Soil seed bank dynamics in relation to topographic position of a mixed-deciduous forest in southern New England,USA

We examined the floristic significance of soil seed banks in relation to valley, midslope and ridge sites in a 70–90-year old forest in northeastern Connecticut. A-horizon mineral soils were collected to 5 cm depth in the early spring from forest understory sites across the topography. Samples from each of the sites were exposed to full sun within a greenhouse. Records of germination were made at regular intervals over a 60-day period. These showed significant differences among sites in number of species and total number of germinants. Greatest numbers of species and germinants were recorded from valley sites and these progressively declined from midslope to ridgetop. Twenty-five different species were identified. Species were grouped into growth habits – graminoids, herbs, shrubs, trees, and vines. Over 61% of all germinants across all sites were graminoids. Seventy-four percent of all germinants in the valley sites were graminoids, with over 93% of them represented by two sedge species, Carex glaucodea and C. lupulina. On the ridgetop sites graminoids were more evenly distributed among six different species. The percentage in each growth habit changed rank across topographic position with germinants of graminoids and trees most abundant on valley sites; herbs, on midslopes, and shrubs, on ridgetops. All germinants, except for those of the trees, Carex spp., and two herb species were weedy species that were not characteristic of the existing vegetation. Germinants of the trees, Carex spp., and herbs that were characteristic of the existing vegetation were mostly confined to soils from the valley sites. Two weedy herbs, Plantago major and Verbascum thapsus, are exotic introductions that originally came from Europe. Only one vine, Vitis aestivalis, from a midslope site germinated. Species diversity is higher on midslope sites than valleys and ridgetops. The significance of these findings in relation to site productivity and disturbance history is discussed.     

Comparison of tree growth and undergrowth development in aerially seeded and planted Pinus tabulaeformis forests

Direct seeding is a less expensive practice than planting and has the potential to become a viable alternative to transplanting for afforestation and regeneration purposes. As an effective and a less costly regeneration method, aerial seeding has been applied with several tree species. As early as 1956, Chinese people engaged in aerial seeding and stands with a total of 2.97×10⁷ hm² have been developed up to 2004. Our study tested whether the growth of planted Chinese pine (Pinus tabulaeformis Carr.) seedlings and its undergrowth development in northwest aspects differ from that of aerially sown seedlings on the northern and northwestern aspects of slopes. In 2007, we collected data such as height, diameter at breast height (DBH), clear bole height and canopy widths of trees, abundance, coverage, and frequency of shrubs and herbs from 21-year-old planted Chinese pine stands on a northwestern aspect (PNW), aerially sown stands in a northwest aspect (ANW) and aerially sown stands in a northern aspect (AN). Results showed that the relation of crown area and mean DBH was best fitted by a double inverse model for the ANW and AN forests and by a quadratic model for the PNW forest. There was no difference in the growth between ANW and AN forests, while growth was significantly higher in the PNW forest than in the ANW and AN forests. That was consistent with the Sorenson diversity indices in the shrub and herb layers, indicating that there was a large number of the same species in both aerially seeded stands, although their locations were different. Both the number of species in the undergrowth and the Shannon-Wiener index in the shrub layer were higher in the PNW stands than in the ANW and AN stands. Dominant families for all three stands were Rosaceae and Compositae in the shrub and herb layer, respectively. The dominant species for all three stands was Spiraea pubescens in the shrub layer, while the dominant species was different from each other in the three stands. The discrepancy in diversity and composition of species in the herb layer show that herbs are sensitive to shrubs in the three forests. High mortality and skewed diameter distributions reflect severe competition and too high a density in the aerially seeded forests. Thus, aerial seeding is a viable and effective regeneration technique, but management practices, such as thinning, should be applied to these forests.     

Human impacts drive forest structure and diversity. Insights from Mediterranean mountain forest dominated by <Emphasis Type="Italic">Abies pinsapo</Emphasis> (Boiss.)

