A suite of ecological indicators for evaluating the integrity of structural eco-complexity in Mexican forests

This paper presents a conceptual framework for analyzing forest complexity as the combination of the variety of species and key structures that are associated with the composition, structure, and function of forest stands. Several spatial indicators have been developed to characterize the biodiversity, the structural complexity, and anthropogenic effects that can be observed in Mexican forests. By integrating several stand complexity attributes, the forest condition can be characterized as a function of species composition, stand structural attributes, and forest development. In addition, indicators of anthropogenic effects were also analyzed to identify their influence on forest eco-complexity, and therefore, on the current condition of forests. The results of applying this conceptual framework showed that Mexican forest are ecologically complex, with varying levels of anthropogenic impacts that modify the structural forest characteristics, particularly in tropical forests. The main factor explaining the current eco-complexity condition in tropical forests was associated with early stages of forest development, due to ecological degradation, and showed a generalized loss of attributes, particularly for stand complexity and stand development. In contrast, temperate forests exhibited better eco-complexity conditions, especially for those attributes that define forest stand occupancy and development. Mining activities, forest extraction as selective harvesting, forest fires, land use change, and road openings are critical human activities that directly affect forest structure and, ultimately, modify forest eco-complexity and integrity. This eco-complexity index derived for Mexican forests can be used to integrate measures of forest structure and functioning, and thereby better inform decision making and policy development.     

Legacy of Pre-Disturbance Spatial Pattern Determines Early Structural Diversity following Severe Disturbance in Montane Spruce Forests

Background

Severe canopy-removing disturbances are native to many temperate forests and radically alter stand structure, but biotic legacies (surviving elements or patterns) can lend continuity to ecosystem function after such events. Poorly understood is the degree to which the structural complexity of an old-growth forest carries over to the next stand. We asked how pre-disturbance spatial pattern acts as a legacy to influence post-disturbance stand structure, and how this legacy influences the structural diversity within the early-seral stand.

Methods

Two stem-mapped one-hectare forest plots in the Czech Republic experienced a severe bark beetle outbreak, thus providing before-and-after data on spatial patterns in live and dead trees, crown projections, down logs, and herb cover.

Results

Post-disturbance stands were dominated by an advanced regeneration layer present before the disturbance. Both major species, Norway spruce (Picea abies) and rowan (Sorbus aucuparia), were strongly self-aggregated and also clustered to former canopy trees, pre-disturbance snags, stumps and logs, suggesting positive overstory to understory neighbourhood effects. Thus, although the disturbance dramatically reduced the stand’s height profile with ~100% mortality of the canopy layer, the spatial structure of post-disturbance stands still closely reflected the pre-disturbance structure. The former upper tree layer influenced advanced regeneration through microsite and light limitation. Under formerly dense canopies, regeneration density was high but relatively homogeneous in height; while in former small gaps with greater herb cover, regeneration density was lower but with greater heterogeneity in heights.

Conclusion

These findings suggest that pre-disturbance spatial patterns of forests can persist through severe canopy-removing disturbance, and determine the spatial structure of the succeeding stand. Such patterns constitute a subtle but key legacy effect, promoting structural complexity in early-seral forests as well as variable successional pathways and rates. This influence suggests a continuity in spatial ecosystem structure that may well persist through multiple forest generations.     

Epiphytic macrolichens in managed and natural forest landscapes: a comparison at two spatial scales

To maintain biodiversity in managed forests we must understand how forestry affects various organisms across a wide range of spatial and temporal scales. We compared landscape structure, forest structure, and species richness and abundance of epiphytic macrolichens in three pairs of natural and managed boreal forest landscapes. Study landscapes (2500 ha) were located within and adjacent to three of the largest forest reserves in Sweden (Reivo, Muddus, Jelka). The structural heterogeneity within landscapes was higher in managed forests whereas within-stand structural heterogeneity was higher in natural landscapes. Species richness of macrolichens at the stand level (sample plot) was 23% higher in natural forests but there was no difference at the landscape level. Most (86%) of the common species were more frequent in natural landscapes. Lichen abundance (estimated by lichen litter) was two times higher in natural than in managed landscapes, 5.6 and 2.7 kg ha^-1 forest (pooled data), respectively. Both species richness and abundance were negatively related to cutting level (number and basal area of cut stumps) and positively related to stand variables (stand age, stem density and basal area). Lichen-rich forest stands were more numerous but covered a smaller area and were more isolated in managed landscapes. This may in turn have important consequences for dispersal of lichen propagules to second-growth forests. In conclusion, the results suggest that effects of forestry on epiphyte diversity and abundance are strongly related to the spatial scale (stand or landscape). To enhance biodiversity in managed forests we must increase structural heterogeneity at the whole range of spatial and temporal scales.     

