Understory abundance,species diversity and functional attribute response to thinning in coniferous stands

Alternative strategies for stand density management in even-aged coniferous forests may increase plant species and functional diversity. We examined the effects of fixed and variable density thinning on tree seedling regeneration as well as on abundance (indexed by cover) and richness of understory vascular plants 11 years after harvesting 45- to 66-year old forests dominated by Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) or western hemlock (Tsuga heterophylla (Raf.) Sarg.) at three sites in western Oregon. Each site contained an unthinned control (CON), and thinning treatments selected to enhance overstory structural diversity and spatial variability within stands (HD, high density treatment at 300 trees ha⁻¹; MD, moderate density treatment at 200 trees ha⁻¹; VD300, VD200 and VD100, variable density treatments at 300, 200 and 100 trees ha⁻¹). Leave islands are included in HD and the other thinning treatments contain both leave islands and gap openings. Tree seedling regeneration was highly variable and generally increased with thinning. Cover of all understory species was greater in VD100 than in the control whereas richness was greater in HD and MD. Cover and richness of early seral species were greater in most thinning treatments than in the control. Understory plant communities were overwhelmingly dominated by native species. In general, vegetation dynamics was accelerated by thinning, especially in variable density treatments. Cover of N-fixing understory species was greater in VD200 than in the other treatments, and in MD and VD300 than in the control, whereas richness of understory N-fixing species increased in all thinning treatments. Cover of understory species with intermediate soil water requirements was greater in MD, VD200 and VD100 than in the control, whereas richness of these species increased in VD200 compared to the control, HD and VD300. Thinning promoted higher diversity of understory conditions without reducing density and species richness of crop tree regeneration, and seemed to increase functional effect and response diversity.     

Understory Response to Repeated Thinning in Douglas-fir Forests of Western Oregon

This study investigated effects of a second-thinning entry on understory vegetation and tree regeneration development and understory vegetation composition. Study sites were located in the Coast Range and Cascade Range mountains of western Oregon and were dominated by Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) forests. Stands were initially thinned between 1975 and 1982 and parts of these same stands were thinned again approximately 20 yr later. Thinning stands a second time resulted in greater amounts of fern, graminoid, and open-site species, but the abundance of tree regeneration was not affected. Despite different site conditions, compositional patterns in the understory consistently shifted toward open-site early seral species following the second thinnings. These results suggest that the initial impacts of a second thinning are not simply predictable from studies in which only a single thinning was implemented. It is important to consider that vegetation trends were already influenced by the previous thinnings, and impacts of the second thinning are thus not as easily detectable. Within these limitations, repeated thinnings may be an effective management tool to maintain early seral species in older forests, while other aspects of understory vegetation and tree regeneration are less influenced in the short term.     

Effects of initial stand states on optimal thinning regime and rotation of Picea abies stands

Abstract

This study analysed the effects of young stand characteristics on optimal thinning regime and length of rotation periods for even-aged Norway spruce [Picea abies (L.) Karst.] stands. Stand development was based on a distance-independent, individual-tree growth model. The young stand data were collected from 12 well-stocked Norway spruce stands in southern Finland. Results showed that optimal thinning regimes and rotation period depend on site quality and initial stand characteristics. At the first thinning, optimal thinning type depended on initial density. Thinning from both ends of the diameter distribution turned out to be optimal for initially dense stands. At the second and subsequent thinnings, thinning from above was clearly superior. At a low interest rate, thinning from below was optimal for the first thinning regardless of stocking level. For the study data, optimal rotation periods varied from 61 to 92 years at 3% interest rate. The high variation in length of rotation period was due to the sensitivity of optimal length of rotation period to site qualities, initial stand structure and density.     

