Ambient and substrate energy influence decomposer diversity differentially across trophic levels.

The species-energy hypothesis predicts increasing biodiversity with increasing energy in ecosystems. Proxies for energy availability are often grouped into ambient energy (i.e., solar radiation) and substrate energy (i.e., non-structural carbohydrates or nutritional content). The relative importance of substrate energy is thought to decrease with increasing trophic level from primary consumers to predators, with reciprocal effects of ambient energy. Yet, empirical tests are lacking. We compiled data on 332,557 deadwood-inhabiting beetles of 901 species reared from wood of 49 tree species across Europe. Using host-phylogeny-controlled models, we show that the relative importance of substrate energy versus ambient energy decreases with increasing trophic levels: the diversity of zoophagous and mycetophagous beetles was determined by ambient energy, while non-structural carbohydrate content in woody tissues determined that of xylophagous beetles. Our study thus overall supports the species-energy hypothesis and specifies that the relative importance of ambient temperature increases with increasing trophic level with opposite effects for substrate energy.

Revegetation through seeding or planting: A worldwide systematic map.

Roughly 2 billion ha of land are degraded and in need of ecological restoration worldwide. Active restoration frequently involves revegetation, which leads to the dilemma of whether to conduct direct seeding or to plant nursery-grown seedlings. The choice of revegetation method can regulate plant survival and performance, with economic implications that ultimately feed back to our capacity to conduct restoration. We followed a peer-reviewed protocol to develop a systematic map that collates, describes and catalogues the available studies on how seeding compares to planting in achieving restoration targets. We compiled a database with the characteristics of all retrieved studies, which can be searched to identify studies of particular locations and habitats, objectives of restoration, plant material, technical aspects, and outcomes measured. The search was made in eight languages and retrieved 3355 publications, of which 178 were retained. The systematic map identifies research gaps, such as a lack of studies in the global South, in tropical rainforests, and covering a long time period, which represent opportunities to expand field-based research. Additionally, many studies overlooked reporting on important technical aspects such as seed provenance and nursery cultivation methods, and others such as watering or seedling protection were more frequently applied for planting than for seeding, which limits our capacity to learn from past research. Most studies measured outcomes related to the target plants but avoided measuring general restoration outcomes or economic aspects. This represents a relevant gap in research, as the choice of revegetation method is greatly based on economic aspects and the achievement of restoration goals goes beyond the establishment of plants. Finally, we identified a substantial volume of studies conducted in temperate regions and over short periods (0-5 y). This research cluster calls for a future in-depth synthesis, potentially through meta-analysis, to reveal the overall balance between seeding and planting and assess whether the response to this question is mediated by species traits, environmental characteristics, or technical aspects. Besides identifying research clusters and gaps, the systematic map database allows managers to find the most relevant scientific literature on the appropriateness of seeding vs. planting for particular conditions, such as certain species or habitats.

Education and knowledge determine preference for bark beetle control measures in El Salvador.

Extensive outbreaks of bark beetles have affected not only large parts of coniferous forests in the Northern Hemisphere, but also - largely absent from global attention - native pine forests of Central America. As such outbreaks frequently spark management debates among residents, land managers, forest owners and the public, the social acceptance of bark beetle control measures has become crucial for modern land management. However, the sociological and psychological determinants of the preference for specific bark beetle control measures outside protected areas remain unclear. To determine the acceptability of bark beetle control measures in El Salvador, we assessed how demographic variables, attitude towards the bark beetle, education, and self-reported knowledge affected the preference for different bark beetle control measures in a survey of government employees and local forest owners using a quantitative questionnaire survey. Cumulative link mixed models revealed that the general preference for control measures increased with increasing self-reported knowledge about the bark beetle but decreased with increasing level of respondent education and an increasing positive attitude towards the bark beetle. Respondents generally preferred beetle control measures on small areas than on large areas. Preferences for control measures did not differ between government employees and forest owners, with controlled burning and chemical control significantly less accepted than stand thinning or salvage logging. We discuss the most preferred control measures considering recent scientific evidence of their efficacy and conclude that the current bark beetle outbreak should be controlled through logging of pines weakened by fire in the short-term and by stand thinning in the medium-term to prevent further outbreaks.