Changes in variance components of forest structure along a chronosequence in a wave-regenerated forest

We explored structural mean and variation along a chronosequence in a wave-regenerated forest on Mt. Shimagare, Japan, and examined differences in the components of variation between developmental stages. Because there are several strips of stands at the same developmental stage in the forest, between- and within-strip variances (i.e., variance components) were analyzed at each stage. Three strips were selected for each of three developmental stages (one for saplings to three for mature trees), and four plots were established in each of the strips. Stem densities and stand basal areas for trees taller than 20 cm were computed for all plots. Six Bayesian ANOVA models were compared to examine the temporal patterns of stage means, total variance, and the variance components for the two stand structural parameters. The results of the model comparison suggested that the variation in stem densities decreased from the sapling stage to the mature stage, but variation in stand basal areas remained roughly constant throughout forest development. For both density and basal area, the relative magnitudes of each variance component compared to the total variance differed between stages; the ratios of between-strip variation to within-strip variation were highest at the mature stage. Our results suggest that forest structure exhibited higher variability during development of the wave-regenerated forest than was assumed in previous studies and that the variance components for the forest structure varied temporally among developmental stages. This study thus illustrated a potential application of the chronosequence approach to infer the dynamics of a spatially heterogeneous and hierarchically structured ecosystem.     

不同采伐强度对阔叶红松林主要树种空间分布格局和物种空间关联性的影响

分析采伐后森林群落中物种的空间格局有助于认识该格局形成的生态学过程、种群的生物学特性及其与环境因子之间的相互关系,并对制定可持续的森林经营方案具有重要意义。以长白山地区经历不同采伐方式形成的阔叶红松林次生林为研究对象,利用空间点格局分析的研究方法,探讨了采伐对阔叶红松林主要树种空间分布格局、种间关联性以及更新的影响。研究结果显示:较低强度的择伐对阔叶红松林主要树种的空间分布格局、种间关联性的改变较小,群落可以在较短时间内恢复。中等强度的择伐减少了成年树种对幼树的抑制作用,可以促进森林的天然更新。皆伐后,森林的群落结构,物种的空间分布格局、种间关联性都发生显著变化,尽管更新状况良好,但要恢复到伐前水平仍需要较长时间。择伐不仅通过改变主要树种的密度对阔叶红松林群落结构产生影响,还通过改变物种空间关联性改变群落的结构动态。因此,在制定森林生态系统经营管理方案时,不仅要选择适合的采伐强度,还要综合考虑采伐时物种的选择以及种间关系。     

Fire disturbance and forest structure in old‐growth mixed conifer forests in the northern Sierra Nevada,California

Question: This study evaluates how fire regimes influence stand structure and dynamics in old‐growth mixed conifer forests across a range of environmental settings. Location: A 2000‐ha area of mixed conifer forest on the west shore of Lake Tahoe in the northern Sierra Nevada, California. Methods: We quantified the age, size, and spatial structure of trees in 12 mixed conifer stands distributed across major topographic gradients. Fire history was reconstructed in each stand using fire scar dendrochronology. The influence of fire on stand structure was assessed by comparing the fire history with the age, size, and spatial structure of trees in a stand. Results: There was significant variation in species composition among stands, but not in the size, age and spatial patterning of trees. Stands had multiple size and age classes with clusters of similar aged trees occurring at scales of 113 ‐ 254 m². The frequency and severity of fires was also similar, and stands burned with low to moderate severity in the dormant season on average every 9–17 years. Most fires were not synchronized among stands except in very dry years. No fires have burned since ca. 1880. Conclusions: Fire and forest structure interact to perpetuate similar stand characteristics across a range of environmental settings. Fire occurrence is controlled primarily by spatial variation in fuel mosaics (e.g. patterns of abundance, fuel moisture, forest structure), but regional drought synchronizes fire in some years. Fire exclusion over the last 120 years has caused compositional and structural shifts in these mixed conifer forests.     

