Long-term trajectories of non-native vegetation on islands globally.

Human-mediated changes in island vegetation are, among others, largely caused by the introduction and establishment of non-native species. However, data on past changes in non-native plant species abundance that predate historical documentation and censuses are scarce. Islands are among the few places where we can track human arrival in natural systems allowing us to reveal changes in vegetation dynamics with the arrival of non-native species. We matched fossil pollen data with botanical status information (native, non-native), and quantified the timing, trajectories and magnitude of non-native plant vegetational change on 29 islands over the past 5000 years. We recorded a proportional increase in pollen of non-native plant taxa within the last 1000 years. Individual island trajectories are context-dependent and linked to island settlement histories. Our data show that non-native plant introductions have a longer and more dynamic history than is generally recognized, with critical implications for biodiversity baselines and invasion biology.

The human dimension of biodiversity changes on islands.

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.

Resilience: nitrogen limitation, mycorrhiza and long-term palaeoecological plant-nutrient dynamics.

Ecosystem dynamics are driven by both biotic and abiotic processes, and perturbations can push ecosystems into novel dynamical regimes. Plant-plant, plant-soil and mycorrhizal associations all affect plant ecosystem dynamics; however, the direction and magnitude of these effects vary by context and their contribution to ecosystem resilience over long time periods remains unknown. Here, using a mathematical framework, we investigate the effects of plant feedbacks and mycorrhiza on plant-nutrient interactions. We show evidence for strong nutrient controlled feedbacks, moderation by mycorrhiza and influence on ecological resilience. We use this model to investigate the resilience of a longitudinal palaeoecological birch-δ15N interaction to plant-soil feedbacks and mycorrhizal associations. The birch-δ15N system demonstrated high levels of resilience. Mycorrhiza were predicted to increase resilience by supporting plant-nitrogen uptake and immobilizing excess nitrogen; in contrast, long-term enrichment in available nitrogen by plant-soil feedbacks is expected to decrease ecological resilience.

Plant controls on Late Quaternary whole ecosystem structure and function.

Plants and animals influence biomass production and nutrient cycling in terrestrial ecosystems; however, their relative importance remains unclear. We assessed the extent to which mega-herbivore species controlled plant community composition and nutrient cycling, relative to other factors during and after the Late Quaternary extinction event in Britain and Ireland, when two-thirds of the region's mega-herbivore species went extinct. Warmer temperatures, plant-soil and plant-plant interactions, and reduced burning contributed to the expansion of woody plants and declining nitrogen availability in our five study ecosystems. Shrub biomass was consistently one of the strongest predictors of ecosystem change, equalling or exceeding the effects of other biotic and abiotic factors. In contrast, there was relatively little evidence for mega-herbivore control on plant community composition and nitrogen availability. The ability of plants to determine the fate of terrestrial ecosystems during periods of global environmental change may therefore be greater than previously thought.

Prescribed fire and its impacts on ecosystem services in the UK.

The impacts of vegetation fires on ecosystems are complex and varied affecting a range of important ecosystem services. Fire has the potential to affect the physicochemical and ecological status of water systems, alter several aspects of the carbon cycle (e.g. above- and below-ground carbon storage) and trigger changes in vegetation type and structure. Globally, fire is an essential part of land management in fire-prone regions in, e.g. Australia, the USA and some Mediterranean countries to mitigate the likelihood of catastrophic wildfires and sustain healthy ecosystems. In the less-fire prone UK, fire has a long history of usage in management for enhancing the productivity of heather, red grouse and sheep. This distinctly different socioeconomic tradition of burning underlies some of the controversy in recent decades in the UK around the use of fire. Negative public opinion and opposition from popular media have highlighted concerns around the detrimental impacts burning can have on the health and diversity of upland habitats. It is evident there are many gaps in the current knowledge around the environmental impacts of prescribed burning in less fire-prone regions (e.g. UK). Land owners and managers require a greater level of certainty on the advantages and disadvantages of prescribed burning in comparison to other techniques to better inform management practices. This paper addresses this gap by providing a critical review of published work and future research directions related to the impacts of prescribed fire on three key aspects of ecosystem services: (i) water quality, (ii) carbon dynamics and (iii) habitat composition and structure (biodiversity). Its overall aims are to provide guidance based on the current state-of-the-art for researchers, land owners, managers and policy makers on the potential effects of the use of burning and to inform the wider debate about the place of fire in modern conservation and land management in humid temperate ecosystems.