An ecosystem services approach to the ecological effects of salvage logging: valuation of seed dispersal.

Forest disturbances diminish ecosystem services and boost disservices. Because post-disturbance management intends to recover the greatest possible value, selling timber often prevails over other considerations. Ecological research has shown diverse effects of salvage logging, yet such research has focused on the biophysical component of post-disturbance ecosystems and lacks the link with human well-being. Here we bridge that gap under the ecosystem services framework by assessing the impact of post-fire management on a non-timber value. By employing the replacement cost method, we calculated the value of the post-fire natural regeneration of Holm oaks in southern Spain under three post-fire management options by considering the cost of planting instead. The value of this ecosystem service in non-intervention areas doubled that of salvage-logged stands due to the preference for standing dead trees by the main seed disperser. Still, most of the value resulted from the resprouting capacity of oaks. The value of this and other ecosystem services should be added to traditional cost/benefit analyses of post-disturbance management. We thus call for a more holistic approach to salvage logging research, one that explicitly links ecological processes with human well-being through ecosystem services, to better inform decision-makers on the outcomes of post-disturbance management.

Effective nut dispersal by magpies (Pica pica L.) in a Mediterranean agroecosystem.

Scatter-hoarding animals such as corvids play a crucial role in the dispersal of nut-producing tree species. This interaction is well known for some corvids, but remains elusive for other species such as the magpie (Pica pica), an abundant corvid in agroecosystems and open landscapes of the Palearctic region. In addition, the establishment of the individual dispersed seeds-a prerequisite for determining seed-dispersal effectiveness-has never before been documented for the interaction between corvids and nut-producing trees. We analyzed walnut dispersal by magpies in an agroecosystem in southern Spain. We used several complementary approaches, including video recording nut removal from feeders, measuring dispersal distance using radio tracking (with radio transmitters placed inside nuts), and monitoring the fate of dispersed nuts to the time of seedling emergence. Magpies were shown to be highly active nut dispersers. The dispersal distance averaged 39.6 ± 4.5 m and ranged from 4.1 to 158.5 m. Some 90% of the removed walnuts were cached later, and most of these (98%) were buried in the soil or hidden under plant material. By the time of seedling emergence, ca. 33% of nuts remained at the caching location. Finally, 12% of the cached nuts germinated and 4% yielded an emerged seedling, facilitating the transition to the next regeneration stage. The results demonstrate for the first time that magpies can be an effective scatter-hoarding disperser of a nut-producing tree species, suggesting that this bird species may play a key role in the regeneration and expansion of broadleaf forests in Eurasia.

Shifting demographic conflicts across recruitment cohorts in a dynamic post-disturbance landscape.

Seed dispersal effectiveness, which measures the number of adult plant individuals produced by seed dispersal, is the product of the number of seeds dispersed and the probability a seed produces an adult. Directed dispersal to certain habitat types may enhance some stages of recruitment but disfavor others, generating demographic conflicts in plant ontogeny. We asked whether temporal changes in habitat features may affect the distribution of seedlings recruited from dispersed acorns, and whether this could induce shifts in the life-stage conflicts experienced by successive cohorts of naturally recruited plants. As early successional habitats are characterized by rapid change, we used a burnt pine stand in southern Spain to monitor the recruitment and performance of a major tree species (Quercus ilex) across 7 yr in four types of post-fire habitats. These differed in structure and included patches of unburnt forest and three management alternatives of burnt trees: logging, partial cutting, and nonintervention. Young oaks that resprouted after the fire were mainly located near acorn sources, while new seedlings initially emerged mostly in habitats with standing snags due to habitat selection by European jays, Garrulus glandarius, for dispersal. The dead pines gradually collapsed and attracted less dispersal, so subsequent seedling cohorts mainly recruited within patches of unburnt pines. These live pines enhanced the survival of the oaks located beneath their canopy but greatly reduced their growth as compared to the other post-fire habitats, thus representing a demographic conflict that was absent elsewhere. As a consequence of the directional shift in the habitat where seedlings recruited, successive seedling cohorts experienced a gradual improvement in their likelihood of survival but a reduction in growth. The progressive intensification of this life-stage conflict hinged on the reduction of vertical structures in the habitat with standing burnt pines. Recruitment success thus involved temporal variation in the habitat where recruitment occurred, likely resulting from changes in the direction of seed dispersal, and spatial variation in habitat suitability for seedling establishment and growth. Temporal changes in habitat structure can indirectly change the environment in which recruitment occurs, and consequently seed dispersal effectiveness, by shifting the direction of seed dispersal.

