Hervé Jactel

Plantation forests and biodiversity: oxymoron or opportunity?

Losses of natural and semi-natural forests, mostly to agriculture, are a significant concern for biodiversity. Against this trend, the area of intensively managed plantation forests increases, and there is much debate about the implications for biodiversity. We provide a comprehensive review of the function of plantation forests as habitat compared with other land cover, examine the effects on biodiversity at the landscape scale, and synthesise context-specific effects of plantation forestry on biodiversity. Natural forests are usually more suitable as habitat for a wider range of native forest species than plantation forests but there is abundant evidence that plantation forests can provide valuable habitat, even for some threatened and endangered species, and may contribute to the conservation of biodiversity by various mechanisms. In landscapes where forest is the natural land cover, plantation forests may represent a low-contrast matrix, and afforestation of agricultural land can assist conservation by providing complementary forest habitat, buffering edge effects, and increasing connectivity. In contrast, conversion of natural forests and afforestation of natural non-forest land is detrimental. However, regional deforestation pressure for agricultural development may render plantation forestry a ‘lesser evil’ if forest managers protect indigenous vegetation remnants. We provide numerous context-specific examples and case studies to assist impact assessments of plantation forestry, and we offer a range of management recommendations. This paper also serves as an introduction and background paper to this special issue on the effects of plantation forests on biodiversity.

The global spectrum of plant form and function

Earth is home to a remarkable diversity of plant forms and life histories, yet comparatively few essential trait combinations have proved evolutionarily viable in today’s terrestrial biosphere. By analysing worldwide variation in six major traits critical to growth, survival and reproduction within the largest sample of vascular plant species ever compiled, we found that occupancy of six-dimensional trait space is strongly concentrated, indicating coordination and trade-offs. Three-quarters of trait variation is captured in a two-dimensional global spectrum of plant form and function. One major dimension within this plane reflects the size of whole plants and their parts; the other represents the leaf economics spectrum, which balances leaf construction costs against growth potential. The global plant trait spectrum provides a backdrop for elucidating constraints on evolution, for functionally qualifying species and ecosystems, and for improving models that predict future vegetation based on continuous variation in plant form and function.

The influences of forest stand management on biotic and abiotic risks of damage

Abstract• This article synthesizes and reviews the available information on the effects of forestry practices on the occurrence of biotic and abiotic hazards, as well as on stand susceptibility to these damaging agents, concentrating on mammal herbivores, pest insects, pathogenic fungi, wind and fire.• The management operations examined are site selection, site preparation, stand composition, regeneration method, cleaning and weed control, thinning and pruning, and harvesting. For each of these operations we have examined how they influence the occurrence of biotic and abiotic damaging agents, the susceptibility of European forests, and describe the ecological processes that may explain these influences.• Overall, we find that the silvicultural operations that have the largest influence on both biotic and abiotic risks to European forest stands are closely related to species composition and the structure of the overstorey. Four main processes that drive the causal relationships between stand management and susceptibility have been identified: effect on local microclimate, provision of fuel and resources to biotic and abiotic hazards, enhancement of biological control by natural enemies and changes in individual tree physiology and development.• The review demonstrates an opportunity to develop silvicultural methods that achieve forest management objectives at the same time as minimising biotic and abiotic risks.Résumé• Cette revue bibliographique s’intéresse aux effets de la sylviculture sur la sensibilité des peuplements forestiers aux principaux agents de dégâts biotiques et abiotiques que sont les mammifères herbivores, les insectes ravageurs, les champignons pathogènes, le feu et les vents forts.• Les pratiques forestières analysées sont la sélection et la préparation des sites de reboisement, la définition de la composition en essences et le choix du matériel génétique, les méthodes de régénération et d’entretien, les modalités d’éclaircie et d’élagage, le mode de récolte finale. L’influence de chacune de ces opérations sur l’occurrence des agents de dégâts biotiques et abiotiques et sur la sensibilité des peuplements est examinée ainsi que les processus écologiques sous-jacents.• Les opérations sylvicoles qui se révèlent les plus déterminantes pour la sensibilité des forêts en Europe sont celles qui affectent la composition et la structure de la strate arborée. Quatre principaux processus écologiques semblent expliquer la relation entre sylviculture et sensibilité des peuplements : la modification du micro-climat, l’apport de ressources ou de combustible aux agents de dégâts, l’amélioration du contrôle biologique par les ennemis naturels et l’altération de la physiologie et du développement des arbres.• Cette revue permet donc d’envisager le développement de méthodes de gestion des peuplements forestiers qui permettent d’atteindre les objectifs de production tout en minimisant les risques de dégâts sanitaires.

