A meta‐analysis of functional group responses to forest recovery outside of the tropics

Both active and passive forest restoration schemes are used in degraded landscapes across the world to enhance biodiversity and ecosystem service provision. Restoration is increasingly also being implemented in biodiversity offset schemes as compensation for loss of natural habitat to anthropogenic development. This has raised concerns about the value of replacing old‐growth forest with plantations, motivating research on biodiversity recovery as forest stands age. Functional diversity is now advocated as a key metric for restoration success, yet it has received little analytical attention to date. We conducted a meta‐analysis of 90 studies that measured differences in species richness for functional groups of fungi, lichens, and beetles between old‐growth control and planted or secondary treatment forests in temperate, boreal, and Mediterranean regions. We identified functional‐group–specific relationships in the response of species richness to stand age after forest disturbance. Ectomycorrhizal fungi averaged 90 years for recovery to old‐growth values (between 45 years and unrecoverable at 95% prediction limits), and epiphytic lichens took 180 years to reach 90% of old‐growth values (between 140 years and never for recovery to old‐growth values at 95% prediction limits). Non‐saproxylic beetle richness, in contrast, decreased as stand age of broadleaved forests increased. The slow recovery by some functional groups essential to ecosystem functioning makes old‐growth forest an effectively irreplaceable biodiversity resource that should be exempt from biodiversity offsetting initiatives.     

The Potential for Species Conservation in Tropical Secondary Forests

Abstract: In the wake of widespread loss of old‐growth forests throughout the tropics, secondary forests will likely play a growing role in the conservation of forest biodiversity. We considered a complex hierarchy of factors that interact in space and time to determine the conservation potential of tropical secondary forests. Beyond the characteristics of local forest patches, spatial and temporal landscape dynamics influence the establishment, species composition, and persistence of secondary forests. Prospects for conservation of old‐growth species in secondary forests are maximized in regions where the ratio of secondary to old‐growth forest area is relatively low, older secondary forests have persisted, anthropogenic disturbance after abandonment is relatively low, seed‐dispersing fauna are present, and old‐growth forests are close to abandoned sites. The conservation value of a secondary forest is expected to increase over time, as species arriving from remaining old‐growth forest patches accumulate. Many studies are poorly replicated, which limits robust assessments of the number and abundance of old‐growth species present in secondary forests. Older secondary forests are not often studied and few long‐term studies are conducted in secondary forests. Available data indicate that both old‐growth and second‐growth forests are important to the persistence of forest species in tropical, human‐modified landscapes.     

Recovery of Faunal Communities During Tropical Forest Regeneration

Abstract:  As mature tropical forests are cleared, secondary forests may play an important role in the conservation of animal species, depending on how fast animal communities recover during forest regeneration. I reviewed published studies on the recovery of animal species richness and composition during tropical forest regeneration. In 38 of the 39 data sets I examined, conversion of forest to agriculture or pasture substantially reduced species richness. Given suitable conditions for forest recovery, the species richness of the animal taxa considered can be predicted to resemble that of mature forests roughly 20–40 years after land abandonment. At least for ants and birds, however, recovery of species composition appears to take substantially longer than recovery of species richness. Because species richness for many taxa appears to recover relatively rapidly in secondary forests, conservation of secondary forests may be an effective investment in future diversity. The slower recovery of species composition indicates, however, that some species will require stands of mature forest to persist.     

Restoration of ecosystems for biodiversity and carbon sequestration: Simulating growth dynamics of brigalow vegetation communities in Australia