Environmental factors and land-use are likely interacting to drive forest structure and species diversity, making it difficult to disentangle their separate impacts. Both components influence the resilience and stability of mountain forests in the Mediterranean Basin, since secular land-use has shaped stands structure and forests dynamics are constrained to the environment. Approaches covering environmental factors and contrasting land-uses are still needed to understand their implications on forest dynamics. We investigated patterns of environmental variables, stand structure and biodiversity in forest dominated by Abies pinsapo for 61 stands over their whole range. In addition, since land-use by local inhabitant could be a key factor, logging intensity was quantified by stumps basal area and recent land-use history was investigated. Partial redundancy analysis (pRDA) was conducted to assess to which degree the structural attributes are related to environmental characteristics and/or the land-use in the main A. pinsapo forests. Environmental characteristics accounted for 13% of the total variance, while the contrasting land-use (Spain vs. Morocco) and the stumps basal area, as indicator of human logging, accounted for 23% of the total variance. Trees species diversity was near 5 times higher in Moroccan stands than in stands from south Spain. According to recent human logging, Spanish stands have been strictly protected, but Moroccan stands showed widespread axe-logging by local inhabitant over the smaller tree sizes. Our results suggest that the preservation of a minor perturbation regimen by local populations enhance and help to preserve the whole biodiversity of this relict habitat.     

Managing forests infested by spruce beetles in south-central Alaska: Effects on nitrogen availability,understory biomass,and spruce regeneration

In Alaska, an outbreak of spruce beetles (Dendroctonus rufipennis) recently infested over one million hectares of spruce (Picea spp.) forest. As a result, land management agencies have applied different treatments to infested forests to minimize fire hazard and economic loss and facilitate forest regeneration. In this study we investigated the effects of high-intensity burning, whole-tree harvest, whole-tree harvest with nitrogen (N) fertilization, and conventional harvest of beetle-killed stands 4 years after treatment, as well as clear-cut salvage harvest 6 years after treatment. We measured available soil ammonium and nitrate and estimated N loss from leaching using in situ cation and anion resin exchange capsules. We also assessed spruce regeneration and responses of understory plant species. Availability and losses of N did not differ among any of the management treatments. Even a substantial application of N fertilizer had no effect on N availability. Spruce regeneration significantly increased after high-intensity prescribed burning, with the number of seedlings averaging 8.9 m⁻² in burn plots, as compared to 0.1 m⁻² in plots that did not receive treatment. Biomass of the pervasive grass bluejoint (Calamagrostis canadensis) was significantly reduced by burning, with burn plots having 9.5% of the C. canadensis biomass of plots that did not receive treatment. N fertilization doubled C. canadensis biomass, suggesting that N fertilization without accompanying measures to control C. canadensis is the least viable method for promoting rapid spruce regeneration.     

Identification of boreal forest stands with high herbaceous plant diversity using airborne laser scanning

Boreal forest stands with high herbaceous plant species diversity have been found to be one of the main habitats for many endangered species, but the locations and sizes of these herb-rich forest stands are not well known in many areas. Better identification of the stands could improve both their conservation and management. A new approach is proposed here for locating the mature herb-rich forest stands using airborne laser scanner (ALS) data and logistic regression, or the k-NN classifier. We show that ALS technology is capable of distinguishing the ecologically important herb-rich forests from those growing on less fertile site types, mainly on the basis of unique but quantifiable crown structure and vertical profile that characterise forests on high fertility sites. The study site, Koli National Park, is located on the border of the southern and middle boreal vegetation zones in Finland, and includes 63 herb-rich forest stands of varying sizes. The model and test data comprised 274 forest stands belonging to five forest site types varying from very fertile to poor. The best overall classification accuracy achieved with the k-NN method was 88.9%, the herb-rich forests being classified correctly in 65.0% of cases and the other forest site types in 95.7%. The best overall classification accuracy achieved with logistic regression was 85.6%, being 55.0% for the herb-rich forests and 94.3% for the other forest site types. Both methods demonstrated promising potential for separating herb-rich forests from other forest site types, although slightly better results were obtained with the non-parametric k-NN method, which was capable of utilising a higher number of explanatory variables. It is concluded that ALS-based data analysis techniques are applicable to the detection of mature boreal herb-rich forests in large-scale forest inventories.     