Post-fire salvage logging alters species composition and reduces cover, richness, and diversity in Mediterranean plant communities.

An intense debate exists on the effects of post-fire salvage logging on plant community regeneration, but scant data are available derived from experimental studies. We analyzed the effects of salvage logging on plant community regeneration in terms of species richness, diversity, cover, and composition by experimentally managing a burnt forest on a Mediterranean mountain (Sierra Nevada, S Spain). In each of three plots located at different elevations, three replicates of three treatments were implemented seven months after the fire, differing in the degree of intervention: "Non-Intervention" (all trees left standing), "Partial Cut plus Lopping" (felling 90% of the trees, cutting the main branches, and leaving all the biomass in situ), and "Salvage Logging" (felling and piling the logs, and masticating the woody debris). Plant composition in each treatment was monitored two years after the fire in linear point transects. Post-fire salvage logging was associated with reduced species richness, Shannon diversity, and total plant cover. Moreover, salvaged sites hosted different species assemblages and 25% lower cover of seeder species (but equal cover of resprouters) compared to the other treatments. Cover of trees and shrubs was also lowest in Salvage Logging, which could suggest a potential slow-down of forest regeneration. Most of these results were consistent among the three plots despite plots hosting different plant communities. Concluding, our study suggests that salvage logging may reduce species richness and diversity, as well as the recruitment of woody species, which could delay the natural regeneration of the ecosystem.

The effect of natural disturbances on forest biodiversity: an ecological synthesis.

Disturbances alter biodiversity via their specific characteristics, including severity and extent in the landscape, which act at different temporal and spatial scales. Biodiversity response to disturbance also depends on the community characteristics and habitat requirements of species. Untangling the mechanistic interplay of these factors has guided disturbance ecology for decades, generating mixed scientific evidence of biodiversity responses to disturbance. Understanding the impact of natural disturbances on biodiversity is increasingly important due to human-induced changes in natural disturbance regimes. In many areas, major natural forest disturbances, such as wildfires, windstorms, and insect outbreaks, are becoming more frequent, intense, severe, and widespread due to climate change and land-use change. Conversely, the suppression of natural disturbances threatens disturbance-dependent biota. Using a meta-analytic approach, we analysed a global data set (with most sampling concentrated in temperate and boreal secondary forests) of species assemblages of 26 taxonomic groups, including plants, animals, and fungi collected from forests affected by wildfires, windstorms, and insect outbreaks. The overall effect of natural disturbances on α-diversity did not differ significantly from zero, but some taxonomic groups responded positively to disturbance, while others tended to respond negatively. Disturbance was beneficial for taxonomic groups preferring conditions associated with open canopies (e.g. hymenopterans and hoverflies), whereas ground-dwelling groups and/or groups typically associated with shady conditions (e.g. epigeic lichens and mycorrhizal fungi) were more likely to be negatively impacted by disturbance. Across all taxonomic groups, the highest α-diversity in disturbed forest patches occurred under moderate disturbance severity, i.e. with approximately 55% of trees killed by disturbance. We further extended our meta-analysis by applying a unified diversity concept based on Hill numbers to estimate α-diversity changes in different taxonomic groups across a gradient of disturbance severity measured at the stand scale and incorporating other disturbance features. We found that disturbance severity negatively affected diversity for Hill number q = 0 but not for q = 1 and q = 2, indicating that diversity-disturbance relationships are shaped by species relative abundances. Our synthesis of α-diversity was extended by a synthesis of disturbance-induced change in species assemblages, and revealed that disturbance changes the ß-diversity of multiple taxonomic groups, including some groups that were not affected at the α-diversity level (birds and woody plants). Finally, we used mixed rarefaction/extrapolation to estimate biodiversity change as a function of the proportion of forests that were disturbed, i.e. the disturbance extent measured at the landscape scale. The comparison of intact and naturally disturbed forests revealed that both types of forests provide habitat for unique species assemblages, whereas species diversity in the mixture of disturbed and undisturbed forests peaked at intermediate values of disturbance extent in the simulated landscape. Hence, the relationship between α-diversity and disturbance severity in disturbed forest stands was strikingly similar to the relationship between species richness and disturbance extent in a landscape consisting of both disturbed and undisturbed forest habitats. This result suggests that both moderate disturbance severity and moderate disturbance extent support the highest levels of biodiversity in contemporary forest landscapes.

