Shrub species affect distinctively the functioning of scattered Quercus ilex trees in Mediterranean open woodlands

A low tree stand density has been showed as necessary to thrive with summer drought in semiarid Mediterranean open woodlands. Shrub encroachment of these open woodlands is currently recommended to guarantee the persistence of the system, due to the nursery effect of shrubs on tree seedling. However, the increase in abundance and cover of a shrub understory in these water limited woodlands could bring consequences to tree overstory functioning. The present study analyzes the physiological status of scattered Quercus ilex L. trees in paired adjacent plots with and without the presence of a shrubby understory in CW Spain. Two contrasting shrub strategies were addressed in order to take into account possible species-specific effects: a dense-shallow rooting shrub (Cistus landanifer L.) and a sparse-deep rooting shrub (Retama sphaerocarpa (L.) Boiss). Leaf water potential (at predawn and midday), leaf gas exchange parameters (net photosynthetic rate and stomatal conductance), leaf nitrogen content and chlorophyll fluorescence transients (maximum photochemical efficiency and performance index, sensuStrasser et al., 2004) were measured during three consecutive summers. Trees growing with Cistus as understory showed significant lower leaf water potential, leaf gas exchange parameters, leaf nitrogen content and chlorophyll photochemical efficiency than trees growing without shrub competence. However, the presence of the legume Retama did not affect significantly the physiological state of Q. ilex. Thus, we conclude that the presence of a shrubby understory has the potential to modify the functioning of scattered trees, but these effects are species-specific.     

Effects of the overstory on the diversity of the herb and shrub layers of Anatolian black pine forests

Understory plants are important components of forests because they are responsible for the majority of the vascular plant diversity of forest ecosystems. The richness and composition of understory communities are closely related to the tree layer diversity, structure and composition. The aim of this study was to examine the understory diversity of Anatolian black pine (Pinus nigra Arnold subsp. pallasiana (Lamb.) Holmboe)-dominated forests on the Kazda?? Mountains of West Turkey. To describe the overstory structure and composition in a numerically and quantitatively well-defined manner, cumulative abundance profiles (CAPs) of the tree species were used. The resemblance of the sampling plots was classified into five stand types assessing the CAP through the Fuzzy C-Means clustering method. A permutational multivariate analysis of variance (PERMANOVA) was performed to test the variance of the community ecological distance between the five stand types, and the results showed significant differences in these clusters. Many shade-tolerant plants were associated with the mixed stands of Anatolian black pine–Kazda?? fir. The composition of the herb and shrub layer could not be explained by the environmental variables but by differences in the overstory structure of the stands. Pure or nearly pure Anatolian Black pine stands were more diverse than mixed oak–Anatolian black pine and Kazda?? fir–Anatolian black pine stands. However, although dense and young pure Anatolian black pine stands had the most diverse plant species in the shrub layer, they were ranked third in terms of the herb layer diversity. The Anatolian black pine–Kazda?? fir mixed stands had the lowest herb and shrub layer diversity. These results allow us to comprehend the relationship between the overstory structure and composition, and the understory diversity. Understanding this relationship is important for the conservation of understory plant diversity in the management of forest ecosystems.     

Understory vegetation response after 30 years of interval prescribed burning in two ponderosa pine sites in northern Arizona,USA

Southwestern USA ponderosa pine (Pinus ponderosa C. Lawson var. scopulorum Engelm.) forests evolved with frequent surface fires and have changed dramatically over the last century. Overstory tree density has sharply increased while abundance of understory vegetation has declined primarily due to the near cessation of fires. We examined effects of varying prescribed fire-return intervals (1, 2, 4, 6, 8, and 10 years, plus unburned) on the abundance and composition of understory vegetation in 2007 and 2008 after 30+ years of fall prescribed burning at two ponderosa pine sites. We found that after 30 years, overstory canopy cover remained high, while understory plant canopy cover was low, averaging <12% on all burn intervals. We attributed the weak understory response to a few factors – the most important of which was the high overstory cover at both sites. Graminoid cover and cover of the major grass species, Elymus elymoides (squirreltail), increased on shorter fire-return intervals compared to unburned plots, but only at one site. Community composition differed significantly between shorter fire-return intervals and unburned plots at one site, but not the other. For several response variables, precipitation levels appeared to have a stronger effect than treatments. Our findings suggest that low-severity burn treatments in southwestern ponderosa pine forests, especially those that do not decrease overstory cover, are minimally effective in increasing understory plant cover. Thinning of these dense forests along with prescribed burning is necessary to increase cover of understory vegetation.     