Disturbance history and stand dynamics in tall open forest and riparian rainforest in the Central Highlands of Victoria

Abstract Spatial heterogeneity in the intensity of past disturbances has directly influenced the structure and composition of present‐day forests around the world. In south‐eastern Australia infrequent, high‐intensity wildfires are a major part of the historical disturbance regime. While these fires are often assumed to produce even‐aged stands, spatial heterogeneity in fire intensity due to highly variable topography may lead to more complex forest age structures. Our study describes the influence of disturbance on the age structure and dynamics of a mosaic of tall, open eucalypt forest, cool temperate rainforest and mixed species forest surrounding Bellel Creek in the Central Highlands of Victoria using dendrochronological techniques. We were particularly interested in the impacts of the 1939 Black Friday fire and its effects on forest age structure and subsequent stand development patterns. Within our study site tall open forest displayed two distinct age cohorts: (i) trees that established immediately after the 1939 fire and accounted for the majority of individuals in the forest, and (ii) scattered groups of older trees estimated to be approximately 200–250 years old. Cool temperate rainforest and mixed forest were also dominated by the post‐1939 fire age cohort. However, a greater proportion of trees in these forest types survived the 1939 fire relative to the tall open forest. The impact of the 1939 fire on the growth of surviving trees was highly variable but generally short‐lived. In most cases growth decreased after the 1939 fire, but generally returned to prefire levels within 1–3 years. Non‐fire disturbances were limited to small‐scale branch‐ and tree‐fall events, although the extreme snowstorm of 1977 appears to have caused extensive damage to rainforest communities. Our study demonstrates the opportunities for dendroecological studies to reconstruct historical dynamics and disturbance patterns in Australian forests and provides important insights into variation in landscape‐scale fire impacts and their effect on subsequent forest development patterns.     

A comparison of structural characteristics between old-growth and postfire second-growth hemlock–hardwood forests in Adirondack Park, New York, U. S. A.

1 This study compares the structural characteristics of 12 old‐growth and six postfire second‐growth hemlock–northern hardwood stands in north central Adirondack Park, New York, in order to test the null hypothesis that there are no differences in species composition, size structure, age structure and attributes such as dead wood and canopy gaps between old‐growth stands and this type of second‐growth forest. 2 The second‐growth forests of this study regenerated following widespread logging‐related fires in either 1903 or 1908; the old growth and second growth have similar environmental settings. 3 Estimates of stand ages, derived from an increment core of the oldest tree in each stand, range from 88 to 390 years. 4 Structural attributes are related to stand age (i.e. stage of development). In comparison with the second‐growth forests of this study, older stands are characterized as (a) a larger average diameter of canopy trees; (b) a greater basal area of trees; (c) a lower density of canopy trees and of all trees ≥ 10 cm d.b.h.; (d) a higher density of eastern hemlock (Tsuga canadensis (L.) Carrière) trees; (e) a higher density of large trees (≥ 50 cm d.b.h.); (f) larger canopy gaps; and (g) a greater volume of coarse woody debris (both logs ≥ 20 cm d.b.h. and snags ≥ 10 cm d.b.h.). 5 Despite differences between old growth and second growth, especially in species composition, it appears from observations of the 18 stands that second‐growth forests are developing some structural characteristics of old growth. 6 Structural attributes of the old‐growth forests are similar to characteristics of the same forest type in geographically distant areas in eastern USA.     