Silviculture as a disturbance regime: the effects of clear-cutting,planting and thinning on polypore communities in mixed forests

The structure of modern forest landscapes is profoundly affected by human-caused disturbances, particularly forest management; however, the effects and prospects of individual silvicultural techniques are insufficiently understood. This study distinguishes the effects of clear-cutting, planting and thinning on species richness and community composition of polypore fungi. In 2008–2009, 181 forest compartments (ranging from naturally regenerated deciduous stands to planted Picea abies stands and 0–137 years post clear-cutting) were explored in a hemiboreal landscape subjected to even-aged management. Altogether 104 polypore species were recorded. For species richness, time since clear-cutting was the most influential factor at both stand and landscape scales, followed by thinning. Clear-cuts had distinct polypore communities (including several red-listed species) whose species richness declined in time. Following 20 years post clear-cutting, species richness started to increase along different community–composition pathways determined by regeneration type. The communities developed after planting were moderately species rich at stand scale but homogeneous over larger areas. Thus, at landscape scale, mature unmanaged naturally regenerated stands hosted most species; thinning reduced species richness by approximately 15%, and among thinned stands, planted areas had a further 9–22% fewer species than naturally regenerated areas. In such variably managed landscape, silviculture appeared to create particularly distinct communities in young stands on nutrient-rich soils, which naturally provide polypores with a rich supply of small deciduous snags absent from stands artificially planted with P. abies and intensively thinned.     

Variable-density thinning promotes differential recruitment and development of shade tolerant conifer species after 17 years

Promoting patchy recruitment of shade tolerant tree species into the midstory is an important step in developing structural diversity in second-growth stands. Variable-density thinning (VDT) has been proposed as a strategy for accelerating structural diversity, as its combination of within-stand treatments (harvest gaps, thinning, and non-harvested skips) should create variable overstory and understory conditions. Here we report on western hemlock (Tsuga heterophylla (Raf.) Sarg.) seedling and sapling densities in five mixed-conifer stands and Sitka spruce (Picea sitchensis (Bong.) Carr.) seedling and sapling densities in two stands in western Washington at 3,7, 10, and 16–17 years after VDT. Additionally, we report on western hemlock advance regeneration growth and survival in two stands over 14 years. Western hemlock seedling density was highest in the thinned treatment but only significantly so in Year 10. In contrast, the gaps contained significantly more western hemlock saplings in Years 7 and 10 and significantly greater growth of western hemlock advance regeneration through Year 10. Skips embedded within the VDT did not differ significantly from unharvested reserves in terms of seedling or sapling densities of either species. Sitka spruce seedling density was highest in the gap and thinned treatments, but saplings were uncommon in all treatments. Collectively, these results indicate that our variant of VDT promoted patchy, midstory recruitment of western hemlock but failed to recruit Sitka spruce saplings in either stand where it established. Consequently, more intensive variants of VDT may be required to promote midstory recruitment of species less tolerant of shade than western hemlock.

     

Effects of Various Silvicultural Systems on Regeneration in Mixed Hardwood Stands of the Southern Appalachians

This study investigated the effects of clear-cutting and several other commonly used silvicultural systems on regeneration at seven sites in the Appalachian Mountains of Virginia and West Virginia. These even-aged oak dominated stands ranged in age from 63 to 100 yr and were located on medium quality oak sites (site indices from 18 to 23 m, base age 50). The treatments evaluated included a clear-cut, commercial harvest, shelterwood, leave-tree, group selection, preharvest herbicide, and control.

Mixed model ANOVA was used to analyze treatment response in five species groups: (a) oak (Quercus spp.); (b) maple (Acer spp.); (c) black cherry (Prunus serotina Ehrh.), and yellow-poplar (Liriodendron tulipifera L.); (d) miscellaneous overstory hardwoods; and (e) midstory species that typically do not occupy main canopy positions in mature stands. Response variables included stem density, importance value, average height for all regeneration, and the tallest 365 stems ha^?1. Comparisons were made among treatments, species groups, and between regeneration of sprout and seedling origin.

Alternative systems with residual trees reduced the regenerations overall mean height growth compared to the clear-cut by 0.34 to 0.74 m. Current conditions indicate oak will be a lesser component of the future stand, represented mostly through stump sprouting, and maple will likely increase in proportion among all treatments.     

