One hundred questions of importance to the conservation of global biological diversity.

We identified 100 scientific questions that, if answered, would have the greatest impact on conservation practice and policy. Representatives from 21 international organizations, regional sections and working groups of the Society for Conservation Biology, and 12 academics, from all continents except Antarctica, compiled 2291 questions of relevance to conservation of biological diversity worldwide. The questions were gathered from 761 individuals through workshops, email requests, and discussions. Voting by email to short-list questions, followed by a 2-day workshop, was used to derive the final list of 100 questions. Most of the final questions were derived through a process of modification and combination as the workshop progressed. The questions are divided into 12 sections: ecosystem functions and services, climate change, technological change, protected areas, ecosystem management and restoration, terrestrial ecosystems, marine ecosystems, freshwater ecosystems, species management, organizational systems and processes, societal context and change, and impacts of conservation interventions. We anticipate that these questions will help identify new directions for researchers and assist funders in directing funds.

The dispersal of vascular plants in a forest mosaic by a guild of mammalian herbivores.

Endozochorous seed dispersal by herbivores can affect plant spatial dynamics and macroecological patterns. We have investigated the number and species composition of viable seeds deposited in faeces of a full guild of macroherbivores (four deer and two lagomorph species) in a forest in eastern Britain. One hundred and one plant species germinated from faecal pellet material, 85 of which were among the 247 vascular plant species recorded in the forest. However, three species - Chenopodium album, Urtica dioica and Agrostis stolonifera - comprised 56% of the seedlings recorded. Of the species recorded in faecal samples, 36% had no recognised dispersal mechanism, while very few (7%) were adapted to endozoochorous dispersal (fleshy fruit or nut). The number of species dispersed by the herbivores was ranked Cervus elaphus and Dama dama (96) > Capreolus capreolus (40) > Muntiacus reevesi (31) > Oryctolagus cuniculus (21) > Lepus europaeus (19), with the other taxa dispersing subsets of those dispersed by C. elpahus and D. dama. The invasive M. reevesi deposited the fewest seeds per gram of faecal pellet material (0.4 g(-1)) and hence fewer seeds per unit area than other deer species despite their numerical dominance, while C. elaphus/D. dama deposited the most (0.43 seeds m(-2) year(-1)). Due to differences in faecal seed density among habitats combined with the ranging behaviour of animals, more seeds were deposited in younger stands, enhancing the potential contribution of macroherbivores to population persistence by dispersal and colonisation in a successional mosaic.

Measuring coral reef decline through meta-analyses.

Coral reef ecosystems are in decline worldwide, owing to a variety of anthropogenic and natural causes. One of the most obvious signals of reef degradation is a reduction in live coral cover. Past and current rates of loss of coral are known for many individual reefs; however, until recently, no large-scale estimate was available. In this paper, we show how meta-analysis can be used to integrate existing small-scale estimates of change in coral and macroalgal cover, derived from in situ surveys of reefs, to generate a robust assessment of long-term patterns of large-scale ecological change. Using a large dataset from Caribbean reefs, we examine the possible biases inherent in meta-analytical studies and the sensitivity of the method to patchiness in data availability. Despite the fact that our meta-analysis included studies that used a variety of sampling methods, the regional estimate of change in coral cover we obtained is similar to that generated by a standardized survey programme that was implemented in 1991 in the Caribbean. We argue that for habitat types that are regularly and reasonably well surveyed in the course of ecological or conservation research, meta-analysis offers a cost-effective and rapid method for generating robust estimates of past and current states.

Amelioration of biodiversity impacts of genetically modified crops: predicting transient versus long-term effects.

It has been suggested that genetically modified herbicide-tolerant crops may benefit biodiversity because spraying of crops may be delayed until later in the growing season, allowing weeds to grow during the early part of the year. This provides an enhanced resource for arthropods, and potentially benefits birds that feed on these. Thus, this technology could enhance biodiversity. Using a review of weed phenologies and a population model, we show that many weeds are unlikely to benefit because spraying is generally delayed insufficiently late in the season to allow most to set seed. The positive effects on biodiversity observed in trials lasting one or two seasons are thus likely to be transient. For one weed of particular significance (Chenopodium album, fat hen) we show that it is unlikely that the positive effects observed could be maintained by inputs of seed during other parts of the rotation. However, we find preliminary evidence that if spraying can be ceased earlier in the season, then a viable population of late-emerging weeds could be maintained. This strategy could benefit weeds in both genetically modified (GM) and non-GM crops, but would probably lead to reduced inputs in GM systems compared with conventional ones.

