Environmental mitigation hierarchy and biodiversity offsets revisited through habitat connectivity modelling.

Biodiversity loss is accelerating because of unceasing human activity and land clearing for development projects (urbanisation, transport infrastructure, mining and quarrying …). Environmental policy-makers and managers in different countries worldwide have proposed the mitigation hierarchy to ensure the goal of "no net loss (NNL) of biodiversity" and have included this principle in environmental impact assessment processes. However, spatial configuration is hardly ever taken into account in the mitigation hierarchy even though it would greatly benefit from recent developments in habitat connectivity modelling incorporating landscape graphs. Meanwhile, national, European and international commitments have been made to maintain and restore the connectivity of natural habitats to face habitat loss and fragmentation. Our objective is to revisit the mitigation hierarchy and to suggest a methodological framework for evaluating the environmental impact of development projects, which includes a landscape connectivity perspective. We advocate the use of the landscape connectivity metric equivalent connectivity (EC), which is based on the original concept of "amount of reachable habitat". We also refine the three main levels of the mitigation hierarchy (impact avoidance, reduction and offset) by integrating a landscape connectivity aspect. We applied this landscape connectivity framework to a simple, virtual habitat network composed of 14 patches of varying sizes. The mitigation hierarchy was addressed through graph theory and EC and several scenarios of impact avoidance, reduction and compensation were tested. We present the benefits of a habitat connectivity framework for the mitigation hierarchy, provide practical recommendations to implement this framework and show its use in real case studies that had previously been restricted to one or two steps of the mitigation hierarchy. We insist on the benefits of a habitat connectivity framework for the mitigation hierarchy and for ecological equivalence assessment. In particular, we demonstrate why it is risky to use a standard offset ratio (the ratio between the amount of area negatively impacted and the compensation area) without performing a connectivity analysis that includes the landscape surrounding the zone impacted by the project. We also discuss the limitations of the framework and suggest potential improvements. Lastly, we raise concerns about the need to rethink the strategy for biodiversity protection. Given that wild areas and semi-natural habitats are becoming scarcer, in particular in industrialised countries, we are convinced that the real challenge is to quickly reconsider the current vision of "developing first, then assessing the ecological damage", and instead urgently adopt an upstream protection strategy that would identify and protect the land that must not be lost if we wish to maintain viable species populations and ecological corridors allowing them the mobility necessary to their survival.

Ecological Equivalence Assessment Methods: What Trade-Offs between Operationality, Scientific Basis and Comprehensiveness?

In many countries, biodiversity compensation is required to counterbalance negative impacts of development projects on biodiversity by carrying out ecological measures, called offset when the goal is to reach "no net loss" of biodiversity. One main issue is to ensure that offset gains are equivalent to impact-related losses. Ecological equivalence is assessed with ecological equivalence assessment methods taking into account a range of key considerations that we summarized as ecological, spatial, temporal, and uncertainty. When equivalence assessment methods take into account all considerations, we call them "comprehensive". Equivalence assessment methods should also aim to be science-based and operational, which is challenging. Many equivalence assessment methods have been developed worldwide but none is fully satisfying. In the present study, we examine 13 equivalence assessment methods in order to identify (i) their general structure and (ii) the synergies and trade-offs between equivalence assessment methods characteristics related to operationality, scientific-basis and comprehensiveness (called "challenges" in his paper). We evaluate each equivalence assessment methods on the basis of 12 criteria describing the level of achievement of each challenge. We observe that all equivalence assessment methods share a general structure, with possible improvements in the choice of target biodiversity, the indicators used, the integration of landscape context and the multipliers reflecting time lags and uncertainties. We show that no equivalence assessment methods combines all challenges perfectly. There are trade-offs between and within the challenges: operationality tends to be favored while scientific basis are integrated heterogeneously in equivalence assessment methods development. One way of improving the challenges combination would be the use of offset dedicated data-bases providing scientific feedbacks on previous offset measures.

Distinct invasion strategies operating within a natural annual plant system.

Alien plant species are known to have a wide range of impacts on recipient communities, from resident species' exclusions to coexistence with resident species. It remains unclear; however, if this variety of impacts is due to different invader strategies, features of recipient communities or both. To test this, we examined multiple plant invasions of a single ecosystem in southwestern Australia. We used extensive community data to calculate pairwise segregation between target alien species and many co-occurring species. We related segregation to species' positions along community trait hierarchies and identified at least two distinct invasion strategies: 'exploiters' which occupy high positions along key trait hierarchies and reduce local native species diversity (particularly in nutrient-enriched situations), and 'coexisters' who occupy intermediate trait positions and have no discernable impact on native diversity. We conclude that trait hierarchies, linked to measures of competition, can provide valuable insights about the processes driving different invasion outcomes.