Feasibility of implementing thinning in even-aged <Emphasis Type="Italic">Larix olgensis</Emphasis> plantations to develop uneven-aged larch–broadleaved mixed forests

Thinning experiments were conducted in larch (Larix olgensis) plantations to assess the feasibility of converting even-aged plantation stands to uneven-aged forests with more complex stand structures. Stands established in 1965 and 1960 were thinned in 2004 (Regime A, for determining the effect of recent thinning on emergence of seedlings) and 1994 (Regime B, for examining the effects of the past thinning on establishments of recruitments), respectively, at two intensities each. Natural regeneration, together with litter depth, canopy openness and vegetation cover, was surveyed in the thinned plots. Results indicated that larch seedlings started to emerge in May, reached a peak in June, decreased from June through September, and then disappeared in October. No larch seedlings exceeded 1 year old in the thinned plots because of the low levels of light and dense litter and vegetation cover. However, there were many naturally regenerated seedlings (5–50 cm in height) and saplings (50–500 cm in height) of broadleaved tree species such as Acer spp., Fraxinus spp., Cornus controversa, Quercus mongolica, and even the climax tree species, Pinus koraiensis, in the thinned plots. The mean density of regenerated seedlings reached 6.7 and 4.5 stems m⁻² in Regimes A and B, respectively, whilst the mean density of regenerated saplings reached 4,595 stems ha⁻¹ in Regime B. These results suggest that it is impractical to turn even-aged larch plantations to uneven-aged larch forests, but it may be feasible to develop uneven-aged larch-broadleaved forests from even-aged larch plantations through thinning.     

Early regeneration of Fraxinus rhynchophylla in the understorey of Larix kaempferi stands in response to thinning

The effects of thinning were investigated on naturally regeneratedFraxinus rhynchophylla Hance (Korean ash) grown in the understoreyof a Larix kaempferi (Lamb.) Carr. plantation, established ona site that previously supported old growth hardwood forests.The regeneration characteristics were evaluated after thinningof the plantation (about 60 per cent of relative density wasremoved). In the naturally regenerated and mixed stands, Quercusmongolica Fisch. was the most abundant tree species in the overstorey,but the understorey of the thinned plantations was dominatedby F. rhynchophylla with six other minor tree species. Fraxinusrhynchophylla was regenerated by four propagation types includingnatural seedling, stump sprout, root sprout and layering. Beforethinning, the difference of average height growth for each regenerationtype was insignificant. One year after thinning, the stump sproutsshowed the fastest growth among the regeneration types. Heightgrowth of advance regeneration was slow in the year followingthinning but then increased rapidly surpassing the other regenerationtypes 5 years after thinning. Thinning had an effect on thecomposition of regenerated trees also by seedling, stump sprout,root sprout and layering in the understorey. Advance regenerationsoccupied the upper understorey at 5 years after thinning. Alarge number of stump sprouts emerged in the plots 1 year afterthinning, while at 5 years after thinning the occupation ofstump sprouts decreased. Root sprouts and layers faded away.Considering the early growth and the composition of understoreyF. rhynchophylla, its advance regeneration has a high potentialto develop into overstorey trees at this site.     

Belowground competition from overstory trees influences Douglas-fir sapling morphology in thinned stands

We evaluated effects of belowground competition on morphology of naturally established coast Douglas-fir (Pseudotsuga menziesii var. menziesii (Mirb.) Franco) saplings in 60- to 80-year-old thinned Douglas-fir stands in southwestern Washington. We separately quantified belowground competition from overstory and understory sources using trenching and understory removal. In this light-limited environment of 26 ± 16% (std. dev.) full sunlight, 2-year exclusion of tree root competition by trenching increased sapling stem biomass by 18%, total aboveground biomass by 21%, number of interwhorl buds by 68%, total foliar biomass by 33%, and foliar biomass on branch components over 4 years old by 143%. Belowground competition did not influence shoot:root ratio or foliar efficiency (i.e., stem growth per unit foliage biomass). Sapling needle size, specific leaf area, and internodal distance also were not affected by belowground competition; these variables were apparently a function of the low-light environment. The principal source of belowground competition was roots of overstory trees; effects of belowground competition from understory vegetation were minor. Thus, under a partial overstory, morphology of Douglas-fir regeneration was influenced by both belowground and aboveground competition from overstory trees. In this environment, understory vegetation control would not likely influence belowground competition to an extent that would affect sapling morphology.     