Quantifying Tropical Dry Forest Type and Succession: Substantial Improvement with LiDAR

Improved technologies are needed to advance our knowledge of the biophysical and human factors influencing tropical dry forests, one of the world's most threatened ecosystems. We evaluated the use of light detection and ranging (LiDAR) data to address two major needs in remote sensing of tropical dry forests, i.e., classification of forest types and delineation of forest successional status. We evaluated LiDAR‐derived measures of three‐dimensional canopy structure and subcanopy topography using classification‐tree techniques to separate different dry forest types and successional stages in the Guánica Biosphere Reserve in Puerto Rico. We compared the LiDAR‐based results with classifications made from commonly used remote sensing data, including Landsat satellite imagery and radar‐based topographic data. The accuracy of the LiDAR‐based forest type classification (including native‐ and exotic‐dominated forest classes) was substantially higher than those from previously available data (kappa = 0.90 and 0.63, respectively). The best result was obtained when combining LiDAR‐derived metrics of canopy structure and topography, and adding Landsat spectral data did not improve the classification. For the second objective, we observed that LiDAR‐derived variables of vegetation structure were better predictors of forest successional status (i.e., mid‐secondary, late‐secondary, and primary forests) than was spectral information from Landsat. Importantly, the key LiDAR predictors identified within each classification‐tree model agreed with previous ecological knowledge of these forests. Our study highlights the value of LiDAR remote sensing for assessing tropical dry forests, reinforcing the potential for this novel technology to advance research and management of tropical forests in general.     

Relations between forest management,stand structure and productivity across different types of Central European forests

In the past 30 years, many stand structural attributes (SSAs) have been suggested and structural indices have been developed to describe the complex structure of forests. Most studies, however, have explored the potential and limits of structural measures to quantify forest structures by applying multiple measures to one stand or few measures to several stands. However, the interdependencies of multiple structural attributes across many stands of different forest management types and developmental stages have not yet been explored. Using 20 structural attributes and 124 completely inventoried 1 ha sized stands we tested to what extent structural characteristics reflect different stand types and management intensities, and how these characteristics change over time. We found that single SSAs do not show the clear gradients that they were intended to reflect, suggesting that stand structure should be described by multiple structural attributes, and that these should represent different structural aspects (including vertical, and horizontal heterogeneity, density, and diversity). A principal component analysis showed that combining several SSAs, allowed us to distinguish between stand types. The structure of mature stands remained rather constant over the observed period of about 6 years, while that of young stands changed more rapidly due to ingrowth and mortality. The older the stands, the less the large trees contributed to stand growth relative to their size. We conclude that multiple stand structural attributes are needed to characterise stand types, management effects and to explain stand productivity.     

基于结构方程模型分析林分空间结构对草本物种多样性的影响

林分空间结构的改变直接影响林下草本物种的多样性。以针叶纯林、针阔混交林和常绿阔叶林为研究对象,采用结构方程模型研究了林分空间结构对林下草本物种多样性的影响,并探讨了林分水平空间结构,垂直空间结构以及林木竞争态势对林下草本物种多样性的影响的相对重要性。结果表明,林分水平空间结构对林下草本的物种多样性指数和物种均匀性指数均存在极显著的影响（P < 0.001）,影响系数高达0.96和0.89,对林下草本物种丰富度存在显著影响（P < 0.01）,影响系数为0.22;林分垂直空间结对林下草本丰富度和物种均匀性指数均存在极显著的影响（P < 0.001）,影响系数分别为0.86和0.43,对林下草本物种多样性指数存在显著影响（P < 0.01）,影响系数为0.16;林木竞争指数与林下草本丰富度和物种多样性指数也均存在极显著的影响（P < 0.001）,但影响系数较小,分别为-0.47和-0.30,而对林下草本物种均匀性指数未达到显著影响作用（P > 0.05）,影响系数仅为-0.04。整体上看,林分水平空间结构、垂直空间结构和林木竞争态势均对林下草本物种多样性有较强的影响作用,但从影响系数看,林分水平空间结构的影响作用最大,垂直空间结构次之,林木竞争态势的影响作用最小。因此,欲维持或改善林下草本物种多样性,应采取调整林分水平空间结构为主,垂直空间结构调整为辅,并适当降低林木竞争程度的综合经营措施。     

Soil heterogeneity in tree mixtures depends on spatial clustering of tree species

