Macaronesia is a biogeographical region comprising five Atlantic Oceanic archipelagos: the Azores, Madeira, Selvagen (Savage Islands), Canaries and Cape Verde. It has strong affinities with the Atlantic coast of the Iberian Peninsula and the north‐western fringes of Africa. This paper re‐evaluates the biogeographical history and relationships of Macaronesia in the light of geological evidence, which suggests that large and high islands may have been continuously available in the region for very much longer than is indicated by the maximum surface area of the oldest current island (27 Ma) – possibly for as long as 60 million years. We review this literature, attempting a sequential reconstruction of Palaeo‐Macaronesia from 60 Ma to the present. We consider the implications of these geological dynamics for our understanding of the history of colonization of the present islands of Macaronesia. We also evaluate the role of these archipelagos as stepping stones and as both repositories of palaeo‐endemic forms and crucibles of neo‐endemic radiations of plant and animal groups. Our principal focus is on the laurel forest communities, long considered impoverished relicts of the Palaeotropical Tethyan flora. This account is therefore contextualized by reference to the long‐term climatic and biogeographical history of Southern Europe and North Africa and by consideration of the implications of changes in land–sea configuration, climate and ocean circulation for Macaronesian biogeography. We go on to provide a synthesis of the more recent history of Macaronesian forests, which has involved a process of impoverishment of the native elements of the biota that has accelerated since human conquest of the islands. We comment briefly on these processes and on the contemporary status and varied conservation opportunities and threats facing these forests across the Macaronesian biogeographical region.
Aims The 50th anniversary of the publication of the seminal book, The Theory of Island Biogeography, by Robert H. MacArthur and Edward O. Wilson, is a timely moment to review and identify key research foci that could advance island biology. Here, we take a collaborative horizon‐scanning approach to identify 50 fundamental questions for the continued development of the field. Location Worldwide. Methods We adapted a well‐established methodology of horizon scanning to identify priority research questions in island biology, and initiated it during the Island Biology 2016 conference held in the Azores. A multidisciplinary working group prepared an initial pool of 187 questions. A series of online surveys was then used to refine a list of the 50 top priority questions. The final shortlist was restricted to questions with a broad conceptual scope, and which should be answerable through achievable research approaches. Results Questions were structured around four broad and partially overlapping island topics, including: (Macro)Ecology and Biogeography, (Macro)Evolution, Community Ecology, and Conservation and Management. These topics were then subdivided according to the following subject areas: global diversity patterns (five questions in total); island ontogeny and past climate change (4); island rules and syndromes (3); island biogeography theory (4); immigration–speciation–extinction dynamics (5); speciation and diversification (4); dispersal and colonization (3); community assembly (6); biotic interactions (2); global change (5); conservation and management policies (5); and invasive alien species (4). Main conclusions Collectively, this cross‐disciplinary set of topics covering the 50 fundamental questions has the potential to stimulate and guide future research in island biology. By covering fields ranging from biogeography, community ecology and evolution to global change, this horizon scan may help to foster the formation of interdisciplinary research networks, enhancing joint efforts to better understand the past, present and future of island biotas.
Although the role that Pleistocene glacial cycles have played in shaping the present biota of oceanic islands world‐wide has long been recognized, their geographical, biogeographical and ecological implications have not yet been fully incorporated within existing biogeographical models. Here we summarize the different types of impacts that glacial cycles may have had on oceanic islands, including cyclic changes in climate, shifts in marine currents and wind regimes and, especially, cycles of sea level change. The latter have affected geographical parameters such as island area, isolation and elevation. They have also influenced the configurations of archipelagos via island fusion and fission, and cycles of seamount emergence and submergence. We hypothesize that these sea level cycles have had significant impacts on the biogeographical processes shaping oceanic island biotas, influencing the rates and patterns of immigration and extinction and hence species richness. Here we provide a first step toward the development of a glacial‐sensitive model of island biogeography, representing the tentative temporal evolution of those biogeographical parameters during the last glacial cycle. From this reasoning we attempt to derive predictions regarding the imprint of sea level cycles on genetic, demographic or biogeographical patterns within remote island biotas.