Commercial thinning that maintained species diversity of a mixed black spruce–jack pine stand enhanced productivity

Commercial thinning enables forest managers to meet timber production objectives. Thinning reduces tree density to alleviate competition for resources and favour growth of selected tree species. However, in doing so, thinning can homogenize the composition of mixed-species forests and raise biodiversity issues. There is increasing evidence that species richness can lead to higher productivity through a complementarity effect. Hence, thinning that would maintain species diversity of mixed-species forests could enhance stand productivity and help forest managers to reconcile timber production objectives and biodiversity issues. The objective of this study was to compare post-thinning stand production, experimentally over 10 years, in mixed and monospecific stands of black spruce (Picea mariana [Mill.] B.S.P.) and jack pine (Pinus banksiana Lamb.). The post-thinning stand production curve of the mixed stand converged toward that of the unthinned mixed stand while the production curves of the thinned and unthinned monospecific stands remained parallel. The convergent productivity of the mixed stand could be explained by a positive interaction between effects of thinning and niche complementarity. We propose that thinning that maintains species diversity of mixed stands could help forest managers who are implementing ecosystem management to reconcile timber production objectives with biodiversity issues.     

Effects of thinning paper birch on conifer productivity and understory plant diversity

Naturally regenerated paper birch (Betula papyrifera Marsh.) is commonly removed from juvenile interior Douglas-fir (Pseudotsuga menziesii var. glauca [Beissn.] Franco) plantations in southern interior British Columbia, Canada, to increase conifer productivity and create a free-growing stand; however, this practice is expensive and contentious because of possible negative ecological impacts. One solution is to retain an optimal density of birch where growth gains of understory Douglas-fir are balanced against losses to Armillaria ostoyae (Romagn.) Herink and understory plant species diversity. We sought to find this optimal density by comparing four evenly applied birch density reduction treatments (0, 400, 1111, and 4444 retained birch stems ha^?1) and an unthinned control (>7300 retained birch stems ha^?1). The mortality rate of Douglas-fir due to Armillaria root disease increased non-significantly with thinning intensity. Mean diameter increment of surviving Douglas-fir improved the most where birch was completely removed, with little variation among intermediate thinning treatments. Height growth was unaffected by the thinning treatments. Diversity of cryptogams was significantly greater in the control than where all birch was removed. We suggest that the treatment with 4444 retained birch stems ha^?1 provides the best balance for improving Douglas-fir growth while minimizing risk of increased Armillaria root disease and reduced understory plant diversity in young mixed stands.     

香港人工林改造对林下植物自然更新 和物种多样性的早期影响

人工林在保护香港自然环境起着重要作用,但绝大多数人工林为纯林,生态系统较为脆弱。人工林的改造对于增加林地生物多样性、提升人工林的生态服务功能具有重要意义。研究选取20 a 生的尾叶桉（Eucalyptus urophylla）林和马占相思（Acacia mangium）林进行间伐套种处理,套种密度为1665株／hm2;以不间伐林分为对照,比较间伐套种后林分的物种多样性及套种树种的生长表现。结果表明,间伐＋套种3 a后,与未间伐林分相比,尾叶桉间伐林分植物的科、属和种数分别增加了78．6％、116．7％和100％,马占相思间伐林分相反。DCA 分析结果显示,群落中灌木层和草本层物种主要聚集在尾叶桉间伐林分,马占相思未间伐林分物种数次之。与未间伐相比,间伐＋套种措施显著提高了尾叶桉林乔木层 Shannon-Wiener 指数和均匀度指数、灌木层的物种丰富度指数和均匀度指数及草本层的丰富度和 Shannon-Wiener 指数（P ＜0．05）;但马占相思间伐＋套种林分的乔木层、灌木层和草本层的 Shannon-Wiener 指数均显著低于未间伐林分（P ＜0．05）;物种多样性指标总体表现为尾叶桉间伐林＞未间伐林,马占相思未间伐林＞间伐林。套种树种在尾叶桉和马占相思林分中的早期生长表现良好,树种间生长量没有明显差异。     

