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Biological invasions, as a result of human intervention through trade and mobility, are the second biggest cause of biodiversity loss. The impacts of invasive alien species (IAS) on the environment are well known, however, economic impacts are poorly estimated, especially in mega-diverse countries where both economic and ecological consequences of these effects can be catastrophic. Ecuador, one of the smallest mega-diverse countries, lacks a comprehensive description of the economic costs of IAS within its territory. Here, using "InvaCost", a public database that compiles all recorded monetary costs associated with IAS from English and Non-English sources, we investigated the economic costs of biological invasions. We found that between 1983 and 2017, the reported costs associated with biological invasions ranged between US$86.17 million (when considering only the most robust data) and US$626 million (when including all cost data) belonging to 37 species and 27 genera. Furthermore, 99% of the recorded cost entries were from the Galapagos Islands. From only robust data, the costliest identified taxonomic group was feral goats (Capra hircus; US$20 million), followed by Aedes mosquitoes (US$2.14 million) while organisms like plant species from the genus Rubus, a parasitic fly (Philornis downsi), black rats (Rattus rattus) and terrestrial gastropods (Achatina fulica) represented less than US$2 million each. Costs of "mixed-taxa" (i.e. plants and animals) represented the highest (61% of total robust costs; US$52.44 million). The most impacted activity sector was the national park authorities, which spent about US$84 million. Results from robust data also revealed that management expenditures were the major type of costs recorded in the Galapagos Islands; however, costs reported for medical losses related to Aedes mosquitoes causing dengue fever in mainland Ecuador would have ranked first if more detailed information had allowed us to categorize them as robust data. Over 70% of the IAS reported for Ecuador did not have reported costs. These results suggest that costs reported here are a massive underestimate of the actual economic toll of invasions in the country.

The rate of biological invasions is growing unprecedentedly, threatening ecological and socioeconomic systems worldwide. Quantitative understandings of invasion temporal trajectories are essential to discern current and future economic impacts of invaders, and then to inform future management strategies. Here, we examine the temporal trends of cumulative invasion costs by developing and testing a novel mathematical model with a population dynamical approach based on logistic growth. This model characterises temporal cost developments into four curve types (I - IV), each with distinct mathematical and qualitative properties, allowing for the parameterization of maximum cumulative costs, carrying capacities and growth rates. We test our model using damage cost data for eight genera (Rattus, Aedes, Canis, Oryctolagus, Sturnus, Ceratitis, Sus and Lymantria) extracted from the InvaCost database – which is the most up-to-date and comprehensive global compilation of economic cost estimates associated with invasive alien species. We find fundamental differences in the temporal dynamics of damage costs among genera, indicating they depend on invasion duration, species ecology and impacted sectors of economic activity. The fitted cost curves indicate a lack of broadscale support for saturation between invader density and impact, including for Canis, Oryctolagus and Lymantria, whereby costs continue to increase with no sign of saturation. For other taxa, predicted saturations may arise from data availability issues resulting from an underreporting of costs in many invaded regions. Overall, this population dynamical approach can produce cost trajectories for additional existing and emerging species, and can estimate the ecological parameters governing the linkage between population dynamics and cost dynamics.

Despite voluminous literature identifying invasive species impacts, understandings of monetary costs remain limited. Recently, profound impacts have been attributed to invasive crustaceans, but associated monetary costs lack synthesis. Here, we analyse globally reported costs of invasive freshwater crayfish across taxonomic, spatial and temporal descriptors. Moreover, we compare their cost magnitude to other invasive crustaceans — crabs, amphipods and lobsters. Between 2000 and 2020, crayfish caused US$ 1.28 billion in reported costs; the vast majority (95%) attributed to Astacidae (principally the signal crayfish Pacifastacus leniusculus) and the remainder to Cambaridae. According to reports, crayfish costs mostly impacted European economies (US$ 1.23 billion), followed by costs reported for North America and Asia. Despite well-known damages caused by invasive crayfish, costs were unreported elsewhere, highlighting knowledge gaps and challenges in cost quantifications. Invasive crayfish costs increased exponentially in the last two decades, averaging at US$ 61 million per-annum. Invasive crabs caused costs of similar magnitude (US$ 1.25 billion; US$ 53 million per-annum) but were mostly confined to North America (95%). Damage-related costs dominated for both crayfish (83%) and crabs (99%), with management spending lacking. Reported economic impacts from amphipods (US$ 178.8 thousand) and lobsters (US$ 44.6 thousand) were considerably lower. We identify burgeoning economic costs from these invasive groups yet highlight pervasive knowledge gaps at multiple scales. Further cost reporting is required to better-ascertain the true scale of monetary costs caused by invasive aquatic crustaceans.

