PhD Paola Anna Movalli

Guy Duke

Gloria Ramello

Rene Dekker

Al Vrezec

Richard F Shore

Antonio Garcia-Fernandez

Chris Wernham

Oliver Krone

Rene Dekker

Nikiforos Alygizakis

Alexander Badry

Fausto Barbagli

Koos Biesmeijer

Giovanni Boano

Alexander L Bond

Yael Choresh

Jan Bolding Christensen

Alessandra Cincinelli

Sara Danielsson

Andreia Dias

Rune Dietz

Marcel Eens

Silvia Espin

Igor Eulaers

Sylke Frahnert

Georgios Gkotsis

Natalia Glowacka

Pilar Gomez-Ramirez

Marco Grotti

Michel Guiraud

Peter Hosner

Ulf Johansson

Veerle L B Jaspers

Pepijn Kamminga

Jan Koschorreck

Burkhard Knopf

Eero Kubin

Sabrina Lobrutto

Rui Lourenco

Tania Martellini

Emma Martinez-Lopez

Rafael Mateo

Maria-Christina Nika

Varvara Nikolopoulou

Dan Osborn

Olivier Pauwels

Marco Pavia

M Gloria Pereira

Heinz Ruedel

Pablo Sanchez-Virosta

Jaroslav Slobodnik

Christian Sonne

Nikolaos Thomaidis

Fuiz Tibor

Till Toepfer

Gabriele Treu

Risto Vainola

Jari Valkama

Steven Van Der Mije

Didier Vangeluwe

Ben H Warren

René Dekker

Antonio García-Fernández

Silvia Espín

Tibor I Fuiz

Pilar Gómez-Ramírez

Correction to: Progress on bringing together raptor collections in Europe for contaminant research and monitoring in relation to chemicals regulation

Keywords: raptor, apex predator, contaminant, biomonitoring, natural history museum, environmental specimen bank, collections, REACH, chemicals regulation, chemicals management, Europe

à la diffusion de documents scientifiques de niveau recherche, publiés ou non, émanant des établissements d'enseignement et de recherche français ou étrangers, des laboratoires publics ou privés.

 (Trasande et al 2015). Although there are more than 60,000 chemicals marketed in Europe, so far only 600 chemicals have been screened and identified as PBT (persistent, bioaccumulative, toxic), ED (endocrine disrupting) and/or CMR (carcinogenic, mutagenic, toxic for reproduction) (Gkotsis et al 2019). A suite of EU chemicals legislation (EC 2001, EC 2004a, EC 2004b, EC 2004c, EC 2006, EC 2008, EC 2009a, EC 2009b, EC 2012) aims to reduce chemical risks to humans and wildlife, and the EU's Seventh Environment Action Programme (EC 2014) calls for better scientific knowledge towards achieving a non-toxic environment. Movalli et al (2017) noted the value of raptors as sentinels of environmental health in general, and highlighted in particular the potential value of raptor tissue collections, which can be analysed for contaminant exposure to inform risk assessment of chemicals, assess the effectiveness of chemicals risk management measures, and provide early warning of emerging contaminant problems. They argued for development of a framework bringing together raptor tissue collections in Europe -including those of natural history museums (NHMs), environmental specimen banks (ESBs) and others (e.g. in research institutions) -in support of contaminant research and monitoring for better chemicals management. They suggested this framework would involve among other things identifying and digitising relevant collections and developing a searchable database of these collections to make them more visible and accessible to ecotoxicologists and competent authorities.

Here, we outline progress that has been made in this regard through the launch of two mutually reinforcing initiatives, the COST Action European Raptor Biomonitoring Facility' (ERBFacility 41 ), which launched in October 2017and in which 27 COST member countries are participating, and the EU LIFE project Systematic use of contaminant data from apex predators and their prey in chemicals management (LIFE APEX 42 ), which launched in September 2018.