Restoration of abandoned and degraded ecosystems through enhanced management of mature remnant patches and naturally regenerating (regrowth) forests is currently being used in the recovery of ecosystems for biodiversity protection and carbon sequestration. Knowledge of long-term dynamics of these ecosystems is often very limited. Vegetation models that examine long-term forest growth and succession of uneven aged, mixed-species forest ecosystems are integral to the planning and assessment of the recovery process of biodiversity values and biomass accumulation. This paper examined the use of the Ecosystem Dynamics Simulator (EDS) in projecting growth dynamics of mature remnant brigalow forest communities and recovery process of regrowth brigalow thickets. We used data from 188 long-term monitored plots of remnant and regrowth forests measured between 1963 and 2010. In this study the model was parameterised for 34 tree and shrub species and tested with independent long-term measurements. The model closely approximated actual development trajectories of mature forests and regrowth thickets but some inaccuracies in estimating regeneration through asexual reproduction and mortality were noted as reflected in stem density projections of remnant plots that had a mean of absolute relative bias of 46.2 (±12.4)%. Changes in species composition in remnant forests were projected with a 10% error. Basal area values observed in all remnant plots ranged from 6 to 29 m² ha⁻¹ and EDS projections between 1966 and 2005 (39 years) were 68.2 (±10.9)% of the observed basal area. Projected live aboveground biomass of remnant plots had a mean of 93.5 (±5.9) t ha⁻¹ compared to a mean of 91.3 (±8.0) t ha⁻¹ observed in the plots. In regrowth thicket, the model produced satisfactory projections of tree density (91%), basal area (89%), height (87%) and aboveground biomass (84%) compared to the observed attributes. Basal area and biomass accumulation in 45-year-old regrowth plots was approximately similar to that in remnant forests but recovery of woody understorey was very slow. The model projected that it would take 95 years for the regrowth to thin down to similar densities observed in original or remnant brigalow forests. These results indicated that EDS can produce relatively accurate projections of growth dynamics of brigalow regrowth forests necessary for informing restoration planning and projecting biomass accumulation.     

Slow recovery of tropical old‐field rainforest regrowth and the value and limitations of active restoration

There is current debate about the potential for secondary regrowth to rescue tropical forests from an otherwise inevitable cascade of biodiversity loss due to land clearing and scant evidence to test how well active restoration may accelerate recovery. We used site chronosequences to compare developmental trajectories of vegetation between self‐organized (i.e., spontaneous) forest regrowth and biodiversity plantings (established for ecological restoration, with many locally native tree species at high density) in the Australian wet tropics uplands. Across 28 regrowth sites aged 1–59 years, some structural attributes reached reference rainforest levels within 40 years, whereas wood volume and most tested components of native plant species richness (classified by species’ origins, family, and ecological functions) reached less than 50% of reference rainforest values. Development of native tree and shrub richness was particularly slow among species that were wind dispersed or animal dispersed with large (>10 mm) seeds. Many species with animal‐dispersed seeds were from near‐basal evolutionary lineages that contribute to recognized World Heritage values of the study region. Faster recovery was recorded in 25 biodiversity plantings of 1–25 years in which wood volume developed more rapidly; native woody plant species richness reached values similar to reference rainforest and was better represented across all dispersal modes; and species from near‐basal plant families were better (although incompletely) represented. Plantings and regrowth showed slow recovery in species richness of vines and epiphytes and in overall resemblance to forest in species composition. Our results can inform decision making about when and where to invest in active restoration and provide strong evidence that protecting old‐growth forest is crucially important for sustaining tropical biodiversity.     

Directional changes in the species composition of a tropical forest

Long-term studies have revealed that the structure and dynamics of many tropical forests are changing, but the causes and consequences of these changes remain debated. To learn more about the forces driving changes within tropical forests, we investigated shifts in tree species composition over the past 25 years within the 50-ha Forest Dynamics Plot on Barro Colorado Island (BCI), Panama, and examined how observed patterns relate to predictions of (1) random population fluctuations, (2) carbon fertilization, (3) succession from past disturbance, (4) recovery from an extreme El Ni?o drought at the start of the study period, and (5) long-term climate change. We found that there have been consistent and directional changes in the tree species composition. These shifts have led to increased relative representations of drought-tolerant species as determined by the species' occurrence both across a gradient of soil moisture within BCI and across a wider precipitation gradient from a dry forest near the Pacific coast of Panama to a wet forest near its Caribbean coast. These nonrandom changes cannot be explained by stochastic fluctuations or carbon fertilization. They may be the legacy of the El Ni?o drought, or alternatively, potentially reflect increased aridity due to long-term climate change. By investigating compositional changes, we increased not only our understanding of the ecology of tropical forests and their responses to large-scale disturbances, but also our ability to predict how future global change will impact some of the critical services provided by these important ecosystems.     

