restrictions; IUCN range maps were modified into a derivative work according to the IUCN's terms of service.

Environmental resistance predicts the spread of alien species.

The unabating rise in the number of species introduced outside of their native range makes predicting the spread of alien species an urgent challenge. Most predictions use models of the ecological niche of a species to identify suitable areas for invasion, but these predictions may have limited accuracy. Here, using the global alien avifauna, we demonstrate an alternative approach for predicting alien spread based on the environmental resistance of the landscape. This approach does not require any information on the ecological niche of the invading species and, instead, uses gradients of biotic similarity among native communities in the invaded region to predict the most likely routes of spread. We show that environmental resistance predicts patterns of spread better than a null model of random dispersal or a model based on climate matching to the native range of each species. Applying this approach to simulate future spread reveals major regional differences in projected invasion risk, shaped by proximity to existing invasion hotspots as well as gradients in environmental resistance. Our results show how environmental resistance may provide a general and complementary approach for predicting invasion risk that can be rapidly deployed even when information on the niche or the identity of potential invaders is unknown.

A counterfactual approach to measure the impact of wet grassland conservation on U.K. breeding bird populations.

Wet grassland populations of wading birds in the United Kingdom have declined severely since 1990. To help mitigate these declines, the Royal Society for the Protection of Birds has restored and managed lowland wet grassland nature reserves to benefit these and other species. However, the impact of these reserves on bird population trends has not been evaluated experimentally due to a lack of control populations. We compared population trends from 1994 to 2018 among 5 bird species of conservation concern that breed on these nature reserves with counterfactual trends created from matched breeding bird survey observations. We compared reserve trends with 3 different counterfactuals based on different scenarios of how reserve populations could have developed in the absence of conservation. Effects of conservation interventions were positive for all 4 targeted wading bird species: Lapwing (Vanellus vanellus), Redshank (Tringa totanus), Curlew (Numenius arquata), and Snipe (Gallinago gallinago). There was no positive effect of conservation interventions on reserves for the passerine, Yellow Wagtail (Motacilla flava). Our approach using monitoring data to produce valid counterfactual controls is a broadly applicable method allowing large-scale evaluation of conservation impact.

Un Enfoque Hipotético para Medir el Impacto de la Conservación de Pastizales Húmedos sobre Poblaciones Reproductoras de Aves en el Reino Unido Resumen Las poblaciones de aves zancudas en los pastizales húmedos del Reino Unido han declinado gravemente desde 1990. Para ayudar con la mitigación de estas declinaciones, la Real Sociedad para la Protección de las Aves ha restaurado y manejado las reservas naturales en pastizales húmedos de tierras bajas para beneficiar a estas y otras especies. Sin embargo, el impacto de estas reservas sobre las tendencias poblacionales de las aves no ha sido evaluado experimentalmente debido a la falta de poblaciones control. Comparamos las tendencias poblacionales entre 1994 y 2018 de cinco especies de aves de importancia para la conservación, que se reproducen dentro de estas reservas naturales, mediante tendencias hipotéticas creadas a partir de los censos de observación de aves reproductoras emparejadas. Comparamos las tendencias de las reservas con tres casos hipotéticos diferentes basados en diferentes escenarios de cómo las poblaciones de la reserva podrían haberse desarrollado en ausencia de la conservación. Los efectos de las intervenciones de conservación fueron positivos para las cuatro especies focales de aves zancudas: Vanellus vanellus, Tringa totanus, Numenius arquata y Gallinago gallinago. No hubo un efecto positivo de las intervenciones de conservación para la especie paserina Motacilla flava. Nuestra estrategia utilizando datos de monitoreos para producir controles hipotéticos válidos es un método ampliamente aplicable que permite ka evaluación del impacto de la conservación a gran escala.

Invasion costs, impacts, and human agency: response to Sagoff 2020.

Article impact statement: In an era of profound biodiversity crisis, invasion costs, invader impacts, and human agency should not be dismissed.

Projecting the continental accumulation of alien species through to 2050.