Forest natural regeneration and biomass production after slash and burn in a seasonally dry forest in the Southern Brazilian Amazon

This study estimates the aboveground biomass accumulation after forest clearing and slash burning and describes the structure and successional development of the secondary forest in the seasonally dry southern Amazon. The original burn study was conducted in four land clearings in 1997, 1998, and 1999. The size of the clearings varied from 1 to 9 ha. The native forest was felled, allowed to dry for approximately three months and then burned by the end of the dry season. A census was conducted in the central 1-ha forest on each site prior to the area's felling and burn. The aboveground biomass (AGB) and structure were similar to other primary tropical forests. However, the high density of Cecropia spp. before the forest felling and burn treatment indicates past low intensity disturbances. Seven and eight years after the fire, the fallow forests were still in an early successional stage dominated by Cecropia spp. The four areas had a high biomass accumulation during the studied period, varying from 7.5 to 15.0 Mg ha⁻¹ year⁻¹. The lower biomass accumulation in one plot was an effect of a higher fire severity, produced by the one-year difference in time between slash and burn of the forest, slowing the natural regeneration of Cecropia spp. The time needed for this forest to recover to the pre-fire AGB levels ranged from 20 to 30 years, assuming the current AGB accumulation rates are maintained. Considering these results, the maintenance of regenerating secondary forests in the Amazon would be a significant contribution to soil and watershed protection, minimizing biodiversity losses and perhaps mitigating climatic changes effects in the region.     

Acidification of primeval forests in the Ukraine Carpathians: Vegetation and soil changes over six decades

Changes to vegetation and soil were assessed in primeval forests of the Eastern Carpathians after a period of 59-68 years. We hypothesized that forest ecosystems were acidified through the long-distance transport of air pollutants. A total of 141 relevés and 20 soil profiles that had been studied in 1938 in spruce- and beech-dominated forests along an altitudinal gradient ranging from 1085-1575 m a.s.l. were re-surveyed from 1997 to 2006. Relevés were analyzed using multidimensional statistics and plant community characteristics (Shannon-Wiener’s index, equitability, fidelity, Ellenberg indication values - EIV); soil reaction and sorption complex properties were analyzed in soils.A total of 159 vascular plant taxa were recorded in 1938, of which 35 were not found during the repeat survey. During the later survey, 137 taxa were found, of which 13 were new findings. The upper mineral (A) as well as cambic or spodic (B) horizons were considerably acidified in both forest types. Both active and exchange soil reaction decreased by 0.1-0.3 units on average, exchangeable acidity significantly increased and the sum of base cations decreased in both soil horizons and forest types. Base saturation decreased by more than half of original values, with a maximum decrease of 68% found in the B-horizon of spruce forests. Whereas the herb layer developed along with soils in beech-dominated forests, EIV values for soil reaction increased in spruce-dominated forests, probably due to the movement of broadleaf woody species to higher elevations or due to the higher resistance of herb species to soil acidification. Significant changes to EIVs also occurred in the beech- and/or spruce-dominated forests for the factors of nitrogen, light moisture and temperature.There was an expansion of the lower tree and shrub layers, primarily Fagus sylvatica, Picea abies and Sorbus aucuparia in intermediate and higher elevations, which can be explained by reduced cattle grazing. Also, the dissipation of Juniperus communis and marked decline of Abiesalba are interpreted as being related to gradual changes in landscape management along with the effect of acid deposition. Since 1938, all stands have shown a significant increase in nitrophilous taxa such as Rubusidaeus, Athyrium distentifolium, Urtica dioica, Calamagrostis arundinacea, Stellaria nemorum. Significant decreases in the number of species, Shannon-Wiener’s index and equitability were only observed in spruce-dominated forests. Neophyte taxa were not detected in either the 1930s or the 1997-2006 period.     