Forest herbicide influences on floristic diversity seven years after broadcast pine release treatments in central Georgia,USA

Maintenance of biodiversity is becoming a goal of forest management. This study determined effects of broadcast pine release herbicide treatments on plant species richness, diversity, and structural proportions seven years after treatment. Three study blocks were established in central Georgia. Plots 0.6–0.8 ha in size were planted to loblolly pine (Pinus taeda L.) in the Winter of 1982–83 and then treated with imazapyr (Arsenal), glyphosate (Roundup), and hexazinone (Velpar L. and Pronone 10G) in 1985. In 1992, overstory and understory (<1.5 m height) layers were examined utilizing stem and rootstock counts and basal area of overstory species and cover of understory species. ANOVA's were used to test for significance using a randomized complete block model. We found no effect of treatments on species richness. Diversity, measured separately for overstory and understory layers by Shannon-Wiener and Simpson indices, also was not influenced significantly by treatments. Arsenal significantly decreased Diospyros virginiana L. and increased Rubus argutus Link and legumes. Hexazinone treatments generally decreased Quercus nigra L., and Roundup significantly reduced Vaccinium spp. compared to the Check. We concluded that herbicide release treatments did not decrease overstory or understory plant species richness and diversity seven years post-treatment.     

Understory succession in post-agricultural oak forests: Habitat fragmentation affects forest specialists and generalists differently

The herbaceous understory forms the richest stratum in temperate broadleaved forests in terms of plant diversity. Understanding the process of understory succession is thus of critical importance for the development of management guidelines for biodiversity restoration in post-agricultural plantation forests.We studied effects of stand age, forest fragmentation, and soil and canopy conditions on species richness and abundance of four species groups in the understory of post-arable oak plantations in southern Sweden: herbaceous forest specialists, habitat generalists and open-land species, and woody species.The group of forest specialists may approach the richness of continuously forested sites after 60-80 years in non-fragmented plantations, but many forest species were sensitive to habitat fragmentation. Open-land species richness decreased during succession, while the richness of woody species and of generalists remained stable, and was not affected by fragmentation. Abundance of generalists gradually decreased in non-fragmented plantations, probably due to competition from colonizing forest specialists. Soil pH in post-arable stands remained consistently higher than in continuously forested stands, which maintained differences in species composition. The development of a shrub layer seemed to imply a competitive advantage for forest specialists compared to generalist species.For successful recovery of a rich understory, we suggest that post-arable plantations should be established on loamy soils of intermediate to high pH proximate to older forest with source populations, and that a continuous overstory canopy cover of 70-80% is maintained by regular light thinnings and promotion of a shrub layer.     

Small-scale variation of vegetation in a mixed forest understorey is partly controlled by the effect of overstory composition on litter accumulation

We investigated how richness and composition of vascular plant species in the understory of a mixed hardwood forest stand varied with respect to the abundance and composition of the overstory. The stand is in central Spain and represents the southernmost range of distribution of several tree and herbaceous species in Europe. Understory species were identified in 46 quadrats (0.25 m²) where variables litter depth and light availability were measured. In addition, we estimated tree density, basal area, and percent basal area by tree species within 6-m-radius areas around each plot. Species richness and composition were studied using path analysis and scale-dependent geostatistical methods, respectively. We found that the relative abundance of certain trees species in the overstory was more important than total overstory abundance in explaining understory species richness. Richness decreased as soil litter depth increased, and soil litter increased as the relative proportion of Fagus sylvatica in the overstory increased, which accounted for a negative, indirect effect of Fagus sylvatica on richness. Regarding understory species composition, we found that some species distributed preferentially below certain tree species. For example, Melica uniflora was most frequent below Fagus sylvatica and Quercus petraea while the increasing proportion of Q. pyrenaica in the overstory favored the presence of Cruciata glabra, Arenaria montana, Prunus avium, Conopodium bourgaei, Holcus mollis, Stellaria media and Galium aparine in the understory. Overall, these results emphasize the importance of individual tree species in controlling the assemblage and richness of understory species in mixed stands. We conclude that soil litter accumulation is one way through which overstory composition shapes the understory community.     