Aim We report the first analysis of the long‐term ecology of Tenerife, in order to establish a pre‐colonization base‐line and to assess the effect of human activity and the role of climatic variation on vegetation during the Late Holocene. Location A former lake bed in the city of La Laguna (Tenerife, Canary Islands, Spain). Methods A sedimentary sequence of over 2 m was obtained from the former lake bed. Fossil pollen and microfossil charcoal concentrations were analysed. Radiocarbon dating of the sequence indicates that it spans approximately the last 4700 years. The pollen diagram was zoned using optimal splitting within psimpoll 4.25. Results Three pollen zones were differentiated: (1) in Zone L1 (c. 4700–2900 cal. yr bp) a mixed forest was dominated by Quercus, Carpinus, Myrica and Pinus; (2) in Zone L2 (c. 2900–2000 cal. yr bp) the laurel forest taxa increased, while Pinus, Juniperus and Phoenix declined; and (3) Zone L3 (c. 2000–400 cal. yr bp) was characterized by the decline of Carpinus and Quercus and the abundance of laurel forest taxa (e.g. Myrica). Neither Carpinus nor Quercus was hitherto considered to be native to the Canary Islands. Their decline started c. 2000 years ago, coinciding with microfossil charcoal evidence of increased burning and with archaeological evidence for the first human settlement on Tenerife. Main conclusions Between c. 4700 and 2000 cal. yr bp, the composition of the forest in the valley of La Laguna was very different from what it is at present. In particular, Quercus and Carpinus appear to have been significant components, alongside components of the present‐day laurel forest, and the native pine (Pinus canariensis) forest and thermophilous woodland were also more prevalent in the region (but probably not within the lake basin itself) until 3000 cal. yr bp. The subsequent decline of Quercus and Carpinus led to the establishment of the present laurel forest in the region and a shift to more open vegetation types. These changes indicate that the aboriginal inhabitants of the islands, the Guanches, had a far more profound impact on the vegetation of Tenerife than hitherto realized.
Islands are among the last regions on Earth settled and transformed by human activities, and they provide replicated model systems for analysis of how people affect ecological functions. By analyzing 27 representative fossil pollen sequences encompassing the past 5000 years from islands globally, we quantified the rates of vegetation compositional change before and after human arrival. After human arrival, rates of turnover accelerate by a median factor of 11, with faster rates on islands colonized in the past 1500 years than for those colonized earlier. This global anthropogenic acceleration in turnover suggests that islands are on trajectories of continuing change. Strategies for biodiversity conservation and ecosystem restoration must acknowledge the long duration of human impacts and the degree to which ecological changes today differ from prehuman dynamics.
Despite islands contributing only 6.7% of land surface area, they harbor ~20% of the Earth’s biodiversity, but unfortunately also ~50% of the threatened species and 75% of the known extinctions since the European expansion around the globe. Due to their geological and geographic history and characteristics, islands act simultaneously as cradles of evolutionary diversity and museums of formerly widespread lineages—elements that permit islands to achieve an outstanding endemicity. Nevertheless, the majority of these endemic species are inherently vulnerable due to genetic and demographic factors linked with the way islands are colonized. Here, we stress the great variation of islands in their physical geography (area, isolation, altitude, latitude) and history (age, human colonization, human density). We provide examples of some of the most species rich and iconic insular radiations. Next, we analyze the natural vulnerability of the insular biota, linked to genetic and demographic factors as a result of founder events as well as the typically small population sizes of many island species. We note that, whereas evolution toward island syndromes (including size shifts, derived insular woodiness, altered dispersal ability, loss of defense traits, reduction in clutch size) might have improved the ability of species to thrive under natural conditions on islands, it has simultaneously made island biota disproportionately vulnerable to anthropogenic pressures such as habitat loss, overexploitation, invasive species, and climate change. This has led to the documented extinction of at least 800 insular species in the past 500 years, in addition to the many that had already gone extinct following the arrival of first human colonists on islands in prehistoric times. Finally, we summarize current scientific knowledge on the ongoing biodiversity loss on islands worldwide and express our serious concern that the current trajectory will continue to decimate the unique and irreplaceable natural heritage of the world’s islands. We conclude that drastic actions are urgently needed to bend the curve of the alarming rates of island biodiversity loss.