Heterogeneity in soil characteristics promotes and maintains coexistence between a diverse set of species. In forests, trees have species-specific impacts on soil abiotic characteristics and mixing of tree species is being promoted as a tool to ensure high levels of diversity and functioning. Yet, limited knowledge is available on the effect of tree species composition and spatial clustering on heterogeneity in soil characteristics. In this paper we derived heterogeneity of key characteristics of the leaf litterfall, the forest floor and the mineral topsoil (C, N and base cation concentration, C:N ratio and mass) in 53 plots of 7 different tree species compositions. We found that heterogeneity increased from the leaf litterfall, through the forest floor down to the mineral topsoil. Mixing tree species did not lead to an increased heterogeneity in the forest floor and topsoil compared to monocultures. However, we did find that mixed plots where conspecific trees stand in groups are more heterogeneous than plots where species are intimately mixed. Our results imply that heterogeneity in soil characteristics does not necessarily increase with tree diversity, but that within mixed stands the spatial organization of tree species should be considered in relation to the scale at which heterogeneity is desired.     

Structural heterogeneity and tree spatial patterns in an old-growth deciduous lowland forest in Cantabria, northern Spain

Old-growth deciduous forests in western Europe, for the most part, consist of small tracts that often may be atypical due to human disturbance, poor soil productivity or inaccessibility. In addition, very little information on tree age distributions, structural heterogeneity and tree spatial patterns appears to be available for west-European forests. Characterization of the structural features of tree populations in these old-growth stands can provide the basis to design conservation plans and also inform on how present forests might look in the absence of human interference. Four old-growth stands in a deciduous forest in the Cantabrian lowlands, northern Spain, were surveyed to determine forest structure and spatial patterns. Live and dead trees were identified, measured and mapped, and live trees were cored for age estimation. Structural heterogeneity was analyzed by means of the spatial autocorrelation of tree diameter, height and age, and the uni- and bivariate spatial patterns of trees were analyzed. The dominant species, Fagus sylvatica and Quercus robur, showed reverse-J shaped size distributions but discontinuous age distributions, with maximum ages of 255–270 yr. Tree ages suggested that the forest was largely modified by past changes in forest-use, especially by temporal variation in grazing intensity. Spatial autocorrelation revealed that former parkland stands were heterogeneous with respect to tree height only, while high forest stands were composed of patches of even-aged and even-sized trees. Young trees were clumped at varying distances and establishment occurred preferentially in canopy gaps, except for Ilex aquifolium that mainly occurred beneath mature Quercus trees. Surviving trees became less intensely clumped in the dominant species, and more strongly clumped in understorey ones, which may have been due to the effects of intraspecific competition and of canopy trees on tree survival, respectively. The spatial associations between species varied within the forest, probably as a consequence of specific establishment preferences and competitive interactions.     

Gap-scale disturbance processes in secondary hardwood stands on the Cumberland Plateau,Tennessee, USA

Disturbance regimes in many temperate, old growth forests are characterized by gap-scale events. However, prior to a complex stage of development, canopy gaps may still serve as mechanisms for canopy tree replacement and stand structural changes associated with older forests. We investigated 40 canopy gaps in secondary hardwood stands on the Cumberland Plateau in Tennessee to analyze gap-scale disturbance processes in developing forests. Gap origin, age, land fraction, size, shape, orientation, and gap maker characteristics were documented to investigate gap formation mechanisms and physical gap attributes. We also quantified density and diversity within gaps, gap closure, and gap-phase replacement to examine the influence of localized disturbances on forest development. The majority of canopy gaps were single-treefall events caused by uprooted or snapped stems. The fraction of the forest in canopy gaps was within the range reported from old growth remnants throughout the region. However, gap size was smaller in the developing stands, indicating that secondary forests contain a higher density of smaller gaps. The majority of canopy gaps were projected to close by lateral crown expansion rather than height growth of subcanopy individuals. However, canopy gaps still provided a means for understory trees to recruit to larger size classes. This process may allow overtopped trees to reach intermediate positions, and eventually the canopy, after future disturbance events. Over half of the trees located in true gaps with intermediate crown classifications were Acer saccharum, A. rubrum, or Liriodendron tulipifera. Because the gaps were relatively small and close by lateral branch growth of perimeter trees, the most shade-tolerant A. saccharum has the greatest probability of becoming dominant in the canopy under the current disturbance regime. Half of the gap maker trees removed from the canopy were Quercus; however, Acer species are the most probable replacement trees. These data indicate that canopy gaps are important drivers of forest change prior to a complex stage of development. Even in relatively young forests, gaps provide the mechanisms for stands to develop a complex structure, and may be used to explain patterns of shifting species composition in secondary forests of eastern North America.     