Using raptors as particularly appropriate sentinels for PBT compounds (Movalli et ERBFacility has since concluded a review of raptor collections in Europe (Ramello et al in prep). The review, which collated data from over a hundred collections across Europe, revealed that many thousands of raptor specimens arrive annually at these collections, offering a very substantial resource of raptor tissue samples for contaminant monitoring. In the UK, specimens are processed and tissues stored by the Predatory Bird Monitoring Scheme maintained by the Centre for Ecology and Hydrology (Walker et al 2008). In Sweden the lead is taken by the national ESB at the Naturhistorika Rijkmuseet. A number of research collections also exist around Europe, such as that of the Leibniz-Institute for Zoo and Wildlife Research, Germany. However, such specimen banks and research collections, set up specifically for the purposes of contaminant monitoring, are the exception rather than the rule. In most countries, the main recipients of raptor specimens are NHMs. Indeed, the review found that around 75% of responding NHMs across Europe receive fresh raptor/owl specimens. Most NHMs freeze raptor carcasses on arrival and then process and freeze wet tissues; indeed, around a third of responding NHMs are already active in storing frozen raptor tissue samples suitable for contaminant monitoring. 41 At a recent ERBFacility meeting convened at the Royal Belgian Institute of Natural Sciences in Brussels, collections came together to validate the review of raptor collections and consider how to build on this towards development of the European Raptor Specimen Bank. The meeting found that there is strong interest among collections to work together in this regard. However, many collections, particularly in eastern and southern Europe, are stretched for resources, so any new 'ask' (e.g. in terms of expanding tissue collections and/or strengthening standard operating procedures to ensure sample quality) must be tightly specified. Moreover, collections cannot be expected to build a substantial raptor tissue bank unless there is a clear regulatory demand for contaminant data and resources for necessary analyses to be performed.

There exists here something of a Catch 22, in that regulatory demand is likely to grow only once the ability of raptor biomonitoring to provide pan-European assessments pertinent to chemicals management is proven, while proving this ability requires a substantial sample bank to be available in the first place. Overcoming this will probably involve carefully targeted proof of concept work on one or two selected species and tissue matrices for which a substantial pan-European bank of samples already exists, and for one or two priority substances of particular interest to regulators.

There are clear incentives for collections to engage in this work. These include: the opportunity to engage in a major European initiative with key regulatory applications, helping to demonstrate the societal relevance of collections; the opportunity presented by novel collaboration between NHMs, ESBs and research collections; the opportunity to expand research interests into ecotoxicology, and to expand publications output; and the opportunity to know of and exchange samples with other NHMs and thereby broaden individual collections.

Taking forward the concept of a distributed ERSpeB will involve further work under the auspices of the COST Action on a number of issues, building on relevant existing networks such as the Consortium of European Taxonomic Facilities and the network of European Bird Curators. These include: prioritising species, tissue matrices and compounds and carrying out proof of concept work; addressing legal, personnel and freezer constraints to expanding raptor tissue sample collections and shipping samples for analysis; adapting NHM standards and protocols (learning from ESBs) for the gathering, processing and storage of samples, to enhance sample quality for contaminant analysis; developing appropriate policy on access to samples to ensure valuable samples required for contaminant monitoring are not consumed for other purposes; circumscribing the data content of the European samples database and how it will be developed, populated and updated to provide real-time information on raptor tissue samples across Europe. These issues will all be addressed by a sequence of scientific missions and meetings planned by ERBFacility for the next 12-15 months. This paper focuses on work being done in the 'collections arena' under the auspices of ERBFacility, but it is worth noting that ERBFacility also involves work in the 'analytical arena', involving ecotoxicologists, veterinary scientists and chemists, to develop a prioritized European Raptor Biomonitoring Scheme (ERBiomS), and work in the 'field arena' involving field ornithologists and ecologists, to develop a European Raptor Sampling Programme (ERSamP). Advancing work towards a comprehensive European Raptor Biomonitoring Facility, capable of delivering pan-European contaminant data for chemicals management, involves collaboration across these three arenas.

