Centuries-old logging legacy on spatial and temporal patterns in understory herb communities

Understory herb communities in the Southern Appalachians are among the highest biodiversity plant communities in North America. In the mid-1990s, a debate began over whether understory herb communities recover to their pre-disturbance states following logging. Studies showing reduced herb-layer diversity in previously logged forests were criticized for not accounting for intersite environmental heterogeneity. More recent studies have addressed environmental heterogeneity, but have neglected long-term recovery by using “mature forests” as young as 80 years old as the benchmark for diversity comparison, even though old growth stands have disturbance return intervals exceeding 500 years. Here we address concerns clouding previous studies of high-diversity Appalachian herb communities and investigate their long-term recovery by comparing paired sites of old growth forest and forest logged 100–150 years ago. We found that species richness and individual abundance is greater in old growth forests than mature forests and that species composition differed significantly between the two. Turnover in species among old growth and mature forests accounted for 11% of the total species richness and was significantly greater than expected. Species turnover at intermediate (5–50 m) and landscape-scales (>10 km) contributed the most towards total species richness. Herb communities in rich cove forests have successional trajectories that exceed 150 years, with important community changes still occurring long after the forest returns to what has been previously termed a “mature” state. To conserve the diverse herb layer, we conclude that mature forest stands are too young to serve as baselines for recovery, landscape-scale preservation of multiple forest stands is needed to maximize species richness, and maintaining 100–150-year logging rotations will likely lead to loss of biodiversity.     

Predicting canopy macrolichen diversity and abundance within old-growth inland temperate rainforests

Windward slopes of the inland mountain ranges in British Columbia support a unique inland temperate rainforest (ITR) ecosystem. Increasing fragmentation and the loss of old ITR stands have highlighted the need for determining conservation biology priorities among remaining old forest stands. We have addressed this issue by surveying foliose macrolichens within 53 old ITR stands in British Columbia's 135,000 ha very wet-cool interior-cedar hemlock (ICHvk2) biogeoclimatic subzone in the upper Fraser River watershed. Study plots were stratified by leading tree species and by “wet” versus “dry” relative soil moisture conditions. Other plot variables included: temperature, precipitation, incident solar loading, and canopy openness. Ordination plots showed a distinct assemblage of foliose cyanolichens, including Lobaria pulmonaria, Lobaria retigera, Sticta fuliginosa, Nephroma isidiosum, Nephroma occultum, and Pseudocyphellaria anomala, whose abundance was correlated with increasing relative soil moisture, temperature, canopy openness, precipitation, and basal area of spruce. Logistic regression models similarly identified relative soil moisture and temperature in all parsimonious models. Leading tree species, in combination with “wet” relative soil moisture and/or temperature, were important factors explaining the presence or absence of five (Cavernularia hultenii, L. retigera, N. occultum, Platismatia norvegica, and Sticta oroborealis) of the eight modeled old-growth associate lichen species. This combination of conditions favouring the development of canopy lichen communities in old forests was best expressed in low elevation water receiving sites. We hypothesize that groundwater availability in these sites promotes species richness and abundance of canopy lichens by creating more favourable conditions for growth, and by reducing fire return intervals which allows for the accumulation of rare species over time. Historically, forests in these wet “toe-slope” positions were disproportionately targeted for logging. Their conservation should now be a high priority, given their disproportionate significance to maintaining canopy lichen diversity in the present-day landscapes.     

Can native vegetation recover after slash pine cultivation in the Brazilian Savanna?