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.     

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.     

Water relations of pine seedlings in contrasting overstory environments

Overstory conditions influence understory microclimate and resource availability, leading to gradients in evaporative demand and moisture availability that influence seedling water relations. Partial canopies may either reduce seedling moisture stress by ameliorating environmental conditions, or increase moisture stress by reducing soil moisture availability. This study used stable isotope ratios of oxygen (δ¹⁸O) and carbon (δ¹³C) and mass-based foliar nitrogen concentrations to investigate changes in transpiration (E), stomatal conductance (g_s) and intrinsic water use efficiency (iWUE) of pine seedlings across an overstory gradient from open canopy gap environments to closed canopy forest. Foliar δ¹⁸O increased sharply from basal areas of 0–10 m² ha⁻¹ in Pinus banksiana, Pinus resinosa, and Pinus strobus seedlings, followed by a more gradual increase with further increases in basal area. Foliar δ¹³C followed a similar, but less pronounced pattern in P. banksiana and P. strobus seedlings, and had no apparent relationship with overstory basal area in P. resinosa seedlings. The slope of the δ¹⁸O:δ¹³C relationship was positive for every species. Foliar nitrogen concentrations were not correlated with overstory basal area. These results suggest seedling E declined as overstory basal area increased due to reductions in g_s, while iWUE increased slightly from open gaps to partial canopy environments. Open gap environments appear to provide sufficient moisture to sustain high leaf-level gas exchange rates in the species we studied, while relatively small increases in overstory basal area apparently promote rapid declines in g_s, leading to greatly reduced seedling water loss and small increases in iWUE.     

Stand characteristics and downed woody debris accumulations associated with a mountain pine beetle (Dendroctonus ponderosae Hopkins) outbreak in Colorado

Lodgepole pine (Pinus contorta Dougl. ex Loud.)-dominated ecosystems in north-central Colorado are undergoing rapid and drastic changes associated with overstory tree mortality from a current mountain pine beetle (Dendroctonus ponderosae Hopkins) outbreak. To characterize stand characteristics and downed woody debris loads during the first 7 years of the outbreak, 221 plots (0.02 ha) were randomly established in infested and uninfested stands distributed across the Arapaho National Forest, Colorado. Mountain pine beetle initially attacked stands with higher lodgepole pine basal area, and lower density and basal area of Engelmann spruce (Picea engelmannii [Parry]), and subalpine fir (Abies lasiocarpa (Hook.) Nutt. var. lasiocarpa) compared to uninfested plots. Mountain pine beetle-affected stands had reduced total and lodgepole pine stocking and quadratic mean diameter. The density and basal area of live overstory lodgepole declined by 62% and 71% in infested plots, respectively. The mean diameter of live lodgepole pine was 53% lower than pre-outbreak in infested plots. Downed woody debris loads did not differ between uninfested plots and plots currently infested at the time of sampling to 3 or 4–7 years after initial infestation, but the projected downed coarse wood accumulations when 80% of the mountain pine beetle-killed trees fall indicated a fourfold increase. Depth of the litter layer and maximum height of grass and herbaceous vegetation were greater 4–7 years after initial infestation compared to uninfested plots, though understory plant percent cover was not different. Seedling and sapling density of all species combined was higher in uninfested plots but there was no difference between infested and uninfested plots for lodgepole pine alone. For trees ≥2.5 cm in diameter at breast height, the density of live lodgepole pine trees in mountain pine beetle-affected stands was higher than Engelmann spruce, subalpine fir, and aspen, (Populus tremuloides Michx.), in diameter classes comprised of trees from 2.5 cm to 30 cm in diameter, suggesting that lodgepole pine will remain as a dominant overstory tree after the bark beetle outbreak.     