Plant apparency, an overlooked driver of associational resistance to insect herbivory

Summary Herbivore regulation is one of the services provided by plant diversity in terrestrial ecosystems. It has been suggested that tree diversity decreases insect herbivory in forests, but recent studies have reported opposite patterns, indicating that tree diversity can trigger associational resistance or susceptibility. The mechanisms underlying the tree diversity–resistance relationship thus remain a matter of debate. We assessed insect herbivory on pedunculate oak saplings (Quercus robur) in a large-scale experiment in which we manipulated tree diversity and identity by mixing oaks, birch and pine species. Tree diversity at the plot scale had no effect on damage due to leaf chewers, but abundance of leaf miners decreased with increasing tree diversity. The magnitude of this associational resistance increased with host dilution, consistent with the ‘resource concentration hypothesis’. At a smaller scale, we estimated tree apparency as the difference in total height between focal oak saplings and their nearest neighbouring trees. Levels of oak infestation with leaf miners decreased significantly with decreasing tree apparency. As the probability of having taller neighbours increased with tree diversity, notably due to the increase in the proportion of faster growing nonhost trees, such as birches and pines, tree apparency may be seen as a ‘hidden’, sampling effect of tree diversity. Synthesis. These findings suggest that greater host dilution and lower tree apparency contribute to associational resistance in young trees. They also highlight the importance of taking plant size into account as a covariate, to avoid misleading interpretations about the biodiversity–resistance relationship.

Comparative responses of bird, carabid, and spider assemblages to stand and landscape diversity in maritime pine plantation forests

ABSTRACT We carried out a multi-taxa study to investigate the responses of species assemblages to stand and landscape variables in maritime pine plantation forests in the Landes de Gascogne (southwestern France). Breeding birds, carabid beetles, and ground-dwelling spiders were sampled simultaneously in a balanced set of 27 stands varying in tree species composition, age, and vertical structure. The composition and structure of surrounding landscapes were quantified within a 500-m radius around the stands. Species responses to a combination of 12 stand and landscape variables were estimated using multiple stepwise regression. Bird, carabid, and spider assemblages were related to the same explanatory variables using a set of co-inertia analyses. Tree height was the best predictive variable for all taxa at species richness, species assemblage, and individual species levels. Landscape variables were important as secondary factors, especially landscape fragmentation, spatial heterogeneity, and shape and spatial distribution of deciduous forest patches in the surrounding landscape. The occurrence of deciduous forest patches and an increasing landscape heterogeneity appeared to be beneficial to forest birds and spiders but not to carabids, probably because forest carabids are generally poor dispersers, more sensitive to habitat fragmentation.

Novel insect-tree associations resulting from accidental and intentional biological 'invasions': a meta-analysis of effects on insect fitness.

The translocation of species beyond their native range is a major threat to biodiversity. Invasions by tree-feeding insects attacking native trees and the colonization of introduced trees by native insects result in new insect-tree relationships. To date there is uncertainty about the key factors that influence the outcome of these novel interactions. We report the results of a meta-analysis of 346 pairwise comparisons of forest insect fitness on novel and ancient host tree species from 31 publications. Host specificity of insects and phylogenetic relatedness between ancient and novel host trees emerged as key factors influencing insect fitness. Overall, fitness was significantly lower on novel host species than on ancient hosts. However, in some cases, fitness increased on novel hosts, mainly in polyphagous insects or when close relatives of ancient host trees were colonized. Our synthesis enables greatly improved impact prediction and risk assessment of biological invasions.

Unraveling plant-animal diversity relationships: a meta-regression analysis.

In the face of unprecedented loss of biodiversity, cross-taxon correlates have been proposed as a means of obtaining quantitative estimates of biodiversity for identifying habitats of important conservation value. Habitat type, animal trophic level, and the spatial extent of studies would be expected to influence the strength of such correlations. We investigated these effects by carrying out a meta-analysis of 320 case studies of correlations between plant and animal species richnesses. The diversity of arthropods, herps, birds, and mammals significantly increased with plant diversity regardless of species habitat. However, correlations were stronger when plant and animal species richnesses were compared between habitats (gamma diversity) than within single habitats (alpha diversity). For arthropods, both the coefficient of correlation and the slope of the regression line were also greater for primary than for secondary consumers. These findings substantiate the use of plant species richness as an indicator of the diversity of animal taxa over space.