While this work is progressing under ERBFacility, LIFE APEX is working to demonstrate the systematic use of contaminant data from raptors and other apex predators for chemicals management. Like ERBFacility, LIFE APEX is building collaborations between collections and analytical laboratories. But while the focus of ERBFacility is on building networks across the collections, analysis and field arenas and developing the overall Facility, the focus of LIFE APEX is on demonstrating that contaminant data from apex predators has useful regulatory applications, notably in relation to the REACH and Biocides regulations [START_REF] Alygizakis | Determination of emerging substances in top predators and their prey using state-of-the-art chemical monitoring methods[END_REF][START_REF] Alygizakis | Determination of emerging substances in top predators and their prey using state-of-the-art chemical monitoring methods[END_REF][START_REF] Alygizakis | Determination of emerging substances in top predators and their prey using state-of-the-art chemical monitoring methods[END_REF]. LIFE APEX will in particular demonstrate the use of wide-scope target screening (of more than 2,400 contaminants of emerging concern -CECs), wide-scope suspect screening (of more than 40,000 CECs) and non-target screening of samples from raptors (in the terrestrial environment) and other apex predators (in the freshwater and marine environments) to help the European Chemicals Agency (ECHA) prioritise substances (among the many tens of thousands currently used) for more strenuous PBT assessments and identify predominant chemical mixtures in biota. LIFE APEX will also demonstrate the power of targeted analyses of raptor matrices to assess the effectiveness of EU chemicals risk management measures. Proof of concept work on this latter demonstrator is under way by the Naturalis Biodiversity Center with the UK Centre for Ecology and Hydrology (which hosts the UK Predatory Bird Monitoring Scheme).

Taken together, the growing collaborations across the collections, analysis and field arenas under ERBFacility and LIFE APEX offer considerable promise to enhance early warning of emerging contaminant problems, for more timely identification and management of persistent and bio-accumulative substances in the environment, for putting risk management measures in place more quickly, for better understanding predominant chemical mixtures in the environment, and to ascertain whether risk management measures (and by extension chemicals regulations) are actually delivering on their purpose in reducing negative impacts on human and wildlife health. This will complement ongoing work to monitoring impacts of contaminants on human health. 44 Life APEX is financed by the European Union through the program LIFE17 ENV/SK/000355 "Systematic use of contaminant data from apex predators and their prey in chemicals management".

  Europe in support of contaminant monitoring for better chemicals management. We outline recent progress in this regard under the COST Action 'European Raptor Biomonitoring Facility' (ERBFacility). A review of 116 collections reveals that many thousands of raptor specimens arrive annually. In most countries, the main recipients are natural history museums, many of which store frozen samples suitable for contaminant monitoring. There is strong interest among collections to collaborate and enhance this tissue resource through development of a 'distributed European Raptor Specimen Bank'. Developing the Bank will involve: prioritising species, tissue matrices and compounds; proof of concept work; addressing legal and other constraints; enhancing standards and protocols for gathering, processing and storing samples; developing policy on access to samples; and developing a samples database. The Bank will be integrated with a complementary Biomonitoring Scheme (specifying priority contaminant analyses) and Sampling Programme (enhancing specimen supply). The resulting European Raptor Biomonitoring Facility will be capable of delivering pan-European contaminant data for chemicals management.Kamminga 1 , Jan Koschorreck 10 , Burkhard Knopf 28 , Eero Kubin 29 , Sabrina LoBrutto 30 , Rui Lourenco 31 , Tania Martellini 32 , Emma Martínez-López 6 , Rafael Mateo 33 , Maria-Christina Nika 22 , Varvara Nikolopoulou 22 , Dan Osborn 34 , Olivier Pauwels 35 , Marco Pavia, M. Glória Pereira 5 , Heinz Rüdel 36 , Pablo Sanchez-Virosta 6 , Jaroslav Slobodnik 9 , Christian Sonne 18 , Nikolaos Thomaidis 22 , Till Töpfer 37 , Gabriele Treu 10 , Risto Väinölä 38 , Jari Valkama 38 , Steven Van der Mije 1 , Didier Vangeluwe 35 , Ben H. Warren 39 and Friederike Woog 40