Invertebrates and Boreal Forest Management

Logging has ecological effects on invertebrates in Fennoscandian boreal forest. Especially affected are species associated with micro-habitats of natural old-growth forest, such as coarse woody debris, large deciduous trees, and patches of wet swamp-forest. Furthermore, the effective control of forest fires has adversely affected many species that require burned substrates. Although old-growth specialists tend to disappear from clear-cuts, local invertebrate species richness (α-diversity) may increase as forest generalists persist and numerous open-habitat species appear. This seems true at least for taxa (e.g., ground-beetles and spiders) that are not directly associated with the characteristic micro-habitats of old growth mentioned above. However, at the landscape or biogeographical scale intensive logging tends to homogenize forest habitats and lead to declines of sensitive species. Based on these findings three complementary approaches appear useful for the maintenance of boreal biodiversity while harvesting timber. (1) Undisturbed old-growth forests must be set aside to sustain specialist species and to serve as sources for recolonization. (2) Because these reserves are not, and will probably not be, large and regionally representative enough, ecologically sound silvicultural practices must be developed using natural disturbance regimes as guidelines. (3) Restoration of habitats and the process of natural regeneration through fire is imperative for the recovery of associated species assemblages. However, the process of recovery presupposes that colonists are available, which may not always be the case.     

Predicting landscape-scale biodiversity recovery by natural tropical forest regrowth

Natural forest regrowth is a cost-effective, nature-based solution for biodiversity recovery, yet different socioenvironmental factors can lead to variable outcomes. A critical knowledge gap in forest restoration planning is how to predict where natural forest regrowth is likely to lead to high levels of biodiversity recovery, which is an indicator of conservation value and the potential provisioning of diverse ecosystem services. We sought to predict and map landscape-scale recovery of species richness and total abundance of vertebrates, invertebrates, and plants in tropical and subtropical second-growth forests to inform spatial restoration planning. First, we conducted a global meta-analysis to quantify the extent to which recovery of species richness and total abundance in second-growth forests deviated from biodiversity values in reference old-growth forests in the same landscape. Second, we employed a machine-learning algorithm and a comprehensive set of socioenvironmental factors to spatially predict landscape-scale deviation and map it. Models explained on average 34% of observed variance in recovery (range 9–51%). Landscape-scale biodiversity recovery in second-growth forests was spatially predicted based on socioenvironmental landscape factors (human demography, land use and cover, anthropogenic and natural disturbance, ecosystem productivity, and topography and soil chemistry); was significantly higher for species richness than for total abundance for vertebrates (median range-adjusted predicted deviation 0.09 vs. 0.34) and invertebrates (0.2 vs. 0.35) but not for plants (which showed a similar recovery for both metrics [0.24 vs. 0.25]); and was positively correlated for total abundance of plant and vertebrate species (Pearson r = 0.45, p = 0.001). Our approach can help identify tropical and subtropical forest landscapes with high potential for biodiversity recovery through natural forest regrowth.     

The Value of Primary, Secondary, and Plantation Forests for a Neotropical Herpetofauna

Abstract:  Plantation forests and second-growth forests are becoming dominant components of many tropical forest landscapes. Yet there is little information available concerning the consequences of different forestry options for biodiversity conservation in the tropics. We sampled the leaf-litter herpetofauna of primary, secondary, and Eucalyptus plantation forests in the Jari River area of northeastern Brazilian Amazonia. We used four complementary sampling techniques, combined samples from 2 consecutive years, and collected 1739 leaf-litter amphibians (23 species) and 1937 lizards (30 species). We analyzed the data for differences among forest types regarding patterns of alpha and beta diversity, species-abundance distributions, and community structure. Primary rainforest harbored significantly more species, but supported a similar abundance of amphibians and lizards compared with adjacent areas of second-growth forest or plantations. Plantation forests were dominated by wide-ranging habitat generalists. Secondary forest faunas contained a number of species characteristic of primary forest habitat. Amphibian communities in secondary forests and Eucalyptus plantations formed a nested subset of primary forest species, whereas the species composition of the lizard community in plantations was distinct, and was dominated by open-area species. Although plantation forests are relatively impoverished, naturally regenerating forests can help mitigate some negative effects of deforestation for herpetofauna. Nevertheless, secondary forest does not provide a substitute for primary forest, and in the absence of further evidence from older successional stands, we caution against the optimistic claim that natural forest regeneration in abandoned lands will provide refuge for the many species that are currently threatened by deforestation .     

