Influence of boreal forest succession and dead wood qualities on saproxylic beetles

1 Saproxylic insects, a functional group dominated by beetles, are dependent on dead or moribund trees as habitat elements. 2 Although there are few studies of saproxylic insects from the North American boreal zone, European studies demonstrate that forest harvest can lead to a biologically significant decrease in saproxylic beetle diversity. 3 We studied saproxylic beetles in the North American boreal mixedwood forest using flight intercept traps established on naturally dead and girdled trembling aspen and spruce trees along a successional gradient of undisturbed stands from deciduous to coniferous overstory trees. 4 Composition and diversity of beetle assemblages differed among forest successional types. 5 Snag age class was an important determinant of composition for saproxylic beetle assemblages. 6 Multivariate regression analysis of these data indicated that saproxylic beetles are responding to changes in coarse woody debris, and not to the relative densities of canopy tree species, although these variables are strongly correlated. 7 Coarse woody debris management should be a primary concern in forest management plans seeking to conserve saproxylic organisms and the critical ecosystem functions (i.e. nutrient cycling) in which they participate.     

Effects of Retention Felling on Epixylic Species in Boreal Spruce Forests in Southern Finland

We studied the effects of patch retention felling and soil scarification by harrowing on the coverage and species richness of epixylic species in boreal Norway spruce (Picea abies) forests in Southern Finland. The epixylics were investigated from both the retention tree groups (RTGs) and the surrounding felling areas before and after fellings and after scarification on consecutive years. The cover percentage of all included species groups was shown to decrease after the felling, especially in the felling areas (vascular plants, ?0.4%; mosses, ?27.8%; liverworts, ?4.0%; and lichens,?2.1%). The decrease was considerable also in the RTGs. The amount of dead moss increased in both the areas indicating microclimatic changes. Species richness also declined rapidly after the first year, especially in the felling areas (vascular plants, ?2.2%; mosses, ?27.3%; liverworts, ?30.3%; and lichens, ?22.9%). Scarification also decreases covers and species richness of bryophytes. After the second year, the covers of the species groups generally started to regain, especially in the untreated RTGs. The size of RTG was in positive correlation with the total species number. Another main result indicates that it is possible to maintain much higher initial vegetation abundance and diversity in the RTGs than in the felling areas. Coarse woody debris formed by the frequent tree uprootings may also enhance the long‐term survival of epixylics over forest regeneration period. RTGs should be at least 10 times larger than the size used in current Finnish forestry, so that they could function as species refugia.     

Species–area relationships of red-listed species in old boreal forests: a large-scale data analysis

Aim Species–area relationships are often applied, but not generally approved, to guide practical conservation planning. The specific species group analysed may affect their applicability. We asked if species–area curves constructed from extensive databases of various sectors of natural resource administration can provide insights into large‐scale conservation of boreal forest biodiversity if the analyses are restricted only to red‐listed species. Location Finland, northern Europe. Methods Our data included 12,645 records of 219 red‐listed Coleoptera and Fungi from the whole of Finland. The forest data also covered the entire country, 202,761 km². The units of species–area analyses were 224 municipalities where the red‐listed forest species have been observed. We performed a hierarchical partitioning analysis to reveal the relative importance of different potential explanatory variables. Based on the results, for all red‐listed species, species associated with coniferous trees and for Fungi, the area of economically over‐aged forests explained the best the variation in data. For species associated with deciduous trees and Coleoptera, the forest area explained better variation in data than the area of old forests. In the subsequent log–log species–area regression analyses, we used the best variables as the explanatory variable for each species group. Results There was a strong relationship between the number of all red‐listed species and the area of old forests remaining, with a z‐value of 0.45. The area explained better the number of species associated with conifer trees and Fungi than the number of species associated with deciduous trees and Coleoptera. Main conclusions The high z‐values of species–area curves indicate that the remaining old‐growth patches constitute a real archipelago for the conifer‐associated red‐listed species, since lower values had been expected if the surrounding habitat matrix were a suitable habitat for the species analysed.     

