Compositional gradients in the forest vegetation of Finland

Abstract. Compositional gradients of forest vegetation in Finland were determined using detrended correspondence analysis, and ecologically interpreted with regression analysis. The data were collected in association with the 8th National Forest Inventory, and they comprised 1514 samples and 472 species, covering the whole forested land area of Finland. In the ordination space the first, very stable axis (DCA1) was much stronger than the following three, more unstable axes (DCA2 - 4). Site fertility (reflected by the forest site type series) and latitude had strongest covariation with the ordination space, followed by a number of variables primarily related to forest age or tree stand volume. Fertility variables were most closely parallel with DCA1, latitudinal ones with DCA1 and DCA2, and successional variables with DCA3. The results indicate that compositional overlap of sites classified into different Caj ander site types is considerable.     

Understorey vegetation development in North Finnish Picea forests after disturbance: re‐analysis of Sirén's data

Abstract. Sirén (1955) studied understorey species composition, tree stand properties and humus‐layer thickness in 64 unlogged forest stands on topographically and pedologically comparable sites. The stands were of even age (6 – 300 yr), stocked with the first or second tree generation after wildfire. The view of Sirén and several authors after him, that the vegetation of old‐growth boreal Picea forests is homogeneous on a broad scale, was examined by applying, in parallel, the partial variants of two ordination methods (DCA and PCA) to Sirén's vegetation data. Two main vegetation gradients were found: a major gradient running from recently burnt plots with prominence of pioneer species to plots with stand age > 100 yr, a well stocked tree layer and a thick humus layer, dominance of feather‐mosses and ample occurrence of shade‐tolerant as well as light‐preferring vascular plant species, and a second gradient along which first‐ and second‐generation plots segregate. The more prominent element of Betula trees in first‐ than in second‐generation stands < 100 yr contributed to the latter. A minor third gradient related to humus‐layer thickness was recovered by partial DCA only. The main vegetation gradient reappeared in separate ordinations of data from 47 mature forest stands (> 100 yr), but without being correlated with forest age. Variation among mature‐forest stands in the importance of pioneer species is considered mainly to be brought about by fine‐scale disturbance processes such as tree uprooting. Increasing importance of factors operating on within‐stand scales [development of a varied gap structure and stronger gradients in tree influence (radiation at ground level), soil moisture, soil depth and nutrient availability] with time is also reflected in the second and third mature‐forest ordination axes. Possible implications of the results for conservation of biological diversity and monitoring of changes in boreal forests are discussed.     

Conservation value of timber quality versus associated non-timber quality stands for understory diversity in Nothofagus forests

Conservation strategies of forested landscapes must consider biodiversity of the included site types, i.e. timber-quality forests and associated non-timber-quality stands. The objectives were to characterize forest overstory structure in timber-quality versus associated non-timber-quality stands; and to compare their understory communities. Six forest types were sampled in Nothofagus forests of Tierra del Fuego (Argentina): two timber-quality N. pumilio forests, and four associated non-timber-quality stands (edge, N. antarctica, wetlands and streamside forests). Overstory structure and understory vegetation (species richness, frequencies, cover and biomass) were characterized during spring and summer seasons. Analysis of variance and multivariates were carried out. Overstory structure differed across the site types, with higher tree size, canopy closure and tree volume in timber-quality stands. Fifty-one understory plant species were observed, but understory variables varied with site types, especially wetlands (highest native and exotic richness, cover and biomass, and 25% of exclusive species). Site types were grouped in three: N. antarctica stands, streamside stands and the other N. pumilio forests according to multivariate analysis. Forty three percent of plants were distributed in all site types, and all timber-quality forest understory species were present in some associated non-timber-quality stands. Timber-quality N. pumilio forests have a marginal value for understory conservation compared to associated non-timber-quality stands, because these last include all the plants observed in timber-quality forests and also possess many exclusive species. Therefore, protection of associated non-timber-quality stands during forest management planning could increase understory conservation at landscape level, and these could be better reserves of understory diversity than retentions of timber-quality stands.     

Temperate tree expansion into adjacent boreal forest patches facilitated by warmer temperatures

Temperate and boreal forests are forecast to change in composition and shift spatially in response to climate change. Local‐scale expansions and contractions are most likely observable near species range limits, and as trees are long‐lived, initial shifts are likely to be detected in the understory regeneration layers. We examined understory relative abundance patterns of naturally regenerated temperate and boreal tree species in two size classes, seedlings and saplings, and across two spatial scales, local stand‐scale ecotones (tens of meters) and the regional temperate–boreal transition zone (?250 km) in central North America, to explore indications of climate‐mediated shifts in regeneration performance. We also tested for the presence of strong environmental gradients across local ecotones that might inhibit species expansion. Results showed that tree regeneration patterns across ecotones varied by species and size class, and varied across the regional summer temperature gradient. Temperate tree species regeneration has established across local ecotones into boreal forest patches and this process was facilitated by warmer temperatures. Conversely, boreal conifer regeneration exhibited negative responses to the regional temperature gradient and only displayed high abundance at the boreal end of local ecotones at cool northern sites. The filtering effects of temperature also increased with individual size for both boreal and temperate understory stems. Observed regeneration patterns and the minor environmental gradients measured across local ecotones failed to support the idea that there were strong barriers to potential temperate tree expansion into boreal forest patches. Detectable responses, consistently in the directions predicted for both temperate and boreal species, indicate that summer temperature is likely an important driver of natural tree regeneration in forests across the temperate–boreal transition zone. Regeneration patterns point toward temperate expansion and reduced but continued boreal presence in the near‐future, resulting in local and regional expansions of mixed temperate‐boreal forests.     

