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For Peer Review 19 15 4Department of Biology, University of Missouri at St Global Ecology and Biogeography New Directions in Isla nd Biogeography Journal: Global Ecology and Biogeography ManuscriptFor ID GEB-2016-0004.R1 Peer Review Manuscript Type: Research Reviews Date Submitted by the Author: n/a Complete List of Authors: Santos, Ana; Museo Nacional de Ciencias Naturales (CSIC), Department of Biogeography & Global Change; Universidade dos Açores , Centre for Ecology, Evolution and Environmental Changes (cE3c)/Azorean Biodiversity Group Field, Richard; University of Nottingham, School of Geography; Ricklefs, Robert; University of Missouri-St,. Louis, Biology Climatic niche, evolutionary processes, General Dynamic Model, Invasive species, marine environments, Natural laboratories, Species-area Keywords: relationship, species interactions, Equilibrium Theory of Island Biogeography, Community Assembly Page 9 of 61 Global Ecology and Biogeography 1 2 3 1 Manuscript type: Research Review 4 2 5 3 New Directions in Island Biogeography 6 4 7 1,2, 3 4 8 5 Ana M. C. Santos *, Richard Field & Robert E. Ricklefs 9 6 10 7 1 Department of Biogeography & Global Change, Museo Nacional de Ciencias Naturales 11 8 (CSIC), C/ José Gutiérrez Abascal 2, 28006 Madrid, Spain. Email: 12 9 [email protected] 13 2 14 10 Centre for Ecology, Evolution and Environmental Changes (cE3c)/Azorean Biodiversity 15 11 Group and Universidade dos Açores – Departamento de Ciências Agrárias, 9700-042 Angra 16 12 do Heroísmo, Açores, Portugal 17 13 3School of Geography, University of Nottingham, NG7 2RD, UK. Email: 18 14 [email protected] Peer Review 19 15 4Department of Biology, University of Missouri at St. Louis, One University Boulevard, St. 20 21 16 Louis, MO 63121 USA. Email: [email protected] 22 17 23 18 *Author for correspondence 24 19 25 20 Keywords : community assembly, climatic niche, Equilibrium Theory of Island Biogeography, 26 27 21 evolutionary processes, general dynamic model, invasive species, marine environments, 28 22 natural laboratories, species–area relationship, species interactions. 29 23 30 24 Running title : New Directions in Island Biogeography 31 25 32 26 Number of words in abstract: 301 33 34 27 35 28 Number of words in main text (Introduction to end of Conclusion): 6397 words. 36 29 37 30 Number of Tables: 1 38 31 39 32 40 Number of Boxes: 1 41 33 42 34 Number of in-paper Appendices: 1 (data sources) 43 35 44 36 Supplementary material: 1 table 45 37 46 47 38 48 39 Number of references: 142 49 40 50 51 52 53 54 55 56 57 58 59 60 Global Ecology and Biogeography Page 10 of 61 1 2 3 41 Abstract 4 42 5 43 Aim: Much of our current understanding of ecological and evolutionary processes comes 6 7 44 from island research. With increasing availability of data on distributions and phylogenetic 8 45 relationships, and new analytical approaches to understanding the processes that shape 9 46 species’ distributions and interactions, a prospective on this ever-interesting topic is timely. 10 47 11 48 Location: Islands globally. 12 49 13 14 50 Methods: We start by arguing that the reasons island research has achieved so much in the 15 51 past also apply to the future. We then critically assess the current state of island 16 52 biogeography, focusing on recent changes in emphasis, including research featured in this 17 53 special issue of Global Ecology and Biogeography . Finally, we suggest promising themes for 18 54 the future. We coverFor both ecological Peer and evolutiona Reviewry topics, although the greater emphasis 19 20 55 on island ecology reflects our own backgrounds and interests. 21 56 22 57 Results: Much ecological theory has been directly or indirectly influenced by research on 23 58 island biotas. Currently, island biogeography is renascent, with research focusing on, among 24 59 other things, patterns and processes underlying species interaction networks, species 25 60 coexistence and the assembly of island communities through ecological and evolutionary 26 27 61 time. Continuing island research should provide additional insight into biological invasions 28 62 and other impacts of human activities, functional diversity and ecosystem functioning, 29 63 extinction and diversification, species pools and more. Deeper understanding of the 30 64 similarities and differences between island and mainland systems will aid transferability of 31 65 island theory to continental regions. 32 33 66 34 67 Main conclusions: As research in biogeography and related fields expands in new directions, 35 68 islands continue to provide opportunities for developing insights, both as natural 36 69 laboratories for ecology and evolution and because of the exceptions islands often present 37 70 to the usual ‘rules’ of ecology. New data-collection initiatives are needed on islands 38 71 worldwide and should be directed towards filling gaps in our knowledge of within-island 39 40 72 distributions of species, as well as island species’ functional traits and phylogenetic 41 73 relationships. 