Effects of treefall gaps created by windthrow on bat assemblages in a temperate forest

Determining the way in which spatial distribution and diversity of forest-dwelling mammals varies with natural disturbance is essential to understanding the spatial dynamics of mammal assemblages in forests. Bats are the only forest-dwelling mammals capable of true flight. At a local scale, bat flight ability, which may be related to ecomorphological traits, is an important factor influencing spatial distribution. We tested two postulates: (1) the spatial distribution of bats is affected by sizes of forest gaps created by natural disturbances and (2) species-specific responses can be predicted from bat ecomorphological traits (aspect ratio (AR) and wingtip shape index (WT)) that influence bat flight ability. We found that sizes of forest gaps affected the occurrence of each bat species and species richness of bats at local scales; species-specific responses were related to the ecomorphological traits of individual species. Bat species with high AR and low WT were not affected by variation in canopy gap size. In contrast, bat species with low AR and high WT responded negatively to gap size, and those with intermediate AR and WT responded positively to canopy gap size at sites with small-sized gaps but responded negatively to large-sized gaps. Overall bat species richness responded negatively to gap size. Thus, ecomorphological traits may be important determinants of bat spatial distributions and species diversity at local scales in disturbed habitats. In this study, forest edges might have been undersampled due to the location of bat detectors. However, this potential undersampling should not have affected the interpretation of occurrence patterns of bat species responding to gap size. Our results imply that bat conservation efforts in forest lands should take into consideration specific responses related to ecomorphological traits of species inhabiting an area. The results also suggest that quantifying the effects of natural disturbances on bat assemblages may provide a knowledge base for forest management to minimize the impacts of unavoidable anthropogenic disturbances on bat species diversity. Rare or infrequent natural disturbances can provide models for forest management aimed at maintaining bat species diversity.     

Effects of gaps on regeneration of woody plants: a meta-analysis

Forest gaps, openings in the canopy caused by death of one or more trees, have a profound effect on forest regeneration and drive the forest growth cycle. It is therefore necessary to understand the effects of forest gaps on regeneration for modern forest management. In order to provide a quantitative assessment of the effects of forest gaps on regen-eration of woody plants, we conducted this review of gap effects on woody plant regeneration on the basis of 527 observations from 42 indi-vidual papers, and reported the results of these data in a meta-analysis. Overall, densities of regenerated woody plants were significantly greater （359%） in forest gaps than on the closed-canopy forest floor. The regen-eration density in gaps of plantation forests was significantly greater （P＆lt;0.05） than that of natural forest because the regeneration in gaps of plan-tation forests was improved by both gap effects and experimental meas-ures. Similarly, in comparison to natural gaps, regeneration was better enhanced in artificial gaps. Regeneration density exhibited a significantly positive correlation with gap size, but a negative correlation with gap age because the gap size decreased with increasing gap age. Shade tolerance of woody plants affected regeneration density in gaps and understory. Average regeneration density of shade-tolerant species exhibited a sig-nificantly positive response to gaps but densities remained lower in total than those of intermediate and shade-intolerant species. Gap effects on regeneration decreased in response to increasing temperature and pre-cipitation because of the limiting effects of lower temperature and moisture on woody plant regeneration. In summary, forest gaps enhance woody plant regeneration, and the effects of gaps varied by forest type, gap characteristics, environmental factors and plant traits. The results of this meta-analysis are useful for better understanding the effects and roles of gaps on forest regeneration and forest management.     

The response of the pseudoannual species Trientalis europaea L. to forest gap dynamics in a near-natural spruce forest

The objective of this study was to test relationships between gap size, gap age and performance of the pseudoannual forest herb layer species Trientalis europaea. We also tested for a potential covariation of performance variables with light as the putative driving factor of gap size and gap age effects. The study took place in the core zone of the Harz National Park in a near-natural spruce forest at Mt. Brocken, Germany. We established 70 randomly distributed plots of three different gap age classes (<15 years, >15 and <60 years, >60 years) and undisturbed forest stands. We recorded growth variables of Trientalis (e.g. height of ramets, number of flowers and fruits per ramet and number of daughter tubers per ramet) and measured PPFD (photosynthetic photon flux density). Gap sizes were assessed with GPS, ranged from 131 m² to 16400 m² and were independent from gap age. Population density of T. europaea was neither affected by gap size nor by gap age. However, gap age had an effect on the fitness of Trientalis ramets. In gaps, the species produced a higher number of flowers and fruits. In general, the average proportion of flowering and fruiting was very low, with 13% and 4%, respectively. In contrast, light intensity had a significant positive effect on ramet density, while the number of daughter tubers differed between 0.8 and 0.5 tubers per individual under lowest light and full light intensity, respectively. The general conclusion is that gap size had no effects on the performance of T. europaea, while gap age had both direct effects and indirect effects mediated by light.     

