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.

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.

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.

Dissecting the null model for biological invasions: A meta-analysis of the propagule pressure effect.

A consistent determinant of the establishment success of alien species appears to be the number of individuals that are introduced to found a population (propagule pressure), yet variation in the form of this relationship has been largely unexplored. Here, we present the first quantitative systematic review of this form, using Bayesian meta-analytical methods. The relationship between propagule pressure and establishment success has been evaluated for a broad range of taxa and life histories, including invertebrates, herbaceous plants and long-lived trees, and terrestrial and aquatic vertebrates. We found a positive mean effect of propagule pressure on establishment success to be a feature of every hypothesis we tested. However, establishment success most critically depended on propagule pressures in the range of 10-100 individuals. Heterogeneity in effect size was associated primarily with different analytical approaches, with some evidence of larger effect sizes in animal rather than plant introductions. Conversely, no variation was accounted for in any analysis by the scale of study (field to global) or methodology (observational, experimental, or proxy) used. Our analyses reveal remarkable consistency in the form of the relationship between propagule pressure and alien population establishment success.

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.

Global rise in emerging alien species results from increased accessibility of new source pools.

Our ability to predict the identity of future invasive alien species is largely based upon knowledge of prior invasion history. Emerging alien species-those never encountered as aliens before-therefore pose a significant challenge to biosecurity interventions worldwide. Understanding their temporal trends, origins, and the drivers of their spread is pivotal to improving prevention and risk assessment tools. Here, we use a database of 45,984 first records of 16,019 established alien species to investigate the temporal dynamics of occurrences of emerging alien species worldwide. Even after many centuries of invasions the rate of emergence of new alien species is still high: One-quarter of first records during 2000-2005 were of species that had not been previously recorded anywhere as alien, though with large variation across taxa. Model results show that the high proportion of emerging alien species cannot be solely explained by increases in well-known drivers such as the amount of imported commodities from historically important source regions. Instead, these dynamics reflect the incorporation of new regions into the pool of potential alien species, likely as a consequence of expanding trade networks and environmental change. This process compensates for the depletion of the historically important source species pool through successive invasions. We estimate that 1-16% of all species on Earth, depending on the taxonomic group, qualify as potential alien species. These results suggest that there remains a high proportion of emerging alien species we have yet to encounter, with future impacts that are difficult to predict.

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.

The Global Distribution and Drivers of Alien Bird Species Richness.

Alien species are a major component of human-induced environmental change. Variation in the numbers of alien species found in different areas is likely to depend on a combination of anthropogenic and environmental factors, with anthropogenic factors affecting the number of species introduced to new locations, and when, and environmental factors influencing how many species are able to persist there. However, global spatial and temporal variation in the drivers of alien introduction and species richness remain poorly understood. Here, we analyse an extensive new database of alien birds to explore what determines the global distribution of alien species richness for an entire taxonomic class. We demonstrate that the locations of origin and introduction of alien birds, and their identities, were initially driven largely by European (mainly British) colonialism. However, recent introductions are a wider phenomenon, involving more species and countries, and driven in part by increasing economic activity. We find that, globally, alien bird species richness is currently highest at midlatitudes and is strongly determined by anthropogenic effects, most notably the number of species introduced (i.e., "colonisation pressure"). Nevertheless, environmental drivers are also important, with native and alien species richness being strongly and consistently positively associated. Our results demonstrate that colonisation pressure is key to understanding alien species richness, show that areas of high native species richness are not resistant to colonisation by alien species at the global scale, and emphasise the likely ongoing threats to global environments from introductions of species.

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.

Developing a list of invasive alien species likely to threaten biodiversity and ecosystems in the European Union.

The European Union (EU) has recently published its first list of invasive alien species (IAS) of EU concern to which current legislation must apply. The list comprises species known to pose great threats to biodiversity and needs to be maintained and updated. Horizon scanning is seen as critical to identify the most threatening potential IAS that do not yet occur in Europe to be subsequently risk assessed for future listing. Accordingly, we present a systematic consensus horizon scanning procedure to derive a ranked list of potential IAS likely to arrive, establish, spread and have an impact on biodiversity in the region over the next decade. The approach is unique in the continental scale examined, the breadth of taxonomic groups and environments considered, and the methods and data sources used. International experts were brought together to address five broad thematic groups of potential IAS. For each thematic group the experts first independently assembled lists of potential IAS not yet established in the EU but potentially threatening biodiversity if introduced. Experts were asked to score the species within their thematic group for their separate likelihoods of i) arrival, ii) establishment, iii) spread, and iv) magnitude of the potential negative impact on biodiversity within the EU. Experts then convened for a 2-day workshop applying consensus methods to compile a ranked list of potential IAS. From an initial working list of 329 species, a list of 66 species not yet established in the EU that were considered to be very high (8 species), high (40 species) or medium (18 species) risk species was derived. Here, we present these species highlighting the potential negative impacts and the most likely biogeographic regions to be affected by these potential IAS.