Invasive alien species are a well-known and pervasive threat to global biodiversity and human well-being. Despite substantial impacts of invasive alien species, quantitative syntheses of monetary costs incurred from invasions in national economies are often missing. As a consequence, adequate resource allocation for management responses to invasions has been inhibited, because cost-benefit analysis of management actions cannot be derived. To determine the economic cost of invasions in Germany, a Central European country with the 4th largest GDP in the world, we analysed published data collected from the first global assessment of economic costs of invasive alien species. Overall, economic costs were estimated at US$ 9.8 billion between 1960 and 2020, including US$ 8.9 billion in potential costs. The potential costs were mostly linked to extrapolated costs of the American bullfrog Lithobates catesbeianus, the black cherry Prunus serotina and two mammals: the muskrat Ondatra zibethicus and the American mink Neovison vison. Observed costs were driven by a broad range of taxa and mostly associated with control-related spending and resource damages or losses. We identified a considerable increase in costs relative to previous estimates and through time. Importantly, of the 2,249 alien and 181 invasive species reported in Germany, only 28 species had recorded economic costs. Therefore, total quantifications of invasive species costs here should be seen as very conservative. Our findings highlight a distinct lack of information in the openly-accessible literature and governmental sources on invasion costs at the national level, masking the highly-probable existence of much greater costs of invasions in Germany. In addition, given that invasion rates are increasing, economic costs are expected to further increase. The evaluation and reporting of economic costs need to be improved in order to deliver a basis for effective mitigation and management of invasions on national and international economies. Wirtschaftliche Kosten invasiver Arten in Deutschland. Invasive gebietsfremde Arten sind eine bekannte und weit verbreitete Bedrohung für die globale Artenvielfalt und das Wohlergehen des Menschen. Trotz erheblicher Auswirkungen invasiver gebietsfremder Arten fehlen häufig quantitative Synthesen der finanziellen Kosten, die durch Invasionen entstehen. Infolgedessen wurde eine angemessene Ressourcenzuweisung für Managementreaktionen auf Invasionen verhindert, da keine Kosten-Nutzen-Analyse von Managementmaßnahmen abgeleitet werden kann. Um die wirtschaftlichen Kosten von Invasionen in Deutschland, einem mitteleuropäischen Land mit dem viertgrößten BIP der Welt, zu ermitteln, haben wir veröffentlichte Daten analysiert, die aus der ersten globalen Bewertung der wirtschaftlichen Kosten invasiver gebietsfremder Arten stammen. Insgesamt wurden die wirtschaftlichen Kosten zwischen 1960 und 2020 auf 9.8 Milliarden US-Dollar geschätzt, einschließlich potenzieller Kosten in Höhe von 8.9 Milliarden US-Dollar. Die potenziellen Kosten waren hauptsächlich auf die extrapolierten Kosten des amerikanischen Ochsenfrosches Lithobates catesbeianus, der Tollkirsche Prunus serotina und zweier Säugetiere zurückzuführen: des Bisamratten Ondatra zibethicus und des amerikanischen Nerz Neovison vison. Die beobachteten Kosten wurden von einer breiten Palette von Taxa getrieben und waren hauptsächlich mit kontrollbezogenen Ausgaben und Ressourcenschäden oder -verlusten verbunden. Wir haben einen erheblichen Anstieg der Kosten im Vergleich zu früheren Schätzungen und im Laufe der Zeit festgestellt. Wichtig ist, dass von den in Deutschland gemeldeten 2.249 gebietsfremden und 181 invasiven Arten nur 28 Arten wirtschaftliche Kosten verzeichneten. Daher sollte die Gesamtquantifizierung der Kosten invasiver Arten hier als sehr konservativ angesehen werden. Unsere Ergebnisse zeigen einen deutlichen Mangel an Informationen in der öffentlich zugänglichen Literatur und in staatlichen Quellen zu Invasionskosten auf nationaler Ebene, was die höchstwahrscheinliche Existenz viel höherer Invasionskosten in Deutschland maskiert. Angesichts der steigenden Invasionsraten werden die wirtschaftlichen Kosten voraussichtlich weiter steigen. Die Bewertung und Berichterstattung der wirtschaftlichen Kosten muss verbessert werden, um eine Grundlage für eine wirksame Eindämmung und Bewältigung der Invasionen in die nationale und internationale Wirtschaft zu schaffen. Costos económicos de las especies invasoras en Alemania. Las especies invasoras representan una conocida amenaza en general para la biodiversidad del planeta y el bienestar humano. A menudo se omite la cuantificación de los costes económicos que provocan las especies invasoras, a pesar de los impactos sustanciales que ellas provocan. En consecuencia, el reparto de los recursos para el manejo de las respuestas es inadecuado ante las invasiones, por lo tanto no es posible obtener un análisis de los costos y beneficios de las acciones de manejo. Para determinar los costos económicos de las invasiones en Alemania, un país Centroeuropeo con el 4to producto interno bruto más grande del planeta, se analizaron datos publicados en la primera evaluación global de los costes económicos de especies invasoras. En general se estimaron costes económicos de US $9.8 mil millones entre los años 1960 y 2020, incluyendo US $8.9 mil millones en costos potenciales. Los costos potenciales se asociaron con los costes extrapolados de la rana toro americana Lithobates catesbeianus, el cerezo negro Prunus serotina y dos mamíferos: la rata almizclera Ondatra zibethicus y el visón americano Neovison vison. Los costes observados se condujeron entre una amplia variedad de taxa de los cuales la mayoría se asociaron con gastos relacionados con el control de las especies y daños a los recursos o pérdidas relacionadas con invasiones. Se identificó un incremento considerable de los costes relacionados con estimaciones previas y a través del tiempo. De manera importante, de las 2,249 especies exóticas y 181 especies invasoras, solo se han reportado costes de 28 especies invasoras en Alemania. Por lo tanto, el total de los costes cuantificados de las especies invasoras que se presentan aquí deben de mostrarse como muy moderados. Se destaca una ausencia de información entre la literatura de acceso libre y las fuentes gubernamentales sobre los costes de las invasiones s a nivel nacional, enmascarando así la alta probabilidad de que existan mayores costes por invasiones en Alemania. Adicionalmente, dado que la tasa de invasión se encuentra en incremento, se espera que los costes económicos sufran un incremento. Se requiere mejorar la evaluación y el reporte de los costes económicos para sentar un adecuado precedente para la mitigación y manejo efectivo de las invasiones en la economía nacional e internacional. Ekonomické náklady na invazivní druhy v Německu. Nepůvodní invazní druhy jsou dobře známou a všudypřítomnou hrozbou pro celosvětovou biodiversitu i blahobyt člověka. Navzdory podstatným dopadům biologických invazí však často postrádáme přehled o jejich nákladovosti pro národní ekonomiky. V důsledku toho byla omezena i adekvátní alokace zdrojů pro management biologických invazí, jelikož nebylo možné vypracovat analýzu nákladů a přínosů takových akcí. Cílem této studie bylo pomocí speciálně vytvořené databáze zhodnotit náklady způsobené invazními druhy v Německu, které je čtvrtou zemí s nejvyšším HDP na světě. Celkově byly ekonomické náklady invazních druhů mezi lety 1960 a 2020 odhadnuty na 9.8 miliardy amerických dolarů. Jednalo se především o potenciální odhadnuté náklady způsobené severoamerickým skokanem volským Lithobates catesbeianus, střemchou pozdní Prunus serotina a dvěma savci – ondatrou pižmovou Ondatra zibethicus a norkem americký Neovison vison. Pozorované náklady byla způsobené různými skupinami organismů a většinou byly spojeny s kontrolou jejich šíření a jimi způsobenými škodami. V průběhu času bylo oproti dřívějším odhadům zaznamenáno značné zvýšení těchto ekonomických škod. V Německu žije 2249 nepůvodních organismů, z nichž je 181 považováno za invazní, avšak pouze u 28 existují ekonomicky vyčíslené škody. Reálná výše těchto škod je tedy očekávatelně podstatně vyšší. Tato zjištění poukazují na značný nedostatek takových údajů v dosažitelných informačních zdrojích, jenž velmi pravděpodobně maskuje existenci podstatně vyšších škod způsobených invazními druhy v Německu. Jelikož počet invazních druhů v zemi roste, lze do budoucna očekávat i růst jimi způsobených škod. Je tedy potřeba zlepšit hodnocení a vykazování ekonomických nákladů souvisejících s invazními druhy, aby bylo možné vytvořit podmínky pro jejich eliminaci a management na národní i mezinárodní úrovni. Coûts économiques des espèces envahissantes en Allemagne. Les espèces exotiques envahissantes sont une menace bien connue et omniprésente pour la biodiversité mondiale et le bien-être humain. Malgré les connaissances sur les impacts substantiels de ces espèces, les synthèses quantitatives des coûts monétaires induits par les invasions sur les économies nationales font souvent défaut. De fait, le rapport coûts-avantages des mesures de gestion des invasions biologiques est souvent difficile à obtenir. Cela a nécessairement des conséquences négatives sur l’allocation adéquate de ressources dédiées à ces mesures et actions destinées à prévenir ou contrôler les espèces exotiques envahissantes. Pour déterminer le coût économique des invasions en Allemagne (pays européen avec le quatrième PIB le plus important au monde), nous avons analysé les données publiées collectées à partir de la première compilation mondiale des coûts économiques des espèces exotiques envahissantes. Dans l'ensemble, les coûts économiques ont été estimés à 9,8 milliards de dollars entre 1960 et 2020, dont 8,9 milliards de dollars de coûts potentiels. Les coûts potentiels étaient principalement liés aux coûts extrapolés du ouaouaron d'Amérique Lithobates catesbeianus, du cerisier noir Prunus serotina et de deux mammifères: le rat musqué Ondatra zibethicus et le vison d'Amérique Neovison vison. Les coûts observés étaient attribuables à un large éventail de taxons et principalement associés aux dépenses liées au contrôle et aux dommages ou pertes affectant les ressources humaines. Nous avons identifié une augmentation considérable des coûts au cours du temps, avec des coûts supérieurs aux estimations réalisées précédemment. Il est important de noter que sur les 2249 espèces exotiques et 181 espèces envahissantes signalées en Allemagne, seules 28 espèces ont des coûts économiques reportés dans la base de données considérée. Par conséquent, les coûts quantifiés ici doivent être interprétés avec prudence. Nos résultats mettent donc en évidence un manque flagrant d'informations dans la littérature librement accessible et les sources gouvernementales sur les coûts des invasions au niveau national, masquant l'existence hautement probable de coûts beaucoup plus élevés en Allemagne. De plus, il ne fait aucun doute que l’augmentation toujours croissante des phénomènes d'invasion biologique sera liée à l’augmentation concomitante des coûts économiques associés. L'évaluation et la communication des coûts économiques doivent nécessairement être améliorées pour contribuer à l’établissement et l’implémentation de mesures de gestion efficaces des invasions aux échelles nationale et internationale.

International audience; Even if there were problems with definitions of nativeness and non-nativeness, it would not at all affect the

Biological invasions are increasing worldwide, damaging ecosystems and socioeconomic sectors. Two decades ago, the “100 of the world’s worst” invasive alien species list was established by the IUCN to improve communications , identifying particularly damaging ‘flagship’ invaders globally (hereafter, worst). Whilst this list has bolstered invader awareness, whether worst species are especially economically damaging and how they compare to other invaders (hereafter, other) remain unknown. Here, we quantify invasion costs using the most comprehensive global database compiling them (InvaCost). We compare these costs between worst and other species against sectorial, taxonomic and regional descriptors, and examine temporal cost trends. Only 60 of the 100 worst species had invasion costs considered as highly reliable and actually observed estimates (median: US$ 43 million). On average, these costs were significantly higher than the 463 other invasive species recorded in InvaCost (median: US$ 0.53 million), although some other species had higher costs than most worst species. Damages to the environment from the worst species dominated, whereas other species largely impacted agriculture. Disproportionately highest worst species costs were incurred in North America, whilst costs were more evenly distributed for other species; animal invasions were always costliest. Proportional management expenditures were low for the other species, and surprisingly, over twice as low for the worst species. Temporally, costs increased more for the worst than other taxa; however, management spending has remained very low for both groups. Nonetheless, since 40 species had no robust and/or reported costs, the “true” cost of “some of the world’s worst” 100 invasive species still remains unknown.