Disturbance interval modulates the starting point for vegetation succession.

Increased frequency and new types of disturbances caused by global change calls for deepened insights into possible alterations of successional pathways. Despite current interest in disturbance interactions there is a striking lack of studies focusing on the implication of decreasing times between disturbances. We surveyed forest-floor vegetation (vascular plants and bryophytes) in a Pinus sylvestris-dominated, even-aged production forest landscape, unique because of the presence of stands under a precisely dated disturbance interval gradient, ranging from 0 to 123 yr between clearcutting and a subsequent megafire. Despite a dominance of early-successional species in all burned stands 5 yr after fire, progression of succession was linked to time since the preceding clearcutting disturbance. This was most clearly seen in increased frequency with time since clearcutting of the dominant, late-successional dwarf shrub Vaccinium myrtillus, with surviving rhizomes as an important mechanism for postfire recovery. Our results demonstrate the role of legacy species as significant drivers of succession. We conclude that the starting point for succession is modulated by disturbance interval, so that shortened intervals risk reducing development towards late-successional stages. We suggest that a decrease in long successional sequences caused by more frequent disturbances may represent a general pattern, relevant also for other forest types and ecosystems.

Restoring oak forests through direct seeding or planting: Protocol for a continental-scale experiment.

The choice of revegetating via direct seeding or planting nursery-grown seedlings influences the potential stresses suffered by seedlings such as herbivory and drought. The outcome of the balance between both revegetation methods may ultimately depend on how species identity and traits such as seed and seedling size interact with environmental conditions. To test this, we will conduct a continental-scale experiment consisting of one mini-experiment replicated by multiple participants across Europe. Each participant will establish a site with seeded and planted individuals of one or more native, locally growing oak (Quercus) species; the selection of this genus aims to favour continental-scale participation and to allow testing the response of a widely distributed genus of broad ecological and economic relevance. At each site, participants will follow the present protocol for seed collection, seeding in the field, nursery cultivation, outplanting, protection against herbivores, site maintenance, and measurement of seedling performance and environmental variables. Each measurement on each species at each site will produce one effect size; the data will be analysed through mixed-effects meta-analysis. With this approach we will assess the main effect of revegetation method, species, plant functional traits, and the potential effect of site-specific effect moderators. Overall, we will provide a continental-scale estimate on the seeding vs. planting dilemma and analyse to what extent the differences in environmental conditions across sites, seed size, functional traits, and the phylogenetic relatedness of species can account for the differences in the effect of revegetation method on seedling performance across study sites and species.

Temporal variation in effect sizes in a long-term, split-plot field experiment.