The influence of canopy patch mosaics on understory plant community composition in boreal mixedwood forest

We assessed the composition of understory vascular plant communities in relation to the mosaic of canopy patch types, and their associated structure and environment, within unmanaged, mature boreal mixedwood forests in western Canada. Within a 30 km² area, we sampled patches of four different canopy types: conifer-dominated, broadleaf-dominated, mixed conifer-broadleaf, and canopy gaps (total n = 98). There were significant differences in understory composition among the four patch types (based on multi-response permutation procedure (MRPP)) and these were mainly due to differences in relative abundances of understory species. The understory communities of conifer patches were characterized by low abundances of shade intolerant species while shade-tolerant and evergreen species were indicators (based on an indicator species analysis (ISA)). Understory communities under gap and broadleaf patches were characterized by higher abundances of grasses and shade intolerant species. Gap, broadleaf, and mixed patches had higher abundances of certain shrub species than did conifer patches. The patch types also differed in terms of their environmental conditions. Conifer patches had drier, cooler soils and the lowest understory light. Broadleaf patches had the warmest soils while understory light during the leaf-off period was similar to that of canopy gaps. Gap patches had the lowest litter cover and PO₄⁻ availability and the highest light. Seven environmental variables (soil moisture, soil temperature, total light during the leaf-off period, cover of coarse and fine downed woody material, and availability of NH₄⁺ and Ca²⁺) were significantly related to understory species composition (in a constrained ordination by means of a distance-based redundancy analysis (db-RDA); 16.5% of variation in understory community data explained). Even within a single patch type, there was substantial environmental variation that was related to understory species composition. Our study suggests that the mosaic of canopy patches within mixedwood forests supports coexistence of both early and late successional understory plant species in mixedwood stands. Maintaining the mixture of canopy patch types within mixedwood stands will be important for conserving the natural patterns of understory plant composition in boreal mixedwood forests.     

Identification and description of forested vegetation communities available to woodland caribou: relating wildlife habitat to forest cover data

Multivariate analyses were applied to forest canopy and understory data collected in 70 forest stands in the Mid-Boreal Upland Ecoregion in Saskatchewan. Random sampling of various types of forest focused on vegetation strata shown elsewhere to be of importance to woodland caribou (Rangifer tarandus caribou Gmelin). Cluster analyses revealed seven community types based on summer vegetation, and six community types when considering vegetation present during winter. Ordination by nonmetric multidimensional scaling suggested that two of the seven summer communities be combined, and that six community types be recognized in each season. Stands dominated by Populus tremuloides Michx. were all placed in one community type as were those dominated by Picea glauca (Moench) Voss. Young stands of Pinus banksiana Lamb. were placed in one community type, while the division of older stands was related to degree of canopy closure. The primary division among Picea mariana (Mill.) B.S.P. dominated stands was also based on the degree of canopy closure. Canonical correspondence analysis revealed strong relationships between the vegetation communities and data on canopy characteristics contained in the provincial forest inventory. In general, the techniques we employ provide a framework for objectively deriving vegetation communities relevant to wildlife species, and measuring their correspondence with readily available forest cover data. Specifically, we describe community types available to woodland caribou in our study area and their relationship to the provincial forest inventory.     

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.     

Oak restoration in water-limited pine plantations: interactive effects of overstory light interception and water availability on understory oak performance