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.     

Stand carbon stocks and soil carbon and nitrogen storage for riparian and upland forests of boreal lakes in northeastern Ontario

The establishment of shoreline reserves (buffer strips) has guided riparian forest management in Ontario for many years. A riparian area is defined as the transitional zone between the aquatic and terrestrial environments and therefore is also known as the aquatic/terrestrial ecotone. While many functions of riparian forests have been recognized and well studied, less is known about their potential to sequester C and whether this potential differs from other areas in the boreal forest landscape. Increased harvesting pressure due to decreased wood supply in Ontario and debate about the effectiveness of the current reserve guidelines has resulted in a renewed interest in harvesting riparian forests. In this study riparian and upslope forest C and soil C and N storage were quantified for 21 lakes shorelines at the Esker Lakes Research Area, a boreal forest ecosystem in northeastern Ontario, Canada. Objectives were to compare the C and N storage potential of riparian forests with those of adjacent upland forests, and to examine the potential impacts of harvesting on C stocks in riparian zones of the boreal forest.Riparian forests did not differ from upslope stands in terms of total aboveground overstory C storage although there were significant differences in stocking density and species composition. However, a greater proportion of total site C in riparian areas was stored in the overstory tree layer (>5 cm dbh) compared to upslope areas. Forest floor layers were deeper and stored more C and N in riparian forest stands in comparison to upslope stands. In contrast, mineral soil in upslope stands had greater C and N storage than mineral soil horizons within the riparian forest. As a result, the riparian organic horizons comprise a larger percentage of the overall soil storage of C and N than upslope layers. Currently practiced full-tree harvesting would result in a removal of approximately 76% of total aboveground C (17% of the ecosystem C) in upslope stands compared to 98% of total aboveground C (35% of the ecosystem C) in riparian forests. Selective or modified harvesting in riparian zones could decrease C removal to levels equal to that obtained by full-tree harvesting in upslope areas.     

Productivity of temperate broad-leaved forest stands differing in tree species diversity

? Understanding the effects of tree species diversity on biomass and production of forests is fundamental for carbon sequestration strategies, particularly in the perspective of the current climate change. However, the diversity-productivity relationship in old-growth forests is not well understood.

? We quantified biomass and above-ground production in nine forest stands with increasing tree species diversity from monocultures of beech to stands consisting of up to five deciduous tree species (Fagus sylvatica, Fraxinus excelsior, Tilia spp., Carpinus betulus, Acer spp.) to examine (a) if mixed stands are more productive than monospecific stands, (b) how tree species differ in the productivity of stem wood, leaves and fruits, and (c) if beech productivity increases with tree diversity due to lower intraspecific competition and complementary resource use.

? Total above-ground biomass and wood production decreased with increasing tree species diversity. In Fagus and Fraxinus, the basal area-related wood productivity exceeded those of the co-occurring tree species, while Tilia had the highest leaf productivity. Fagus trees showed no elevated production per basal area in the mixed stands.

? We found no evidence of complementary resource use associated with biomass production. We conclude that above-ground productivity of old-growth temperate deciduous forests depend more on tree species-specific traits than on tree diversity itself.

     

Comparison of potential non-timber forest products in intensively managed young stands and mature/old-growth forests in south-central British Columbia