There is a widespread view that forest plantations with exotic species are green deserts, unable to sustain biodiversity. Few studies have demonstrated, however, that planted stands of exotic trees have a greater negative effect on the plant diversity of savanna vegetation. We compared the native woody flora under four stands of slash pine of about 45 years old with four stands where the previously existing native Cerrado vegetation was preserved and protected from disturbances for the same period, has changed into dense vegetation - the “cerradão”, at Assis municipality, São Paulo State, Brazil. Aiming at understanding the potential ecological filters driving these communities, we assessed air and soil humidity, light availability and classified the native species on the basis of shade tolerance, dispersal syndrome and biomes in which they occur (Atlantic Forest or Cerrado). We recorded an average of 70 (±13) species under pine stands and 54 (±16) species in cerradão. Of the total of 136 species recorded, 78 occurred in both habitats, eight were exclusive to the “cerradão” (shade tolerant and also occurring in forest ecosystems) and 18 were recorded only under pine stands (82% heliophytic, exclusive to the Cerrado biome). Among the functional attributes and abiotic variables analyzed, only light availability explained the floristic differences found. Since richness was higher under pine, we refuted the hypothesis that exotic species constrain the establishment of the native species richness in the understory. On the other hand, the dark environment under the closed-canopy of the “cerradão” acts as a filter inhibiting the establishment of typical Cerrado species. Since pine stands, if managed in long cycle, maintain a reasonable pool of Cerrado endemic species in the understory pine plantations may be a good starting point for savanna restoration.     

Epiphytic lichen diversity in old-growth and managed Picea abies stands in Alpine spruce forests

In the last decades, a large body of literature has grown to evaluate the impact of forest management on epiphytic lichens in boreal coniferous forests. However, information is still lacking on coniferous forests of the Alps. This study compares lichen diversity between spruce forest stands of four successional stages: (1) young, (2) intermediate, (3) mature forests managed for timber production with a rotation cycle of 120–180 years, and (4) old-growth protected forests. The emphasis was placed on the occurrence of nationally rare and calicioid species (lichens and fungi traditionally referred to as Caliciales, known to be indicative of forest age and continuity). For each forest successional stage, four plots were selected. In each plot, 7 spruce individuals were surveyed for epiphytic lichens according to a standardised sampling method. Species richness increased from young to mature stands, while no difference was detected between mature and old-growth stands. This pattern was also confirmed for rare and calicioid species which are, however, more frequent in old-growth stands. Differences in species composition were also found between the different forest successional stages. Mature and old-growth plots slightly overlap, indicating that to some extent comparable lichen assemblages could be found in these stands. A nested pattern of species assemblages was found, old-growth stands hosting most of the species which were also found in stands belonging to the previous forest successional stages. Our results support the hypothesis that the management regime applied to spruce forests of the Italian Alps renders mature stands managed for timber production somewhat similar to old-growth stands as lichen habitat. However, we found a higher complexity in old-growth forests, and many species of conservation concern clearly preferred old-growth stands. In this perspective, a further prolongation of the normal cycle it is likely to be a most favourable conservation-oriented management to be recommended at least within protected areas and Natura 2000 sites, where conservation purposes should receive a high priority.     

Disturbance changes arbuscular mycorrhizal fungal phenology and soil glomalin concentrations but not fungal spore composition in montane rainforests in Veracruz and Chiapas,Mexico

Mexican montane rainforests and adjacent disturbed areas were studied for disturbance-related spatio-temporal changes to the arbuscular mycorrhizal fungal (AMF) community and soil glomalin concentration. The AMF community functions to both improve plant growth and soil conditions and is thus an important component to the restoration of this forest type to disturbed areas. The study areas included mature rainforests that were converted to pine forests, milpas, pastures and shrub/herbaceous plant communities via burning and logging. Seasonal patterns in AMF spore species richness and sporulation significantly differed across disturbance types at two of the three sites surveyed. Contrasting patterns of sporulation among AMF families across different disturbance types helped to explain how species richness and composition were maintained despite dramatic changes to the host plant community. Meaning, in most cases, disturbance induced changes in when different AMF taxa sporulated but not what taxa sporulated. Only conversion from mature pine–oak–Liquidambar–Persea forests to pine-dominated stands severely reduced AMF spore richness and total sporulation. Surprisingly, in pine-dominant stands no concomitant negative impacts on soil glomalin (MAb32B11 immunoreactive soil protein) concentrations were detected. However, soils of mature forests containing no pines had the highest concentration of glomalin. Conversion to pasture and milpa (diverse cornfield) had a strong negative impact on the concentration of soil glomalin concentrations. In sharp contrast, the same disturbance types improved AMF sporulation and AMF spore richness. It appears that disturbance type, and not AMF community measures used herein, best predicts changes in soil glomalin concentration.     