Ecological field experiments initiate successional and evolutionary changes among resident species, yet effect sizes are often reported as if they were constants. Few ecological studies have addressed their questions through long-term, experimental approaches, and many questions remain unanswered regarding temporal patterns in ecological effect sizes. We document temporal variation in effect sizes in response to pulse and press manipulations in a long-term factorial field experiment at Nash's Field, England. The experiment comprises seven treatments applied in a split-plot design to test the single and interactive effects of herbivory by insects, molluscs, and rabbits, liming, nutrient limitation (applied as press experiments), competition (exclusion of grasses or herbs with specific herbicides), and seed limitation (pulse experiments) on plant community dynamics. The response of all vascular plant species was followed for two decades. High species richness was positively related to the minus-grass herbicide in the first decade and negatively related to both nitrogen addition and the abundance of dominant species in both decades. Many significant effects appeared quickly, but some large effects were not detected until year 15. Press experiments produced some long-lasting effects, but effect sizes changed due to both idiosyncratic "year effects" and secular trends. For pulse experiments, most effects, including positive and negative responses to herbicide application and the invasion of most of the sown species, disappeared quickly. However, some endured or grew monotonically, such as the invasion of two sown species that benefited from particular combinations of the press treatments. The fastest effects to appear were the responses from established species. Many of these responses were negative, likely resulting from reduced niche dimensionality and competitive exclusion by new dominant species. Contrarily, one of the largest community-level effects took well over a decade to appear: the natural invasion by one species, which responded to a four-way interaction between experimental treatments. The insights gained from individual effects increased with the duration of the lag before their first appearance, drawing attention to the importance of long-term, manipulative field experiments. This experiment also reinforces the point that factorial experiments are the most insightful way to explore ecological interactions.

Estimating retention benchmarks for salvage logging to protect biodiversity.

Forests are increasingly affected by natural disturbances. Subsequent salvage logging, a widespread management practice conducted predominantly to recover economic capital, produces further disturbance and impacts biodiversity worldwide. Hence, naturally disturbed forests are among the most threatened habitats in the world, with consequences for their associated biodiversity. However, there are no evidence-based benchmarks for the proportion of area of naturally disturbed forests to be excluded from salvage logging to conserve biodiversity. We apply a mixed rarefaction/extrapolation approach to a global multi-taxa dataset from disturbed forests, including birds, plants, insects and fungi, to close this gap. We find that 75 ± 7% (mean ± SD) of a naturally disturbed area of a forest needs to be left unlogged to maintain 90% richness of its unique species, whereas retaining 50% of a naturally disturbed forest unlogged maintains 73 ± 12% of its unique species richness. These values do not change with the time elapsed since disturbance but vary considerably among taxonomic groups.

Impacts of salvage logging on biodiversity: a meta-analysis.

Logging to "salvage" economic returns from forests affected by natural disturbances has become increasingly prevalent globally. Despite potential negative effects on biodiversity, salvage logging is often conducted, even in areas otherwise excluded from logging and reserved for nature conservation, inter alia because strategic priorities for post-disturbance management are widely lacking.A review of the existing literature revealed that most studies investigating the effects of salvage logging on biodiversity have been conducted less than 5 years following natural disturbances, and focused on non-saproxylic organisms.A meta-analysis across 24 species groups revealed that salvage logging significantly decreases numbers of species of eight taxonomic groups. Richness of dead wood dependent taxa (i.e. saproxylic organisms) decreased more strongly than richness of non-saproxylic taxa. In contrast, taxonomic groups typically associated with open habitats increased in the number of species after salvage logging.By analysing 134 original species abundance matrices, we demonstrate that salvage logging significantly alters community composition in 7 of 17 species groups, particularly affecting saproxylic assemblages.Synthesis and applications. Our results suggest that salvage logging is not consistent with the management objectives of protected areas. Substantial changes, such as the retention of dead wood in naturally disturbed forests, are needed to support biodiversity. Future research should investigate the amount and spatio-temporal distribution of retained dead wood needed to maintain all components of biodiversity.