As a critical stage in the life cycle of ant colonies, nest establishment depends on external and internal factors. This study investigates the effect of the number of queens on queen and worker behavior during nest establishment in invasive Argentine ants (Linepitema humile) and native Mediterranean Tapinoma nigerrimum. We set up experimental colonies with the same number of workers but with one or six queens. At different time points, we recorded the positions of queens and workers inside and outside the nest. Our results highlight the influence of the number of queens on the position of queens and workers with between-species differences. Queens of both species entered the nests more quickly when there were six queens. During nest establishment, more workers were inside nests with six queens for both species, with this effect being greater for T. nigerrimum. Once nests were established, fewer workers of both species were engaged in nest maintenance and feeding in nests with six queens, T. nigerrimum had fewer workers engaged in patrolling. These results suggest that the number of queens is a key factor driving queen and worker behavior during and after nest establishment with different species responses.

Within ant communities, the biotic resistance of native species against invasive ones is expected to be rare, because invasive species are often highly dominant competitors. The invasive Argentine ant (Linepithema humile (Mayr)) often demonstrated numerical dominance against its opponents, increased aggressiveness, and ability to quickly recruit to food. The present study aimed to assess the behavioral mechanisms involved in the interspecific competition between L. humile, facing either an invasive species (Lasius neglectus Van Loon, Boomsma and Andr&aacute, sfalvy) or a native dominant species (Lasius niger (Linnaeus)). The resource exploitation by the Argentine ant was investigated during one-hour competitive interactions using 10 dead Drosophila flies as prey. When facing La. niger, L. humile exploratory behavior was strongly inhibited, it brought very few prey resources, and killed few opponents. Conversely, La. neglectus had a low impact on L. humile. Contrarily to expectations, the invasive La. neglectus lacked the ability to hinder L. humile resource exploitation, whereas the native La. niger did. These results suggest that La. niger could impact invasive populations of L. humile by interference competition, perhaps better so than some invasive species. While L. humile has become invasive in Southern Europe, the invasion process could be slowed down in the northern latitudes by such native dominant species.

International audience; Biological invasions have been unambiguously shown to be one of the major global causes of biodiversity loss. Despite the magnitude of this threat and recent scientific advances, this field remains a regular target of criticism – from outright deniers of the threat to scientists questioning the utility of the discipline. This unique situation, combining internal strife and an unaware society, greatly hinders the progress of invasion biology. It is crucial to identify the specificities of this discipline that lead to such difficulties. We outline here 24 specificities and problems of this discipline and categorize them into four groups: understanding, alerting, supporting, and implementing the issues associated with invasive alien species, and we offer solutions to tackle these problems and push the field forward.

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

In our recent Discussion paper, we presented our view that the only real distinction between biological invasions and natural colonisations is the human element. We agree that invasion science is a very important science, not only to better understand the role that human mediation plays for colonisation, but also for many other science fields. We agree with all invasion researchers that the human influence can result in spectacular differences, including in rates of species movement, rates of successful colonisation, the particular species being moved, the biogeography of dispersal pathways and rates of any resulting ecological disturbance and biodiversity loss. Our deep point is that that species dispersed by human-mediation or natural colonisation are all subject to the same basic laws and rules of ecology, identical to many other phenomenon that occur naturally and can be greatly influenced by people. The human dimension is merely a mechanistic distinction, albeit important because it exposes insights about the colonisation process that cannot be seen by the study of natural colonisations alone. We provide 10 hypotheses that can be scientifically tested to determine whether biological invasions and natural colonisations are two separate processes or the same process being influenced by different mechanisms.

More than US$21 billion is spent annually on biodiversity conservation. Despite their importance for preventing or slowing extinctions and preserving biodiversity, conservation interventions are rarely assessed systematically for their global impact. Islands house a disproportionately higher amount of biodiversity compared with mainlands, much of which is highly threatened with extinction. Indeed, island species make up nearly two-thirds of recent extinctions. Islands therefore are critical targets of conservation. We used an extensive literature and database review paired with expert interviews to estimate the global benefits of an increasingly used conservation action to stem biodiversity loss: eradication of invasive mammals on islands. We found 236 native terrestrial insular faunal species (596 populations) that benefitted through positive demographic and/or distributional responses from 251 eradications of invasive mammals on 181 islands. Seven native species (eight populations) were negatively impacted by invasive mammal eradication. Four threatened species had their International Union for the Conservation of Nature (IUCN) Red List extinction-risk categories reduced as a direct result of invasive mammal eradication, and no species moved to a higher extinction-risk category. We predict that 107 highly threatened birds, mammals, and reptiles on the IUCN Red List-6% of all these highly threatened species-likely have benefitted from invasive mammal eradications on islands. Because monitoring of eradication outcomes is sporadic and limited, the impacts of global eradications are likely greater than we report here. Our results highlight the importance of invasive mammal eradication on islands for protecting the world's most imperiled fauna.

Recent global efforts in biodiversity accounting, such as those undertaken through the Convention on Biological Diversity (CBD) and Intergovernmental Science-Policy Platform on Biodiversity and Ecosystem Services (IPBES), are vital if we are to track conservation progress, ensure that we can address the challenges of global change, and develop powerful and scientifically sound indicators. Schlaepfer [1] proposes that we should work toward inventories of biodiversity that account for native and non-native species regardless of species origin and ecological context. We strongly disagree with the approach of combining counts of native and non-native species because this will reduce our capacity to detect the effects of non-native species on native biodiversity with potentially devastating consequences. Compelling and abundant evidence demonstrates that some non-native species can become invasive and produce major ecosystem disruptions and even native species extinction. Unfortunately, we still cannot be certain which non-native species will be the most detrimental (e.g., [2]). Combining native and non-native species together into a single biodiversity index would not only inflate biodiversity estimates and risk promoting the spread of invasive non-native species but would also ignore the fundamental ecological differences between the two groups. The critical differences that should be considered when assessing biodiversity include the following. Fil: Pauchard, Aníbal. Universidad de Concepción; Chile Fil: Meyerson, Laura A.. University of Rhode Island; Estados Unidos Fil: Bacher, Sven. University of Fribourg; Suiza Fil: Blackburn, Tim M.. Colegio Universitario de Londres; Reino Unido. The Zoological Society of London; Reino Unido Fil: Brundu, Giuseppe. University of Sassari; Italia Fil: Cadotte, Marc W.. University of Toronto; Canadá Fil: Courchamp, Franck. Université Paris Sud; Francia Fil: Essl, Franz. Universidad de Viena; Austria Fil: Genovesi, Piero. Institute for Environmental Protection and Research and Chair IUCN SSC Invasive Species Specialist Group; Italia Fil: Haider, Sylvia. Institute of Biology/Geobotany and Botanical Garden; Alemania Fil: Holmes, Nick D.. Island Conservation; Estados Unidos Fil: Hulme, Philip E.. Lincoln University; Nueva Zelanda Fil: Jeschke, Jonathan M.. Leibniz - Institute of Freshwater Ecology and Inland Fisheries; Alemania. Freie Universität Berlin; Alemania Fil: Lockwood, Julie L.. Rutgers University; Estados Unidos Fil: Novoa, Ana. Czech Academy of Sciences; República Checa Fil: Nuñez, Martin Andres. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte. Instituto de Investigaciones en Biodiversidad y Medioambiente. Universidad Nacional del Comahue. Centro Regional Universidad Bariloche. Instituto de Investigaciones en Biodiversidad y Medioambiente; Argentina Fil: Peltzer, Duane A.. Manaaki Whenua Landcare Research; Nueva Zelanda Fil: Pysek, Petr. Charles University; República Checa Fil: Richardson, David M.. Stellenbosch University; Sudáfrica Fil: Simberloff, Daniel. University of Tennessee; Estados Unidos Fil: Smith, Kevin. International Union for Conservation of Nature; Reino Unido Fil: van Wilgen, Brian W.. Stellenbosch University; Sudáfrica Fil: Vilà, Montserrat. Consejo Superior de Investigaciones Científicas. Estación Biológica de Doñana; España Fil: Wilson, John R. U.. Stellenbosch University; Sudáfrica Fil: Winter, Marten. German Centre for Integrative Biodiversity Research; Alemania Fil: Zenni, Rafael D.. University of Lavras; Brasil

The global loss of biodiversity can be attributed to numerous threats. While pioneer studies have investigated their relative importance, the majority of those studies are restricted to specific geographic regions and/or taxonomic groups and only consider a small subset of threats, generally in isolation despite their frequent interaction. Here, we investigated 11 major threats responsible for species decline on islands worldwide. We applied an innovative method of network analyses to disentangle the associations of multiple threats on vertebrates, invertebrates, and plants in 15 insular regions. Biological invasions, wildlife exploitation, and cultivation, either alone or in association, were found to be the three most important drivers of species extinction and decline on islands. Specifically, wildlife exploitation and cultivation are largely associated with the decline of threatened plants and terrestrial vertebrates, whereas biological invasions mostly threaten invertebrates and freshwater fish. Furthermore, biodiversity in the Indian Ocean and near the Asian coasts is mostly affected by wildlife exploitation and cultivation compared to biological invasions in the Pacific and Atlantic insular regions. We highlighted specific associations of threats at different scales, showing that the analysis of each threat in isolation might be inadequate for developing effective conservation policies and managements.