Wood Density as a Conservation Tool: Quantification of Disturbance and Identification of Conservation‐Priority Areas in Tropical Forests

Abstract: Inventories of tree species are often conducted to guide conservation efforts in tropical forests. Such surveys are time consuming, demanding of expertise, and expensive to perform and interpret. Approaches to make survey efforts simpler or more effective would be valuable. In particular, it would be good to be able to easily identify areas of old‐growth forest. The average density of the wood of a tree species is closely linked to its successional status. We used tree inventory data from eastern Borneo to determine whether wood density can be used to quantify forest disturbance and conservation importance. The average density of wood in a plot was significantly and negatively related to disturbance levels, with plots with higher wood densities occurring almost exclusively in old‐growth forests. Average wood density was unimodally related to the diversity of tree species, indicating that the average wood density in a plot might be a better indicator of old‐growth forest than species diversity. In addition, Borneo endemics had significantly heavier wood than species that are common throughout the Malesian region, and they were more common in plots with higher average wood density. We concluded that wood density at the plot level could be a powerful tool for identifying areas of conservation priority in the tropical rain forests of Southeast Asia.     

Effects of New Forestry Practices on Rare Epiphytic Macrolichens

Abstract:  The preservation of key habitats is included in modern forest-management practices as one of the main means to preserve biodiversity in northern European production forests. I examined the distribution patterns and persistence of occurrence of rare epiphytic macrolichens in a predominantly unprotected forest landscape with an area of 278 km² in southern Finland. Occurrence data of macrolichens in two key habitats, rock outcrops and mires, were compared with data from production forests. The density of populations of rare macrolichens was 50-fold higher on rock outcrops and 25-fold higher on mires than in production forests. Most rare species of epiphytic macrolichens were found only in key habitats. In modern, intensively managed forest landscapes, rock outcrops appeared to represent the main habitats for rare macrolichens as a result of the long-term continuity of old trees. Most macrolichen occurrences represented very small populations with a high extinction risk. Of the populations found in 1989–1995 in key habitats, 51% had disappeared, by 2000–2001, mainly as a result of forestry activities. The disappearance of populations significantly exceeded the rate of establishment of new populations. My results emphasize the importance of key habitats for declining forest species. There is also a need to improve and sharpen the guidelines for delimiting and managing key habitats in order to halt the continued decline of epiphytic macrolichens.     

Effects of Timber Harvesting on Southern Appalachian Salamanders

We compared the species richness and abundance of salamanders on six recent clearcuts (< 10 years old) with that of salamanders on 34 mature forest stands (>50 years old) in southern Appalachian forests in western North Carolina, U.S.A. Catches of salamanders from plots in mature forest stands were about five times higher than those on recent clearcuts. Almost all species and major taxonomic groups of salamanders were adversely affected by timber removal. Mean number of species collected per plot was about twice as great in mature forest stands as in clearcuts. Analyses of stand age versus salamander catch for 47 plots indicate that 50–70 years are required for populations to return to predisturbance levels following cutting. We conservatively estimate that clearcutting in U.S. national forests in western North Carolina results in a loss of nearly 14 millton salamanders annually. It also is chronically reducing regional populations by more than a quarter of a billion salamanders (9%) below that which could be sustained if mature forests were not cut.     