Development of understory vegetation has been influenced by the many densely stocked second-growth forest stands in North America, which have an extended stem exclusion successional stage. Understory composition and structure is important for ecosystem functioning, while also having social and economic value through the harvest of certain herb and shrub species. The potential for co-management of young and mature, managed and unmanaged stands for timber and non-timber forest products (NTFPs) was assessed in two separate replicated experiments. Experiment A examined pole-sized lodgepole pine (Pinus contorta) stands that had been pre-commercially thinned (PCT) to target densities of 250, 500, 1000, and 2000 stems/ha. Half of each of these four thinning units was repeatedly fertilized, resulting in eight experimental units. Experiment B examined six different stand types: young plantations, pole-sized lodgepole pine stands either PCT, PCT plus repeated fertilization, or unthinned, mature, and old growth. Fifty-four herb and shrub species were identified as potential NTFPs, with the responses of individual species, as well as mean total herb, shrub, berry-producing and overall total NTFPs being assessed. In Experiment A, mean total abundance (crown volume index) of NTFPs, as well as mean total herb NTFPs were significantly greater in fertilized than in unfertilized stands. Thinning treatments did not significantly affect NTFP volume, however, fertilization treatments produced five significant responses by individual species (Achillea millefolium, Epilobium angustifolium, Taraxacum officinale, Arctostaphylos uva-ursi, Rubus idaeus). In Experiment B, four of the six species responses that were significant had greater abundance in young, managed stands (young plantation, thinned, or thinned-fertilized) than in the unmanaged stands. Mean total NTFP volume and mean total herb NTFP volume also followed this pattern. A. uva-ursi, E. angustifolium, Lonicera involucrata, Sorbus sitchensis and Thalictrum occidentale all had significantly higher abundance in young, managed stands than in all other treatments. Results suggest that co-management for timber and NTFPs is possible in this ecosystem, with further research needed to evaluate livelihood values of these crops.     

Links between plant diversity, carbon stocks and environmental factors along a successional gradient in a subalpine coniferous forest in Southwest China

In all, 48 sites of subalpine coniferous forest that had undergone natural regeneration for 5-310 years were selected as study locations in the Southwest China. We compared species richness (S), plant diversity (Shannon-Wiener index, H′; Margalef index, R), and above- and below-ground ecosystem carbon (C) pools of six plant communities along a chronosequence of vegetation restoration, and we also examined evidence for a functional relationship between plant diversity and C storage. Our results showed that above-ground C increased significantly (over 52-fold), mainly due to the increase of C in aboveground living plants and surface litter. Soil organic carbon (SOC) content increased from the herb community type (dominated by Deyeuxia scabrescens, P1) to mixed forest type (dominated by Betula spp. and Abies faxoniana, P4), which constituted the main C pool of the system (63-89%), but decreased thereafter (communities P5-P6). The mean C stock in the whole ecosystem - trees, litter layer and mineral soil - ranged from 105 to 730 Mg C ha⁻¹ and was especially high in the spruce forest community type (dominated by Picea purpurea, P6). On the other hand, the relationships between C stocks (soil, aboveground) and mean annual temperature or altitude were generally weak (P > 0.05). Moreover, we did not detect a relationship between S and aboveground C storage, while we found a significant negative relationship between H′, R and aboveground C storage. In addition, our experiment demonstrated that total root biomass and litter C/N ratio were significant functional traits influencing SOC, while S, R, and H′ had little effect. Path analysis also revealed that litter C/N ratio predominantly regulated SOC through changes in the quantity of microorganisms and soil invertase enzyme activity.     

Grapevine (Vitis spp.) dynamics in association with manual tending,physiography, and host tree associations in temperate deciduous forests