			Friederike Woog
		Order of Authors Secondary Information:
	1 2	Funding Information:	European Cooperation in Science and Technology (CA16224)	Not applicable
	3 4		EU LIFE Programme (LIFE17/ENV/SK/000355)	Not applicable
	5 32 31 30 6 7 8 9 10 11 12 13 14 15 17 18 19 20 21 22 23 24 25 26 27 28 29 16	Abstract: tissue collections in Suggested Reviewers: Movalli et al (2017) argued for development of a framework bringing together raptor Jose Tarazona Responsible for biomonitoring of PPPs at EFSA jose.tarazona@efsa.europa.eu European Food Safety Authority a Corresponding author: paola.movalli@naturalis.nl Environmental contaminants impose multi-billion Euro costs on human and wildlife health in Europe
	33		Keith Bildstein
	34		Hawk Mountain Sanctuary
	35 36		bildstein@hawkmountain.org Was involved in previous EURAPMON network
	37 38 39 40 41 42		Peter Korytar European Commission peter.korytar@ec.europa.eu Responsible for EC's IPChem platform for data on environmental contaminants -has attended ERBFacility meeting.
	43	Opposed Reviewers:
	44 45	Additional Information:
	46	Question	Response
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Simultaneously, the project LIFE APEX is demonstrating systematic use of contaminant data from raptors (and other apex predators) for chemicals management. Applications include prioritization of substances for PBT assessments, identifying predominant chemical mixtures in biota, and assessing the effectiveness of EU chemicals risk management measures. Together, ERBFacility and LIFE APEX offer considerable promise to reduce contaminant risks to human and wildlife health in Europe.

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  al 2018), ERBFacility will help answer what the environmental risks are for specific chemicals, whether there are emerging contaminant problems needing remedial action and whether legislation is at all effective in reducing environmental exposure to contaminants in Europe. ERBFacility builds on ten years of work involving ecotoxicologists and ornithologists leading up to and under the European Science Foundation's research networking programme EURAPMON (Research and Monitoring for and with Raptors in Europe) (see, for example, Movalli et al 2008, Gómez-Ramírez et al 2015, Espín et al 2016, Derlink et al 2018, Duke et al 2018). 43 At its first General Meeting in Ciudad Real, Spain in February 2018, ERBFacility set in motion work to bring together NHMs, ESBs and other research collections towards development of a European Raptor Specimen Bank (ERSpeB).

  www.erbfacility.eu 42 www.lifeapex.eu 43 www.eurapmon.eu

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  EU 2012. Regulation (EU) No 528/2012 of the European Parliament and of the Council of 22 May 2012 concerning the making available on the market and use of biocidal products. EU (2014) General Union Environment Action Programme to 2020. Living well, within the limits of our planet. Luxembourg: Publications Office of the European Union. doi:10.2779/66315 Gkotsis G, Alygizakis N, Cincinelli A, Dekker R, Duke G, Glowacka N, Knopf B, Koschorreck J, Martellini T, Movalli P, Nika MC, Nikolopoulou V, Ruedel H, Shore R, Thomaidis NS, Treu G. Slobodnik J (2019) Determination of emerging contaminants in apex predators and their prey from European Specimen Banks and Natural History Museums by High Resolution Mass Spectrometry Techniques. 17th International Conference on Chemistry and the Environment, which will be held in Thessaloniki, 16-20 June 2019. Gómez-Ramírez, P., Shore, R. F., Van Den Brink, N. W., Van Hattum, B., Bustnes, J. O., Duke, G., ... & Krone, O. (2014). An overview of existing raptor contaminant monitoring activities in Europe. Environ Internat 67, 12-21. Koschorreck J, Alygizakis N, Cincinelli A, Dekker R, Duke G, Glowacka N, Knopf B, Martellini T, Movalli P, Nika MC, Ruedel H, Shore R, Thomaidis NS, Treu G. Slobodnik J. (2019) LIFE APEX -Systematic use of contaminant data from apex predators and their prey in chemicals management. SETAC Europe 29th Annual Meeting in Helsinki, Finland (26-30 May, 2019) Movalli P, Dekker R, Koschorreck J, Treu G (2017) Bringing together raptor collections in Europe for contaminant research and monitoring in relation to chemicals regulations. Environ. Sci. Pollut. Res. DOI: 10.1007/s11356-017-0096-s Movalli P, Krone O, Osborn D, Pain D (2018) Monitoring contaminants, emerging diseases and environmental change with raptors and links to human health. Bird Study 64:sup1, 596-5109. Ramello G, Dekker R, Duke G, van der Mije S, Movalli P. (in prep.) A review of European Raptor Tissue Collections. Trasande L, Zoeller, RT, Hass U, Kortenkamp A, Grandjean P, Myers JP, DiGangi J, Bellanger M, Hauser R, Legler J, Skakkebaek NE, Heindel J (2015) Estimating burden and disease costs of exposure to endocrine-disrupting chemicals in Europe. J Clin Endocrinol Metab 100(4): 1245-1255. Walker, LA, Shore RF, Turk A, Pereira MG, Best J (2008) The Predatory Bird Monitoring Scheme (PBMS) -identifying chemical risks to top predators in Britain. Ambio 37, 466-471.