A unified classification of alien species based on the magnitude of their environmental impacts.

Species moved by human activities beyond the limits of their native geographic ranges into areas in which they do not naturally occur (termed aliens) can cause a broad range of significant changes to recipient ecosystems; however, their impacts vary greatly across species and the ecosystems into which they are introduced. There is therefore a critical need for a standardised method to evaluate, compare, and eventually predict the magnitudes of these different impacts. Here, we propose a straightforward system for classifying alien species according to the magnitude of their environmental impacts, based on the mechanisms of impact used to code species in the International Union for Conservation of Nature (IUCN) Global Invasive Species Database, which are presented here for the first time. The classification system uses five semi-quantitative scenarios describing impacts under each mechanism to assign species to different levels of impact-ranging from Minimal to Massive-with assignment corresponding to the highest level of deleterious impact associated with any of the mechanisms. The scheme also includes categories for species that are Not Evaluated, have No Alien Population, or are Data Deficient, and a method for assigning uncertainty to all the classifications. We show how this classification system is applicable at different levels of ecological complexity and different spatial and temporal scales, and embraces existing impact metrics. In fact, the scheme is analogous to the already widely adopted and accepted Red List approach to categorising extinction risk, and so could conceivably be readily integrated with existing practices and policies in many regions.

Alien species as a driver of recent extinctions.

We assessed the prevalence of alien species as a driver of recent extinctions in five major taxa (plants, amphibians, reptiles, birds and mammals), using data from the IUCN Red List. Our results show that alien species are the second most common threat associated with species that have gone completely extinct from these taxa since AD 1500. Aliens are the most common threat associated with extinctions in three of the five taxa analysed, and for vertebrate extinctions overall.

Magnitude and variation of prehistoric bird extinctions in the Pacific.

The largest extinction event in the Holocene occurred on Pacific islands, where Late Quaternary fossils reveal the loss of thousands of bird populations following human colonization of the region. However, gaps in the fossil record mean that considerable uncertainty surrounds the magnitude and pattern of these extinctions. We use a Bayesian mark-recapture approach to model gaps in the fossil record and to quantify losses of nonpasserine landbirds on 41 Pacific islands. Two-thirds of the populations on these islands went extinct in the period between first human arrival and European contact, with extinction rates linked to island and species characteristics that increased susceptibility to hunting and habitat destruction. We calculate that human colonization of remote Pacific islands caused the global extinction of close to 1,000 species of nonpasserine landbird alone; nonpasserine seabird and passerine extinctions will add to this total.

The influence of numbers on invasion success.

The process by which a species becomes a biological invader, at a location where it does not naturally occur, can be divided into a series of sequential stages (transport, introduction, establishment and spread). A species' success at passing through each of these stages depends, in a large part, on the number of individuals available to assist making each transition. Here, we review the evidence that numbers determine success at each stage of the invasion process and then discuss the likely mechanisms by which numbers affect success. We conclude that numbers of individuals affect transport and introduction by moderating the likelihood that abundant (and widespread) species are deliberately or accidentally translocated; affect establishment success by moderating the stochastic processes (demographic, environmental, genetic or Allee) to which small, introduced populations will be vulnerable; and affect invasive spread most likely because of persistent genetic effects determined by the numbers of individuals involved in the establishment phase. We finish by suggesting some further steps to advance our understanding of the influence of numbers on invasion success, particularly as they relate to the genetics of the process.

Horizon scanning for invasive alien species with the potential to threaten biodiversity in Great Britain.