Assessing above and belowground recovery from ammonium sulfate addition and wildfire in a lowland heath: mycorrhizal fungi as potential indicators

Atmospheric pollution containing soil-nitrifying ammonium sulfate ((NH₄)₂SO₄) affects semi-natural ecosystems worldwide. Long-term additions of (NH₄)₂SO₄ to nitrogen (N)-limited habitats, including heathlands, increase climate stress affecting recovery from wildfires. Although heathland vegetation largely depends on ericoid mycorrhizal fungi (ErM) to access soil N, we lack detailed understanding of how prolonged exposure to (NH₄)₂SO₄ may alter ErM community composition and host plants' reliance on fungal partners following wildfire and affect recovery. Simulation of atmospheric pollution ((NH₄)₂SO₄) occurred bi-weekly for 5 years after a 2006 wildfire in a UK heathland. Ten years after treatments ceased, we measured vegetation structure, lichen and lichen photobiont composition, soil characteristics, ErM colonization, ErM diversity in roots and soil, and assessed ErM potential as novel recovery indicators. Heather height and density, and moss groundcover, were greater in N-enriched plots. Lichen community indices showed significant treatment effects, but without differences in photobionts. Soil pH and Mg were significantly lower in treated plots while soil cation exchange capacity was significantly higher. There were no detectable differences in ErM composition and keystone ErM taxa between control and treated plots. Soil carbon stock measures were variable. Our results indicate atmospheric pollution following fire can have significant lingering effects above- and belowground. ErM diversity and root colonization were not assessed in the original N-addition experiment; we advocate for their inclusion in future studies as an integral part of the recovery assessment toolkit. We show that mycorrhizal fungi diversity is a viable ecological tool and summarize key steps for ErM identification.     

Sampling the rarest: threatened beetles in boreal forest biodiversity inventories

Presence or absence of threatened species in samples is information that is widely used in designing and implementing conservation actions. We explored the effectiveness of beetle (Coleoptera) inventories and contribution of different sampling methods in revealing occurrences of threatened and near threatened species in boreal forests. The number of species caught using traps in a particular area proved to be a useful indicator of the representativeness of data, the relationship between total number of species and the number of threatened and near threatened species being almost exponential. Samples containing less than 200 trapped species (or 2000 individuals) are almost useless in surveying threatened and near threatened species. The probability of finding such species increases considerably when the number of trapped species exceeds 400. Window traps attached directly on the trunks of dead trees proved to be the most efficient sampling method in trapping threatened beetles, whereas many other standard methods gave relatively poor results. We suggest that the best alternative in surveying threatened species in boreal forests is a combination of intensive direct searching and trunk window traps. Finding threatened beetles with rigorous probability requires very large sample sizes, even if the most effective sampling methods are used. For example, ranking 10 boreal forest areas to be protected according to the occurrence of threatened species with some reliability may require trapping of over 100000 beetle individuals. Collecting and identifying these large samples routinely in conservation actions is not feasible, which means that shortcuts (indicators etc.) are necessary. However, a lot of good-quality inventories with appropriate sampling efforts are needed before these shortcuts can be identified and elaborated. Such inventories are also crucial for the improvement of the classification of threatened species and full assessment on how past forest management has eventually affected the biota.     

北方森林土壤呼吸和木质残体分解释放出的CO2通量

北方森林因其面积大、土壤碳储量高以及对全球暖化响应敏感而在全球碳平衡和气候系统中起着至关重要的作用。土壤呼吸和木质残体分解释放出的 CO2 通量是北方森林生态系统输入大气圈的最主要的碳源。量化这个通量并深刻理解其中的机理过程 ,是评价和预测北方森林在全球变化中的作用必不可少的内容。综述了北方森林生态系统土壤呼吸和木质残体分解释放出的 CO2 通量随生态系统类型及环境条件而变化的一般格局以及自养呼吸和异氧呼吸在土壤表面 CO2 通量中的相对贡献 ;分析了影响北方森林土壤呼吸的主要生物物理因子 ;讨论了该领域研究存在的问题和今后的研究方向 ;并强调木质残体分解释放出的 CO2 通量虽然在以往的森林生态系统碳平衡研究中常被忽略 ,但在火灾频繁的北方森林中不容忽视     

Natural and anthropogenic influences on coarse woody debris stocks in Nothofagus–Araucaria forests of northern Patagonia,Argentina