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Fraunhofer Institute for Molecular Biology and Applied Ecology, 57392 Schmallenberg, Germany

Acknowledgments

European Raptor Biomonitoring Facility COST Action (CA16224) is supported by COST (European Cooperation in Science and Technology) and funded by the Horizon 2020 Framework Programme of the European Union.


D Gravel

N Mouquet

B H Warren

R E Ricklefs

How Consideration of Islands Has Inspired Mainstream Ecology: Links

  L'archive ouverte pluridisciplinaire

related ideas [START_REF] Macarthur | The theory of island biogeography[END_REF]. We were involved in one such initiative, a working group on island biogeography funded by the Centre for the Synthesis and Analysis of Biodiversity (CESAB). A major output of the group was a collective perspective on how key attributes of islands surrounded by water provide prospects for improving our understanding of the ecology and evolution of biological communities in general [START_REF] Warren | Islands as model systems in ecology and evolution: prospects fifty years after MacArthur-Wilson[END_REF] see Box 1 for a detailed distinction of IBT and Core IBT). In the process of organising our ideas, interesting issues arose that were tangential to this aim. One example is that over the decades since 1967, IBT has inspired numerous ecological theories and schools of thought. Although IBT is focussed on islands surrounded by water, even in the opening chapter of the 1967 monograph, MacArthur and Wilson emphasized that such island biomes are an extreme example of fragmented or insular environments (isolated by geography or ecology) that are common worldwide. Consistent with this view, not only has IBT proven relevant to a wide variety of insular environments that are not islands surrounded by water (e.g., [START_REF] Brown | Mammals on mountaintops: nonequilibrium insular biogeography[END_REF][START_REF] Drake | The biology of insularity: an introduction[END_REF][START_REF] Wagner | Cichlid species-area relationships are shaped by adaptive radiations that scale with area[END_REF], but it also has links to important ecological theories and concepts, the implications of which stretch far beyond such islands.

Here we highlight some of the main links between IBT and subsequent developments in ecology and conservation biology.

The elegance of Core IBT stands in its ability to model complex phenomena with few parameters. Diversity dynamics on an island incorporate a simple birth and death process in which births represent species arriving through immigration and deaths represent local (i.e., island-wide) extinctions; Core IBT predicts that diversity will tend towards a "dynamic equilibrium" in which these processes are balanced. These ideas predict considerable variation in local community composition due to the stochasticity of immigration and local extinction, and the time taken to reach equilibrium (see also Warren et al., 2015, Box 6, for related situations with unattained equilibria). Nonetheless, Core IBT provides a highly deterministic perspective on species richness based on the assumption that the rates of immigration and extinction are determined by the geographic context: near islands have a higher rate of immigration than far islands, and small islands have a higher rate of extinction than large islands. Accordingly, other things being equal, diversity should be highest on islands that are large and close to the mainland, and lowest on islands that are small and far.