Cultivation of Non‐timber Forest Products Alters Understory Light Availability in a Humid Tropical Forest in Mexico1

The planting of non‐timber forest products (NTFPs) in the understory of tropical forests is promoted in many regions as a strategy to conserve forested lands and meet the economic needs of rural communities. While the forest canopy is left intact in most understory plantations, much of the midstory and understory vegetation is removed in order to increase light availability for cultivated species. We assessed the extent to which the removal of vegetation in understory plantations of Chamaedorea hooperiana Hodel (Arecaceae) alters understory light conditions. We also examined how any changes in light availability may be reflected by changes in the composition of canopy tree seedlings regenerating in understory plantations. We employed a blocked design consisting of four C. hooperiana plantation sites; each site was paired with an adjacent, unmanaged forest site. Hemispherical canopy photographs were taken and canopy tree seedlings were identified and measured within 12 3 × 2 m randomly placed plots in each site for a total of 96 plots (4 blocks × 2 sites × 12 plots). Plantation management did not affect canopy openness or direct light availability but understory plantations had a higher frequency of plots with greater total and diffuse light availability than unmanaged forest. Comparisons of canopy tree seedling composition between understory plantations and unmanaged forest sites were less conclusive but suggest that management practices have the potential to increase the proportion of shade‐intolerant species of tree seedlings establishing in plantations. Given the importance of advanced regeneration in gap‐phase forest dynamics, these changes may have implications for future patterns of succession in the areas of forest where NTFPs are cultivated.     

Forest plant community as a driver of soil biodiversity: experimental evidence from collembolan assemblages through large‐scale and long‐term removal of oak canopy trees Quercus petraea

Plant–soil interactions are increasingly recognized to play a major role in terrestrial ecosystems functioning. However, few studies to date have focused on slow dynamic ecosystems such as forests. As they are vertically stratified by multiple vegetation strata, canopy tree removal by thinning operations could alter forest plant community through tree canopy opening. Very little is known about cascading effects on soil biodiversity. We conducted a large‐scale, multi‐site assessment of collembolan assemblage response to long‐term canopy tree removal in sessile oak Quercus petraea temperate forests. A total of 33 experimental plots were studied covering a large gradient of canopy tree basal area, stand age and local abiotic contexts. Collembolan abundance strongly declined with canopy tree removal in early forest successional stage and this was mediated by negative effect of understory plant community composition changes, i.e. shift from moss and forb to tree seedling, fern, shrub and grass species. Negative effect of this composition shift on collembolan species richness was largely offset by positive effect of the increase in understory plant species richness. This gives support to both the plant mass‐ratio and functional diversity hypotheses. Collembolan functional groups had contrasting response patterns, which were mediated by different ecological factors. Epedaphic (r‐strategist) abundance and species richness increased with canopy tree removal in relation with the increase in understory plant species richness. In contrast, euedaphic (K‐strategist) abundance and species richness declined with canopy tree removal in early forest successional stage in relation with changes in understory plant community composition and species richness, as well as microclimatic conditions. Overall, our study provides experimental evidence that forest plant community can be a strong driver of collembolan assemblages. It also emphasizes the role of trees as foundation species of forest ecosystems that can shape soil biodiversity through their regulation of understory plant community and ecosystem abiotic conditions.     

Overstory-Understory Relationships along Forest Type and Environmental Gradients in the Spring Mountains of Southern Nevada, USA

Isolated forested mountains in deserts have numerous ecological and societal values, but land-management practices (e.g., fire-regime alteration) and climate change can affect forest composition. We analyzed tree overstory-understory relationships on 123 sites in the Spring Mountains within the Mojave Desert near Las Vegas, Nevada, USA to assess three hypotheses. We hypothesized that: the tree species comprising understories are less tolerant of fire than species in overstories, reflecting land-management practices of fire exclusion; mid-elevation forests have the lowest overstory:understory similarity because this zone could have maximum species mixing; and overstory:understory similarity is correlated with environmental gradients (consisting of 14 topographic and soil variables). We found that Pinus monophylla comprised greater relative canopy cover in understories of juniper (32% relative cover) and pinyon-juniper (78%) forests than it did in overstories of these forests (0% and 53%). Similarly, fire-intolerant Abies concolor had 6-fold greater understory than overstory cover in forests with overstories dominated by the fire-tolerant Pinus ponderosa. Overstory:understory S?rensen similarity averaged 43%?77% among six forest types, and there was little support for the supposition that similarity was lowest in mid-elevation forests. Distributions of individual overstory and understory species more closely corresponded with environmental gradients than did overstory:understory similarity. Results suggest that there is high potential for change in at least two of the six dominant forest types of the Spring Mountains. The direction of change (species of moist, higher elevation sites establishing in understories of drier forests) is the opposite of what would be expected for forest adaptation to the warmer, drier, more fire-prone conditions projected for the next century in the southwestern USA.     