Diminished soil functions occur under simulated climate change in a sup-alpine pasture, but heterotrophic temperature sensitivity indicates microbial resilience.

The pressure of climate change is disproportionately high in mountainous regions, and small changes may push ecosystem processes beyond sensitivity thresholds, creating new dynamics of carbon and nutrient cycling. Given that the rate of organic matter decomposition is strongly dependent upon temperature and soil moisture, the sensitivity of soil respiration to both metrics is highly relevant when considering soil-atmosphere feedbacks under a changing climate. To assess the effects of changing climate in a mountain pasture system, we transplanted turfs along an elevation gradient, monitored in situ soil respiration, incubated collected top-soils to determine legacy effects on temperature sensitivity, and analysed soil organic matter (SOM) to detect changes in quality and quantity of SOM fractions. In situ transplantation down-slope reduced soil moisture and increased soil temperature, with concurrent reductions in soil respiration. Soil moisture acted as an overriding constraint to soil respiration, and significantly reduced the sensitivity to temperature. Under controlled laboratory conditions, removal of the moisture constraint to heterotrophic respiration led to a significant respiration-temperature response. However, despite lower respiration rates down-slope, the response function was comparable among sites, and therefore unaffected by antecedent conditions. We found shifts in the SOM quality, especially of the light fraction, indicating changes to the dynamics of decomposition of recently deposited material. Our findings highlighted the resilience of the microbial community to severe climatic perturbations, but also that soil moisture stress during the growing season can significantly reduce soil function in addition to direct effects on plant productivity. This demonstrated the sensitivity of subalpine pastures under climate change, and possible implications for sustainable use given reductions in organic matter turnover and consequent feedbacks to nutrient cycling.

Differential allelopathic effects of Japanese knotweed on willow and cottonwood cuttings used in riverbank restoration techniques.

Using bioengineering techniques to restore areas invaded by Fallopia japonica shows promising results. Planting tree cuttings could allow both rapidly re-establishing a competitive native plant community and reducing F. japonica performance. However, F. japonica has been shown to affect native plant species through different mechanisms such as allelopathy. This article investigates the phytotoxic effect of F. japonica on the resprouting capacity and the growth of three Salicaceae species with potential value for restoration. An experimental design which physically separates donor pots containing either an individual from F. japonica or bare soil from target pots containing cuttings of Populus nigra, Salix atrocinerea or Salix viminali was used. Leachates from donor pots were used to water target pots. The effects of leachates were evaluated by measuring the final biomass of the cuttings. F. japonica leachates inhibited the growth of cuttings, and this effect is linked to the emission of polyphenol compounds by F. japonica. Leachates also induced changes in soil nitrogen composition. These results suggest the existence of allelopathic effects, direct and/or indirect, of F. japonica on the growth of Salicaceae species cuttings. However, the three species were not equally affected, suggesting that the choice of resistant species could be crucial for restoration success.

Transplantation of subalpine wood-pasture turfs along a natural climatic gradient reveals lower resistance of unwooded pastures to climate change compared to wooded ones.

Climate change could impact strongly on cold-adapted mountain ecosystems, but little is known about its interaction with traditional land-use practices. We used an altitudinal gradient to simulate a year-round warmer and drier climate for semi-natural subalpine grasslands across a landscape of contrasting land-use management. Turf mesocosms from three pasture-woodland land-use types-unwooded pasture, sparsely wooded pasture, and densely wooded pasture-spanning a gradient from high to low management intensity were transplanted downslope to test their resistance to two intensities of climate change. We found strong overall effects of intensive (+4 K) experimental climate change (i.e., warming and reduced precipitation) on plant community structure and function, while moderate (+2 K) climate change did not substantially affect the studied land-use types, thus indicating an ecosystem response threshold to moderate climate perturbation. The individual land-use types were affected differently under the +4 K scenario, with a 60% decrease in aboveground biomass (AGB) in unwooded pasture turfs, a 40% decrease in sparsely wooded pasture turfs, and none in densely wooded ones. Similarly, unwooded pasture turfs experienced a 30% loss of species, advanced (by 30 days) phenological development, and a mid-season senescence due to drought stress, while no such effects were recorded for the other land-use types. The observed contrasting effects of climate change across the pasture-woodland landscape have important implications for future decades. The reduced impact of climate change on wooded pastures as compared to unwooded ones should promote the sustainable land use of wooded pastures by maintaining low management intensity and a sparse forest canopy, which buffer the immediate impacts of climate change on herbaceous vegetation.

Light availability prevails over soil fertility and structure in the performance of Asian knotweeds on riverbanks: new management perspectives.