The "island biome" (islands surrounded by water) is an obvious choice for studying the effects of area and isolation on species richness since insularity is acute; an island's boundaries frequently provide longstanding limits to the distributions of species or populations (see [START_REF] Warren | Islands as model systems in ecology and evolution: prospects fifty years after MacArthur-Wilson[END_REF] for further discussion, including key attributes of islands). Nonetheless, in addition to the predominance of other insular and fragmented environments worldwide, small habitat islands are among those most easily studied and manipulated. Such practical constraints are reflected in the range of systems in which the equilibrium predictions of Core IBT have been tested [START_REF] Schoener | The MacArthur-Wilson Equilibrium Model: a chronicle of what it said and how it was tested[END_REF], which include fragmented crops and intertidal habitats, as well as islands surrounded by water. Application of Core IBT to insular systems that are not "true islands" arguably reached an extreme in the design of protected areas, such as the debate over whether a single large, or several small, reserves is better for ensuring the persistence of species ("SLoSS" for short). Problems with this approach include the fact that Core IBT does not generate clear predictions in this matter [START_REF] Simberloff | Refuge design and island biogeographic theory: effects of fragmentation[END_REF], and also that for habitat islands, unlike "true islands", an organism's ability to inhabit, or disperse through, the intervening matrix is a critical factor. In the end, ecologists concluded that Core IBT provides only limited practical lessons for conservation managers [START_REF] Soulé | What do genetics and ecology tell us about the design of nature reserves[END_REF]. Nonetheless, by forcing decision makers to think about population turnover within habitat fragments, as well as the influence of fragment size and isolation on colonization and extinction, a highly diversified field of fragmentation research can be traced back to Core IBT [START_REF] Laurence | Beyond island biogeography theory: understanding habitat fragmentation in the real world[END_REF].

A more direct example of the relevance to IBT of insular systems that are not "true islands" comes from an extension to the Core IBT model to improve its ability to predict species diversity. At low isolation, the frequent arrival of propagules protects insular populations from extinction. This "rescue effect" was first demonstrated empirically for arthropods on isolated plants (thistles; [START_REF] Brown | Turnover rates in insular biogeography: effect of immigration and extinction[END_REF]. Other examples of modifications to Core IBT include adding the effect of island habitat heterogeneity on colonization and extinction dynamics (e.g., [START_REF] Johnson | Ecological parameters and plant species diversity[END_REF][START_REF] Whitehead | Small islands and equilibrium theory of insular biogeography[END_REF][START_REF] Kadmon | Integrating the effects of area, isolation, and habitat heterogeneity on species diversity: A unification of island biogeography and niche theory[END_REF], as well as the effects of changes in the physical geography of islands over time [START_REF] Whittaker | A general dynamic theory of oceanic island biogeography[END_REF]. Recent developments have extended this legacy by showing how, without adding much complexity to the original model, species interactions could easily be incorporated with important consequences for the shape of the species-area relationship [START_REF] Gravel | Trophic theory of island biogeography[END_REF] or even for the number of trophic levels expected in fragmented systems [START_REF] Holt | Trophic rank and the species-area relationship[END_REF][START_REF] Calcagno | Constraints on food chain length arising from regional metacommunity dynamics[END_REF]. For example, predator-prey interactions may slow down species accumulation with area, while mutualistic interactions may accelerate it.

Realising such multi-trophic extensions to Core IBT was key in the development of recent theories of community assembly and co-distribution in isolated habitats [START_REF] Cazelles | On the integration of biotic interaction and environmental constraints at the biogeographical scale[END_REF][START_REF] Cazelles | A theory for species co-occurrence in interaction networks[END_REF]. Furthermore, some of the main parameters of Core IBT have sometimes been modified to provide new interpretations, as in [START_REF] Wright | Species-energy theory: an extension of species-area theory[END_REF] replacing area with available energy to develop a species-energy theory.