Nonmanipulative determination of plant community dynamics.

Knowledge of the strengths of interactions between species in plant communities is of fundamental importance to our understanding of how communities are structured, although they are notoriously difficult to quantify. Techniques have recently been developed that allow the detailed enumeration of the strength of interactions between plant species within unmanipulated multispecies communities. Nonlinear regression analysis is used to fit competition models to long-term census data using natural variations in plant densities in lieu of manipulation. The models generated have been used to infer the intensity and importance of interactions as well as to analyse the effects of spatial and temporal variability. Theoretical work has begun to look at how different techniques for measuring competition perform in a range of systems, highlighting the importance of spatial scale. The lessons learned from applying these methods will enable improved estimation of the strength of competition in natural communities.

The depletion of algal beds by geese: a predictive model and test.

The short-term impact of herbivores on plant productivity has been well studied. Demonstrating long-term effects of grazing on plant populations is much more difficult, but knowledge of such long-term effects is vital in understanding plant-herbivore interactions. We address this issue in a relatively simple plant herbivore system involving the dark-bellied brent goose Branta bernicla bernicla and two marine macroalgae, Enteromorpha spp. and Ulva lactuca, on which the geese graze. In 3 years of sampling, goose grazing was responsible for depleting between 23% and 60% of the algae in the autumn, while wave damage caused between 15% and 70% depletion. The degree of depletion in autumn had no effect on the biomass of algae present the following summer, suggesting no long-term consequences of grazing for the population dynamics of the algae. A model simulating the change in mean algal biomass over the autumn and winter, incorporating changes due to depletion by geese, wave action and productivity, successfully described the date at which geese abandoned the algal bed in six different years. These years varied in numbers of geese, strength and timing of storms and initial biomass available. The most important factor determining the date of abandonment of the algal bed was a tradeoff between the timing of storms and the numbers of geese. When storms occur early, most depletion is due to wave action and geese must abandon the bed early, regardless of the numbers grazing there. As the algae become depleted, the geese switch to feeding on saltmarsh, pastures or arable crops. The rate of depletion of algae affects the timing of this movement, and the model presented here enables the switch to be predicted.

Predictions of biodiversity response to genetically modified herbicide-tolerant crops.

We simulated the effects of the introduction of genetically modified herbicide-tolerant (GMHT) crops on weed populations and the consequences for seed-eating birds. We predict that weed populations might be reduced to low levels or practically eradicated, depending on the exact form of management. Consequent effects on the local use of fields by birds might be severe, because such reductions represent a major loss of food resources. The regional impacts of GMHT crops are shown to depend on whether the adoption of GMHT crops by farmers covaries with current weed levels.

Determinants of the abundance of invasive annual weeds: community structure and non-equilibrium dynamics.

The dynamics of an annual pasture community are described from a five-year experimental and monitoring study. The community was dominated by two grasses (Lolium rigidum and Vulpia bromoides) and a legume (Trifolium subterraneum). Fits of population dynamic models to per capita rates of population change indicate that interactions between the grasses were generally strong, while interactions between the grasses and legumes were weaker. Most, but not all, of the net effects of competition on population growth could be attributed to interactions occurring during plant growth. Phase-plane analysis indicated that, for a constant environment, a joint equilibrium of the two grasses is unstable since interspecific competition between Lolium and Vulpia is stronger than intraspecific competition. Consequently, the community will tend to a mixture of only one or other of the grass species and T. subterraneum, depending on the founding composition of the pasture. Analysis of data taken from a year in which a drought occurred (1993-1994) demonstrated profound effects on all three species. Modelling of the long-term impacts of the effects of repeated droughts showed that disturbance of this form overrides the founder effect observed under constant conditions. Consequently, Vulpia is ultimately able to invade any mixture of the other species in environments where stochastic disturbances occur.

Multi-functionality and biodiversity in arbuscular mycorrhizas.