Allee effects have important implications for many aspects of basic and applied ecology. The benefits of aggregation of conspecific individuals are central to Allee effects, which have led to the widely held assumption that social species are more prone to Allee effects. Robust evidence for this assumption, however, remains rare. Furthermore, previous research on Allee effects has failed to adequately address the consequences of the different levels of organisation within social species' populations. Here, we review available evidence of Allee effects and model the role of demographic and behavioural factors that may combine to dampen or strengthen Allee effects in social species. We use examples across various species with contrasting social structure, including carnivores, bats, primates and eusocial insects. Building on this, we provide a conceptual framework that allows for the integration of different Allee effects in social species. Social species are characterised by nested levels of organisation. The benefits of cooperation, measured by mean individual fitness, can be observed at both the population and group levels, giving rise to "population level" and "group level" Allee effects respectively. We also speculate on the possibility of a third level, reporting per capita benefits for different individuals within a group (e.g. castes in social insects). We show that group size heterogeneity and intergroup interactions affect the strength of population-level demographic Allee effects. Populations with higher group size heterogeneity and in which individual social groups cooperate demonstrate the weakest Allee effects and may thus provide an explanation for why extinctions due to Allee effects are rare in social species. More adequately accounting for Allee effects in social species will improve our understanding of the ecological and evolutionary implications of cooperation in social species.

Miconia calvescens D.C. appears in the list ''100 of the world's worst invasive alien species'', devised by the IUCN. It is considered the worst plant pest in Hawaii and French Polynesia. This species has also invaded the rain forest of Australia, New Cale- donia and Sri Lanka, where it is extremely difficult to eradicate. To assess the susceptibility to invasion by M. calvescens in new areas, we investigated the current and future suitable areas for this aggressive invader worldwide. We also assessed the protected areas currently at risk of invasion by considering botanic gardens as a proxy for likelihood of introduc- tion, since most successful invasions by M. calvescens originated from private or public garden escapees. Our results predict that about 7.2 % of total landmass is currently suitable for M. calvescens, with 54.8 % outside the native range including 44.5 % within tropical forests in the southern hemisphere. We identified 91 countries, 400 islands, and up to 364 protected areas with suitable environments outside of M. calvescens native range. By the 2080s, worldwide land suitable for M. calvescens is predicted to be reduced by up to half due to climate change. This decrease is mainly predicted to occur in M. calvescens native ranges as well as in countries where the presence of the species has not yet been reported. In contrast, the invaded range is predicted to slightly decrease, showing an interesting example of a double negative effect of climate change on the distribution of an invader. Our work provides information for land managers and stakeholders that can help to avert the introduction and spread of M. calvescens in their territories. We also emphasize the importance of risk assessments on the living collections of botanic gardens, as a common source of escapees of invasive plants.

virtualspecies is a freely available package for R designed to generate virtual species distributions, a procedure increasingly used in ecology to improve species distribution models. This package combines the existing methodological approaches with the objective of generating virtual species distributions with increased ecological realism. The package includes 1) generating the probability of occurrence of a virtual species from a spatial set of environmental conditions (i.e. environmental suitability), with two different approaches; 2) converting the environmental suitability into presence-absence with a probabilistic approach; 3) introducing dispersal limitations in the realised virtual species distributions and 4) sampling occurrences with different biases in the sampling procedure. The package was designed to be extremely flexible, to allow users to simulate their own defined species-environment relationships, as well as to provide a fine control over every simulation parameter. The package also includes a function to generate random virtual species distributions. We provide a simple example in this paper showing how increasing ecological realism of the virtual species impacts the predictive performance of species distribution models. We expect that this new package will be valuable to researchers willing to test techniques and protocols of species distribution models as well as various biogeographical hypotheses.

James C. Russell,∗† Holly P. Jones,‡ Doug P. Armstrong,§ Franck Courchamp,∗∗ Peter J. Kappes,†† Philip J. Seddon,‡‡ Steffen Oppel,§§ Mark J. Rauzon,∗∗∗ Phil E. Cowan,††† Gerard Rocamora,† Piero Genovesi,‡‡‡ Elsa Bonnaud,∗∗ Bradford S. Keitt,§§§ Nick D. Holmes,§§§ and Bernie R. Tershy∗∗∗∗ ∗School of Biological Sciences, University of Auckland, Private Bag 92019, Auckland, 1142, New Zealand, email j.russell@auckland.ac.nz †Island Biodiversity & Conservation Center, University of Seychelles, P.O. Box 1348, Anse Royale, Republic of Seychelles ‡Department of Biological Sciences and Institute for the Study of the Environment, Northern Illinois University, DeKalb, IL 60115, U.S.A. §Wildlife Ecology Group, Institute of Natural Resources, Massey University, Private Bag 11 222, Palmerston North 4442, New Zealand ∗∗Ecologie Systematique Evolution, Universite Paris-Sud, CNRS, AgroParisTech, Universite Paris-Saclay, 91400 Orsay, France ††Oregon Cooperative Fish and Wildlife Research Unit, Department of Fisheries and Wildlife, Oregon State University, Corvallis, OR 97331, U.S.A. ‡‡Department of Zoology, University of Otago, P.O. Box 56, Dunedin 9054, New Zealand §§RSPB Centre for Conservation Science, Royal Society for the Protection of Birds, The Lodge, Sandy, Bedfordshire SG19 2DL, United Kingdom ∗∗∗Geography Department, Laney College, Oakland, CA 94607, U.S.A. †††Landcare Research, P.O. Box 69040, Lincoln 7640, New Zealand ‡‡‡Institute for Environmental Protection and Research, Via V. Brancati 48, Rome I-00144, Italy §§§Island Conservation, 2161 Delaware Avenue Suite A, Santa Cruz, CA 95060, U.S.A. ∗∗∗∗University of California Santa Cruz, 100 Shaffer Road, Santa Cruz, CA 95060, U.S.A.

International audience; Predictive studies are of paramount importance for biological invasions, one of the biggest threats for biodiversity. To help and better prioritize management strategies, species distribution models (SDMs) are often used to predict the potential invasive range of introduced species. Yet, SDMs have been regularly criticized, due to several strong limitations, such as violating the equilibrium assumption during the invasion process. Unfortunately, validation studies–with independent data–are too scarce to assess the predictive accuracy of SDMs in invasion biology. Yet, biological invasions allow to test SDMs usefulness, by retrospectively assessing whether they would have accurately predicted the latest ranges of invasion. Here, we assess the predictive accuracy of SDMs in predicting the expansion of invasive species. We used temporal occurrence data for the Asian hornet Vespa velutina nigrithorax, a species native to China that is invading Europe with a very fast rate. Specifically, we compared occurrence data from the last stage of invasion (independent validation points) to the climate suitability distribution predicted from models calibrated with data from the early stage of invasion. Despite the invasive species not being at equilibrium yet, the predicted climate suitability of validation points was high. SDMs can thus adequately predict the spread of V. v. nigrithorax, which appears to be—at least partially–climatically driven. In the case of V. v. nigrithorax, SDMs predictive accuracy was slightly but significantly better when models were calibrated with invasive data only, excluding native data. Although more validation studies for other invasion cases are needed to generalize our results, our findings are an important step towards validating the use of SDMs in invasion biology.

SUMMARYAnts are among the worst invasive species, and can have tremendous negative impacts on native biodiversity, agriculture, estates, property and human health. Invasive ants are extremely difficult to control, and thus early detection is essential to prevent ant invasions, in particular through surveillance efforts at ports of entry. This paper assesses the potential distribution of 14 of the worst invasive ant species in France, under current and future climatic conditions. Consensus species distribution models, using five different modelling techniques, three global climate models and two CO2 emission scenarios, indicated that France presented suitable areas for 10/14 species, including five listed on the Invasive Species Specialist Group's selection of the world's 100 worst invasive species. Among these 10 species, eight were predicted to increase their potential range with climate change. Areas with the highest concentration of potential invaders were mainly located along the coastline, especially in the south-west of France, but all departments appeared to be climatically suitable for at least two invasive species. A ranking of climatic suitability per species for 17 major airports and 14 maritime ports indicated that the ports of entry with the highest suitability were located in Biarritz, Toulon and Nice, and the species with the greatest potential distribution in France were Lasius neglectus and Linepithema humile, followed by Solenopsis richteri, Pheidole megacephala and Wasmannia auropunctata.