Diversity and succession of adventive and indigenous vascular understorey plants in Pinus radiata plantation forests in New Zealand

The vegetation of Pinus radiata plantation forests in New Zealand was studied to examine how the indigenous flora has responded to this novel habitat. A chronosequence of stands about 5, 16 and 27 years was assessed in each of four different biogeographic regions to test the effects of several stand and site factors on the succession of vascular understorey plant communities. A total of 202 indigenous and 70 adventive vascular plant species were found across all study areas, with considerable geographic variation among forests in species composition, species richness (range 48–135 species), and the percentage of indigenous species (50–86%). Both richness and cover of adventive species decreased significantly over time, whereas richness and cover of indigenous species was highest in the oldest stands, and overall species richness was lowest at mid-rotation. The guild composition changed from dominance of grasses and forbs in young stands to dominance of ferns and understorey trees in mature stands. These temporal changes were accompanied by a decrease in light-demanding pioneer species and an increase in shade tolerant, later seral species adapted to a forest environment. Measurements of the degree of canopy closure in stands with low or high stocking and modelling of temporal changes of canopy closure indicated that these understorey plant dynamics are influenced by changes in light availability as stands age. Despite the successional changes within forests, geographic variation more strongly influenced understorey communities because stands within a forest area were grouped together in DCA and TWINSPAN analyses, along rainfall and temperature gradients. Although the canopy species of such intensively managed plantation forests is an alien element in the New Zealand flora, the sheltered forest environment of older stands allows the establishment of a mostly indigenous forest understorey community with considerable similarities to indigenous forests located nearby.     

Epiphytic lichen diversity in late-successional Pinus sylvestris forests along local and regional forest utilization gradients in eastern boreal Fennoscandia

Pinus sylvestris-dominated forests have been heavily utilized across all of boreal Fennoscandia and the remaining natural forests are generally highly fragmented. However, there are considerable local and regional differences in the intensity and duration of past forest utilization. We studied the impact of human forest use on the diversity of epiphytic and epixylic lichens in late-successional Pinus sylvestris-dominated forests by assessing species richness and composition along both local and regional gradients in forest utilization. The effects of local logging intensity were analysed by comparing three types of stands: (i) near-natural, (ii) selectively logged (in the early 20th century) and (iii) managed stands. The effects of regional differences in duration and intensity of past forest use were analysed by comparing stands in two contrasting regions (Häme and Kuhmo–Viena). The species richness of selectively logged stands was as high as that of near-natural stands and significantly higher in these two stand categories than in managed stands. Species richness increased with the density of small understorey Picea, which correlated strongly with decreasing intensity of local forest use and increasing structural complexity of selectively logged and near-natural stands. Stands in the Häme region hosted a lower number of species, and were less likely to host many old-growth indicator species than the Kuhmo–Viena region, suggesting that species have been lost from stands in the Häme region due to a longer history of intensive forest use. We conclude that selectively logged stands, along with near-natural stands, are valuable lichen habitats particularly for species confined to old-growth structures such as coarse trees and deadwood. In landscapes where natural forests have become fragmented, the management or restoration of the remaining late-successional Pinus-dominated forests, e.g. through the use of fire, should be carefully planned to avoid adverse effects on lichen species richness.     