A review of the roles of forest canopy gaps

Treefall gap, canopy opening caused by the death of one or more trees, is the dominant form of disturbance in many forest systems worldwide. Gaps play an important role in forest ecology helping to pre- serve bio- and pedo-diversity, influencing nutrient cycles, and maintain- ing the complex structure of the late-successional forests. Over the last 30 years, numerous reviews have been written describing gap dynamics. Here we synthesize current understanding on gap dynamics relating to tree regeneration with particular emphasis on gap characteristics consid- ered critical to develop ecologically sustainable forest management sys- tems and to conserve native biodiversity. Specifically, we addressed the question： how do gaps influence forest structure？ From the literature re- viewed, the size of gaps induces important changes in factors such as light intensity, soil humidity and soil biological properties that influence tree species regeneration and differ in gaps of different sizes. Shade- tolerant species can colonize small gaps; shade-intolerant species need large gaps for successful regeneration. Additionally, gap dynamics differ between temperate, boreal, and tropical forests, showing the importance of climate differences in driving forest regeneration. This review summa- rizes information of use to forest managers who design cutting regimes that mimic natural disturbances and who must consider forest structure, forest climate, and the role of natural disturbance in their designs.     

Treefall gaps of certified,conventionally managed and natural forests as regeneration sites for Neotropical timber trees in northern Honduras

Forest certification programmes promote sustainable management of the world's forests. Certification can be given to a forest management system that fulfils certain environmental, social and economic criteria.     

A pre-harvest method for predicting understory light levels created by artificial canopy gaps

A new method for predicting understory light levels around an artificial canopy gap prior to the selective cutting is presented. In two even-aged coppice stands, hemispherical photographs were taken at a height of 1.3 m, and crowns of each canopy stem on the photographs were identified by field observations. To predict light levels after cutting, we used computerized images of the hemispherical photographs, in which we digitally erased all crowns to be cut (named ‘erasing method’). Light levels were calculated by an analyzing program; HEMIPHOT. Light levels predicted by the ‘erasing method’ were compared with that calculated by the ordinary photograph-analyzing method using photographs taken after the actual cutting. The correlations between the two methods were highly significant in the two stands. The ‘erasing method’ is eminently effective for the prediction of understory light levels in an artificial canopy gap.     

Effects of harvest management practices on forest biomass and soil carbon in eucalypt forests in New South Wales,Australia: Simulations with the forest succession model LINKAGES

Understanding long-term changes in forest ecosystem carbon stocks under forest management practices such as timber harvesting is important for assessing the contribution of forests to the global carbon cycle. Harvesting effects are complicated by the amount, type, and condition of residue left on-site, the decomposition rate of this residue, the incorporation of residue into soil organic matter and the rate of new detritus input to the forest floor from regrowing vegetation. In an attempt to address these complexities, the forest succession model LINKAGES was used to assess the production of aboveground biomass, detritus, and soil carbon stocks in native Eucalyptus forests as influenced by five harvest management practices in New South Wales, Australia. The original decomposition sub-routines of LINKAGES were modified by adding components of the Rothamsted (RothC) soil organic matter turnover model. Simulation results using the new model were compared to data from long-term forest inventory plots. Good agreement was observed between simulated and measured above-ground biomass, but mixed results were obtained for basal area. Harvesting operations examined included removing trees for quota sawlogs (QSL, DBH >80 cm), integrated sawlogs (ISL, DBH >20 cm) and whole-tree harvesting in integrated sawlogs (WTH). We also examined the impact of different cutting cycles (20, 50 or 80 years) and intensities (removing 20, 50 or 80 m³). Generally medium and high intensities of shorter cutting cycles in sawlog harvesting systems produced considerably higher soil carbon values compared to no harvesting. On average, soil carbon was 2–9% lower in whole-tree harvest simulations whereas in sawlog harvest simulations soil carbon was 5–17% higher than in no harvesting.     