Genetic aspects of artificial regeneration of Douglas-fir (Pseudotsuga menziesii) in Bavaria

Douglas-fir has been planted in Bavaria for over 100 years. Its high growth potential and good wood quality combined with its silvicultural characteristics have increased interest in the species which can easily be integrated into existing natural ecosystems. In order to be successful in the establishment of Douglas-fir one must select the best provenance (only the so called “green” or viridis coastal type form is suitable for Bavaria), maintain a broad genetic base and carry out proper silviculture. The goal of the study was to check the origin/provenance (coastal or interior type) of older Bavarian Douglas-fir stands, the respective genetic variation of the stands and also the natural regeneration, the collected seed and the plants raised in the nursery. Possible changes in the genetic composition and diversity due to selective processes in the nursery phase were also investigated. Isoenzyme gene markers were used for the study. The majority of the investigated older stands apparently belong to the coastal type Douglas-fir; however, we also found mixed stands and stands solely of the interior (unsuitable) type. Older stands show large variation in their genetic diversity (v _gam between 19.9 and 90.1). Some of the stands of unsuitable provenance have natural regeneration and others have been selected as seed collecting stands. Through natural regeneration the genetic structure of the older stands is passed on to the following generation. The same is true for reproductive material collected from such stands. In comparison to seed collected from stands in the USA (from stands selected by the EU for seed collection), seed from Bavarian stands is genetically very heterogeneous in respect to the allele variants and the diversity. No significant changes in the genetic population structure were found in the nursery due to sowing or transplanting. The critical stage for obtaining high quality reproductive material of Douglas-fir is the selection of seed collecting stands and the harvesting procedure. The resulting consequences for seed collection and procurement of suitable reproductive material for Douglas-fir are discussed.     

Flowering of understory herbs following thinning in the western Cascades,Oregon

Thinning of young forest stands encourages development of the understory layer by increasing the levels of light and belowground resources. High-intensity thinning, with associated ground disturbance and high light levels, can lead to dominance by early seral species or by a few species of shrubs, while low-intensity thinning may not increase resource levels enough to encourage forest herbs. Changes in herb-layer abundance can be hard to detect because forest-floor herbs are often slow growing, but flowering increases rapidly in response to high resource levels. This study examined flowering of the understory herb community before and 5 years after low-intensity thinning. Flowering of 10 herb species was sampled in 62 nine-meter radius plots in six treatment units within the H.J. Andrews Experimental Forest. Thinning proved to be the most important determinant of the composition of the flowering assemblage (that is of which plants were flowering in a given plot). Old-growth species and release species (those specializing in large forest openings) showed higher numbers of flowering ramets following thinning. Release species also showed significant positive linear relationships with the plot-level degree of reduction in Douglas-fir (Pseudotsuga menziesii) basal area (BA), while old-growth species showed no significant relationships and forest generalist species showed significant negative linear relationships with reduction in BA. Plot-level reduction in BA explained little of the variation in numbers of flowering ramets per plot for most species except for the release species Hieracium albiflorum and the forest generalist Trillium ovatum. The overall lack of strong linear relationships between herb flowering and reduction in BA makes it difficult to predict optimal thinning intensities for these species. However, the results of this study suggest that even when low-intensity thinning does not significantly change herb abundance it could still influence the ecology of the understory herb community by increasing carbon allocation to sexual reproduction.     

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.     