Aim To quantify the influence of past archipelago configuration on present‐day insular biodiversity patterns, and to compare the role of long‐lasting archipelago configurations over the Pleistocene to configurations of short duration such as at the Last Glacial Maximum (LGM) and the present‐day. Location 53 volcanic oceanic islands from 12 archipelagos worldwide—Azores, Canary Islands, Cook Islands, Galápagos, Gulf of Guinea, Hawaii, Madeira, Mascarenes, Pitcairn, Revillagigedo, Samoan Islands and Tristan da Cunha. Time period The last 800 kyr, representing the nine most recent glacial–interglacial cycles. Major taxa studied Land snails and angiosperms. Methods Species richness data for land snails and angiosperms were compiled from existing literature and species checklists. We reconstructed archipelago configurations at the following sea levels: the present‐day high interglacial sea level, the intermediate sea levels that are representative of the Pleistocene and the low sea levels of the LGM. We fitted two alternative linear mixed models for each archipelago configuration using the number of single‐island endemic, multiple‐island endemic and (non‐endemic) native species as a response. Model performance was assessed based on the goodness‐of‐fit of the full model, the variance explained by archipelago configuration and model parsimony. Results Single‐island endemic richness in both taxonomic groups was best explained by intermediate palaeo‐configuration (positively by area change, and negatively by palaeo‐connectedness), whereas non‐endemic native species richness was poorly explained by palaeo‐configuration. Single‐island endemic richness was better explained by intermediate archipelago configurations than by the archipelago configurations of the LGM or present‐day. Main conclusions Archipelago configurations at intermediate sea levels—which are representative of the Pleistocene—have left a stronger imprint on single‐island endemic richness patterns on volcanic oceanic islands than extreme archipelago configurations that persisted for only a few thousand years (such as the LGM). In understanding ecological and evolutionary dynamics of insular biota it is essential to consider longer‐lasting environmental conditions, rather than extreme situations alone.
Summary1. Garajonay National Park in La Gomera (Canary Islands) contains one of the largest remnant areas of a forest formation once widespread throughout Europe and North Africa. Here, we aim to address the long-term dynamics (the last 9600 cal. years) of the monteverde forest (laurel forest and Morella-Erica heath) located close to the summit of the National Park (1487 m a.s.l.) and determine past environmental and human impacts. 2. We used palaeoecological (fossil pollen, microscopic and macroscopic charcoal) and multivariate ecological techniques to identify compositional change in the monteverde forest in relation to potential climatic and human influences, based on the analysis of a core site at 1250-m elevation. 3. The regional mid-Holocene change towards drier conditions was matched in this system by a fairly rapid shift in representation of key forest elements, with declines in Canarian palm tree (Phoenix canariensis), Canarian willow (Salix canariensis) and certain laurel forest taxa and an increase in representation of the Morella-Erica woody heath. 4. Charcoal data suggest that humans arrived on the island between about 3000 and 1800 years ago, a period of minimal vegetation change. Levels of burning over the last 800 years are among the lowest of the entire 9600 years. 5. Synthesis. A rapid climatic-induced shift of forest taxa occurred 5500 years ago, with a decrease in hygrophilous species in the pollen record. In contrast, we found no evidence of a significant response to human colonization. These findings support the idea that Garajonay National Park is protecting a truly ancient relict, comprising a largely natural rather than cultural legacy.
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