Calculation method for sunshine duration in canopy gaps and its application in analyzing gap light regimes

Understory light is essential to the growth and survival of plants, yet the light varies temporally and spatially within forest gaps. Measurement of understory light levels using light sensors is both time and labor intensive. Sunshine duration (SD) has a strong correlation with solar radiation and has been the variable most widely used for estimating solar radiation. The power of SD-based methods for estimating solar radiation lies in its ability to quickly estimate light levels. Although several calculation methods for SD within canopy gaps are available, all the methods oversimplify canopy gaps by classifying them as cylindrical or ellipsoid and thus have a relatively low level of accuracy. In this study we developed a calculation method for SD at any given point within natural canopy gaps and we used SD to estimate solar radiation within 12 canopy gaps on Mt. Taibai in China based on the Angström-Prescott model. We then evaluated the SD-based solar radiation by the total and direct solar radiation derived from a gap light index based on hemispherical photographs (HP). Finally, we analyzed the spatial-temporal characteristics of light levels within these 12 gaps by using the solar radiation derived from SD at hundreds of simulated points in each gap. Our results showed that (1) SD-based solar radiation was not statistically different from HP-based direct solar radiation and had a strong linear correlation with HP-based total solar radiation; (2) growing-season daily mean solar radiation within the 12 gaps varied from 0.08 to 13.28 MJ m⁻² day⁻¹ with an average of 4.13 MJ m⁻² day⁻¹; (3) solar radiation had a positive correlation with the ratio of the square root of the canopy gap area to the mean canopy height. This relationship was significant but solar radiation had no correlation with the canopy gap area; (4) among most gaps solar radiation was greatest in May while potential SD was longest in June 2008. From these results we can conclude that the SD-based method for estimating solar radiation developed in this study can quickly and accurately estimate light levels at any specified point within canopy gaps. SD-based solar radiation appears to be a good choice for studies on the spatial-temporal characteristics of gap light levels.     

Permanence of resilience and protection efficiency in mountain Norway spruce forest stands: A simulation study

An objective of mountain forest management is to increase the ability of forest stands to protect human activities against natural hazards such as rock-falls and snow avalanches in a sustainable way. The challenge is to find a compromise between efficient instantaneous protection, favoured by dense stands, and continuous renewal, minimizing time periods of low protection efficiency. We used a Norway spruce stand dynamics model to compare the respective advantages of individual tree and gap selection silviculture in this context. We simulated stand dynamics over 800 years with either individual tree or gap thinning every 20 years with several thinning intensities. At each time step, we evaluated stand resilience, protection efficiency against rock-falls, protection efficiency against avalanches, and structural complexity with four indicators based on stand structure. Every scenario produced short time periods with low stand resilience and protection efficiency. Such periods can be tolerated if they are sufficiently rare compared to the local disturbance regime. We characterized the permanence of resilience and protection of a forest stand as its ability to remain within boundary values of the different indicators, without going out of them during continuous time periods longer than fixed maximum durations. Permanence of resilience and permanence of protection decreased with thinning intensity. Efficient protection against rock-falls was obtained with gap thinning of intermediate intensity while protection against avalanches was obtained only for very low thinning intensities. For our ecological context, the best compromise between resilience and protection was obtained with three 10 m radius gaps per hectare every 20 years (9.5% of the area of a stand). This strategy led to uneven-aged stand structures with a high diversity of diameters classes. Our results suggest that small gap silviculture may be a good way to combine forest renewal and protection efficiency in mountain regions.     