Plant roots in natural ecosystems are typically colonized by a wide range of fungi. Some of these are pathogenic, others appear to be opportunistic and have no apparent impact, while mycorrhizal fungi are generally regarded as mutualistic. Of the various types of mycorrhizal fungi, the arbuscular mycorrhizal (AM) association is by far the most abundant and widespread. While the most widely accepted model of AM function depends upon plants benefiting from the facilitation of phosphorus uptake, recent data from field-based studies in temperate ecosystems indicate that only plant species with poorly branched root systems benefit from AM fungi in this way: species with highly branched root systems may benefit in other ways, such as by being protected against root pathogenic fungi. These two responses apparently represent extremes along a continuum of AM benefit determined by root system architecture.

Rhizosphere and root-infecting fungi and the design of ecological field experiments.

As part of a wider study into the role of soil fungi in the ecology of the winter annual grass, Vulpia ciliata ssp. ambigua (Le Gall) Stace & Auquier, we applied the fungicides benomyl and prochloraz to three natural populations of the grass growing in East anglia, United Kingdom. The rhizosphere and rootinfecting fungi associated with the three populations were analysed each month between February and May 1992 when plants set seed. There were marked differences between the fungal floras associated with each of the three populations of V. ciliata, despite the fact that associated plant species and soil nutrient status were broadly similar between sites. This was attributed to wide differences in soil pH between the three populations. Prochloraz did not affect fungal abundance, but benomyl decreased the isolation frequencies of Fusarium oxysporum from roots and the frequencies of Penicillium and Trichoderma spp. isolated from rhizosphere soil, and increased the frequency of isolation of Mucor hiemalis from the rhizosphere of V. ciliata. There were also significant increases in the isolation frequencies of F. oxysporum from roots and M. hiemalis, Trichoderma spp. and Phoma fimeti from the rhizosphere of V. ciliata as plants matured. The significance of these results for the design of ecological field experiments are discussed in light of a previous study which has shown that asymptomatic root-infecting fungi can affect plant fecundity and hence abundance in natural populations of V. ciliata. We propose that differences in microbial communities between sites, controlled in part by soil chemistry, are a major factor determining plant performance under field conditions.

Habitat switching by dark-bellied brent geese Branta b. bernicla (L.) in relation to food depletion.

Seasonal changes in the distribution and feeding behaviour of dark-bellied brent geese Branta b. bernicla (L.) and the biomass of their food plants were studied in three successive winters on the Norfolk coast. The data was used, in conjunction with published information, to show how depletion, productivity and mortality of food plants drive the pattern of habitat switching in this species. It is then possible to explain the habitat shifts observed over the last 35 years and predict future changes. On arrival, geese fed first on algal beds and then on salt marsh, grass and arable fields before returning to feed entirely on the salt marsh in spring. The biomass of green algae, and subsequently the salt marsh vegetation, declined during the autumn and this could be attributed to depletion through goose grazing and natural mortality. As depletion occurred the geese fed more intensively, for a greater percentage of time and with an increasing pace rate, the net result, however, was a declining intake rate (as measured by defaecation rate). The algal biomass at which the geese switched from the algal beds to salt marsh was consistent between years, with heavy storm-induced loss of algae in one year resulting in an earlier switch. That the timing of habitat switches may be explained by depletion of food plants was further supported by historical data: the number of brent geese wintering at the site has increased dramatically over the last 30-35 years and the time of switching from algal beds to salt marsh and from salt marsh to salt marsh and fields has become progressively earlier, as expected from the increased depletion. The expected further increase in brent goose numbers will increase the rate of depletion of intertidal vegetation so that the switches between habitats will be more rapid and the geese will move inland earlier and remain inland longer. The expected increase in the brent goose population will thus result in a disproportionate increase in the levels of conflict between brent geese and agriculture.

Ecotypic differentiation of response to enhanced CO2 and temperature levels in Arabidopsis thaliana.

Five ecotypes of Arabidopsis thaliana, from widely dispersed origins, were grown under combinations of ambient and elevated atmospheric CO2 concentrations and ambient and elevated temperatures within solardomes. Total above-ground plant biomass was measured when the majority of plants across all ecotypes and treatments had formed seed pods. There were substantial differences in biomass between the ecotypes across all treatments. Temperature had no effect on biomass whilst CO2 had a significant effect both alone and in interaction with ecotype. The CO2 x ecotype interaction was mostly due to the enhancement of a single ecotype from the Cape Verde Islands.

The role of the hemiparasitic annual Rhinanthus minor in determining grassland community structure.