Island biodiversity conservation needs palaeoecology.

The discovery and colonization of islands by humans has invariably resulted in their widespread ecological transformation. The small and isolated populations of many island taxa, and their evolution in the absence of humans and their introduced taxa, mean that they are particularly vulnerable to human activities. Consequently, even the most degraded islands are a focus for restoration, eradication, and monitoring programmes to protect the remaining endemic and/or relict populations. Here, we build a framework that incorporates an assessment of the degree of change from multiple baseline reference periods using long-term ecological data. The use of multiple reference points may provide information on both the variability of natural systems and responses to successive waves of cultural transformation of island ecosystems, involving, for example, the alteration of fire and grazing regimes and the introduction of non-native species. We provide exemplification of how such approaches can provide valuable information for biodiversity conservation managers of island ecosystems.

The ecological consequences of megafaunal loss: giant tortoises and wetland biodiversity.

The giant tortoises of the Galápagos have become greatly depleted since European discovery of the islands in the 16th Century, with populations declining from an estimated 250 000 to between 8000 and 14 000 in the 1970s. Successful tortoise conservation efforts have focused on species recovery, but ecosystem conservation and restoration requires a better understanding of the wider ecological consequences of this drastic reduction in the archipelago's only large native herbivore. We report the first evidence from palaeoecological records of coprophilous fungal spores of the formerly more extensive geographical range of giant tortoises in the highlands of Santa Cruz Island. Upland tortoise populations on Santa Cruz declined 500-700 years ago, likely the result of human impact or possible climatic change. Former freshwater wetlands, a now limited habitat-type, were found to have converted to Sphagnum bogs concomitant with tortoise loss, subsequently leading to the decline of several now-rare or extinct plant species.

Ecosystem resilience and threshold response in the Galápagos coastal zone.

BACKGROUND: The Intergovernmental Panel on Climate Change (IPCC) provides a conservative estimate on rates of sea-level rise of 3.8 mm yr(-1) at the end of the 21(st) century, which may have a detrimental effect on ecologically important mangrove ecosystems. Understanding factors influencing the long-term resilience of these communities is critical but poorly understood. We investigate ecological resilience in a coastal mangrove community from the Galápagos Islands over the last 2700 years using three research questions: What are the 'fast and slow' processes operating in the coastal zone? Is there evidence for a threshold response? How can the past inform us about the resilience of the modern system? METHODOLOGY/PRINCIPAL FINDINGS: Palaeoecological methods (AMS radiocarbon dating, stable carbon isotopes (δ(13)C)) were used to reconstruct sedimentation rates and ecological change over the past 2,700 years at Diablas lagoon, Isabela, Galápagos. Bulk geochemical analysis was also used to determine local environmental changes, and salinity was reconstructed using a diatom transfer function. Changes in relative sea level (RSL) were estimated using a glacio-isostatic adjustment model. Non-linear behaviour was observed in the Diablas mangrove ecosystem as it responded to increased salinities following exposure to tidal inundations. A negative feedback was observed which enabled the mangrove canopy to accrete vertically, but disturbances may have opened up the canopy and contributed to an erosion of resilience over time. A combination of drier climatic conditions and a slight fall in RSL then resulted in a threshold response, from a mangrove community to a microbial mat. CONCLUSIONS/SIGNIFICANCE: Palaeoecological records can provide important information on the nature of non-linear behaviour by identifying thresholds within ecological systems, and in outlining responses to 'fast' and 'slow' environmental change between alternative stable states. This study highlights the need to incorporate a long-term ecological perspective when designing strategies for maximizing coastal resilience.

When is an invasive not an invasive? Macrofossil evidence of doubtful native plant species in the Galápagos Islands.

The Galápagos Islands are globally renowned for their ecological value and as a world symbol of scientific discovery; however the native biodiversity of this unique region is currently under threat. One of the primary concerns is the detrimental impact of approximately 750 nonnative plants introduced over the last 500 years of human presence in the archipelago. In addition to these known introduced species, there are an additional 62 vascular plants classified as "doubtful natives," where native status remains unclear. To help address the questions of provenance regarding these doubtfully native species and their impact on highland ecosystems over the past 500-1000 years, we analyzed plant macrofossils in sedimentary records. Appropriate species classification (native or introduced) was determined using baseline data of species presence on the islands. We confirmed that six plants (Ageratum conyzoides, Solanum americanum, Ranunculus flagelliformis, Brickellia diffusa, Galium canescens, and Anthephora hermaphrodita) once considered doubtful natives or introduced are actually native to the Galápagos flora. These results have relevance not just for the Galápagos but also many other oceanic islands in demonstrating the application of palaeobotanical data to conserving and restoring native biodiversity.