Influence of fire and mountain pine beetle on the dynamics of lodgepole pine stands in British Columbia,Canada

An outbreak of the mountain pine beetle (Dendroctonus ponderosae Hopkins; MPB), currently affecting over 10.1 million hectares of lodgepole pine forests (Pinus contorta Dougl.) in British Columbia, Canada, is the largest in recorded history. We examined the dynamics of even-aged lodgepole pine forests in southern British Columbia, which were undergoing this MPB outbreak. Using dendroecology and forest measurements we reconstructed the stand processes of stand initiation, stand disturbances, tree mortality, and regeneration, and explained the current stand structure and the potential MPB impacts in selected stands. Our results indicate that stand-replacing fires initiated even-aged seral lodgepole pine stands in this region. In the absence of fire in the 20th century, multiple MPB disturbances, which each resulted in partial canopy removal, modified the simple one-layer structure of the fire-origin stands by the initiation of post-MPB disturbance regeneration layers, transforming the stands into complex, multi-aged stands. Despite high overstory mortality due to the current MPB outbreak, regeneration layers, which are likely to survive the current outbreak, will provide important ecological legacies and will contribute to mid-term timber supply.     

Fire-free natural regeneration in water limited Pinus halepensis forests: a silvicultural approach

We aimed to elucidate environmental and silvicultural factors that determine the extent of fire-free natural regeneration in east Mediterranean Pinus halepensis forests. The specific aims were to study the potential and identify bottlenecks for natural regeneration and examine the effects of overstory thinning and site preparation treatments. We integrated four experiments conducted in diverse ecoregions in Israel ranging from semiarid to subhumid. Seedling emergence and survival were traced for 2–4 years in Mishmar Ha’Emek Forest (MHF) with annual rainfall of 604 mm; Shaharia Forest (SF)—435 mm; and Yatir Forest (YF)—280 mm. Effects of thinning treatments: clearcut, 100 tree, 200 tree ha^?1 and control—310 tree ha^?1, were examined in MHF. Effects of site preparation treatments: soil scarification and herb clipping were examined in SF. Density of emerged seedlings varied among ecoregions and was linked to average annual rainfall. Within ecoregions, emerged seedling density was higher on north-facing slopes than on south-facing slopes and was strongly related to stand characteristics (e.g., stem basal area) and seed rain. Seedling survival rate of about 20 % was recorded in MHF while no seedling survival was observed in YF and SF during the study years. Thinning treatments reduced seed rain and emerging seedling density, but increased seedling survival and growth. Recruitment 4 years after thinning was highest in the 100 tree ha^?1 treatment and lowest in the control. In SF, seedling survival was limited mainly by herbaceous vegetation and was improved by soil scarification and herb clipping. Silvicultural implications are discussed.     

Responses of community structure,diversity, and abundance of understory plants and insect assemblages to thinning in plantations

Identifying effective management operations for plantations is important for conservation of biodiversity in a plantation-dominated landscape. We tested whether pre-commercial thinning influenced community structures and could be an effective strategy for increasing diversity and abundance of plants and animals in plantations. We designated thinned and unthinned study stands in Japanese cedar (Cryptomeria japonica) plantations and compared the communities of understory vegetation, bees, butterflies, hoverflies, and longhorn beetles 1 and 3 years after thinning. The analyses of vegetation showed that pre-commercial thinning affected community structure, but species richness and vegetation abundance were not significantly affected. Thinning affected insect community structure, and both species richness and abundance of all insect groups increased 1 year after thinning. However, 3 years after thinning, significant differences only remained in the species richness of bees and the abundance of bees, butterflies, and hoverflies. These results suggest that pre-commercial thinning in plantation stands influences the community structure of understory vegetation and can be an effective way to increase the diversity and abundance of some insect groups in the short term. However, the results also suggest that the duration of the operational effects of pre-commercial thinning varies among insect groups; thus, the variable effects of pre-commercial thinning should be carefully considered in the conservation-based management of plantation stands.