Changes in Forest Use Value through Ecological Succession and Their Implications for Land Management in the Peruvian Amazon

Abstract:  Research on local use values of forests across an ecological succession informs land-use decisions and conservation planning. I evaluated use values of three age classes of secondary forest: fallow fields (<5 years old, $8.20/ha/year), young secondary forest (5–20 years old, $20.60/ha/year), and old secondary forest (>20 years old, $6.80/ha/year). I quantified daily forest product use and calculated use values in dollars per hectare per year for three communities in the northern Peruvian Amazon. I made three comparisons between forest types: number of useful species, value based on different use categories, and overall use values. Old secondary forest had the greatest number of total species present and species collected. Wood, food, and medicine were the three most valuable use categories. The value different families extracted from local forests varied enormously, but median forest values were lower for all forest types than potential gains from agricultural land use (e.g., coffee $167/ha/year). Values of different-aged stands on privately owned lands in two communities did not differ significantly, whereas in the third community, young secondary forest had a significantly greater value than other forest types. Old secondary forests were the most valuable source of wood products, and wood was the only use category in which there was any difference in the value of products extracted from different-aged forest stands. The value of all three forest types on open-access (nonprivate) lands was minimal (mean in each forest type, $0/ha/year). Local people can utilize the valuation results to develop land-use strategies that balance forest product use, agricultural productivity, and biodiversity conservation.     

Rethinking Tropical Forest Conservation: Perils in Parks

Abstract: According to some conservationists, large, pristine, uninhabited parks are the defining criterion of success in conserving tropical forests. They argue that human residents in tropical forests inevitably deplete populations of large animals through hunting, which triggers a chain reaction of ecological events that greatly diminish the conservation value of these forests. Hence, they believe that removal of people from tropical forests is an essential step in the creation of successful parks and in the conservation of nature in the tropics. This approach can lead to undesirable consequences, however. Forest residents—and rural people generally—are potent political actors in tropical forest regions and an essential component of the environmental political constituencies that are necessary for the long-term conservation of tropical forests. In Amazonia and elsewhere, rural people are defending far bigger areas of tropical forest from unfettered deforestation and logging than are parks, thereby conserving the ecological services provided by these forests and the majority of their component plant and animal species. Moreover, the data are too sparse to judge the effects of forest peoples on populations of large forest animals. The establishment of pristine, tropical forest parks is an important conservation goal, but the exclusive pursuit of this goal undermines the broader objectives of conservation when it identifies forest residents and other rural people as the enemies of nature.     

Ecological consequences of forest elephant declines for Afrotropical forests

Poaching is rapidly extirpating African forest elephants (Loxodonta cyclotis) from most of their historical range, leaving vast areas of elephant‐free tropical forest. Elephants are ecological engineers that create and maintain forest habitat; thus, their loss will have large consequences for the composition and structure of Afrotropical forests. Through a comprehensive literature review, we evaluated the roles of forest elephants in seed dispersal, nutrient recycling, and herbivory and physical damage to predict the cascading ecological effects of their population declines. Loss of seed dispersal by elephants will favor tree species dispersed abiotically and by smaller dispersal agents, and tree species composition will depend on the downstream effects of changes in elephant nutrient cycling and browsing. Loss of trampling and herbivory of seedlings and saplings will result in high tree density with release from browsing pressures. Diminished seed dispersal by elephants and high stem density are likely to reduce the recruitment of large trees and thus increase homogeneity of forest structure and decrease carbon stocks. The loss of ecological services by forest elephants likely means Central African forests will be more like Neotropical forests, from which megafauna were extirpated thousands of years ago. Without intervention, as much as 96% of Central African forests will have modified species composition and structure as elephants are compressed into remaining protected areas. Stopping elephant poaching is an urgent first step to mitigating these effects, but long‐term conservation will require land‐use planning that incorporates elephant habitat into forested landscapes that are being rapidly transformed by industrial agriculture and logging.     