DIATOMS (BACILLARIOPHYTA) OF ISOLATED ISLANDS: NEW TAXA IN THE GENUS NAVICULA SENSU STRICTO FROM THE GALáPAGOS ISLANDS(1).

The diatoms (Bacillariophyta) from a coastal lagoon from the Diablas wetlands (Isla Isabela, the Galápagos Islands) were studied in material from surface samples and a sediment core spanning the past 2,700 years in order to examine evidence of diatom evolution under geographic isolation. The total number of taxa found was ∼100. Ultrastructural variation in valve morphology between members of Galápagos taxa was used to describe 10 species from the genus Navicula sensu stricto, which are new to science. Four taxa: N. isabelensis, N. isabelensoides, N. isabelensiformis, and N. isabelensiminor, shared several key characteristics that may be indicative of a common evolutionary heritage; these species therefore provide possible evidence for the in situ evolution of diatoms in the Galápagos coastal lagoons. Shared morphological characteristics include: (i) stria patterning in the central area, (ii) an elevated and thickened external raphe-sternum, (iii) external central raphe endings that are slightly deflected toward the valve primary side, and (iv) an arched valve surface. To explain these findings, two models were proposed. The first suggested limited lateral diatomaceous transport of Navicula species between the Galápagos and continental South America. Alternatively, these new species may be ecological specialists arising from the unique environmental conditions of the Galápagos coastal lagoons, which restrict the colonization of common diatom taxa and enable the establishment of novel, rare species. The Diablas wetlands are an important site for diatom research, where local-scale environmental changes have combined with global-scale biogeographic processes resulting in unique diatom assemblages.

Fossil pollen as a guide to conservation in the Galapagos.

Paleoecological evidence from the past 8000 years in the Galápagos Islands shows that six presumed introduced or doubtfully native species (Ageratum conyzoides, Borreria laevis/Diodia radula-type, Brickellia diffusa, Cuphea carthagenensis, Hibiscus diversifolius, and Ranunculus flagelliformis) are in fact native to the archipelago. Fossil pollen and macrofossils from four sites in the highlands of Santa Cruz Island reveal that all were present thousands of years before the advent of human impact, refuting their classification as introduced species. These findings have substantial implications not only for conservation in Galápagos but for the management of introduced species and pantropical weeds in general.

How can a knowledge of the past help to conserve the future? Biodiversity conservation and the relevance of long-term ecological studies.

This paper evaluates how long-term records could and should be utilized in conservation policy and practice. Traditionally, there has been an extremely limited use of long-term ecological records (greater than 50 years) in biodiversity conservation. There are a number of reasons why such records tend to be discounted, including a perception of poor scale of resolution in both time and space, and the lack of accessibility of long temporal records to non-specialists. Probably more important, however, is the perception that even if suitable temporal records are available, their roles are purely descriptive, simply demonstrating what has occurred before in Earth's history, and are of little use in the actual practice of conservation. This paper asks why this is the case and whether there is a place for the temporal record in conservation management. Key conservation initiatives related to extinctions, identification of regions of greatest diversity/threat, climate change and biological invasions are addressed. Examples of how a temporal record can add information that is of direct practicable applicability to these issues are highlighted. These include (i) the identification of species at the end of their evolutionary lifespan and therefore most at risk from extinction, (ii) the setting of realistic goals and targets for conservation 'hotspots', and (iii) the identification of various management tools for the maintenance/restoration of a desired biological state. For climate change conservation strategies, the use of long-term ecological records in testing the predictive power of species envelope models is highlighted, along with the potential of fossil records to examine the impact of sea-level rise. It is also argued that a long-term perspective is essential for the management of biological invasions, not least in determining when an invasive is not an invasive. The paper concludes that often inclusion of a long-term ecological perspective can provide a more scientifically defensible basis for conservation decisions than the one based only on contemporary records. The pivotal issue of this paper is not whether long-term records are of interest to conservation biologists, but how they can actually be utilized in conservation practice and policy.