Coarse woody debris (CWD) is an important element driving ecological processes, strengthening ecosystem resilience and for biodiversity within forest ecosystems. However, the abundance and distribution of CWD and their relation to natural and human factors are poorly known in southern South America. In this work we studied the density and volume of CWD types in Nothofagus–Araucaria stands in northern Patagonia (Neuquén – Argentina) and relationships with forest composition and structure. We also studied their relationships with fire history, topography and human‐related variables. Twenty‐three stands with Nothofagus pumilio, Nothofagus antarctica and/or Araucaria araucana were sampled to estimate quantities of logs, snags and dead branches using the planar‐intersect method. CWD density and volume in these forests were moderate and varied across the landscape with a spatial pattern determined by biotic, abiotic and human use–related variables. Mean CWD volume was 52.9 m³ ha^?1 (range: 1.6–143.7) and significantly varied among forest types and watersheds. CWD was positively related to dbh, tree height and slope, but negatively related to tree density. CWD was clearly influenced by composition and structural characteristics of stands, where the tree species traits had an important role. As well, the observed amount and type of CWD, whereby most of the stands showed low levels of old (pre‐disturbance) logs/snags and poor new inputs of deadwood, may be explained by fire frequency. Firewood gathering and livestock grazing negatively affected deadwood stocks and topography counteracts this effect by limiting human access. Fire disturbance history, windthrow and dieback pulses produced by insect outbreaks and human access seemed to be the main causes that best explained CWD spatial distribution and abundance patterns in north‐western Patagonian forests.     

湘南典型次生林粗木质残体生物量、碳储量和养分特征

对九龙江森林公园次生常绿阔叶林粗木质残体(CWD)进行量化研究,了解亚热带典型次生林CWD的本底数据。以九龙江森林公园典型天然次生林中的6个20m×20m的标准样地为对象,调查并分析样地内不同分解等级CWD的生物量、碳储量和养分特征变化。九龙江森林公园亚热带典型次生林的CWD储量在2.8—30.4t/hm~2之间,碳储量在6.10—6.75t/hm~2之间,大量营养元素(N、P、K、 Ca、 Mg)含量相对稳定,化学计量变化不显著,微量营养元素中Pb与Cd元素含量随着分解等级的增加而增加。九龙江森林公园亚热带典型次生林的CWD储量在亚热带常绿阔叶林中处于中等水平,CWD主要以分解中后期的倒木为主,反映出该次生林处于中幼龄林阶段,具有较高的碳储量,养分含量相对稳定,并表现出Pb与Cd在CWD中积累的现象,在中度分解和重度分解的CWD中Pb含量分别增加了62.65%和69.88%,Cd含量则分别增加了33.33%和100%,其内在机理有待进一步深入研究。研究结果有助于进一步了解CWD如何参与森林生态系统养分循环、重金属积累等生态过程,比较不同林分干扰历史下CWD储量、分布及养分特征的异质性...     

Maintaining and restoring biodiversity in European boreal forests by developing natural disturbance regimes

Abstract. A conceptual model is presented as a guide to the maintenance and restoration of ecologically sustainable boreal forest. The model is based on the hypothesis that self-sustained forest ecosystems can be (re-)created, and their biodiversity developed, if forest management can simulate the composition and structure of boreal forest landscapes by introducing and maintaining disturbances leading to naturally dynamic spatial and temporal patterns of forest regeneration. The major explanatory variable in the model is the effect of wildfire on sites with different fuel characteristics and climates found in the European boreal forest. Four levels of fire intensity are distinguished, based on mean fire frequencies. These range from extremely low in some wet tall-herb sites or sites at high altitudes or latitudes with a humid climate, where fire is absent or rare, to dry lichen-rich sites where fire occurs often. The model is called ASIO, after the words Absent, Seldom, Infrequent and Often, indicating the four levels. Three main disturbance regimes are distinguished in the European boreal forest, based on the complex interactions between probabilistic (e.g. mean fire intervals at different site types) and random events (e.g. where and when a fire occurs): (1) gap-phase Picea abies dynamics; (2) succession from young to old-growth mixed deciduous/coniferous forest; and (3) multi-cohort Pinus sylvestris dynamics. The model stems mainly from studies in Fennoscandia, but some studies from outside this region are reviewed to provide support for a more general application of the model. The model has been implemented in planning systems on the landscape level of several large Swedish forest enterprises, and is also used as an educational tool to help private land owners with the location and realization of forest management regimes. Finally, the model can be used to develop an administrative system for the monitoring of biodiversity in boreal forest.     