42 74 43 44 75 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Page 11 of 61 Global Ecology and Biogeography 1 2 3 76 Introduction 4 77 5 78 “In islands we have the facts of distribution often presented to us in their simplest forms, 6 7 79 along with others which become gradually more and more complex; and we are therefore 8 80 able to proceed step by step in the solution of the problems they present.” 9 81 (Wallace, 1880, p. 234) 10 82 11 83 The nature of island biology research 12 84 Wallace (1880) argued that islands offer solutions to key ‘problems’ in biogeography, 13 14 85 ecology, and evolutionary biology. But do biologists still believe this? Or has island 15 86 biogeography become peripheral to the mainstream, pursued by a few enthusiasts mindful 16 87 of early seminal contributions of island studies to these fields? Do islands continue to 17 88 provide key insights and unique settings for research on important biogeographical, 18 89 ecological, and evolutionaryFor questions? Peer We argue Review here that the answer to the last question 19 90 is emphatically “yes”. While island-specific knowledge and understanding continue to grow, 20 21 91 island biogeography is more integrated with mainstream research than ever before, and this 22 92 integration will intensify over the coming decades. 23 93 Current themes in ecology, biogeography and evolutionary biology include the causes 24 94 of variation in species richness, the assembly of ecological communities, mechanisms 25 95 shaping the outcomes of evolutionary processes (including adaptive radiations), and the 26 27 96 impact of global change on biodiversity (e.g. Sutherland et al. , 2013). These issues are 28 97 complex, involving scale-dependent processes, including species interactions, dispersal and 29 98 colonization, gene flow and population dynamics, all played out in a heterogeneous world. 30 99 Addressing such broad and complex issues will require the resolution of underlying 31 100 mechanisms into straightforward hypotheses amenable to testing by experimental or 32 101 comparative approaches. Islands can be powerful allies in such endeavours. 33 34 102 Island systems often present ecological, evolutionary and geological processes on 35 103 timescales (usually so different among these processes) that are relatively similar, and often 36 104 quite recent. On volcanic oceanic islands (Box 1), in particular, geological dynamics may be 37 105 unusually fast and ecological dynamics relatively slow, aligning both quite closely with each 38 106 other and with evolutionary dynamics (Rominger et al. , this issue). Spatial scales tend to be 39 107 40 compressed (e.g., by steep terrain) and spatial structure within archipelagos is discrete; local 41 108 extinctions on oceanic islands are often global extinctions; and population differentiation 42 109 between and within islands provides model systems for studying speciation. These features 43 110 of islands provide a valuable testing ground for theory because they break typical scaling 44 111 relationships that constrain the expression of many key biological processes within larger 45 112 continental regions (Warren et al. , 2015). 46 47 113 The importance of islands thus goes far beyond their shorelines and, ever since the 48 114 seminal works of the great nineteenth century naturalists, much of the understanding 49 115 gained from island-based studies has informed studies of mainland systems, as illustrated by 50 116 many of the references highlighted in Tables 1 and S1. Indeed, during the 19 th century, the 51 117 work of Charles Darwin in the Galápagos Archipelago and of Alfred Russel Wallace in the 52 118 Malay Archipelago, secured island biology in the foundation of contemporary research 53 54 119 across most biological disciplines. Much of our current understanding of the ecological and 55 120 evolutionary processes that shape diversity patterns comes from knowledge gained from 56 121 islands. As Robert MacArthur and Edward O. Wilson (1967: 3) stated, “insularity is … a 57 122 universal feature of biogeography … many of the principles graphically displayed in the 58 123 Galápagos Islands … apply in lesser or greater degree to all natural habitats.” Some discrete 59 60 Global Ecology and Biogeography Page 12 of 61 1 2 3 124 habitats have been studied in the context of island theory, from mountaintops (or ‘sky 4 125 islands’) to forest fragments and lakes (e.g. Brown, 1971, 1978; March & Bass, 1995), and 5 126 island theory has provided the conceptual basis for much research on the ecological impacts 6 127 of habitat fragmentation (Laurance, 2009). Island-like marine environments (e.g., marine 7 8 128 lakes, i.e. pieces of seawater entirely surrounded by land; seamounts) also can be integrated 9 129 within the general theories of island biogeography, according to Dawson (this issue). More 10 130 generally, insularity within the contiguous landmasses of the continents remains under- 11 131 appreciated (but see Steinbauer et al., in press b), and the same may apply to the oceans.
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