The role of lianas (woody vines) in the development and diversity of both tropical and temperate forests under differing management scenarios has not been thoroughly explored. We examined changes in grapevine (Vitis spp.) densities over time in clearcut stands as influenced by manual tending, physiography, and host tree associations. We used data from long-term studies on 66 clearcut stands dominated by temperate deciduous forest tree species on the Hoosier National Forest in south-central Indiana, USA. Fourteen of the stands had grapevines removed manually, approximately during the stem exclusion stage of development. Grapevine densities steadily increased from age 5 until age 15, which coincides with the period of stem exclusion of these stands. Subsequent grapevine mortality may have been related to light competition. Manually treated stands had similar grapevine densities as untreated stands after 20 years across sites, and it appeared that only on the most xeric area was the grapevine treatment effective in reducing grapevine densities. During early stand development, ranging from ages 5 to 17, grapevine density was strongly related to slope position, but as stands developed through the stem exclusion stage, aspect emerged as a stronger factor influencing grapevine density. Black cherry (Prunus serotina Ehrh.), walnut (Juglans spp.), and elm (Ulmus spp.) were the most common grapevine host trees under both treated and untreated scenarios, which may be associated with the crown architecture of these species. Results suggest lianas play a critical role in the early development of disturbed forest sites in temperate deciduous forests. With concerns that lianas are increasing in abundance and distribution in these forest types, understanding their role in forest dynamics, such as host tree associations at different stages of development, competition dynamics on different sites and corresponding influences on tree growth, species composition, and diversity, will be critical to decision-making processes in achieving desired management goals in the future.     

Spatial patterning and population structure of a common woodland herb, Trillium erectum, in primary and post-logging secondary forests

We investigated the population age structure and spatial patterning of a common understory perennial, Trillium erectum var. album, in primary and post-logging secondary (industrially logged in the early 1900s) cove forests of the southern Appalachian Mountains. Within each of 8 T. erectum populations (4 in primary forest and 4 in secondary forest), we mapped the locations of all T. erectum plants in a 10 m × 10 m plot located within the center of each population. We recorded the height and life stage of each plant and excavated a randomly selected subset for age determination. Trillium erectum populations in primary forests displayed a somewhat stronger and more consistent spatial structure than those found in secondary forests. However, aside from being on-average younger due to a higher concentration of juveniles, age structures of secondary forest populations were similar to those of primary forests and both forest types had similar maximum plant ages. Our results, when considered with the life history of the species, suggest that logging did not eliminate T. erectum from secondary forests. Rather, residual plants and patches of plants that survived logging provided the propagules needed to expand and reestablish populations following logging. This relatively rapid recovery stands in stark contrast to the slow recovery observed by others in post-agricultural secondary forest. When considered in the context of other studies of post-disturbance recovery in secondary forests, our results suggest that the full history of a site must be carefully considered when examining the impacts of disturbance on forest perennial herbs.     

Floristic and structural patterns along a chronosequence of secondary forest succession in Argentinean subtropical montane forests

We studied forest structure and composition along a chronosequence of secondary forest succession in Northwest Argentina's montane forests (‘Yungas’) at 27°S, between 700 and 900 m. Early herbaceous stages, forested stages of 11, 25, 45, and 50 years after abandonment, and old-growth forests were surveyed. Secondary forests included stands that originated in abandoned herbaceous crops and in abandoned fruit orchards. Basal area and species composition differed between 50-year-old secondary forests and old-growth forests. In contrast, tree density and species diversity were similar in the 50-year-old and in the old-growth forests. The previous use (herbaceous crops or fruit orchards) was an important influence on secondary forest composition. Whereas stands originating in herbaceous fields were dominated by wind-dispersed native species such as Heliocarpus popayanensis, Tecoma stans, Parapiptadenia excelsa, and Tipuana tipu, fruit-orchard-originated stands were dominated by animal-dispersed species. Among the animal-dispersed species, the exotic tree Morus alba was the most abundant, and its abundance in secondary forests seems to slow the succession toward old-growth forest composition. Overall, after accounting for differences attributable to pre-abandonment conditions, secondary forest succession showed a trend toward compositional convergence, with the rate of succession apparently regulated by the demography of long-lived pioneer species.     