International audience; Biological invasions are among the greatest threats to global biodiversity, but in contrast to most other global threats, they suffer from specific communication issues. Our paper presents the first new addition to the widely cited IUCN list of ''100 of the world's worst invasive species'', a list created a decade ago in response to these communication issues. We briefly present this list, the recent removal of one species from that list, and the rationale to include a novel, 100th species to replace it. The new species of this list, giant salvinia (Salvinia molesta), was chosen by the community of invasion biologists (over 650 experts from over 60 countries). This new addition to the list will draw public attention to the damage caused by invasive alien species and it will help stimulate the necessary discussion of this critical issue in science and policy circles.

International audience; Biological invasion is increasingly recognized as one of the greatest threats to biodiversity. Using ensemble forecasts from species distribution models to project future suitable areas of the 100 of the world's worst invasive species defined by the International Union for the Conservation of Nature, we show that both climate and land use changes will likely cause drastic species range shifts. Looking at potential spatial aggregation of invasive species, we identify three future hotspots of invasion in Europe, northeastern North America, and Oceania. We also emphasize that some regions could lose a significant number of invasive alien species, creating opportunities for ecosystem restoration. From the list of 100, scenarios of potential range distributions show a consistent shrinking for invasive amphibians and birds, while for aquatic and terrestrial invertebrates distributions are projected to substantially increase in most cases. Given the harmful impacts these invasive species currently have on ecosystems, these species will likely dramatically influence the future of biodiversity.

Asclimaticallysuitablerangeprojectionsbecomeincreasinglyusedtoassessdistributionsofspecies, we recommend systematic assessments of the quality of habitat in addition to the classical binary classification of habitat. We devised a method to assess occurrence probability, captured by a climatic suitability index, through which we could determine variations in the quality of potential habitat. This relative risk assessment circumvents the use of an arbitrary suitability threshold. We illustrated our method with 2 case studies on invasive ant species. We estimated invasion potential of the destroyer ant (Monomorium destructor) and the European fire ant (Myrmica rubra) on a global scale currently and by 2080 with climate change. We found that 21.1% of the world's landmass currently has a suitable climate for the destroyer ant and 16% has a suitable climate for European fire ant. Our climatic suitability index showed that both ant species would benefit from climate change, but in different ways. The size of the potential distribution increased by 35.8% for the destroyer ant. Meanwhile, the total area of potential distribution remained the same for the European fire ant (>0.05%), but the level of climatic suitability within this range increased greatly and led to an improvement in habitat quality (i.e., of invasive species' establishment likelihood). Either through quantity or quality of suitable areas, both invasive ant species are likely to increase the extent of their invasion in the future, following global climate change. Our results show that species may increase their range if either more areas become suitable or if the available areas present improved suitability. Studies in which an arbitrary suitability threshold was used may overlook changes in area quality within climatically suitable areas and as a result reach incorrect predictions.

Insects have presented human society with some of its greatest development challenges by spreading diseases, consuming crops and damaging infrastructure. Despite the massive human and financial toll of invasive insects, cost estimates of their impacts remain sporadic, spatially incomplete and of questionable quality. Here we compile a comprehensive database of economic costs of invasive insects. Taking all reported goods and service estimates, invasive insects cost a minimum of US$70.0 billion per year globally, while associated health costs exceed US$6.9 billion per year. Total costs rise as the number of estimate increases, although many of the worst costs have already been estimated (especially those related to human health). A lack of dedicated studies, especially for reproducible goods and service estimates, implies gross underestimation of global costs. Global warming as a consequence of climate change, rising human population densities and intensifying international trade will allow these costly insects to spread into new areas, but substantial savings could be achieved by increasing surveillance, containment and public awareness., Invasive insects impose many economic costs, for example by consuming crops and spreading disease. Here, Bradshaw et al. compile a database of the costs of invasive insects and conservatively estimate that the yearly global cost (in 2014-equivalent US dollars) is at least $70 billion for goods and services and $6.9 billion for human health.

In conservation science, assessments of trends and priorities for actions often focus on species as the management unit. Studies on species coverage in online media are commonly conducted by using species vernacular names. However, the use of species vernacular names for web-based data search is problematic due to the high risk of mismatches in results. While the use of Latin names may produce more consistent results, it is uncertain whether a search using Latin names will produce unbiased results as compared to vernacular names. We assessed the potential of Latin names to be used as an alternative to vernacular names for the data mining within the field of conservation science. By using Latin and vernacular names, we searched for species from four species groups: diurnal birds of prey, Carnivora, Primates and marine mammals. We assessed the relationship of the results obtained within different online sources, such as Internet pages, newspapers and social media networks. Results indicated that the search results based on Latin and vernacular names were highly correlated, and confirmed that one may be used as an alternative for the other. We also demonstrated the potential of the number of images posted on the Internet to be used as an indication of the public attention towards different species.

Data Deficient species (DD) comprise a significant portion of the total number of species listed within the IUCN Red List. Although they are not classified within one of the threat categories, they may still face high extinction risks. However, due to limited data available to infer their extinction risk reliably, it is unlikely that the assessment of the true status of Data Deficient species would be possible before many species decline to extinction. An appropriate measure to resolve these problems would be to introduce a flag of potentially threatened species within the Data Deficient category [i.e., DD(PT)]. Such a flag would represent a temporary Red List status for listed Data Deficient species that are, based on the available direct evidence and/or indirect indices, likely to be assigned to one of the threat categories, but where current data remains insufficient for a complete classification. The use of such a flag could increase the focus of the scientific community and conservation decision-makers on such species, thus avoiding the risk that necessary conservation measures are implemented too late. As such, establishment of the DD(PT) category as a kind of alarm for priority species could be beneficial.

We argue that human-mediated invasions are part of the spectrum of species movements, not a unique phenomenon, because species self-dispersing into novel environments are subject to the same barriers of survival, reproduction, dispersal and further range expansion as those assisted by people. Species changing their distributions by human-mediated and non-human mediated modes should be of identical scientific interest to invasion ecology and ecology. Distinctions between human-mediated invasions and natural colonisations are very valid for management and policy, but we argue that these are value-laden distinctions and not necessarily an appropriate division for science, which instead should focus on distinctions based on processes and mechanisms. We propose an all-encompassing framework of species range expansion. This does not detract from the importance of invasion biology as a discipline, but instead will help bring together research being conducted on multiple taxa, and by multiple disciplines, including epidemiology, that are often focused on an identical phenomenon: colonisation.

Among invasive species, ants are a particularly prominent group with enormous impacts on native biodiversity and ecosystem functioning. Globalization and on-going climate change are likely to increase the rate of ant invasions in the future, leading to simultaneous introductions of several highly invasive species within the same area. Here, we investigate pairwise interactions among four highly invasive species, Linepithema humile, Lasius neglectus, Pheidole megacephala and Wasmannia auropunctata, at the whole colony level, using a laboratory set-up. Each colony consisted of 300 workers and one queen. The number of surviving workers in the competing colonies was recorded daily over 7 weeks. We modelled the survival of each colony during pairwise colony interactions, using a nonlinear model characterizing the survival dynamics of each colony individually. The least dominant species was P. megacephala, which always went extinct. Interactions among the three other species showed more complex dynamics, rendering the outcome of the interactions less predictable. Overall, W. auropunctata and L. neglectus were the most dominant species. This study shows the importance of scaling up to the colony level in order to gain realism in predicting the outcome of multiple invasions., This paper was supported by Région Ile-de-France (03-2010/GV-DIM ASTREA), ANR (2009 PEXT 010 01) and BiodivERsa Eranet grants, FFII Project.

Agence Nationale de la Recherche (Sextinction project) ANR-2010-BLAN-1717 (RARE project) ANR-2009-PEXT-01001 Fondation de Recherche sur la Biodiversite (VORTEX project) APP-IN-2009-052; The Allee effect is a theoretical model predicting low growth rates and the possible extinction of small populations. Historically, studies of the Allee effect have focused on demography. As a result, underlying processes other than the direct effect of population density on fitness components are not generally taken into account. There has been heated debate about the potential of genetic processes to drive small populations to extinction, but recent studies have shown that such processes clearly impact small populations over short time scales, and some may generate Allee effects. However, as opposed to the ecological Allee effect, which is underpinned by cooperative interactions between individuals, genetically driven Allee effects require a change in genetic structure to link the decline in population size with a decrease in fitness components. We therefore define the genetic Allee effect as a two-step process whereby a decrease in population size leads to a change in population genetic structure and, in turn, to a decrease in individual fitness. We describe potential underlying mechanisms and review the evidence for this original type of component Allee effect, using published examples from both plants and animals. The possibility of considering demogenetic feedback in light of genetic Allee effects clarifies the analysis and interpretation of demographic and genetic processes, and the interplay between them, in small populations.