Invasive alien species (IAS) are considered one of the greatest threats to biodiversity, particularly through their interactions with other drivers of change. Horizon scanning, the systematic examination of future potential threats and opportunities, leading to prioritization of IAS threats is seen as an essential component of IAS management. Our aim was to consider IAS that were likely to impact on native biodiversity but were not yet established in the wild in Great Britain. To achieve this, we developed an approach which coupled consensus methods (which have previously been used for collaboratively identifying priorities in other contexts) with rapid risk assessment. The process involved two distinct phases: Preliminary consultation with experts within five groups (plants, terrestrial invertebrates, freshwater invertebrates, vertebrates and marine species) to derive ranked lists of potential IAS. Consensus-building across expert groups to compile and rank the entire list of potential IAS. Five hundred and ninety-one species not native to Great Britain were considered. Ninety-three of these species were agreed to constitute at least a medium risk (based on score and consensus) with respect to them arriving, establishing and posing a threat to native biodiversity. The quagga mussel, Dreissena rostriformis bugensis, received maximum scores for risk of arrival, establishment and impact; following discussions the unanimous consensus was to rank it in the top position. A further 29 species were considered to constitute a high risk and were grouped according to their ranked risk. The remaining 63 species were considered as medium risk, and included in an unranked long list. The information collated through this novel extension of the consensus method for horizon scanning provides evidence for underpinning and prioritizing management both for the species and, perhaps more importantly, their pathways of arrival. Although our study focused on Great Britain, we suggest that the methods adopted are applicable globally.

Effects of recent environmental change on accuracy of inferences of extinction status.

Correctly classifying a species as extinct or extant is of critical importance if current rates of biodiversity loss are to be accurately quantified. Observing an extinction event is rare, so in many cases extinction status is inferred using methods based on the analysis of records of historic sighting events. The accuracy of such methods is difficult to test. However, results of recent experiments with microcosm communities suggest that the rate at which a population declines to extinction, potentially driven by varying environmental conditions, may alter one's ability accurately to infer extinction status. We tested how the rate of population decline, driven by historic environmental change, alters the accuracy of 6 commonly applied sighting-based methods used to infer extinction. We used data from small-scale experimental communities and recorded wild population extirpations. We assessed how accuracy of the different methods was affected by rate of population decline, search effort, and number of sighting events recorded. Rate of population decline and historic population size of the species affected the accuracy of inferred extinction dates; however, faster declines produced more accurate inferred dates of extinction, but only when population sizes were higher. Optimal linear estimation (OLE) offered the most reliable and robust estimates, though no single method performed best in all situations, and it may be appropriate to use a different method if information regarding historic search efforts is available. OLE provided the most accurate estimates of extinction when the number of sighting events used was >10, and future use of this method should take this into account. Data from experimental populations provide added insight into testing techniques to discern wild extirpation events. Care should be taken designing such experiments so that they mirror closely the abundance dynamics of populations affected by real-world extirpation events.

Defining the impact of non-native species.

Non-native species cause changes in the ecosystems to which they are introduced. These changes, or some of them, are usually termed impacts; they can be manifold and potentially damaging to ecosystems and biodiversity. However, the impacts of most non-native species are poorly understood, and a synthesis of available information is being hindered because authors often do not clearly define impact. We argue that explicitly defining the impact of non-native species will promote progress toward a better understanding of the implications of changes to biodiversity and ecosystems caused by non-native species; help disentangle which aspects of scientific debates about non-native species are due to disparate definitions and which represent true scientific discord; and improve communication between scientists from different research disciplines and between scientists, managers, and policy makers. For these reasons and based on examples from the literature, we devised seven key questions that fall into 4 categories: directionality, classification and measurement, ecological or socio-economic changes, and scale. These questions should help in formulating clear and practical definitions of impact to suit specific scientific, stakeholder, or legislative contexts.

A population model for predicting the successful establishment of introduced bird species.

One of the strongest generalities in invasion biology is the positive relationship between probability of establishment and the numbers of individuals introduced. Nevertheless, a number of significant questions remain regarding: (1) the relative importance of different processes during introduction (e.g., demographic, environmental, and genetic stochasticity, and Allee effects); (2) the relative effects of propagule pressure (e.g., number of introductions, size of introductions, and lag between introductions); and (3) different life history characteristics of the species themselves. Here, we adopt an individual-based simulation modeling approach to explore a range of such details in the relationship between establishment success and numbers of individuals introduced. Our models are developed for typical exotic bird introductions, for which the relationship between probability of establishment and the numbers of individuals introduced has been particularly well documented. For both short-lived and long-lived species, probability of establishment decreased across multiple introductions (compared with a single introduction of the same total size), and this decrease was greater when inbreeding depression was included. Sensitivity analyses revealed four predictors that together accounted for >95% of model performance. Of these, R 0 (the average number of daughters produced per female over her lifetime) and propagule pressure were of primary importance, while random environmental effects and inbreeding depression exerted lesser influence. Initial founder size is undoubtedly going to be important for ensuring the persistence of introduced populations. However, we found the demographic traits, which influence how introduced individuals behave, to have the greatest effect on establishment success.