Simulated biomass and soil carbon of loblolly pine and cottonwood plantations across a thermal gradient in southeastern United States

Changes in biomass and soil carbon with nitrogen fertilization were simulated for a 25-year loblolly pine (Pinus taeda) plantation and for three consecutive 7-year short-rotation cottonwood (Populus deltoides) stands. Simulations were conducted for 17 locations in the southeastern United States with mean annual temperatures ranging from 13.1 to 19.4 °C. The LINKAGES stand growth model, modified to include the “RothC” soil C and soil N model, simulated tree growth and soil C status. Nitrogen fertilization significantly increased cumulative cottonwood aboveground biomass in the three rotations from a site average of 106 to 272 Mg/ha in 21 years. The equivalent site averages for loblolly pine showed a significant increase from 176 and 184 Mg/ha in 25 years with fertilization. Location results, compared on the annual sum of daily mean air temperatures above 5.5 °C (growing-degree-days), showed contrasts. Loblolly pine biomass increased whereas cottonwood decreased with increasing growing-degree-days, particularly in cottonwood stands receiving N fertilization. The increment of biomass due to N addition per unit of control biomass (relative response) declined in both plantations with increase in growing-degree-days. Average soil C in loblolly pine stands increased from 24.3 to 40.4 Mg/ha in 25 years and in cottonwood soil C decreased from 14.7 to 13.7 Mg/ha after three 7-year rotations. Soil C did not decrease with increasing growing-degree-days in either plantation type suggesting that global warming may not initially affect soil C. Nitrogen fertilizer increased soil C slightly in cottonwood plantations and had no significant effect on the soil C of loblolly stands.     

Forest land-use history affects the species composition and soil properties of old-aged hillock forests in Estonia

Decisions regarding forest typology, management and protection are often based on the structures of present-day forests, ignoring their successional history. Forests growing on kames, eskers and various moraine hillocks common in regions with Holocene glaciation are good examples of this approach. In Estonia, these forests locally persist as fragments of continuous primary forest, but usually they are situated on former slash-and-burn areas (bushlands) or reforested agricultural land. Our aim was to elucidate the strength of the effect of long-term land-use history on the present-day vegetation compositions of mature hillock forests and their soil chemistry. It appeared that even the mature secondary hillock forests are still distinct from historically continuous stands in terms of species composition. We discovered connections between stand history and species content in hillock forests as well as transformed soil properties. The carbon and nitrogen contents in the humus horizons of secondary forests are lower while their carbon–nitrogen ratios are higher than in continuous forests. The relationship between vegetation and stand history is demonstrated by the higher proportions of anthropophytic and apophytic species in the herb layer of the secondary forests. The presence of species that are tolerant of anthropogenic impact on the secondary hillock forests floor can also be partly explained by the effect of different species in the tree and shrub layers, gaps in the tree canopy, and the boundary effect caused by the small areas of forest patches, neighboring grasslands or fields. The extinction debt in secondary communities should also be considered.     

Toward reference conditions: wildfire effects on flora in an old-growth ponderosa pine forest

Remote ponderosa pine (Pinus ponderosa) forests on the North Rim of Grand Canyon National Park, Arizona, USA provide valuable examples of reference conditions due to their relatively uninterrupted fire regimes, limited grazing history, and protection from logging. Wildfire is an important disturbance agent in upland forests of the Interior West, yet repeated measurements taken before and after lightning-ignited fires are rare. In 1999, a low-severity Wildland Fire Use fire burned 156 ha on Fire Point, a peninsula dominated by old-growth ponderosa pines, which had not burned for at least 76 years. We measured understory plant community and forest floor characteristics in 1998 (1 year before the fire) and 2001 (2 years after the fire) at this site and at nearby reference sites that did not burn in 1999 but have had continuing fire regimes throughout the past century. After the wildfire, the plant community at Fire Point shifted toward higher compositional similarity with the reference sites. Analysis of functional group composition indicated that this change was due primarily to an increase in annual and biennial forbs. Gayophytum diffusum, Polygonum douglasii, Chenopodium spp., Solidago spp., Elymus elymoides, Calochortus nuttallii, Hesperostipa comata, and Lotus spp. were indicative of forests influenced by recent fires. Species richness, plant cover, plant layer density and plant diversity were significantly lower at Fire Point than at the reference sites, possibly due to long-term fire exclusion, but the fire did not increase the rate of change in these variables after 2 years. Few exotic species were present at any site. Forest floor depths at Fire Point were reduced to depths similar to the reference sites, primarily due to consumption of the duff layer. There was a significant inverse relationship between the ratio of duff:litter and species richness. Compared to fire-excluded forests, old-growth ponderosa pine forests influenced by low-intensity surface fires generally have greater plant species richness (especially annual forbs) and lighter fuel loads. This study supports the continued application of the Wildland Fire Use strategy in old-growth montane forests to maintain and improve forest health by altering understory species composition and reducing fuel loads.     