A 10-year evaluation of the functional basis for regeneration habitat preference of trees in an African evergreen forest

The spatial distribution of tree juveniles in relation to light environments may reflect species differences in growth, survival, and functional traits and will shape the nature of forest regeneration. Long-term field experiments are important to evaluate this issue because of the potentially very long juvenile period in trees. Here, we combine a 10-year seedling survival–growth data with the results of community ordination and multivariate analyses of functional traits to ask how observed juvenile light guilds are related to species functional traits and seedling performance. We transplanted seedlings at a standardized height of 11 cm into the shaded understory and quantified their growth and survival for 10-years. Using the community-wide stem distribution data, we categorized 33 species including the focal 11 species to understory vs. gap/edge guilds. Then, we determined differences between the two guilds in seedling survival, growth, as well as seed size, adult height, and a series of leaf traits, including toughness and chemical traits (fiber, protein, phenolics, tannins, alkaloids, saponins). Among the 11 non-pioneer species whose seedlings were planted into the understory, there was no significant difference in 10-year survival between light guilds, but species in gap/edge guild tended to achieve greater height than species in the understory guild. The leaf chemical traits of 33 species did not differ between the two juvenile light guilds, but gap/edge species had smaller seeds, taller adults, and tougher leaves than understory species. We used logistic regression as a complementary approach to assess the extent to which plant traits varied between light guilds and the most parsimonious model based on AIC_c ranking included only leaf toughness and had an Akaike weight of 0.52. In addition, across the 11 species planted as seedlings, these traits were not significantly related to survivorship or growth over 10 years. A Principle Components Analysis illustrated associations among traits. We conclude that light guilds in terms of juvenile stem distribution could not be explained by long-term field performance of post-establishment seedlings alone. Earlier seedling stage or later sapling stage may be more important in differentiation of light guilds. For the species examined difference in growth rates could be linked to seed size and adult stature, but not to the adult leaf chemical traits considered. These results suggest the importance of examining ontogenetic shifts and relationships among functional traits for a better understanding of regeneration strategies of tropical trees.     

Emulating natural disturbance regimes as a basis for forest management: A North American view

It is widely accepted that for many North American forest ecosystems, changes in species composition and structure are associated with departures from natural disturbance regimes (e.g., fire exclusion). It also widely accepted that restoration of species composition and structure, similar to those which existed under natural disturbance regimes, will in turn restore and maintain: (1) ecosystem resilience to a broad range of environmental challenges; (2) important elements of ecosystem function; and (3) disturbance-adapted species assemblages and thus conservation of biodiversity. Much of the research relevant to emulating natural disturbance regimes (ENDR) in North America has focused on natural fire regimes, particularly those characterized by frequent/low-severity fires. Increasingly, attention is being focused on natural disturbance regimes other than fire and on interactions between different disturbance types. However, the greatest challenges to implementation of ENDR are socioeconomic rather than ecological. There is, for example, limited social acceptance of management based on high-severity natural disturbance regimes. And while there is much broader acceptance of ENDR based on frequent/low-severity disturbance regimes, there is nevertheless strong disagreement concerning how ENDR is to be implemented.     

International Demand Equations for Forest Products: A Comparison of Methods

The objective of this study was to compare different methods of estimating the demand equations of global forest sector models. Past applications have shown that model projections depend critically on the elasticity of demand with respect to income and prices. To measure the elasticity coefficients, nine groups of products were examined, ranging from fuelwood to paper and paperboard, in 64 countries. The analysis was done with panel data on consumption, price and income, from 1973 to 1997. Two alternative forms were hypothesized: a static demand model, and a dynamic partial adjustment model. The demand equations were estimated first with classical panel methods: ordinary least squares pooling of country data, least squares with country dummy variables, between group pooling and error-component models. However, rejection of the hypothesis of homogeneity of elasticities across countries led to the investigation of two shrinkage estimators: the Stein rule and the iterative empirical Bayes estimator. Within-sample predictions were carried out to compare the accuracy of each estimator. The Stein rule estimator, based on the static model, gave the smallest root mean square error. Nevertheless, because the Stein-rule estimates often had theoretically wrong signs and large variances, other estimators, although biased, were deemed preferable for policy analysis. In conclusion, pooling by ordinary least squares with country dummy variables was one of the most useful methods to obtain demand equations applicable to many countries. Furthermore, static models predicted demand better than dynamic models, conditional on price and national income.     