Assessing the relationships between stand development and understory vegetation using a 420-year chronosequence

The large-scale conversion of old forests to tree plantations has made it increasingly important to understand how understory vegetation responds to such landscape changes. For instance, in some forest types a reduction in understory richness and cover is thought to result from the development of canopy closure in plantations, although there is a paucity of empirical data demonstrating this relationship. We used a 420-year forest chronosequence as a case study to assess the relationship between stand age, tree canopy cover and understory vascular plant richness and composition in the Siskiyou Mountains of Oregon. The chronosequence consisted of six young managed (age 7–44) and nine older unmanaged (age 90–427) stands. All stands were similar in underlying geology, slope, elevation, and aspect. We found a non-linear relationship between stand age and richness, in which richness was highest in the youngest stands, reached a low in mid-aged stands (∼55 years), then increased in the oldest stands. We also found that percent tree canopy cover was correlated with total understory cover, richness, diversity, and species composition. In general, young stands were characterized by high shrub and graminoid cover and old stands were characterized by an abundant herb layer. Our work suggests that a major component of our study landscape is currently entering the forest stage (canopy closure) characterized by low levels of vascular plant species richness and cover. We use our results to discuss the potential effects of future forest management on understory plants.     

Long-term trends in vegetation changes of managed versus unmanaged Eastern Carpathian beech forests

A vegetation survey of semi-permanent plots was conducted between years 1955–1962 and 2005–2009, and aimed to determine floristic changes occurring in beech forests located on a scarcely populated, dense forest area. The survey encompassed all natural beech forest types within the Polish part of the “Eastern Carpathians” International Reserve of Biosphere. A comparison of trends in changes between unmanaged (Bieszczady National Park) and managed (the remaining part of the Biosphere Reserve) forests was attempted. DCA analysis was used for the determination of changes on the community level. The density of vegetation, species richness, frequency of occurrence and cover, and functional groups of species were determined. Much more profound community changes were determined to have occurred in managed beech forests. However, a similar direction in vegetation changes within both managed and unmanaged forests verifies the existence of a regional pattern of changes in beech forests. This regional pattern involves (1) a decrease in the shares of arborescent species within the shrub layer, (2) disturbance of the canopy layer, (3) acidification of the top layer of the soil, (4) change of light conditions, (5) increase in shares of generalist species and decrease in shares of specialist species. A decreasing anthropogenic pressure, aging of forest stands, functioning of large, dense forest areas and sustainable forest management strategies forestry management system mimicking natural deciduous forest disturbance regime constitute factors shaping the regional changes of the forest vegetation. The undergoing changes can lead to biotic and spatial homogenization of the Eastern Carpathian beech forests.     

Partial cutting does not maintain spider assemblages within the observed range of natural variability in Eastern Canadian black spruce forests

In boreal black-spruce forests of Eastern Canada, a cohort model of ecosystem management has been proposed whereby a combination of both partial and more intensive cutting are used to emulate old-growth stands and the re-establishment of stands following severe wildfire. As with other approaches to coarse filter conservation, partial cutting is hypothesized to maintain and potentially recreate plant and animal assemblages consistent with a range of natural variability. In this study, we used ground-dwelling spiders (Araneae) to evaluate whether partial cutting and a cohort model of ecosystem management are sufficient to preserve biodiversity found in mature and over-mature boreal black spruce stands prone to paludification. We compared the spider fauna (11,628 individuals representing 136 species) in replicated partial cuts, clear cuts and uncut control stands with a chronosequence of mature and over-mature naturally regenerated stands (94-288 years since the last fire) [25 stands in total] in the same region of the northern Clay belt in Québec (Canada). In stands that were old-growth prior to cutting, harvesting had strong repercussions on spider assemblages that were not attenuated by less intensive, partial cutting. The most obvious changes in spider assemblages were related to increased recruitment of species that were nearly absent in uncut stands. Several cosmopolitan species that were widely distributed among stands prior to harvest increased in cut stands. Spider assemblages collected following either cutting methods were not consistent with assemblages found within the chronosequence and thus fell outside the observed range of natural variability. However we did not observe a similar recruitment effect for spider species in younger stands with relatively higher levels of retention. We demonstrate that the interaction between stand age prior to cutting, the degree of paludification as well as remnant basal area are important considerations when evaluating the efficacy of partial cutting to maintain forest spider assemblages and biodiversity at large in black-spruce stands. “We also suggest that retention levels in partial cuts will have to be substantially increased to maintain spider communities within their range of natural variation in managed forested landscapes.” However partial cutting with higher levels of retention, particularly in younger stands, may be useful in coarse filter management particularly for establishing or accelerating spider assemblages towards those found in old-growth stands.     