Rhinanthus minor (yellow-rattle) is a widespread hemiparasitic plant of grassland habitats throughout Great Britain. It is usually considered to be indicative of species-rich grassland, but in a survey of 14 habitats throughout Britain it was found that R. minor at the time of flowering normally occupied relatively low-diversity patches within areas of high diversity as determined by the number of species, Simpson's Index and the Shannon-Wiener Index. Following the death of adult plants of R. minor in the summer it was shown that the pattern of species diversity changed such that by the time R. minor germinated in the following spring the differences between the areas containing and not containing R. minor were much less distinct. A perturbation experiment in which R. minor was removed from four sites indicated that the effect of the removal of R. minor on the development of community structure over the next year was to increase species diversity on three of the sites and decrease it on the fourth. Those species which responded to the removal of R. minor by an increase in abundance were shown to be preferred hosts. All three lines of evidence point to the fact R. minor has a significant effect on the species diversity of the communities in which it grows by selectively parasitizing components of the flora and modifying the competitive relationships between plants. However, as the communities generally responded to the removal of R. minor by an increase in diversity and as the general survey indicated that R. minor is generally associated with areas of low diversity it would appear that the plants which are selectively parasitized are generally not the competitive dominants in the community.

Host selectivity and the mediation of competition by the root hemiparasite Rhinanthus minor.

Rhinanthus minor (Yellow-rattle) was grown in replacement series mixtures with Lolium perenne and Trifolium repens. The hemiparasitic interaction resulted in Relative Yield Totals (the sum of the yields in mixture relative to those in monoculture) considerably above 2. The hemiparasite caused a greater decrease in the yield of the legume and also performed better on the legume, indicating that T. repens was a better host for R. minor than L. perenne under the experimental conditions. When L. perenne and T. repens were grown in binary mixture with or without R. minor the hemiparasite affected considerably the competitive relationship between the two species by selectively parasitizing the legume. The effect of R. minor on competition between the two species was, however, dependent upon the nutrient status of the soil: the higher the level of soil nitrogen the fewer haustorial connections were made with T. repens and the less was the depression in its yield. In another series of experiments in which Festuca rubra, Holcus lanatus and L. perenne were grown in various binary mixtures with or without R. minor it was also shown that the yield of a preferred host was depressed to the advantage of a non-preferred host. It is suggested that the mediation of competition by the hemiparasite provides a mechanism by which it might affect the structure and diversity of plant communities.

The host range and selectivity of a parasitic plant: Rhinanthus minor L.

Rhinanthus minor (Yellow-rattle) is a widespread hemiparasitic plant of grassland habitats throughout Britain. Association analysis of the dune vegetation at Holme-next-the-Sea in eastern England revealed only two potential host plants through positive association. In contrast direct examination of the root systems revealed haustorial connections with 20 host species. The number of species parasitized by one plant ranged from one to seven. Data from another four sites in Britain and one in central Europe indicate that the natural host range of R. minor encompasses at least 50 species from 18 families with 22% in the Leguminosae and 30% in the Gramineae. Comparison of the number of haustorial connections made to each species with the abundance of roots in the soil shows that R. minor is a highly selective parasite, but that the selectivity is not consistent between populations or between plants from different parts of the same population. The reasons for host selectivity are discussed.

On the analysis of competition at the level of the individual plant.

The extent to which some measure of local crowding can account for the performance of individual plants is examined with reference to populations of two species of annual plant. Only a relatively small proportion of the variation in individual plant yield could be accounted for by measures of local crowding. These included the number of close neighbours, an estimate of the area available to each plant and competitive pressure. A multiple regression that took account of both emergence time and local crowding increased the proportion of variance that could be accounted for up to 50%. Computer simulations of the growth of indivudual plants in monoculture were then caried out in order to determine whether the unexplained variation resulted from fundamental flaws in the models or from unaccounted for sources of variation in the field. The results from the simulations again indicated that only a relatively low proportion of the variation in individual plant yield could be accounted for by emergence time and local density, even though these were known to be the only variables present. These findings are discussed in relation to the relative importance of one-sided and two-sided competition, and the complex cross-correlations that occur between individuals in plant populations. These two factors will make it very difficult for field workers to determine accurately what factors determine individual plant yield and in particular to predict the effects of local crowding on the performance of individual plants.