Alternative silvicultural practices with variable retention to improve understory plant diversity conservation in southern Patagonian forests

Understory plants could can act as indicators of temperate forest sustainability, health and conservation status due to their importance in ecosystem function. Harvesting impacts on understory plant diversity depends on their intensity. Variable retention has been proposed to mitigate the harmful effects of timber harvesting, but its effectiveness remains unknown in southern Patagonian Nothofagus pumilio forests. The objectives of this study were to: (i) define a baseline of understory plant diversity in old-growth forests along a site quality gradient and under canopy gaps; (ii) evaluate stands with three different variable retention treatments compared to old-growth forests; and (iii) assess temporal changes during 4 years after harvesting (YAH). A 61 ha N. pumilio forest was selected. Understory plant (Dicotyledonae, Monocotyledonae and Pteridophyta) richness, cover (including woody debris and bare forest floor) and aboveground dry biomass were characterized in summer for 5 years. Before harvesting, baseline samples were conducted along a site quality gradient and outside/inside canopy gaps. Analyzed treatments include a control of old-growth forest (OGF) and three different harvesting treatments with variable retention: (i) dispersed retention (DR) of 30 m² ha⁻¹ (20-30% retention); (ii) aggregated retention (AR) with one aggregate per hectare and clear-cuts (28% retention); and (iii) combined dispersed and aggregated retention (DAR) with one aggregate per hectare and dispersed retention of 10-15 m² ha⁻¹ (40-50% retention). Data analyses included parametric and permutational ANOVAs, multivariate classification and ordinations.Before harvesting, 31 plant species were found, where richness, cover and biomass were directly related to site quality. The presence of canopy gaps did not have a significant impact on the measured variables. After harvesting, 20 new species appeared from adjacent associated environments (two from N. antarctica forests and 18 from grasslands and peatlands). At the stand level, understory values were higher in AR > DR > DAR > OGF. Most (81-95%) plant richness at baseline conditions was conserved in all treatments, where inside the aggregates understory remained similar to OGF. Combination of aggregated and dispersed retention (DAR) better limited exotic species introduction and protected sensitive species, improving conservation in harvested stands. Changes in understory variables were observed after the first YAH in all treatments; greater changes were observed in the harvested areas than in aggregates. Changes stabilized at the fourth YAH. As a conclusion, the location of retention aggregates should be selected to preserve species understory diversity of more speciose and diverse habitats or particularly uncommon stands. Implementation of different kinds (patterns and levels) of retention for improvement of biodiversity conservation in harvested forests should be included in timber and forest management planning.     

坡位对杉木×闽粤栲混交林和杉木纯林养分积累的影响

选择福建省建阳范桥国有林场16年生杉木×闽粤栲混交林与杉木纯林为研究对象,对不同坡位不同林分养分积累量进行分析对比。结果表明:上坡立地条件下,杉木纯林乔木层N、P、K养分积累量均大于杉木×闽粤栲混交林。中坡和下坡林分乔木层养分积累量则表现为混交林大于杉木纯林。不同林分乔木层养分积累量均占整个林分养分积累总量的90%以上,且表现为N>K>P和下坡>中坡>上坡。上坡杉木纯林的灌木层和草本层N、P积累量均大于混交林,下坡林分的灌木层N、P、K养分积累量均表现为杉木×闽粤栲混交林大于杉木纯林。不同林分凋落物的N和P的积累量均大大超过草本层和灌木层的积累量。