Invasive rodents occur on over 80% of the world's island groups, invasions are continuing, and rodent impacts on insular wildlife have been well demonstrated. The extent of this problem calls for tools to aid large-scale prioritisation among the many candidate eradication operations. As conservation funds are limited, biologists have responded with prioritisation systems based on financial cost-effec- tiveness. Instead, we claim that long-term conserva- tion gain should be the primary focus when prioritising islands for invasive rodent eradication. This concept is embodied mainly by invasive rodent reinvasion risk, which we categorise as natural or anthropogenic, based on the mechanism of reinvasion and our ability to mitigate the risk. The result is a first-pass triage system that prioritises eradication programmes by their long-term conservation potential, not their immediate value for money. To construct a prioritiza- tion list, we group islands into units for simultaneous eradication, to minimize inter-island reinvasion risk, and then assign weights to levels of unit reinvasion risk and unit conservation value. The choice of parameter weights may depend on capacity for biosecurity action (i.e. reduction in reinvasion risk) and a choice of tertiary filter variables can further discriminate within priority ranks. We illustrate our prioritization framework with a case study on rodents in New Caledonia but explain how our system can be adapted to suit any invasive rodent species or island configuration.

From insects to mammals, many animals engage in behaviours known to follow cyclic patterns over days (e.g. singing, diving or foraging behaviours). Many of them are regulated by external factors, such as light intensity, and are thus associated with sunrise, sunset or zenith. However, these astronomical events do not occur at the same time everyday: they vary with both the time of the year and the latitude. Logically, therefore, behaviour timing should be recorded relative to these events. Yet, in the field, recording the timing of behaviour is much less difficult with a clock, which is often deemed a suitable common proxy. In this paper, we assess the potential methodological problems associated with analyzing behaviours on the basis of clock time rather than with the actual position of the sun. To demonstrate the important difference between these methods of analysis, we first simulated a behaviour set at sunrise and compared the time of occurrence with the two methods. We then used a dataset, based on a long-term monitoring of hunting behaviour of African wild dogs, Lycaon pictus, to reveal how using clock time can result in erroneous assumptions about behaviour. Finally, we investigated the occurrence of sun time records in published field studies. As a majority of them did not take into account the relevance of astronomical events, it is probable that many result in faulty behavioural timings. The model presented can change clock-recorded time into actual deviation from astronomical events to assist current protocols as well as correct the already recorded datasets.

The size and shape of a trophy constitute major determinants of its value. We postulate that the rarity of a species, whatever its causes, also plays a major role in determining its value among hunters. We investigated a role for an Anthropogenic Allee effect in trophy hunting, where human attraction to rarity could lead to an over-exploitative chain reaction that could eventually drive the targeted species to extinction. We performed an inter-specific analysis of trophy prices of 202 ungulate taxa and quantified to what extent morphological characteristics and their rarity accounted for the observed variation in their price. We found that once location and body mass were accounted for, trophies of rare species attain higher prices than those of more common species. By driving trophy price increase, this rarity effect may encourage the exploitation of rare species regardless of their availability, with potentially profound consequences for populations.

A.M. Kramer & J.M. Drake (2010) Experimental demonstration of population extinction due to a predator-driven Allee effect. Journal of Animal Ecology, 79, 633–639. Experimental evidence of extinction via an Allee effect (AE) is a priority as more species become threatened by human activity. Kramer & Drake (2010) begin the International Year of Biodiversity with the important – but double-edged – demonstration that predators can induce an AE in their prey. The good news is that their experiments help bridge the knowledge gap between theoretical and empirical AEs. The bad news is that this predator-driven AE precipitates the prey extinction via a demographic AE. Although their findings will be sensitive to departures from their experimental protocol, this link between predation and population extinction could have important consequences for many prey species.

International audience; A widespread opinion is that conservation efforts disproportionately benefit charismatic species. However, this doesn’t mean that they are not threatened, and which species are “charismatic” remains unclear. Here, we identify the 10 most charismatic animals and show that they are at high risk of imminent extinction in the wild. We also find that the public ignores these animals’ predicament and we suggest it could be due to the observed biased perception of their abundance, based more on their profusion in our culture than on their natural populations. We hypothesize that this biased perception impairs conservation efforts because people are unaware that the animals they cherish face imminent extinction and do not perceive their urgent need for conservation. By freely using the image of rare and threatened species in their product marketing, many companies may participate in creating this biased perception, with unintended detrimental effects on conservation efforts, which should be compensated by channeling part of the associated profits to conservation. According to our hypothesis, this biased perception would be likely to last as long as the massive cultural and commercial presence of charismatic species is not accompanied by adequate information campaigns about the imminent threats they face.

Summary 1. In Caut, Angulo & Courchamp (2008a) rats were fed with experimental diets of distinct isotopic values (d 13 Ca ndd 15 N) in order to infer the discrimination factors. We showed negative relationships between discrimination factors and diet isotopic values. In Caut, Angulo & Courchamp (2009), our aim was to generalize these relationships to other taxonomic groups with a view to providing ecologists with a general and flexible method to obtain discrimination factors for diet reconstruction studies when difficult to obtain otherwise. 2. Perga & Grey (2010) claims that there is an artefact of experimental design (due to protein contents) and partly reanalyzes our data, taking into account the type of diet (mixed or not; carnivorous, omnivorous or herbivorous). However, (i) the role of protein content through isotopic routing is not clear; (ii) a reanalysis of our data shows that the relationships between discrimination factors and diet isotopic values remain significant even when following Perga’s suggestions. 3. Auerswald et al. (2010) criticized the imprecise definitions, mathematical artefacts and invalid statistical analysis of Caut, Angulo & Courchamp (2009). However, (i) their arguments are not new; they challenge the use of stable isotope in ecological trophic studies in general, and lack both alternative suggestions and ecological perspective; (ii) none of their statistical and mathematical objections has a significant effect on the results. 4. Synthesis and applications. Neither reply reflects the current context of debate. The complementarity between experimental studies and their application in field studies is indispensable for the use of stable isotopes in ecology. Protocols should be very well assessed in order to represent field conditions with accuracy. Further analyses are necessary to remove some of the flaws in the use of stable isotopes (carrying out an in-depth analysis of isotopic routing, turnover, and dynamic during trophic experiments to ensure accurate attainment of equilibrium), but none of the issues raised by Perga & Grey (2010) and Auerswald et al. (2010) alter our main conclusions.

In this paper, we review mate-finding Allee effects from ecological and evolutionary points of view. We define ‘mate-finding’ as mate searching in mobile animals, and also as the meeting of gametes for sessile animals and plants (pollination). We consider related issues such as mate quality and choice, sperm limitation and physiological stimulation of reproduction by conspecifics, as well as discussing the role of demographic stochasticity in generating mate-finding Allee effects. We consider the role of component Allee effects due to mate-finding in generating demographic Allee effects (at the population level). Compelling evidence for demographic Allee effects due to mate-finding (as well as via other mechanisms) is still limited, due to difficulties in censusing rare populations or a failure to identify underlying mechanisms, but also because of fitness trade-offs, population spatial structure and metapopulation dynamics, and because the strength of component Allee effects may vary in time and space. Mate-finding Allee effects act on individual fitness and are thus susceptible to change via natural selection. We believe it is useful to distinguish two routes by which evolution can act to mitigate mate-finding Allee effects. The first is evolution of characteristics such as calls, pheromones, hermaphroditism, etc. which make mate-finding more efficient at low density, thus eliminating the Allee effect. Such adaptations are very abundant in the natural world, and may have arisen to avoid Allee effects, although other hypotheses are also possible. The second route is to avoid low density via adaptations such as permanent or periodic aggregation. In this case, the Allee effect is still present, but its effects are avoided. These two strategies may have different consequences in a world where many populations are being artificially reduced to low density: in the first case, population growth rate can be maintained, while in the second case, the mechanism to avoid Allee effects has been destroyed. It is therefore in these latter populations that we predict the greatest evidence for mate-finding Allee effects and associated demographic consequences. This idea is supported by the existing empirical evidence for demographic Allee effects. Given a strong effect that mate-finding appears to have on individual fitness, we support the continuing quest to find connections between component mate-finding Allee effects (individual reproductive fitness) and the demographic consequences. There are many reasons why such studies are difficult, but it is important, particularly given the increasing number of populations and species of conservation concern, that the ecological community understands more about how widespread demographic Allee effects really are, and why.