Biological invasions have steadily increased over recent centuries. However, we still lack a clear expectation about future trends in alien species numbers. In particular, we do not know whether alien species will continue to accumulate in regional floras and faunas, or whether the pace of accumulation will decrease due to the depletion of native source pools. Here, we apply a new model to simulate future numbers of alien species based on estimated sizes of source pools and dynamics of historical invasions, assuming a continuation of processes in the future as observed in the past (a business-as-usual scenario). We first validated performance of different model versions by conducting a back-casting approach, therefore fitting the model to alien species numbers until 1950 and validating predictions on trends from 1950 to 2005. In a second step, we selected the best performing model that provided the most robust predictions to project trajectories of alien species numbers until 2050. Altogether, this resulted in 3,790 stochastic simulation runs for 38 taxon-continent combinations. We provide the first quantitative projections of future trajectories of alien species numbers for seven major taxonomic groups in eight continents, accounting for variation in sampling intensity and uncertainty in projections. Overall, established alien species numbers per continent were predicted to increase from 2005 to 2050 by 36%. Particularly, strong increases were projected for Europe in absolute (+2,543 ± 237 alien species) and relative terms, followed by Temperate Asia (+1,597 ± 197), Northern America (1,484 ± 74) and Southern America (1,391 ± 258). Among individual taxonomic groups, especially strong increases were projected for invertebrates globally. Declining (but still positive) rates were projected only for Australasia. Our projections provide a first baseline for the assessment of future developments of biological invasions, which will help to inform policies to contain the spread of alien species.

Zoonotic host diversity increases in human-dominated ecosystems.

Land use change-for example, the conversion of natural habitats to agricultural or urban ecosystems-is widely recognized to influence the risk and emergence of zoonotic disease in humans1,2. However, whether such changes in risk are underpinned by predictable ecological changes remains unclear. It has been suggested that habitat disturbance might cause predictable changes in the local diversity and taxonomic composition of potential reservoir hosts, owing to systematic, trait-mediated differences in species resilience to human pressures3,4. Here we analyse 6,801 ecological assemblages and 376 host species worldwide, controlling for research effort, and show that land use has global and systematic effects on local zoonotic host communities. Known wildlife hosts of human-shared pathogens and parasites overall comprise a greater proportion of local species richness (18-72% higher) and total abundance (21-144% higher) in sites under substantial human use (secondary, agricultural and urban ecosystems) compared with nearby undisturbed habitats. The magnitude of this effect varies taxonomically and is strongest for rodent, bat and passerine bird zoonotic host species, which may be one factor that underpins the global importance of these taxa as zoonotic reservoirs. We further show that mammal species that harbour more pathogens overall (either human-shared or non-human-shared) are more likely to occur in human-managed ecosystems, suggesting that these trends may be mediated by ecological or life-history traits that influence both host status and tolerance to human disturbance5,6. Our results suggest that global changes in the mode and the intensity of land use are creating expanding hazardous interfaces between people, livestock and wildlife reservoirs of zoonotic disease.

Animal invaders threaten protected areas worldwide.

Protected areas are the cornerstone of biodiversity conservation. However, alien species invasion is an increasing threat to biodiversity, and the extent to which protected areas worldwide are resistant to incursions of alien species remains poorly understood. Here, we investigate establishment by 894 terrestrial alien animals from 11 taxonomic groups including vertebrates and invertebrates across 199,957 protected areas at the global scale. We find that <10% of protected areas are home to any of the alien animals, but there is at least one established population within 10-100 km of the boundaries of 89%-99% of protected areas, while >95% of protected areas are environmentally suitable for establishment. Higher alien richness is observed in IUCN category-II national parks supposedly with stricter protection, and in larger protected areas with higher human footprint and more recent designation. Our results demonstrate that protected areas provide important protection from biological invasions, but invasions may become an increasingly dominant problem in the near future.

Scientists' warning on invasive alien species.

Biological invasions are a global consequence of an increasingly connected world and the rise in human population size. The numbers of invasive alien species - the subset of alien species that spread widely in areas where they are not native, affecting the environment or human livelihoods - are increasing. Synergies with other global changes are exacerbating current invasions and facilitating new ones, thereby escalating the extent and impacts of invaders. Invasions have complex and often immense long-term direct and indirect impacts. In many cases, such impacts become apparent or problematic only when invaders are well established and have large ranges. Invasive alien species break down biogeographic realms, affect native species richness and abundance, increase the risk of native species extinction, affect the genetic composition of native populations, change native animal behaviour, alter phylogenetic diversity across communities, and modify trophic networks. Many invasive alien species also change ecosystem functioning and the delivery of ecosystem services by altering nutrient and contaminant cycling, hydrology, habitat structure, and disturbance regimes. These biodiversity and ecosystem impacts are accelerating and will increase further in the future. Scientific evidence has identified policy strategies to reduce future invasions, but these strategies are often insufficiently implemented. For some nations, notably Australia and New Zealand, biosecurity has become a national priority. There have been long-term successes, such as eradication of rats and cats on increasingly large islands and biological control of weeds across continental areas. However, in many countries, invasions receive little attention. Improved international cooperation is crucial to reduce the impacts of invasive alien species on biodiversity, ecosystem services, and human livelihoods. Countries can strengthen their biosecurity regulations to implement and enforce more effective management strategies that should also address other global changes that interact with invasions.