Changes in spatial pattern of trees and snags during structural development in Picea mariana boreal forests

Questions : How do gap abundance and the spatial pattern of trees and snags change throughout stand development in Picea mariana forests? Does spatial pattern differ among site types and structural components of a forest? Location : Boreal forests dominated by Picea mariana, northern Quebec and Ontario, Canada. Methods : Data on the abundance, characteristics and spatial location of trees, snags and gaps were collected along 200 m transects at 91 sites along a chronosequence. Spatial analyses included 3TLQV, NLV and autocorrelation analysis. Non‐parametric analyses were used to analyse trends with time and differences among structural components and site types. Results : Gaps became more abundant, numerous and more evenly distributed with time. At distances of 1–4 m, tree cover, sapling density and snag density became more heterogeneous with time. Tree cover appeared to be more uniform for the 10–33 m interval, although this was not significant. Patch size and variance at 1 m were greater for overstorey than for understorey tree cover. Snags were less spatially variable than trees at 1 m, but more so at intermediate distances (4–8 m). Few significant differences were found among site types. Conclusions : During stand development in P. mariana forest, gaps formed by tree mortality are filled in slowly due to poor regeneration and growth, leading to greater gap abundance and clumping of trees and snags at fine scales. At broader scales, patchy regeneration is followed by homogenization of forest stands as trees become smaller with low productivity due to paludification.     

洞庭湖森林生态系统空间结构均质性评价

以东、南、西洞庭湖区域君山岛、龙山、赤山岛龙虎山林场、明朗山、常德林场、和洑林场和桃源县天然次生林为对象,借鉴生态学有关生态系统内部均衡和均质原理,在森林生态系统(斑块)尺度,从空间结构的混交、竞争、空间分布格局3个方面分析提出空间优化的均质性目标、均质性指数及均质性评价方法和评价标准.结果表明:15个洞庭湖湿地调查林分平均均质性评价指数为0.2517.均质性评价的5个等级中处于l级和2级的林分为12个,占80％,处于3级的林分只有3个,仅占20％.说明洞庭湖各林分整体上在空间结构、生长环境和树种优势度等方面不具有明显优势,离理想空间结构差距太大;按照高程不同,洞庭湖湿地从湖沼到丘陵岗地,林分空间结构均质性特征逐渐增强.森林生态系统空间优化均质性目标及均质性指数的提出是森林可持续经营空间途径的拓展,为森林经营的理想空间结构及其表达探索一条新途径.     

The effects of stand structure on ground-floor bryophyte assemblages in temperate mixed forests

The effect of tree species composition, stand structure characteristics and substrate availability on ground-floor bryophyte assemblages was studied in mixed deciduous forests of Western Hungary. Species composition, species richness and cover of bryophytes occurring on the soil and logs were analysed as dependent variables. The whole assemblage and functional groups defined on the basis of substrate preference were investigated separately. Substrate availability (open soil, logs) was the most prominent factor in determining species composition, cover and diversity positively, while the litter of deciduous trees had a negative effect on the occurrence of forest floor bryophytes. Besides, bryophyte species richness increased with tree species and stand structural diversity, and for specialist epiphytic and epixylic species log volume was essential. Sapling density and light heterogeneity were influential on bryophyte cover, especially for the dominant terricolous species. Many variables of the forest floor bryophyte community can be estimated efficiently by examining stand structure in the studied region. Selective cutting increasing tree species diversity, stand structural heterogeneity and dead wood volume can maintain higher bryophyte diversity in this region than the shelter-wood system producing even-aged, monodominant, structurally homogenous stands.     