Two‐Stage Recovery of Amphibian Assemblages Following Selective Logging of Tropical Forests

There is a lack of quantitative information on the effectiveness of selective‐logging practices in ameliorating effects of logging on faunal communities. We conducted a large‐scale replicated field study in 3 selectively logged moist semideciduous forests in West Africa at varying times after timber extraction to assess post logging effects on amphibian assemblages. Specifically, we assessed whether the diversity, abundance, and assemblage composition of amphibians changed over time for forest‐dependent species and those tolerant of forest disturbance. In 2009, we sampled amphibians in 3 forests (total of 48 study plots, each 2 ha) in southwestern Ghana. In each forest, we established plots in undisturbed forest, recently logged forest, and forest logged 10 and 20 years previously. Logging intensity was constant across sites with 3 trees/ha removed. Recently logged forests supported substantially more species than unlogged forests. This was due to an influx of disturbance‐tolerant species after logging. Simultaneously Simpson's index decreased, with increased in dominance of a few species. As time since logging increased richness of disturbance‐tolerant species decreased until 10 years after logging when their composition was indistinguishable from unlogged forests. Simpson's index increased with time since logging and was indistinguishable from unlogged forest 20 years after logging. Forest specialists decreased after logging and recovered slowly. However, after 20 years amphibian assemblages had returned to a state indistinguishable from that of undisturbed forest in both abundance and composition. These results demonstrate that even with low‐intensity logging (≤3 trees/ha) a minimum 20‐year rotation of logging is required for effective conservation of amphibian assemblages in moist semideciduous forests. Furthermore, remnant patches of intact forests retained in the landscape and the presence of permanent brooks may aid in the effective recovery of amphibian assemblages. Recuperación de Ensambles de Anfibios en Dos Etapas Después de la Tala Selectiva de Bosques Tropicales     

Conservation of Epiphyte Diversity in an Andean Landscape Transformed by Human Land Use

Abstract: Epiphytes are diverse and important elements of tropical forests, but as canopy‐dwelling organisms, they are highly vulnerable to deforestation. To assess the effect of deforestation on epiphyte diversity and the potential for epiphyte conservation in anthropogenically transformed habitats, we surveyed the epiphytic vegetation of an Ecuadorian cloud forest reserve and its surroundings. Our study was located on the western slopes of the Andes, a global center of biodiversity. We sampled vascular epiphytes of 110 study plots in a continuous primary forest; 14 primary forest fragments; isolated remnant trees in young, middle‐aged, and old pastures; and young and old secondary forests. It is the first study to include all relevant types of habitat transformation at a single study site and to compare epiphyte diversity at different temporal stages of fragmentation. Epiphyte diversity was highest in continuous primary forest, followed by forest fragments and isolated remnant trees, and lowest in young secondary forests. Spatial parameters of habitat transformation, such as fragment area, distance to the continuous primary forest, or distance to the forest edge from inside the forest, had no significant effect on epiphyte diversity. Hence, the influence of dispersal limitations appeared to be negligible or appeared to operate only over very short distances, whereas microclimatic edge effects acted only in the case of completely isolated trees, but not in larger forest fragments. Epiphyte diversity increased considerably with age of secondary forests, but species assemblages on isolated remnant trees were impoverished distinctly with time since isolation. Thus, isolated trees may serve for recolonization of secondary forests, but only for a relatively short time. We therefore suggest that the conservation of even small patches of primary forest within agricultural landscape matrices is essential for the long‐term maintenance of the high epiphyte diversity in tropical cloud forests.     

Multidimensional trade-offs in species responses to disturbance: implications for diversity in a subtropical forest