Long-term influence of fire and harvesting on boreal forest age structure and forest composition in eastern Québec

In boreal forests, historical variations in the area disturbed by natural disturbances or harvesting have rarely been compared. We measured temporal and spatial variations in areas affected by severe fires and clearcutting throughout the 20th century in a 57, 332 km² section of the eastern Canadian boreal forest. We examined the effects of these disturbances on spatio-temporal variations in the abundance of forests >60 years. Natural variability for the abundance of forests >60 years was estimated from simulations of natural disturbance regimes. We also measured compositional and structural differences between three categories of stands originating from relatively recent disturbances (∼50 years; clearcutting, fires, and clearcutting followed by fires), and one category of stands that were undisturbed for at least 200 years. At the regional level, we observed that forests >60 years gradually became scarcer throughout the 20th century due to a gradual expansion of harvested areas, an effect most pronounced in the southern part of the region, where mature and old forest abundance was clearly outside the range of natural variability at the end of the studied period. At the stand level, forest composition and structure differed between stand-origin categories: clearcutting-origin stands contained more balsam fir (Abies balsamea), fire-origin stands more black spruce (Picea mariana), and fire/clearcutting-origin stands more hardwoods (Betula papyrifera and Populus tremuloides). Overall, we estimate that strict forest management targets based on natural disturbance regimes will be difficult to achieve in eastern North-American boreal forests, most notably because contemporary disturbance rates, including both clearcutting and fire, have gradually become higher than the fire rates observed during the preindustrial period.     

The complex responses of social stingless bees (Apidae: Meliponini) to tropical deforestation

Despite concern over a putative “global pollination crisis”, we still have an incomplete understanding of how bee communities respond to land-use change. I studied the responses of social stingless (or “meliponine”) bees (Hymenoptera: Apidae: Meliponini) to surrounding forest cover and floral resources in 35 sites in a largely deforested landscape in Costa Rica over three years, sampling bees with a standardized netting protocol. I recorded a diverse fauna of meliponines, comprised of 20 species and nine genera. I found that meliponine species richness and abundance are strongly related to forest cover, but not floral resource variables (blooming plant species richness and abundance). The effect of forest on meliponine abundance, but not diversity, disappeared when the most common meliponine species, Trigona fulviventris (which comprised ∼45% of sampled individuals), was excluded from analyses. Meliponine community composition, by contrast, was related most strongly to plant species richness, only weakly to forest cover, and not related to blooming plant abundance. This work differs from past work in the same landscape, which did not find evidence of changes in species richness or abundance of meliponines and forest-related variables (distance to forest or forest fragment size), but did find shifts toward meliponine-dominated communities near forests, especially larger ones. The larger true sample size (i.e. number of sample sites) of the present work likely improved the statistical power to detect these relationships. While meliponines are forest dependent, I recorded some species in the smallest forest fragments in the landscape, and as a group they respond strongly to overall forest cover in the landscape (i.e. including both small and large patches of forest). Both of these observations support arguments for preserving even small fragments of forest in agricultural landscapes. Given the ecological and economic importance of meliponine bees, it is imperative that we better understand their long-term conservation needs in the changing tropical landscapes of the world.     