Coarse woody debris in undisturbed and logged forests in the eastern Brazilian Amazon

Coarse woody debris (CWD) is an important component of the carbon cycle in tropical forests. We measured the volume and density of fallen CWD at two sites, Cauaxi and Tapajós in the Eastern Amazon. At both sites we studied undisturbed forests (UFs) and logged forests 1 year after harvest. Conventional logging (CL) and reduced impact logging (RIL) were used for management on areas where the geometric volumes of logs harvested was about 25–30 m³ ha^?1. Density for five classes of fallen CWD for large material (>10 cm diameter) ranged from 0.71 to 0.28 Mg m^?3 depending upon the degree of decomposition. Density of wood within large fallen logs varied with position relative to the ground and with distance from the center of the log. Densities for materials with diameters from 2 to 5 and 5 to 10 cm were 0.36 and 0.45 Mg m^?3, respectively. The average mass (±SE) of fallen CWD at Cauaxi was 55.2 (4.7), 74.7 (0.6), and 107.8 (10.5) Mg ha^?1 for duplicate UF, RIL, and CL sites, respectively. At Tapajós, the average mass of fallen CWD was 50.7 (1.1) Mg ha^?1 for UF and 76.2 (10.2) Mg ha^?1 for RIL for duplicate sites compared with 282 Mg ha^?1 for live aboveground biomass. Small‐ and medium‐sized material (<10 cm dia.) accounted for 8–18% of the total fallen CWD mass. The large amount of fallen CWD at these UF sites relative to standing aboveground biomass suggests either that the forests have recently been subjected to a pulse of high mortality or that they normally suffer a high mortality rate in the range of 0.03 per year. Accounting for background CWD in UF, CL management produced 2.7 times as much CWD as RIL management. Excess CWD at logging sites would generate a substantial CO₂ emission given the high rates of decay in moist tropical forests.     

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.     

Wood decomposition is more strongly controlled by temperature than by tree species and decomposer diversity in highly species rich subtropical forests

While the number of studies on the role of biodiversity on ecosystem functioning is steadily increasing, a key component of biogeochemical cycling in forests, dead wood decay, has been largely neglected. It remains widely unknown whether and how dead wood decay is affected by diversity loss in forests. We studied the hierarchical effects of tree species diversity on wood decay rates in a subtropical forest landscape in southeast China via its influence on fungal OTU richness and invertebrate diversity using piecewise structural equation models. The experiment was conducted in natural forest plots that span a wide gradient of tree species diversity embedded in a heterogeneous topography. To account for interactions between macro‐invertebrates and fungi, that potentially modify the influence of tree biodiversity and climate on dead wood decay, we compared a macro‐invertebrate exclusion treatment with a control treatment that allowed access to all types of decomposers. Diversity effects of trees on wood decay rates were mostly negative and mediated by the diversity of macro‐invertebrates. However, the effects of tree species diversity or fungal OTU richness and macro‐invertebrate diversity on wood decay rates were comparatively weak. Temperature affected decay rates positively and had the strongest influence in all treatments. While the exclusion of macro‐invertebrates did not lead to a reduction of wood decay rates, our results suggest that they may however have a mediating role in the process. In the presence of invertebrates the predictability of wood decay rates was higher and we observed a tendency of a stronger temperature control. Our results suggest that there is evidence for diversity effects on wood decomposition, but the temperature control is still more important. Thus, an increase in mean annual temperature will increase carbon and nutrient turnover through wood decomposition in subtropical forest irrespective of biotic composition.     