Species employ diverse strategies to cope with natural disturbance, but the importance of these strategies for maintaining tree species diversity in forests has been debated. Mechanisms that have the potential to promote tree species coexistence in the context of repeated disturbance include life history trade-offs in colonization and competitive ability or in species' ability to survive at low resource conditions and exploit the temporary resource-rich conditions often generated in the wake of disturbance (successional niche). Quantifying these trade-offs requires long-term forest monitoring and modeling. We developed a hierarchical Bayes model to investigate the strategies tree species employ to withstand and recover from hurricane disturbance and the life history trade-offs that may facilitate species coexistence in forests subject to repeated hurricane disturbance. Unlike previous approaches, our model accommodates temporal variation in process error and observations from multiple sources. We parameterized the model using growth and mortality data from four censuses of a 16-ha plot taken every five years (1990-2005), together with damage data collected after two hurricanes and annual seed production data (1992-2005). Species' susceptibilities to hurricane damage as reflected by changes in diameter growth and fecundity immediately following a storm were weak, highly variable, and unpredictable using traditional life history groupings. The lower crowding conditions (e.g., high light) generated in the wake of storms, however, led to greater gains in growth and fecundity for pioneer and secondary-forest species than for shade-tolerant species, in accordance with expectation of life history. We found moderate trade-offs between survival in high crowding conditions, a metric of competitive ability, and long-distance colonization. We also uncovered a strong trade-off between mean species fecundity in low crowding conditions, a metric of recovery potential, and competitive ability. Trade-offs in competitive and colonization ability, in addition to successional niche processes, are likely to contribute to species persistence in these hurricane-impacted forests. The strategies species employ to cope with hurricane damage depend on the degree to which species rely on sprouting, repair of adult damage, changes in demographic rates in response to enhanced resource availability after storms, or long-distance dispersal as recovery mechanisms.     

Biodiversity Differences between Managed and Unmanaged Forests: Meta-Analysis of Species Richness in Europe

Abstract: Past and present pressures on forest resources have led to a drastic decrease in the surface area of unmanaged forests in Europe. Changes in forest structure, composition, and dynamics inevitably lead to changes in the biodiversity of forest‐dwelling species. The possible biodiversity gains and losses due to forest management (i.e., anthropogenic pressures related to direct forest resource use), however, have never been assessed at a pan‐European scale. We used meta‐analysis to review 49 published papers containing 120 individual comparisons of species richness between unmanaged and managed forests throughout Europe. We explored the response of different taxonomic groups and the variability of their response with respect to time since abandonment and intensity of forest management. Species richness was slightly higher in unmanaged than in managed forests. Species dependent on forest cover continuity, deadwood, and large trees (bryophytes, lichens, fungi, saproxylic beetles) and carabids were negatively affected by forest management. In contrast, vascular plant species were favored. The response for birds was heterogeneous and probably depended more on factors such as landscape patterns. The global difference in species richness between unmanaged and managed forests increased with time since abandonment and indicated a gradual recovery of biodiversity. Clearcut forests in which the composition of tree species changed had the strongest effect on species richness, but the effects of different types of management on taxa could not be assessed in a robust way because of low numbers of replications in the management‐intensity classes. Our results show that some taxa are more affected by forestry than others, but there is a need for research into poorly studied species groups in Europe and in particular locations. Our meta‐analysis supports the need for a coordinated European research network to study and monitor the biodiversity of different taxa in managed and unmanaged forests.     

The role of canopy structural complexity in wood net primary production of a maturing northern deciduous forest

The even-aged northern hardwood forests of the Upper Great Lakes Region are undergoing an ecological transition during which structural and biotic complexity is increasing. Early-successional aspen (Populus spp.) and birch (Betula papyrifera) are senescing at an accelerating rate and are being replaced by middle-successional species including northern red oak (Quercus rubra), red maple (Acer rubrum), and white pine (Pinus strobus). Canopy structural complexity may increase due to forest age, canopy disturbances, and changing species diversity. More structurally complex canopies may enhance carbon (C) sequestration in old forests. We hypothesize that these biotic and structural alterations will result in increased structural complexity of the maturing canopy with implications for forest C uptake. At the University of Michigan Biological Station (UMBS), we combined a decade of observations of net primary productivity (NPP), leaf area index (LAI), site index, canopy tree-species diversity, and stand age with canopy structure measurements made with portable canopy lidar (PCL) in 30 forested plots. We then evaluated the relative impact of stand characteristics on productivity through succession using data collected over a nine-year period. We found that effects of canopy structural complexity on wood NPP (NPPw) were similar in magnitude to the effects of total leaf area and site quality. Furthermore, our results suggest that the effect of stand age on NPPw is mediated primarily through its effect on canopy structural complexity. Stand-level diversity of canopy-tree species was not significantly related to either canopy structure or NPPw. We conclude that increasing canopy structural complexity provides a mechanism for the potential maintenance of productivity in aging forests.