Oak regeneration within pine monocultures is an opportunity to diversify forest structure. We examined the relationships between overstory (Pinus brutia) light interception and understory oak (Quercus ithaburensis) performance in water-limited forests. The study was performed in a mature pine plantation in Mediterranean Israel. Twenty-year-old oaks differing in location with respect to pine overstory and representing a gradient of light availability, such as open space (irradiance 100 %), interface (17–77 %), and understory (14–23 %), were monitored. Photosynthetic photon flux density (PPFD), leaf gas exchange, and twig water potential (TWP) were measured during the growth season under increasing drought stress. Predawn TWP decreased sharply from early to late spring and was positively related to irradiance during mid-spring only. Predawn to midday TWP gradient was positively related to irradiance mostly so during mid-spring. Daily averages of stomatal conductance (gs), net carbon assimilation rate (A), and transpiration rate (E) were highest in early spring and decreased gradually toward late spring. They were positively related to irradiance though this relationship became less pronounced from early to late spring. Oak height and stem basal area were positively related to irradiance. A/gs ratio was positively related to irradiance throughout the entire growth season. It increased from early to mid-spring but decreased toward late spring. A/PPFD ratio decreased from early to late spring showing a negative relationship with irradiance. We concluded that light availability was mainly responsible for spatial variation in oak performance and proposed that small-scale overstory gaps aiming for direct sunlight exposure during early spring should achieve maximum understory oak performance with minimal pine removal.     

Effectiveness of basal area for estimating canopy cover of ponderosa pine

Basal area is commonly used as a measure of canopy cover when describing overstory-understory relationships in forest ecosystems. We evaluated basal area and a number of other biotic and abiotic variables from 61 stands along the Rocky Mountain Front Range to establish whether this procedure has scientific merit and ascertain whether other models might provide a more precise estimator. Only basal area was correlated to overstory canopy and that relationship broke down above a canopy cover of 60%. Quadratic and piecewise linear models both provided an adequate fit of data over all stands sampled, however, the latter was regarded as better because (1) it reflected the straight-line relationship found below 60% canopy cover and (2) it objectively estimated the change-over point from a positively correlated model to one of no correlation. Basal area explained about 83% of the variation in canopy cover in both models.     

Can we use shelterwoods in Mediterranean pine forests to promote oak seedling development?

The use of shelterwoods to favour the development of natural or underplanted seedlings is common in temperate forests but rare in the pine forests of the Mediterranean area. Our aim was to assess the use of shelterwoods in Aleppo pine (Pinus halepensis) woodlands in southern France to promote the survival and growth of two co-occurring oak species: the deciduous Quercus pubescens and the evergreen Quercus ilex.Twelve Aleppo pine stands were selected and differentially thinned to create a light shelterwood (basal area = 10 m²/ha, irradiance 52%), a medium shelterwood (basal area = 19 m²/ha, irradiance 33%) and a dense shelterwood (basal area = 32 m²/ha, irradiance 13%). A total of 1248 sowing points, half composed of Q. pubescens and half of Q. ilex, were then set up in these three conditions. Seedling survival and growth were monitored for 3 years. Plant stress was assessed by measuring predawn leaf potential and photosynthetic performance through the F_v/F_m ratio. Soil moisture was also recorded at two depths during two growing seasons.Survival was high for both species in all three conditions due to three consecutive wet years. The lowest survival was recorded for Q. pubescens in the dense shelterwoods. Growth in diameter and height increased from the dense to the light shelterwoods. Shrubs developed more strongly in the light shelterwood, and increasing shrub cover enhanced height growth. Photosynthetic performance was lowest for Q. pubescens in dense shelterwoods and highest in light shelterwoods, whereas the reverse was true for Q. ilex. The lowest predawn potentials were recorded in the dense shelterwoods even though higher soil water content values were measured in this treatment during the summer drought.We show that light shelterwoods were more beneficial to growth than denser ones, indicating control mainly by light availability during the 3 years of the study. However, as lower soil moisture at 30-50 cm depth and faster understorey development were also recorded in this condition, more extended observation is needed to determine whether this benefit persists in subsequent years.     