Another important extension of IBT was to consider the dynamics of populations on multiple islands (metapopulations) connected by movements of individuals. Although IBT and metapopulation theory [START_REF] Levins | Some demographic and genetic consequences of environmental heterogeneity for biological control[END_REF][START_REF] Levins | Extinction[END_REF] were not formally connected initially, their relationship can be traced back to an agreement among their authors to work on complementary questions, MacArthur and Wilson's being "how many species does an island contain?" and Levins' being "how many islands does a species occupy?" (R. Levins, pers. comm., 2013). This initial plan included the authors coming together later for a study of continental biogeography, although this was never realised (R. Levins, pers. comm., 2013).

Current thinking in metapopulation ecology does fit the Core ITB perspective perfectly once changes in scale and geography are incorporated [START_REF] Hanski | The theories of island biogeography and metapopulation dynamics: science marches forward, but the legacy of good ideas lasts for a long time[END_REF]. Core IBT becomes a metapopulation model when one excludes the mainland with its permanent source populations of each species and considers migration among multiple islands (e.g., [START_REF] Mouquet | Coexistence in metacommunities: the regional similarity hypothesis[END_REF]. Although [START_REF] Levins | Some demographic and genetic consequences of environmental heterogeneity for biological control[END_REF][START_REF] Levins | Extinction[END_REF] original model focussed on a single species, and "islands" (habitat patches) that did not differ in size or accessibility, numerous later developments have relaxed these simplifying assumptions [START_REF] Hanski | Metapopulation ecology[END_REF][START_REF] Hanski | The theories of island biogeography and metapopulation dynamics: science marches forward, but the legacy of good ideas lasts for a long time[END_REF]. Furthermore, it is clear that much heritage of both IBT and metapopulation theory has percolated into today's study of spatial ecology from metacommunities (Leibold et al., 2004;[START_REF] Leibold | From metapopulations to metacommunities[END_REF][START_REF] Logue | Empirical approaches to metacommunities: a review and comparison with theory[END_REF][START_REF] Massol | Linking community and ecosystem dynamics through spatial ecology[END_REF] to metaecosystems [START_REF] Loreau | Meta-ecosystem: a framework for a spatial ecosystem ecology[END_REF][START_REF] Gravel | Trophic theory of island biogeography[END_REF][START_REF] Massol | Linking community and ecosystem dynamics through spatial ecology[END_REF][START_REF] Leibold | Metacommunity Ecology[END_REF].

Neutral biodiversity theory [START_REF] Bell | Neutral macroecology[END_REF][START_REF] Hubbell | The unified neutral theory of biodiversity and biogeography[END_REF] is perhaps the most striking example of an unexpected legacy of IBT. MacArthur and Wilson apparently did not intend for IBT to be thought of as "neutral" at the species level (that is, species treated as ecologically equivalent). The concave functional forms for immigration and extinction rates in Core IBT were partially motivated by potential ecological differences between species (differences in propensity for immigration and extinction; [START_REF] Macarthur | An equilibrium theory of insular zoogeography[END_REF] for further discussion see [START_REF] Gilpin | On the concavity of island biogeographic rate functions[END_REF][START_REF] Schoener | The MacArthur-Wilson Equilibrium Model: a chronicle of what it said and how it was tested[END_REF]. From a strictly technical perspective, however, species in the classic IBT model are not labelled with their ecological identities.

Rather, they are all treated as identical, and the model is thus neutral at the species level regardless of the ideas that motivated it.

The better-known form of neutral theory is one in which individuals (not species) are ecologically equivalent [START_REF] Hubbell | Tree dispersion, abundance, and diversity in a tropical dry forest[END_REF][START_REF] Hubbell | A unified theory of biogeography and relative species abundance and its application to tropical rain forests and coral reefs[END_REF], and even the earliest work on such models [START_REF] Caswell | Community structure: a neutral model analysis[END_REF]) notes the similarities with IBT. Later development of "The Unified Neutral Theory of Biodiversity and Biogeography" [START_REF] Hubbell | The unified neutral theory of biodiversity and biogeography[END_REF] caused a wave of interest in neutral models and further added to the legacy of MacArthur and Wilson as an inspiration for them. In particular, the unified neutral theory incorporates an explicit distinction between the local community and metacommunity, with both maintained at dynamic equilibria.