Long-term understory recovery after mechanical fuel reduction in Mediterranean cork oak forests

In the Euro-Mediterranean region, mechanical fuel reduction is increasingly used in response to the mounting occurrence of catastrophic wildfires, yet their long-term ecological effects are poorly understood. Although Mediterranean vegetation is resilient to a range of disturbances, it is possible that widespread fuel management at short intervals may threaten forest structural complexity and the persistence of some plant species and functional types, with overall negative consequences for biodiversity. We used a chronosequence approach to infer woody vegetation changes in the first 70 years after understory clearing in upland cork oak (Quercus suber) forests, and to assess how these are affected by treatment frequency. Across the chronosequence there was a shift between plant communities with contrasting composition, structure and functional organization. Understory cover increased quickly after disturbance and a community dominated by pioneer seeder and dry-fruited shrubs (Cistus ladanifer, C. populifolius, Genista triacanthos, and Lavandula stoechas) developed during about 15 years, but this was slowly replaced by a community dominated by resprouters and fleshy-fruited species (Arbutus unedo, Erica arborea) >40 years after disturbance. During the first 15 years there were rapid increases in woody species richness, vertical structural diversity, cover by Q. suber juveniles and saplings, and shrub cover at <1.5 m strata, which levelled off or slightly declined thereafter. In contrast, tree species richness, tree density and density of arboreal A. unedo and E. arborea, vertical structural evenness, and cover at >1.5 m strata increased slowly for >50 years. Treatment frequency showed strongly negative relationships with species richness, structural diversity and evenness, and horizontal and vertical understory cover, particularly that of slowly recovering species. These findings suggest that fuel reduction programs involving widespread and recurrent understory clearing may lead to the elimination at the landscape scale of stands with complex multi-layered understory occupied by large resprouters and fleshy-fruited species, which take a long time to recover after disturbance. Fuel management programs thus need to balance the dual goals of fire hazard reduction and biodiversity conservation, recognizing the value of stands untreated for >50 years to retain ecological heterogeneity in Mediterranean forest landscapes.     

The effects of partial harvest on the understory vegetation of southern Ontario woodlands

We studied the effects of partial cutting on understory vegetation communities within 19 mixed maple forests in an agriculture-dominated landscape in southwestern Ontario. Woodlots that had been recently harvested were grouped according to provincial silviculture guidelines (standard and heavy cuts) and compared to woodlots that had been uncut for at least 24 years (reference stands). We found significant differences in richness, diversity, and quality of understory vegetation in response to harvest indices. More intensive harvesting resulted in increased richness and diversity, but mostly through the addition of habitat generalists and weedy species. However, partial harvest does not appear to drive vegetation community composition, as ordination methods found no clear community differences between the treatments. Use of the single-tree selection system based on basal area and harvest intensity targets will have an effect on the understory plants, but other factors including past management, disturbance history, and site level microclimate features will also play an important role in shaping vegetation communities. We caution against the removal of large volumes of trees ≥38 cm in diameter, and large reductions in canopy cover, as this can reduce the presence of “conservative” (forest dependent specialist) species, despite a general increase in species richness and diversity. Furthermore, we recommend additional research to investigate the potential for incremental degradation to occur on sites with a long-term history of harvesting, as we found that richness of exotics increased on sites with a history of forest management.     

Distribution of saproxylic beetles in a recently burnt landscape of the northern boreal forest of Québec

This study evaluated the importance of burned habitat characteristics as well as the likely dispersal from specific habitats in the distribution of saproxylic beetles the same year as a fire occurred, in burned black spruce stands (Picea mariana [Mill] B.S.P.) in the northern boreal forest of Québec. The distribution of early post-fire saproxylic species was mainly driven by burned habitat attributes at the plot scale (0.04 ha), especially fire severity, suggesting that the effect of environment attributes can act at a relatively fine scale. Some xylophagous and most predaceous species were more abundant in severely burned stands whereas fire severity had the opposite effect on several common mycophagous species. The amount of newly fire-killed trees that could be used as breeding substrates in the burned stands had only a weak positive influence on these functional groups. The great majority of early saproxylic species were weakly associated with the distance from unburned forests or other recently burned patches that could act as potential “source habitats”. Indeed, these variables were of lesser importance than the attributes of the burned habitat. Woody debris that were already present in plots before the fire, potentially serving as local of source-populations for early colonizers, had virtually no influence on the local abundance of species. Many saproxylic species, including some true pyrophilous, clearly showed higher abundance as distance from unburned stands increased. This unexpected relation may reflect that dispersal of insects toward the burnt landscape very shortly after fire could be driven by the higher amount of volatiles released by severely burned forests, which are more likely as distance from unburned forest increased.     