树木根系分布与结构研究方法综述

根系是树木生长发育的基础。开展树木根系分布和结构研究,有利于探究根系与地下环境的相互联系,研究根系分布与生长之间的关系,了解生物地球化学循环中的碳素平衡与循环。文中简要综述了根系研究的历史,分析了树木根系研究的传统方法及其优缺点,介绍了根系无损检测的新方法和新手段。分析认为,根系分布与结构的相关研究方法在不断进步,但新技术和新手段仍存在局限性,还有待进一步改进。     

Tithonia diversifolia as a green manure for soil fertility improvement in western Kenya: A review

Tithonia diversifolia, a shrub in the family Asteraceae, is widely distributed along farm boundaries in the humid and subhumid tropics of Africa. Green biomass of tithonia has been recognized as an effective source of nutrients for lowland rice (Oryza sativa) in Asia and more recently for maize (Zea mays) and vegetables in eastern and southern Africa. This paper reviews the potential of tithonia green biomass for soil fertility improvement based on recent research in western Kenya. Green leaf biomass of tithonia is high in nutrients, averaging about 3.5% N, 0.37% P and 4.1% K on a dry matter basis. Boundary hedges of sole tithonia can produce about 1 kg biomass (tender stems + leaves) m^–1 yr^–1 on a dry weight basis. Tithonia biomass decomposes rapidly after application to soil, and incorporated biomass can be an effective source of N, P and K for crops. In some cases, maize yields were even higher with incorporation of tithonia biomass than with commercial mineral fertilizer at equivalent rates of N, P and K. In addition to providing nutrients, tithonia incorporated at 5 t dry matter ha^–1 can reduce P sorption and increase soil microbial biomass. Because of high labor requirements for cutting and carrying the biomass to fields, the use of tithonia biomass as a nutrient source is more profitable with high-value crops such as vegetables than with relatively low-valued maize. The transfer of tithonia biomass to fields constitutes the redistribution of nutrients within the landscape rather than a net input of nutrients. External inputs of nutrients would eventually be required to sustain production of tithonia when biomass is continually cut and transferred to agricultural land.This revised version was published online in November 2005 with corrections to the Cover Date.     

Intensive,site-specific silviculture: Manipulating resource availability at establishment for improved stand productivity. A review of South African research

An important window of opportunity to increase and sustain productivity in short-rotation plantations is the period from felling through re-establishment to canopy closure. This paper explores the effects, interactions and response mechanisms of intensive silvicultural practices on plantation productivity and sustainability, using five South African case studies (a–e). (a) Land preparation trials showed that complete surface cultivation by ploughing had a significant beneficial effect when afforestation is done for the first time in grasslands, improving basal area growth by 11–52% over pitting only. However, similar treatments have not resulted in significant growth responses under re-establishment conditions. (b) Stand growth suppression resulting chiefly from soil compaction during mechanised harvesting operations is strongly related to soil type, soil textural class and residue management options. Volume growth reduction in short-rotation eucalypt crops ranged from 25% on compaction sensitive loamy soils to less than 2% in resistant sandy soils. (c) The response mechanism whereby vegetation management improves stand productivity is a reduction in both inter-specific and intra-genotypic competition for resources, as well as a decrease in stand variability. Operationally, the most important criteria in a vegetation management programme relate to the timing of control operations across diverse site conditions. In local trials, the primary factors controlling the time taken for competition-induced tree growth suppression to occur were related to altitude, slash burning and the interaction between these factors, which facilitated the development of regional vegetation management strategies. (d) Empirical fertilizer trials in short-rotation hardwood stands have shown significant improvements in final productivity (commonly 20–90 m³ ha⁻¹ in eucalypts and 30–50 m³ ha⁻¹ in Acacia), as well as wood density (15–30 kg m⁻³ for eucalypts) following improvements in early nutrition. Improved nutrition was achieved through fertilization at planting or indirectly through residue management. The response mechanism is primarily due to early canopy development and associated increases in light capture, coupled with a more modest increase in canopy quantum efficiency and above-ground carbon allocation on a dry site. On sites with abundant water supply, increased quantum efficiency is likely to be the dominant response mechanism. (e) A series of operational gains trials tested the interactive effect of genetic tree improvement, site–genotype interaction, stand density and vegetation management + fertilization on eucalypt stand growth across five sites. There were no significant interactions between factors, but importantly, the results were additive, emphasizing the need to optimise each practice in the value chain to achieve maximum productivity.