Leaf area dynamics of a boreal black spruce fire chronosequence

Specific leaf area (SLA) and leaf area index (LAI) were estimated using site-specific allometric equations for a boreal black spruce (Picea mariana (Mill.) BSP) fire chronosequence in northern Manitoba, Canada. Stands ranged from 3 to 131 years in age and had soils that were categorized as well or poorly drained. The goals of the study were to: (i) measure SLA for the dominant tree and understory species of boreal black spruce-dominated stands, and examine the effect of various biophysical conditions on SLA; and (ii) examine leaf area dynamics of both understory and overstory for well- and poorly drained stands in the chronosequence. Overall, average SLA values for black spruce (n = 215), jack pine (Pinus banksiana Lamb., n = 72) and trembling aspen (Populus tremuloides Michx., n = 27) were 5.82 +/- 1.91, 5.76 +/- 1.91 and 17.42 +/- 2.21 m2 x kg-1, respectively. Foliage age, stand age, vertical position in the canopy and soil drainage had significant effects on SLA. Black spruce dominated overstory LAI in the older stands. Well-drained stands had significantly higher overstory LAI (P < 0.001), but lower understory LAI (P = 0.022), than poorly drained stands. Overstory LAI was negligible in the recent (3-12 years old) burn sites and highest in the 70-year-old burn site (6.8 and 3.0 in the well- and poorly drained stands, respectively), declining significantly (by 30-50%) from this peak in the oldest stands. Understory leaf area represented a significant portion (> 40%) of total leaf area in all stands except the oldest.     

Plant and carabid beetle species diversity in relation to forest type and structural heterogeneity

The aim of this study was to evaluate the influence of forest structure (mainly resulting from human uses) and forest type (the identity of the dominant tree species) on biodiversity. We determined the diversity of two taxonomical groups: the understory vegetation and the edaphic carabid beetle fauna. We selected eight types of forest ecosystems (five replicates or stands per forest type): pine (Pinus sylvestris) plantations of three age classes (10, 40 and 80 years since reforestation), an old-growth relict natural pine forest, and four types of oak (Quercus pyrenaica) stands: mature forests with livestock grazing and firewood extraction, mature forests where uses have been abandoned, “dehesa” ecosystems and shrubby oak ecosystems. The results obtained by a global PCA analysis indicated that both tree size and dominant species influenced the ordination of the 40 forest stands. In general, carabids were more sensitive to changes in forest heterogeneity and responded more clearly to the analysed structural variables than the understory vegetation, although the species richness of both groups was significantly correlated and higher in case of oak forests. Pine forest ecosystems were characterised by the lowest species richness for both taxonomical groups, the lowest plant diversity and by the lowest coefficients of variation and, consequently, low structural heterogeneity. As a result, it was very difficult to discriminate the effects of the spatial heterogeneity and the dominant tree species on biodiversity.     

Windthrow and salvage logging in an old-growth hemlock-northern hardwoods forest

Although the initial response to salvage (also known as, post-disturbance or sanitary) logging is known to vary among system components, little is known about longer term forest recovery. We examine forest overstory, understory, soil, and microtopographic response 25 years after a 1977 severe wind disturbance on the Flambeau River State Forest in Wisconsin, USA, a portion of which was salvage logged. Within this former old-growth hemlock-northern hardwoods forest, tree dominance has shifted from Eastern hemlock (Tsuga canadensis) to broad-leaf deciduous species (Ulmus americana, Acer saccharum, Tilia americana, Populus tremuloides, and Betula alleghaniensis) in both the salvaged and unsalvaged areas. While the biological legacies of pre-disturbance seedlings, saplings, and mature trees were initially more abundant in the unsalvaged area, regeneration through root suckers and stump sprouts was common in both areas. After 25 years, tree basal area, sapling density, shrub layer density, and seedling cover had converged between unsalvaged and salvaged areas. In contrast, understory herb communities differed between salvaged and unsalvaged forest, with salvaged forest containing significantly higher understory herb richness and cover, and greater dominance of species benefiting from disturbance, especially Solidago species. Soil bulk density, pH, organic carbon content, and organic nitrogen content were also significantly higher in the salvaged area. The structural legacy of tip-up microtopography remains more pronounced in the unsalvaged area, with significantly taller tip-up mounds and deeper pits. Mosses and some forest herbs, including Athyrium filix-femina and Hydrophyllum virginianum, showed strong positive responses to this tip-up microrelief, highlighting the importance of these structural legacies for understory biodiversity. In sum, although the pathways of recovery differed, this forest appeared to be as resilient to the compound disturbances of windthrow plus salvage logging as to wind disturbance alone, by most vegetative measures.