Lasting the distance: The survival of alien birds shipped to New Zealand in the 19th century.

Invasive alien species are a major threat to biodiversity and human activities, providing a strong incentive to understand the processes by which alien invasion occurs. While it is important to understand the determinants of success at each of several invasion stages-transport, introduction, establishment, and spread-few studies have explored the first of these stages. Here, we quantify and analyze variation in the success of individual animals in surviving the transport stage, based on shipping records of European passerines destined for New Zealand. We mined the original documents of Acclimatisation Societies, established in New Zealand for the purpose of introducing supposedly beneficial alien species, in combination with recently digitized newspaper archives, to produce a unique dataset of 122 ships that carried passerines from Europe to New Zealand between 1850 and 1885. For 37 of these shipments, data on the survival of individual species were available. Using generalized linear mixed models, we explored how survival was related to characteristics of the shipments and the species. We show that species differed greatly in their survival, but none of the tested traits accounted for these differences. Yet, survival increased over time, which mirrors the switch from early haphazard shipments to larger organized shipments. Our results imply that it was the quality of care received by the birds that most affected success at this stage of the invasion process.

The relationship between propagule pressure and establishment success in alien bird populations: a re-analysis of Moulton & Cropper (2019).

A recent analysis by Moulton & Cropper (2019) of a global dataset on alien bird population introductions claims to find no evidence that establishment success is a function of the size of the founding population. Here, we re-analyse Moulton & Cropper's data and show that this conclusion is based on flawed statistical methods-their data in fact confirm a strong positive relationship between founding population size and establishment success. We also refute several non-statistical arguments against the likelihood of such an effect presented by Moulton & Cropper. We conclude that a core tenet of population biology-that small populations are more prone to extinction-applies to alien populations beyond their native geographic range limits as much as to native populations within them.

Location-level processes drive the establishment of alien bird populations worldwide.

Human-mediated translocation of species to areas beyond their natural distribution (which results in 'alien' populations1) is a key signature of the Anthropocene2, and is a primary global driver of biodiversity loss and environmental change3. Stemming the tide of invasions requires understanding why some species fail to establish alien populations, and others succeed. To achieve this, we need to integrate the effects of features of the introduction site, the species introduced and the specific introduction event. Determining which, if any, location-level factors affect the success of establishment has proven difficult, owing to the multiple spatial, temporal and phylogenetic axes along which environmental variation may influence population survival. Here we apply Bayesian hierarchical regression analysis to a global spatially and temporally explicit database of introduction events of alien birds4 to show that environmental conditions at the introduction location, notably climatic suitability and the presence of other groups of alien species, are the primary determinants of successful establishment. Species-level traits and the size of the founding population (propagule pressure) exert secondary, but important, effects on success. Thus, current trajectories of anthropogenic environmental change will most probably facilitate future incursions by alien species, but predicting future invasions will require the integration of multiple location-, species- and event-level characteristics.

Risks of Biological Invasion on the Belt and Road.

China's Belt and Road Initiative (BRI) is an unprecedented global development program that involves nearly half of the world's countries [1]. It not only will have economic and political influences, but also may generate multiple environmental challenges and is a focus of considerable academic and public concerns [2-6]. The Chinese government expects BRI to be a sustainable development, paying equal attention to economic development and environmental conservation [7]. However, BRI's high expenditure on infrastructure construction, by accelerating trade and transportation, is likely to promote alien species invasions [5], one of the primary anthropogenic threats to global biodiversity [8]. BRI countries may have different susceptibilities to invasive species due to different financial and response capacities [9]. Moreover, these countries overlap 27 of 35 recognized global biodiversity hotspots [10]. Identifying those areas with high-invasion risks, and species with high invasive potentials within BRI countries, is therefore of vital importance for the sustainable implementation of the BRI, and the development of early, economical, and effective biosecurity strategies [11]. In response, we present here a comprehensive study to evaluate invasion risks by alien vertebrates within BRI. We identified a total of 14 invasion hotspots, the majority of which fall along the six proposed BRI economic corridors, with the proportion of grid cells in invasion hotspots 1.6 times higher than other regions. Based on our results, we recommend the initiation of a project targeting early prevention, strict surveillance, rapid response, and effective control of alien species in BRI countries to ensure that this development is sustainable.