Asian knotweeds (Fallopia spp.) are considered one of the world's most invasive species. Restoring habitats dominated by these exotic species requires a better understanding of the importance of abiotic factors controlling the invasive knotweeds performance. We used observational data obtained on the embankment of the Isère River (France) to study the performance of Fallopia spp. under different soil, light, and disturbance conditions. On the Isère riverbanks, light intensity assessed by light quantity transmitted through canopy was the most important factor explaining the variability observed on knotweed performance expressed as above-ground biomass per square meter. Asian knotweeds were more productive under intensive light conditions. Alternatively other factors such as mowing (twice a year), soil fertility, soil texture, position on the bank or exposure to the sun had no significant effect on knotweed biomass production. We conclude that decreasing light resources, for example, by increasing competitive pressure on sites dominated by Asian knotweeds could be included in management plans to control the populations of this invasive taxon.

Arbuscular mycorrhizal fungi reduce the differences in competitiveness between dominant and subordinate plant species.

In grassland communities, plants can be classified as dominants or subordinates according to their relative abundances, but the factors controlling such distributions remain unclear. Here, we test whether the presence of the arbuscular mycorrhizal (AM) fungus Glomus intraradices affects the competitiveness of two dominant (Taraxacum officinale and Agrostis capillaris) and two subordinate species (Prunella vulgaris and Achillea millefolium). Plants were grown in pots in the presence or absence of the fungus, in monoculture and in mixtures of both species groups with two and four species. In the absence of G. intraradices, dominants were clearly more competitive than subordinates. In inoculated pots, the fungus acted towards the parasitic end of the mutualism-parasitism continuum and had an overall negative effect on the growth of the plant species. However, the negative effects of the AM fungus were more pronounced on dominant species reducing the differences in competitiveness between dominant and subordinate species. The effects of G. intraradices varied with species composition highlighting the importance of plant community to mediate the effects of AM fungi. Dominant species were negatively affected from the AM fungus in mixtures, while subordinates grew identically with and without the fungus. Therefore, our findings predict that the plant dominance hierarchy may flatten out when dominant species are more reduced than subordinate species in an unfavourable AM fungal relationship (parasitism).

Calcium induces long-term legacy effects in a subalpine ecosystem.

Human activities have transformed a significant proportion of the world's land surface, with profound effects on ecosystem processes. Soil applications of macronutrients such as nitrate, phosphorus, potassium or calcium are routinely used in the management of croplands, grasslands and forests to improve plant health or increase productivity. However, while the effects of continuous fertilization and liming on terrestrial ecosystems are well documented, remarkably little is known about the legacy effect of historical fertilization and liming events in terrestrial ecosystems and of the mechanisms involved. Here, we show that more than 70 years after the last application of lime on a subalpine grassland, all major soil and plant calcium pools were still significantly larger in limed than in unlimed plots, and that the resulting shift in the soil calcium/aluminium ratio continues to affect ecosystem services such as primary production. The difference in the calcium content of the vegetation and the topmost 10 cm of the soil in limed vs. unlimed plots amounts to approximately 19.5 g m(-2), equivalent to 16.3% of the amount that was added to the plots some 70 years ago. In contrast, plots that were treated with nitrogen-phosphorus-potassium fertilizer in the 1930s did not differ from unfertilized plots in any of the soil and vegetation characteristics measured. Our findings suggest that the long-term legacy effect of historical liming is due to long-term storage of added calcium in stable soil pools, rather than a general increase in nutrient availability. Our results demonstrate that single applications of calcium in its carbonated form can profoundly and persistently alter ecosystem processes and services in mountain ecosystems.

Long-term effects of short-term perturbation in a subalpine grassland.

Theoretical advances and short-term experimental studies have furthered our understanding of how ecosystems respond to perturbation. However, there are few well-replicated experimental studies that allow an assessment of long-term responses. Results from a controlled, large-scale field experiment in a subalpine grassland near Interlaken, Switzerland, show that 2-4 years of liming (Ca: 40 g x m(-2) x yr(-1)) still significantly affected the composition of the vegetation and the soil microbial community nearly 70 years after the treatments were imposed, whereas NPK fertilization (8 g x m(-2) x yr(-1)) only marginally affected vegetation composition. The exchangeable content of Ca ions and soil pH were higher in limed plots but were unaffected in fertilized plots. Plant species and PLFAs (phospholipid fatty acids) indicating low pH values were found in higher abundance in the unlimed plots, suggesting that the long-lasting effects of liming on the above- and belowground communities were mediated through changes in soil pH. The results of this long-term study indicate that the resilience of mountain ecosystems may be particularly low in response to perturbations that substantially alter soil pH or other key determinants of belowground processes.