Individual-based neutral theory has since accurately predicted a range of ecological data both in mainstream ecology and in the more specific domain of island biogeography [START_REF] Bell | Neutral macroecology[END_REF][START_REF] Chave | Neutral theory and community ecology[END_REF][START_REF] Alonso | The merits of neutral theory[END_REF]. Although neutral theory is also faced with contrary data (e.g., [START_REF] Ricklefs | A comment on Hubbell's zero-sum ecological drift model[END_REF][START_REF] Ricklefs | The unified neutral theory of biodiversity: do the numbers add up?[END_REF][START_REF] Mcgill | Empirical evaluation of neutral theory[END_REF], it is widely viewed as a useful model to compare with empirical data, highlighting key roles for dispersal limitation, speciation, and ecological drift [START_REF] Jabot | Analyzing tropical forest tree species abundance distributions using a nonneutral model and through Approximate Bayesian Inference[END_REF][START_REF] Rosindell | The unified neutral theory of biodiversity and biogeography at age ten[END_REF][START_REF] Rosindell | The case for ecological neutral theory[END_REF].

While the influence of IBT in ecology is predominantly linked to Core IBT, it is important to note that the legacy also extends to other elements of the 1967 monograph. For example, Chapter 7 predicts evolutionary changes following island colonization. [START_REF] Macarthur | The theory of island biogeography[END_REF] reasoned that, upon first arriving on an island, a species' population density is much lower relative to the carrying capacity of the environment than it was at its mainland source. They argued that selection on such newly-arriving immigrants should initially favour adaptations that produce high rates of population growth, even if resources are used inefficiently. They envisaged a contrasting situation for species in the mainland source for which population density is already at the environmental limit; in such source populations, selection should favour the ability to survive and reproduce with limited resources (i.e., efficient resource use). Borrowing notation from standard population models, they coined the terms r-selection and K-selection for these alternative regimes, respectively. They also argued that once a colonist attains its maximum population size on an island, it will tend to experience a switch back towards a K-selection regime (and indeed one that is stronger than on the mainland, due to a temporally more stable climate). [START_REF] Macarthur | The theory of island biogeography[END_REF] ideas were further developed by [START_REF] Pianka | On r-and K-selection[END_REF] to make explicit predictions for how individual life-history traits would evolve in response to r-and K-selection, forming the basis of what became known as life-history strategies. However, the r-and K-selection paradigm has since been criticised on numerous grounds, in particular that it has been applied in the absence of data on population dynamics, oversimplifies the process of natural selection, and is unable to make precise predictions about phenotypic change (see [START_REF] Reznick | r-and K-selection revisited: the role of population regulation in life-history evolution[END_REF] for a review, and more recent related paradigms). Despite such shortcomings, the r-and Kselection paradigm has been useful in defining fitness under density-dependent population regulation [START_REF] Roff | The evolution of life histories: theory and analysis[END_REF], and retains considerable popularity (e.g., [START_REF] Saether | Evidence for r-and K-selection in a wild bird population: a reciprocal link between ecology and evolution[END_REF]. Furthermore, consistent with the low species diversity of small and remote islands predicted by Core IBT, such islands provide particularly striking examples of rapid change in selection gradients following the arrival of new immigrants (e.g., [START_REF] Losos | Predator-induced behaviour shifts and natural selection in field-experimental lizard populations[END_REF][START_REF] Losos | Rapid temporal reversal in predator-driven natural selection[END_REF].

IBT is a major component of the foundation of modern ecology. Even though our working group included participants with a range of profiles, including island evolutionary biologists and ecological theoreticians, the number of important links with other ecological theories and schools of thought took many of us by surprise. From an ecological perspective, IBT has left a legacy that has (i) forced ecologists to connect local processes to regional patterns, and (ii) enabled theoretical ecologists to propose simple models that capture a significant portion of ecological complexity on a biogeographic scale.