Emulating natural disturbances: the role of silviculture in creating even-aged and complex structures in the black spruce boreal forest of eastern North America

Ecosystem-based forest management is based on the principle of emulating regional natural disturbance regimes with forest management. An interesting area for a case study of the potential of ecosystem-based forest management is the boreal forest of north-western Québec and north-eastern Ontario, where the disturbance regime creates a mosaic of stands with both complex and simple structures. Old-growth stands of this region have multi-storied, open structures, thick soil organic layers, and are unproductive, while young post-fire stands established following severe fires that consumed most of the organic soil show dense and even-sized/aged structures and are more productive. Current forest management emulates the effects of low severity fires, which only partially consume the organic layers, and could lead to unproductive even-aged stands. The natural disturbance and forest management regimes differ in such a way that both young productive and old-growth forests could ultimately be under-represented on the landscape under a fully regulated forest management regime. Two major challenges for ecosystem-based forest management of this region are thus to: (1) maintain complex structures associated with old-growth forests, and (2) promote the establishment of productive post-harvest stands, while at the same time maintaining harvested volume. We discuss different silvicultural approaches that offer solutions to these challenges, namely the use of (1) partial harvesting to create or maintain complex structures typical of old-growth stands, and (2) site preparation techniques to emulate severe soil burns and create productive post-harvest stands. A similar approach could be applied to any region where the natural disturbance regime creates a landscape where both even-aged stands established after stand-replacing disturbances and irregular old-growth stands created by smaller scale disturbances are significant.     

Potential effects of climate change on insect herbivores in European forests—General aspects and the pine processionary moth as specific example

The nature of “climate change” will differ with geographical regions and its final impact on ecosystems vary with the extent of temperature increase, changes in irradiance and levels of UVB, amount and patterns of precipitation and humidity, and alterations in the incidence and nature of abiotic disturbances. Despite many uncertainties, there is consensus in the fact that global warming already has and will have impact on the temporal and spatial dynamics of insect herbivores. Ectothermic organisms are affected by the changes in environmental conditions directly in dispersal, reproduction, development and mortality, and indirectly through altered plant nutritional quality, resistance and via community interactions. Ambiguous consequences are to be expected depending on the individual host plant and herbivore species, probably involving altered incidence and intensity of pest outbreaks and changes in distributional ranges. Regions that represent northern or upper limits of occurrence, such as the Alps or the boreal zone, are likely to be affected most by an increase in stability and population density of certain pest species, such as defoliating insects or bark beetles. At the same time, temperature increase and drought will render areas of distribution in southern and continental parts of Europe less suitable for heat susceptible species, which will probably not only result in northwards shifts, but range contractions. The review is based on chapters of the “Study on impacts of climate change on European forests and options for adaptation” led by the European Forest Institute (Efi) and on results of the EU project “PROMOTH - Global change and pine processionary moth: a new challenge for integrated pest management”. Thaumetopoea pityocampa serves as illustrative example for insect herbivores whose latitudinal and altitudinal distribution is mainly controlled by temperature and already modified by global warming.     