Species variables or environmental variables as indicators of forest biodiversity: a case study using saproxylic beetles in Maritime pine plantations

Abstract• To assess the sustainability of plantation forest management we compare two types of biodiversity indicators. We used the species richness of saproxylic beetles as a case study to test the “species” and “environmental” indicator approaches. We compared single species abundance or occurrence and deadwood volume or diversity as predictor variables.• Beetles were sampled with flight interception traps in 40 Maritime pine plantation stands. The volume and diversity of deadwood was estimated with line intersect and plot sampling in the same stands. Predictive models of species richness were built with simple linear or Partial Least Square regressions.• Deadwood variables appeared to be good predictors of saproxylic beetle richness at the stand-scale with at least 75% of variance explained. Deadwood diversity variables consistently provided better predictive models than volume variables. The best environmental indicator was the diversity of deadwood elements larger than 15 cm in diameter.• By contrast, the use of “species variables” appeared to be less relevant. To reach the quality of prediction obtained with “environmental variables”, the abundance or occurrence of 6 to 7 species — some of which are difficult to identify — had to be used to build the indicator.Résumé• Pour améliorer le suivi de la gestion durable des forêts de plantation, nous avons cherché à développer des indicateurs de biodiversité en prenant pour exemple la richesse en Coléoptères saproxyliques. Nous avons comparé l’approche directe, basée sur l’abondance ou l’occurrence de certaines espèces de coléoptères saproxyliques, et l’approche indirecte, basée sur les caractéristiques du bois mort comme variables prédictives.• Les Coléoptères ont été inventoriés à l’aide de pièges à interception dans 40 peuplements de pin maritime. Le volume et la diversité des pièces de bois mort ont été estimés à l’aide d’un échantillonnage par transects ou par placettes dans les mêmes plantations. Les modèles prédictifs ont été construits à l’aide de régressions linéaires simples ou PLS.• Les variables locales de bois mort apparaissent comme de bons prédicteurs de la richesse spécifique en coléoptères saproxyliques, avec jusqu’à 75% de la variance expliquée. Les variables de diversité du bois mort semblent de meilleurs prédicteurs que les variables volumiques. Le meilleur indicateur « indirect » pourrait être la diversité des pièces de bois mort de diamètre supérieur à 15 cm.• En revanche, le recours à des indicateurs « directs » semble moins prometteur. Pour obtenir une qualité de prédiction équivalente à l’approche indirecte, il faudrait prendre en compte l’abondance ou l’occurrence d’au moins 6 ou 7 espèces dont certaines sont difficilement identifiables.

Importance of semi-natural habitats for the conservation of butterfly communities in landscapes dominated by pine plantations

While the area of plantation forests continues to increase worldwide, their contribution to the conservation of biodiversity is still controversial. There is a particular concern on the central role played by natural habitat remnants embedded within the plantation matrix in conserving species-rich insect communities. We surveyed butterflies in maritime pine plantation landscapes in south-western France in 83 plots belonging to seven habitat types (five successional stages of pine stands, native deciduous woodlands and herbaceous firebreaks). The effect of plot, habitat and landscape attributes on butterfly species richness, community composition and individual species were analysed with a General Linear Model (GLM), partial Canonical Correspondence Analysis (CCA) and the IndVal method. The most important factors determining butterfly diversity and community composition were the presence of semi-natural habitats (deciduous woodlands and firebreaks) at the landscape scale and the composition of understorey vegetation at the plot scale. Pure effects of plot variables explained the largest part of community variation (12.8%), but landscape factors explained an additional, independent part (6.7%). Firebreaks were characterized by a higher species richness and both firebreaks and deciduous woodlands harboured species not or rarely found in pine stands. Despite the forest-dominated landscape, typical forest butterflies were rare and mainly found in the deciduous woodlands. Threatened species, such as Coenonympha oedippus and Euphydryas aurinia, were found in pine stands and in firebreaks, but were more abundant in the latter. In the studied plantation forest, the conservation of butterflies depends mainly on the preservation of semi-natural habitats, an adequate understorey management and the maintenance of soil moisture levels.

Genetic Diversity Increases Insect Herbivory on Oak Saplings

A growing body of evidence from community genetics studies suggests that ecosystem functions supported by plant species richness can also be provided by genetic diversity within plant species. This is not yet true for the diversity-resistance relationship as it is still unclear whether damage by insect herbivores responds to genetic diversity in host plant populations. We developed a manipulative field experiment based on a synthetic community approach, with 15 mixtures of one to four oak (Quercus robur) half-sib families. We quantified genetic diversity at the plot level by genotyping all oak saplings and assessed overall damage caused by ectophagous and endophagous herbivores along a gradient of increasing genetic diversity. Damage due to ectophagous herbivores increased with the genetic diversity in oak sapling populations as a result of higher levels of damage in mixtures than in monocultures for all families (complementarity effect) rather than because of the presence of more susceptible oak genotypes in mixtures (selection effect). Assemblages of different oak genotypes would benefit polyphagous herbivores via improved host patch location, spill over among neighbouring saplings and diet mixing. By contrast, genetic diversity was a poor predictor of the abundance of endophagous herbivores, which increased with individual sapling apparency. Plant genetic diversity may not provide sufficient functional contrast to prevent tree sapling colonization by specialist herbivores while enhancing the foraging of generalist herbivores. Long term studies are nevertheless required to test whether the effect of genetic diversity on herbivory change with the ontogeny of trees and local adaptation of specialist herbivores.