Summary 1The use of stable isotopic techniques to study animal diets and trophic levels requires a priori estimates of discrimination factors (Δ13C and Δ15N, also called fractionation factors), which are the differences in isotopic composition between an animal and its diet. Previous studies have shown that these parameters depend on several sources of variation (e.g. taxon, environment, tissue) but diet as a source of variation still needs assessment. 2We conducted an extensive review of the literature (66 publications) concerning estimates of animal-diet Δ13C (n = 290) and Δ15N (n = 268). We analysed this data set to test the effect of diet isotopic ratio on the discrimination factor, taking into account taxa, tissues, environments and lipid extraction treatments. Our results showed differences among taxonomic classes for Δ13C, but not for Δ15N, and significant differences among tissues for both Δ13C and Δ15N. We found a significant negative relationship between both, Δ13C and Δ15N, with their corresponding diet isotopic ratios. This relationship was found also within taxonomic classes for mammals (Δ13C and Δ15N), birds (Δ13C), fishes (Δ13C and Δ15N) and invertebrates (Δ13C and Δ15N). From these relationships, we propose a method to calculate discrimination factors based on data on diet isotope ratios (termed the ‘Diet-Dependent Discrimination Factor’, DDDF). 3To investigate current practice in the use of discrimination factors, we reviewed studies that used multi-resource isotopic models. More than 60% of models used a discrimination factor coming from a different species or tissues, and in more than 70% of models, only one Δ13C or Δ15N was used for all resources, even if resources had very different isotopic ratios. Also, we estimated DDDFs for the studies that used isotopic models. More than 40% used Δ15N values and more than 33% used Δ13C values differing > 2‰ from estimated DDDFs. 4Synthesis and applications. Over the last decade, applied ecologists have discovered the potential of stable isotopes for animal diet reconstruction, but the successful adoption of the method relies on a good estimation of discrimination factors. We draw attention to the high variability in discrimination factors, advise caution in the use of single discrimination factors in isotopic models, and point to a method for obtaining adequate values for this parameter when discrimination factors cannot be measured experimentally. Future studies should focus on understanding why discrimination factors vary as a function of the isotopic value of the diet.

Eradications of invasive alien species have generally benefited biodiversity. However, without sufficient planning, successful eradications can have unexpected and unwanted consequences for native species and ecosystems. In particular, the “surprise effect” is the rapid increase of hitherto unnoticed species following the sudden removal of an invasive alien that was exerting an ecological force on those species (predation, competition or herbivory, for example). The only way to prevent these undesired outcomes is to adapt the control programme following the characterization of the trophic relationships between the invasive alien species and the invaded communities, that is, to view the control with a holistic perspective. Here, we illustrate this point with the study of the role of the ship rat (Rattus rattus), which invaded a tropical pacific atoll, Surprise Island, New Caledonia. We assessed the risk of surprise effects during a pre-eradication phase of several years, and then adapted our eradication strategy accordingly.

Summary 1Naturally occurring stable isotopes in resources and their consumer allow the estimation of nutritional flows between the two and have been much used to improve our understanding of the nutritional ecology of free-living animals. 2The difference in isotopic composition between an animal and its diet is represented by a discrimination factor. Carbon and nitrogen flows are estimated by calculating the discrimination factors in stable isotope ratios (δ15N and δ13C), which are presumed to be c. 3‰ and 1‰ heavier in the consumer tissues than those in their resources, respectively. 3The discrimination factor is known to vary according to species, tissue, age, growth rates and food quality, but the estimation of discrimination factors is difficult and a fixed discrimination factor is usually used in diet reconstruction. It has also been suggested that discrimination factors could vary linearly with the diet isotopic ratio. If this linear relationship could be demonstrated using regression, this would provide an adequate method for the estimation of discrimination factors. In order to understand how diet isotopic ratios affect the discrimination factor, we investigated the pattern of its change in nitrogen (Δ15N) and carbon (Δ13C) in different tissues (liver, muscle and hair) of an omnivore species, the rat Rattus rattus. We fed captive rats with diets of the same nutritional quality but on different isotopic ratios. 4First, discrimination factors for Δ15N and Δ13C showed great variability, ranging from –1·46‰ to 4·59‰ and from –8·79‰ to 0·64‰, respectively. Discrimination factors depended on both diet isotopic ratio and tissue. 5We also show that isotope ratios in shaved hairs showed a turnover during the first month, and then stabilized during the second month. Using shaved hairs has the potential to be an effective non-lethal method for determining resource shifts in non-specialist consumers. 6Finally, we demonstrated, for all tissues, a decrease of Δ15N and Δ13C with an increased values of δ15N and δ13C, respectively. These relationships allow us to propose a framework to estimate discrimination factors from diet isotopic ratios by means of regression models.

Introduced vertebrate predators are one of the most important threats to endemic species throughout a range of ecosystems, in particular on islands in biodiversity hot spots. Consequently, the reduction of predator numbers is considered a key conservation action in the management of many native vertebrates vulnerable to predators. It is now established that control attempts may affect non-target species through trophic interactions, but little is known concerning their consequences on competitive relationships. We study a mathematical model mimicking the effects of controlling introduced species in the presence of their competitors. We used two competing rodents to illustrate our study: black rats, Rattus rattus, and mice, Mus musculus. Analyses of the model show that control of only one introduced species logically results in the dramatic increase of the overlooked competitor. We present empirical data that confirm our theoretical predictions. Less intuitively, this process, which we term ‘the competitor release effect’, may also occur when both introduced competitors are simultaneously controlled. In our setting, controlling both predators can promote their coexistence. This occurs as soon as the inferior competitor benefits from the differential effect of the simultaneous control of both competitors, that is, when the indirect positive effect of control (the removal of their competitors) exceeds its direct negative effect (their own removal). Both control levels and target specificity have a direct influence on the extent of this process: counter-intuitively, the stronger and more specific the control, the greater the effect. The theoretical validation of the competitor release effect has important implications in conservation, especially for control management.

Allee effects, strongly related to the extinction vulnerability of populations and gradually becoming acknowledged by both theoretically oriented and applied ecologists, have already been shown to have important roles in the dynamics of many populations. Although not yet widely recognized, two or more Allee effects can occur simultaneously in the same population. Here, we review the evidence for multiple Allee effects and show that their interactions can take several forms, many of which are far from inconsequential. We suggest that more research is needed to assess the prevalence and interactions of multiple Allee effects, as failing to take them into account could have adverse conse- quences for the management of threatened or exploited populations.

International audience; Ants figure prominently among the worst invasive species because of their enormous ecological and economic impacts. However, it remains to be investigated which species would be behaviourally dominant when confronted with another invasive ant species, should two species be introduced in the same area. In the future, many regions might have suitable environmental conditions for several invasive ant species, as predicted under climate change scenarios. Here, we explored interactions among several highly invasive ant species, which have been shown to have overlapping suitable areas. The aim of this study was to evaluate the performance in interference competition of seven of the world's worst invasive ant species (Anoplolepis gracilipes, Paratrechina longicornis, Myrmica rubra, Linepithema humile, Lasius neglectus, Wasmannia auropunctata and Pheidole megacephala). We conducted pairwise confrontations, testing the behaviour of each species against each of the six other species (in total 21 dyadic confrontations). We used single worker confrontations and group interactions of 10 versus 10 individuals to establish a dominance hierarchy among these invasive ant species. We discovered two different behavioural strategies among these invasive ants: three species displayed evasive or indifferent behaviour when individuals or groups were confronted (A. gracilipes, Pa. longicornis, M. rubra), while the four remaining species were highly aggressive during encounters and formed a linear dominance hierarchy. These findings contrast with the widespread view that invasive ants form a homogeneous group of species displaying the `invasive syndrome', which includes generally aggressive behaviour. The dominance hierarchy among the four aggressive species may be used to predict the outcome of future competitive interactions under some circumstances. Yet, the existence of several behavioural strategies renders such a prediction less straightforward.

International audience; Ants are among the most problematic invasive species. They displace numerous native species, alter ecosystem processes, and can have negative impacts on agriculture and human health. In part, their success might stem from a departure from the discovery-dominance trade-off that can promote co-existence in native ant communities, that is, invasive ants are thought to be at the same time behaviorally dominant and faster discoverers of resources, compared to native species. However, it has not yet been tested whether similar asymmetries in behavioral dominance, exploration, and recruitment abilities also exist among invasive species. Here, we establish a dominance hierarchy among four of the most problematic invasive ants (Linepithema humile, Lasius neglectus, Wasmannia auropunctata, Pheidole megacephala) that may be able to arrive and establish in the same areas in the future. To assess behavioral dominance, we used confrontation experiments, testing the aggressiveness in individual and group interactions between all species pairs. In addition, to compare discovery efficiency, we tested the species' capacity to locate a food resource in a maze, and the capacity to recruit nestmates to exploit a food resource. The four species differed greatly in their capacity to discover resources and to recruit nestmates and to dominate the other species. Our results are consistent with a discovery-dominance trade-off. The species that showed the highest level of interspecific aggressiveness and dominance during dyadic interactions.