Within‐stand spatial structure and relation of boreal canopy and understorey vegetation

Question: How do spatial patterns and associations of canopy and understorey vegetation vary with spatial scale along a gradient of canopy composition in boreal mixed‐wood forests, from younger Aspen stands dominated by Populus tremuloides and P. balsamifera to older Mixed and Conifer stands dominated by Picea glauca? Do canopy evergreen conifers and broad‐leaved deciduous trees differ in their spatial relationships with understorey vegetation? Location: EMEND experimental site, Alberta, Canada. Methods: Canopy and understorey vegetation were sampled in 28 transects of 100 contiguous 0.5 m × 0.5 m quadrats in three forest stand types. Vegetation spatial patterns and relationships were analysed using wavelets. Results: Boreal mixed‐wood canopy and understorey vegetation are patchily distributed at a range of small spatial scales. The scale of canopy and understorey spatial patterns generally increased with increasing conifer presence in the canopy. Associations between canopy and understorey were highly variable among stand types, transects and spatial scales. Understorey vascular plant cover was generally positively associated with canopy deciduous tree cover and negatively associated with canopy conifer tree cover at spatial scales from 5–15 m. Understorey non‐vascular plant cover and community composition were more variable in their relationships with canopy cover, showing both positive and negative associations at a range of spatial scales. Conclusions: The spatial structure and relation of boreal mixed‐wood canopy and understorey vegetation varied with spatial scale. Differences in understorey spatial structure among stand types were consistent with a nucleation model of patch dynamics during succession in boreal mixed‐wood forests.     

Drought-induced mortality affects understory vegetation: release after death

In recent decades, severe droughts have become an important cause of canopy disturbance in forests, and have shown potential to cause rapid and pronounced vegetation shifts. Under dead canopy, undamaged understory could influence the nature of resource limitation for seedling growth and survival, limiting forest regeneration. We assessed the release response of understory vegetation after a severe drought event in temperate forests of northern Patagonia. Growth trends of dominant tree saplings, and changes in vegetation biodiversity and cover were compared between drought-dead and unaffected canopy. Nothofagus dombeyi undergo growth release after the climatic event in affected forests, and the response was evidenced immediately after the disturbance. For Austrocedrus chilensis, the growth release response was less evident, due mainly to a difference in age structure. In the understory the release response was barely discernable for some components. There was a tendency towards higher cover of the shrub layer in the understory of drought-affected forests, and an important presence of the exotic shrub Rosa rubiginosa. However, the clearest biotic response following drought mortality was the release in growth of understory dominant tree component. Those results strongly suggest that the environment under drought-dead canopy, and the die-off in woody sapling cohorts in a self-thinning process, could favor crown expansion and growth release of understory species that could help predict future forest trajectories in the context of the influence of climatic extreme events.     

Factors influencing the distribution and growth of dwarf bamboo, <Emphasis Type="Italic">Fargesia nitida</Emphasis>, in a subalpine forest in Wolong Nature Reserve,southwest China

Dwarf bamboos impose intense resource competition in subalpine coniferous forests, and their exclusive densities have crucial impacts on tree regeneration and understory species diversity. We studied the factors influencing the distribution and growth of dwarf bamboo, Fargesia nitida, in a subalpine forest in southwest China. TWINSPAN, based on an attribute matrix, could divide the subalpine forest into 11 sub-associations, and more clearly reflected ecological functional features of the subalpine forest than analysis based on a species matrix. TWINSPAN was also generally consistent with DCA ordination based on the attribute matrix. DCA and DCCA ordination showed relationships between the distribution of F. nitida population and environment factor. The first DCCA axis showed topography and disturbance gradients (except fallen trees and broken branches); the second DCCA axis showed canopy density and composition gradients. Stepwise multiple regression analyses showed that distribution (culm density and coverage) of F. nitida decreased significantly with landslide and slope aspect, and increased significantly with soil status. The light condition had positive effects on growth and size of bamboo. A stable environment in the northern slope and more broadleaved species dominating in canopy would increase the dwarf bamboo biomass. Thus, the disturbance regimes, the slope aspect and the BA of evergreen conifer trees can provide useful guidelines for the control and management of F. nitida populations, and in helping to understand the succession and regeneration of subalpine forest in this region.