Species and structural diversity affect growth of oak,but not pine,in uneven-aged mature forests

The effects of mixing tree species on tree growth and stand production have been abundantly studied, mostly looking at tree species diversity effects while controlling for stand density and structure. Regarding the shift towards managing forests as complex adaptive systems, we also need insight into the effects of structural diversity. Strict forest reserves, left for spontaneous development, offer unique opportunities for studying the effects of diversity in tree species and stand structure. We used data from repeated inventories in ten forest reserves in the Netherlands and northern Belgium to study the growth of pine and oak. We investigated whether the diversity of a tree's local neighbourhood (i.e., species and structural diversity) is important in explaining its basal area growth. For the subcanopy oak trees, we found a negative effect of the tree species richness of the local neighbours, which – in the studied forests – was closely related to the share of shade-casting tree species in the neighbourhood. The growth of the taller oak trees was positively affected by the height diversity of the neighbour trees. Pine tree growth showed no relation with neighbourhood diversity. Tree growth decreased with neighbourhood density for both species (although no significant relationship was found for the small pines). We found no overall diversity-growth relationship in the studied uneven-aged mature forests; the relationship depended on tree species identity and the aspect of diversity considered (species vs. structural diversity).     

Biomass Stock and Productivity of Primeval and Production Beech Forests: Greater Canopy Structural Diversity Promotes Productivity

Our knowledge of temperate broadleaf forest ecology is based mostly on the study of production forests, which lack the terminal stage of forest development and have a simpler stand structure than old-growth and primeval forests. How primeval and production forests differ in net primary production (NPP) is not well known. In three primeval and three nearby production forests of European beech (Fagus sylvatica) in the Slovakian Carpathians, we measured aboveground biomass stocks (live and dead), aboveground NPP (ANPP) and parameters characterizing canopy structural diversity (leaf area index and its spatial variation). Our study aims were (1) to explore the role of canopy structural diversity for ANPP and (2) to assess evidence of a productivity decline in the terminal stage. While aboveground live biomass stocks were on average 20% greater in the primeval forests (386 vs. 320 Mg ha^?1; insignificant difference at two sites), deadwood mass stocks were on average four times larger than in the production forests (86 vs. 19 Mg ha^?1). ANPP was similarly high in the primeval and production forests (10.0 vs. 9.9 Mg ha^?1 y^?1) and did not decrease towards the terminal stage. Production models indicate that, in the primeval forests, about 10% of ANPP (ca. 1 Mg ha^?1 y^?1) was generated by effects related to leaf area heterogeneity, evidencing a positive effect of structural diversity on forest productivity, even though species diversity was low. This study helps to better understand the impact of forest management on the productivity and carbon storage in temperate woodlands.     

Stand characteristics and biodiversity indicators along the productivity gradient in boreal forests: Defining a critical set of indicators for the monitoring of habitat nature quality

Abstract

We quantified the effects of anthropogenic disturbances on the structure and biodiversity of boreal forests on acidic soils and created a statistically supported rational set of indicators to monitor the stand “naturalness”. For that, we surveyed various traits of tree layer, understory, herb layer, forest floor and several widely accepted biodiversity epiphytic indicators in 252 old‐aged boreal stands in Estonia, mostly dominated by Scots pine or Norway spruce. Multifactorial general linear model analyses showed that many forest characteristics and potential indicators were confounded by the gradient of soil productivity (reflected by the forest site type), local biogeographic gradients and also by stand age. Considering confounding effects, boreal forests in a near‐natural state have more large‐diameter trees (diameter at breast height >40 cm) and larger variety of diameter classes, higher proportion of spruce or deciduous trees, a larger amount of coarse woody debris in various stages, a more closed tree canopy and denser understory than managed mature forests. By increasing light availability above the field layer, forest management indirectly increases the coverage of herbs and lichens on the forest floor but reduces the alpha‐ and beta‐diversity of herbs and the proportion of graminoids. Human disturbances reduce the relative incidence of many commonly accepted biodiversity indicators such as indicator lichens, woodpeckers, wood‐dwelling insects or fungi on trees. The test for the predictive power of characteristics reacting on disturbance revealed that only a fraction of them appeared to be included in a diagnostic easy‐to‐apply set of indicators to assess the nature quality of boreal forest: the amount of dead wood, the proportion of deciduous trees, the presence of specially shaped trees and woodpeckers and, as an indicator of disturbances, the forest herb Melampyrum pratensis. Many of these indicators have already been implemented in practice.