Valery et al. recently proposed to end the field of Invasion Biology on the grounds that it is based on an inadequate definition of the concept of biological invasion and that, as exotic species, native species should also be called invasive whenever they outbreak. We argue, on the contrary, that the sudden demographic dominance of native species cannot be termed invasion. Moreover, we claim that the suggestion of ending a fruitful and useful discipline because it does not conform to a subjective definition or because it still encompasses some debatable ideas and unresolved questions is both irrelevant and excessive. We believe that the thousands of researchers working in this discipline do not perform nonsensical science, and that their efforts to understand and limit biological invasions are compatible with debating on the key concept of that field.

The IUCN Red List of Threatened Species is often advocated as a tool to assist decision-making in conservation investment and research focus. It is frequently suggested that research efforts should prioritize species in higher threat categories and those that are Data Deficient (DD). We assessed the linkage between IUCN listing and research effort in DD and Critically Endangered (CR) species, two groups generally advocated as research priorities. The analysis of the change in the research output following species classification indicated a listing effect in DD species, while such effect was observed in only a minority of CR species groups. DD species, while chronically understudied, seem to be recognized as research priorities, while research effort for endangered species appears to be driven by various factors other than the IUCN listing. Optimized conservation research focus would require international science planning efforts, harmonized through international mechanisms and promoted by financial and other incentives.

The painted hunting dog or African wild dog, Lycaon pictus, is one of the most endangered large carnivores in Africa, with extinction predicted within a few decades if their dramatic decline is not stopped. It has recently been hypothesized that because of their constraining need for helpers, group size was of major importance in obligate cooperative breeding species, and that the resulting likely existence of a threshold number of adults could create an Allee effect, increasing the group extinction risk. One example where the importance for a critical number of adults may have major repercussions for painted hunting dogs concerns baby-sitting, or pup-guarding, a behavior typical of obligate cooperative breeders. We propose that, as forgoing this behavior is costly because pup guards have the potential to decrease pup mortality, its use is costly too, especially in small packs, because helpers are strongly needed for their cooperative foraging (hunting, protecting the kill and bringing back food to the pups). We present a simple model showing how pup-guarding imposes a cost because it implies that less food per hunt is brought back to more individuals at the den. We complete these analyses with empirical tests of the effect of pack size on the probability of pup-guarding, from field data from the Hwange population in Zimbabwe. Our model, as well as our 5 years of empirical data, both suggest a critical threshold at a size of about five individuals. Key words: African wild dog conservation, Allee effect, cooperative hunting, inverse density dependence, Lycaon pictus, painted hunting dog, pup-guarding. [Behav Ecol 13:20–27 (2002)]

Rabbits are considered by biologists to be among the worst invasive species. Their impact on invaded ecosystems, epitomized in Australia and on islands worldwide (e.g. North, Bullock & Dulloo, 1994; Williams et al., 1995; Burbidge & Moris, 2001; Garzon-Machado et al., 2010), has earned them a well-deserved place in the infamous Invasive Species Specialist Group’s ‘100 of the Worst Invasive Alien Species’ list. Rabbits are capable of profoundly altering communities, ecosystems and landscapes by sapping biodiversity and biomass from the basic trophic levels of often fragile ecosystems. However, they also have more complex, and hitherto seldom documented effects on higher trophic levels (Zedler & Black, 1992; Priddel, Carlile & Wheeler, 2000; Oliver, LuqueLarena & Lambin, 2009; Denyer, Hartley & John, 2010). In the context of a few, but loud voices attempting to question the importance of alien invasive species (but see Lambertini et al., 2011; Lockwood, Hoopes & Marchetti, 2011), the paper by Brodier et al. (2011) is well timed. It reminds us that restoration ecology and the fight against biodiversity loss are possible, in particular on islands, as they generally have many endemic and threatened species. This paper also highlights two very important, yet previously overlooked points. The first is the importance of longterm follow-up after restoration programmes. The second is the need to consider the indirect (and consequently often hidden) effects that alien invasive species can have on seemingly unconnected species. By monitoring the populations, several petrel species and their avian predator (the brown skua, Catharacta skua) following the removal of rabbits, Brodier et al. (2011) show that rabbits compete for space with blue petrels, Halobaena caerulea. Following rabbit eradication, the competition for burrows was removed and the blue petrel population increased up to eightfold in a few generations. Brown skua also increased their survivorship in response to the increase in its main prey, the blue petrel. Interestingly, the population of another petrel, the Antarctic prion, Pachyptila desolata, was also impacted by the rabbits. Following the eradication, the Antarctic prion population declined fourfold (to exclusion) in areas of deep soil and remained stable where the soil was too shallow for blue petrel (and rabbits). The resulting system, illustrated in Fig. 1, shows that one unexpected, and up to now unreported, effect of rabbits was to erode the niche partitioning of nesting seabirds on the island. Removing the rabbits led to both a competitor release (of the blue petrel, freed from the competition with rabbits) and to an apparent competitor release (of the shared predator, the brown skua, boosted by increased blue petrel availability). The combination of direct and indirect interactions restricted Antarctic prions to shallow soil areas, less favoured by the blue petrel. This neat system shows how alien invasive species can have effects beyond the trophic level constituting their resources. The contrast between the spectacular recovery of the blue petrel, the increased survival but stable number of skua breeding pairs and the decrease of the Antarctic prion also illustrates well the difficulties to forecast outcomes that conservation program may face even in relatively simple trophic webs. This is especially so in the current context of global climate change, which already affects subantarctic islands (Chapuis, Frenot & Lebouvier, 2004; Dowding et al., 2009), and which could have played a role in the system, for example, in the delayed recovery of the skua. Monitoring population densities, survival and predator diet, every year, on this remote, far reaching island, for long after the rabbit eradication might have been seen as a risky bet. As it turned out, the bet paid off, in the best currency ever: knowledge and understanding. Deciphering the indirect effects a terrestrial herbivore can have on a community of seabirds with which they have few interactions if any, is crucial in our battle against alien invasive species (Bergstrom et al., 2009; Genovesi, 2011). It demonstrates very well that if we were to ‘judge alien species only on their effects’, we would miss all but the tip of the iceberg and would allow important but unapparent, indirect effects of alien invasive species on native ecosystems. One regret from this study is that the monitoring effort conducted after rabbit eradication was not also implemented Animal Conservation. Print ISSN 1367-9430

Island ecosystems are particularly vulnerable to exotic species. Here we show how an introduced prey has led to the wholesale restructuring of an island food web, including the near extinction of an endemic carnivore. Introduced pigs, by providing abundant food, enabled golden eagles to colonize the California Channel Islands. Eagles preyed heavily on the island fox, whose resulting decline toward extinction released populations of the competitively inferior island skunk. The presence of exotic pigs led to major ecosystem shifts by indirectly causing predation to replace competition as the dominant force shaping these island communities.

Although the massive organized slaughter of African wild dogs, Lycaon pictus, largely ended several decades ago, this endangered canid continues to decline and faces extinction. Several lines of evidence suggest that this arises from obligate cooperative breeding, which makes Lycaon more sensitive to anthropogenic mortality. A number of behaviours in this species are characterized by a reliance on helpers. These include cooperative hunting, defence from kleptoparasitism, pup feeding and baby-sitting. As a result, there are strong, positive relationships between pack size and the production and survival of pups, and pairs of wild dogs are often unsuccessful at raising offspring without the assistance of helpers. Consequently, a pack in which membership drops below a critical size may be caught in a positive feedback loop: poor reproduction and low survival further reduce pack size, culminating in failure of the whole pack. Here, we review the literature to reveal the importance of pack size in the African wild dog. Most importantly, we argue that there is a critical minimum threshold, below which packs face an increasing probability of extinction - an Allee effect with consequences for the conservation of this species, and of other obligate cooperators.

Summary 1. Feral cats Felis catus introduced onto oceanic islands pose a major ecological threat to endemic vertebrates, but their control is difficult. Immunocontraception has not been considered previously as a method for their control or eradication, and therefore we used a modelling approach to assess whether virus-vectored immunocontraception (VVIC) might be effective. 2. We compared the relative efficiency of cat control/eradication using immunocontraception and three different disseminating techniques, i.e. baits, genetically modified viral vectors, or both. We accounted for several forms of dynamic compensation likely to arise in a population with artificially reduced fertility. 3. We conclude that, under the assumptions of our model, immunocontraception can control or eradicate feral cats on oceanic islands. VVIC was found to be a more efficient dissemination technique than baits, but an integrated method involving viral-infected baits was the most likely to lead to eradication. 4. We advocate field trials of this VVIC technique, when available, under island conditions where any risks to non-target fauna would be minimal.