Positive association between understory species richness and a dominant shrub species (Corylus avellana) in a boreonemoral spruce forest

Old growth stands of boreonemoral spruce (Picea abies) forests frequently have a shrub layer dominated by hazel (Corylus avellana) – a species which is generally excluded in intensively managed forests due to clearcutting activities. We sampled understory species composition, richness and biomass, as well as environmental variables beneath these two species and also within forest ‘gaps’ in order to determine the effect of overstory species on understory vegetation. Species richness and biomass of herbaceous plants was significantly greater under Corylus compared with plots under Picea and in forest gaps. Indicator species analysis found that many species were significantly associated with Corylus. We found 45% of the total species found under woody plants occurred exclusively under Corylus. Light availability in spring and summer was higher in gaps than under forest cover but no difference was found between plots under Corylus and Picea. Hence, reductions in light availability cannot explain the differences in species composition. However, Ellenberg indicator values showed that more light demanding species were found under Corylus compared to Picea, but most light demanding species were found in gaps. The litter layer under Picea was three times thicker than under Corylus and this may be an important mechanism determining differences in understory composition and richness between the woody species. The presence of Corylus is an important factor enhancing local diversity and small-scale species variation within coniferous stands. Hence, management should maintain areas of Corylus shrubs to maintain understory species diversity in boreal forests.     

Understorey vegetation in young naturally regenerated and planted birch (Betula spp.) stands on abandoned agricultural land

The abandonment of agricultural lands in Northern and Eastern Europe increases the area covered by first generation forests, which are either formed as an outcome of secondary succession or established as plantations. However, questions remain as to how these new stands develop and what kind of species they favour, which in turn has impacts on their ecological and economical value. Our aim was to compare understorey vascular plant and bryophyte vegetation characteristics between naturally regenerated and planted birch stands on abandoned agricultural sites in Estonia, focusing on the aspects of species richness and forest understorey recovery. Species richness and diversity of vascular plants were similar in both stand types but the number of forest vascular plant species was significantly higher in naturally regenerated stands. The bryophyte layer of naturally regenerated stands had a higher species richness, diversity, and number of forest bryophyte species. The higher number of forest vascular plant and bryophyte species in naturally regenerated stands can be explained by the longer undisturbed succession period. The recovery of the forest understorey was unaffected by former agricultural land use (crop field or grassland). The influence of soil properties on the recovery of the forest understorey was not detected, but the number of vascular plant species that grow in forests as well as in grasslands was negatively correlated with distance from forest. Overall, understorey vegetation of natural and planted birch stands did not reveal substantial differences. However, in the case of vigorous natural birch regeneration in the vicinity of forest land, unassisted reforestation should be favoured.     

Understory vegetation in old-growth and second-growth Tsuga canadensis forests in western Massachusetts

We compared the understory communities (herbs, shrubs, and tree seedlings and saplings) of old-growth and second-growth eastern hemlock forests (Tsuga canadensis) in western Massachusetts, USA. Second-growth hemlock forests originated following clear-cut logging in the late 1800s and were 108–136 years old at the time of sampling. Old-growth hemlock forests contained total ground cover of herbaceous and shrub species that was approximately 4 times greater than in second-growth forests (4.02 ± 0.41%/m² versus 1.06 ± 0.47%/m²) and supported greater overall species richness and diversity. In addition, seedling and sapling densities were greater in old-growth stands compared to second-growth stands and the composition of these layers was positively correlated with overstory species composition (Mantel tests, r > 0.26, P < 0.05) highlighting the strong positive neighborhood effects in these systems. Ordination of study site understory species composition identified a strong gradient in community composition from second-growth to old-growth stands. Vector overlays of environmental and forest structural variables indicated that these gradients were related to differences in overstory tree density, nitrogen availability, and coarse woody debris characteristics among hemlock stands. These relationships suggest that differences in resource availability (e.g., light, moisture, and nutrients) and microhabitat heterogeneity between old-growth and second-growth stands were likely driving these compositional patterns. Interestingly, several common forest understory plants, including Aralia nudicaulis, Dryopteris intermedia, and Viburnum alnifolium, were significant indicator species for old-growth hemlock stands, highlighting the lasting legacy of past land use on the reestablishment and growth of these common species within second-growth areas. The return of old-growth understory conditions to these second-growth areas will largely be dependent on disturbance and self-thinning mediated changes in overstory structure, resource availability, and microhabitat heterogeneity.