天山雪岭云杉林粗木质残体储量特征

粗木质残体(coarse woody debris,CWD)在天山雪岭云杉林生态系统中起着重要的结构性和生物地球化学作用,解释其储量特征是研究CWD的基础,但尚未有大尺度研究见诸报道。以天山雪岭云杉8 hm~2森林动态监测样地为研究对象,采用野外调查、室内试验以及数据分析相结合的方法,调查了样地内CWD的储量组成、径级以及分解等级分布格局等基本特征及其影响因子。结果表明:(1)天山雪岭云杉8 hm~2森林动态监测样地内共有直径≥10 cm的CWD 936株,CWD的密度、体积、储量分别为117株/hm~2,15.13 m~3/hm~2,4.52 t/hm~2;其中倒木是CWD的主要贡献者,占CWD总储量的52.21%;(2)样地内各径级CWD的数量呈典型的倒"J"型结构,直径30 cm的CWD个体占全部CWD的83%;(3)样地内CWD总体上处于以Ⅱ、Ⅲ分解等级为主的中度分解状态,CWD径级越大,分解程度越高;(4)林分密度、郁闭度和海拔是影响天山雪岭云杉林CWD储量特征的主要因素。研究可为天山雪岭云杉林的可持续发展与经营提供科学依据。     

Effects of Forest Use on Aphyllophoraceous Fungal Community Structure in Sarawak, Malaysia

Aphyllophoraceous fungi are expected to reflect changes in the environmental conditions caused by forest use. To reveal the effects of forest uses on the fungal community structure, we performed a 3‐month survey of aphyllophoraceous species in five forest types (undisturbed primary forest, isolated patches of primary forest, old and young fallow forest, and rubber plantations) in Sarawak, Malaysia in 2005. We used a canonical correspondence analysis (CCA) to reveal the relationships between fungal community composition and the environmental variables (canopy openness, soil water potential, amount and composition of coarse woody debris, litter mass, basal area, plant species composition). A total of 155 samples from 67 species were collected during the study period. The fungal species density represented by the number of species in a transect differed significantly among forest types. The fungal species density increased significantly with increasing number of pieces of coarse woody debris (CWD), but decreased significantly with increasing the scores of second axis of principal component analysis (PCA) for plant species composition. In the CCA ordination, automatic forward selection revealed that only the number of pieces of CWD significantly affected the fungal species composition. The occurrences of Flabellophora licmophora, Coriolopsis retropicta, Microporus vernicipes, and Amauroderma subrugosum were positively correlated with the number of pieces of CWD. Our study clearly demonstrated that forest use negatively affected aphyllophoraceous fungal diversity and suggest that the quantity of CWD would be an important determinant of fungal diversity and composition.     

The response of lichens inhabiting exposed wood of spruce logs to post-hurricane disturbances in Western Carpathian forests

We investigated which of the following environmental factors: the number of years since the windthrow of the tree (the age of dead wood), the phytocenosis (the type of forest community), altitude, exposure, wood hardness and the spatial scale of forest disturbances (small gaps with a few fallen spruces vs large-area windthrows) contributed to the diversity and abundance of lichens inhabiting the exposed wood of windthrown spruce trees in Polish Western Carpathian forests. Both Shannon H index and sum of coverage coefficients rose with increasing age of the wood, levelling off after 11–14 y (diversity) and 14–17 y (abundance). This factor appeared to be the most important for this group of lichens, but the significant positive impact of large-area windthrows on the lichen abundance was also demonstrated by using a GLM model. The age of the wood we precisely determined on the basis of data on Norway spruce mortality collected annually in permanent plots of the Gorce National Park since 2000. Using the Shore durometer we linked the course of the wood-inhabiting lichen succession with wood decay more precisely than before. The largest number of species was associated with medium hard wood, i.e., 51 < x ≤ 80 on the Shore scale. Based on the NMDS analysis, we distinguished four age groups of logs, differing in lichen abundance and defined by the dominance of distinctive species. A large number of usually corticolous lichen species used the wood of windthrown spruce logs as an optional habitat to survive large-scale, post-hurricane forest disturbances.     

Fire and site type effects on the long-term carbon and nitrogen balance in pristine Siberian Scots pine forests

Effects of fire and site type on carbon (C) and nitrogen (N) balances were determined by following the change of total and component C and N pools along four chronosequences of fire-prone Siberian Scots pine ecosystems. These differed in the mean return interval of surface fires (unburned – moderately burned, 40 years – heavily burned, 25 years) and site quality (lichen versus Vaccinium site type). Of the Vaccinium site type (higher site quality) only a moderately burned chronosequence was studied. A total of 22 even-aged stands were investigated with stand ages ranging from 2 to 383 years. The C balance was dominated by the opposing dynamics of coarse woody debris (CWD) and biomass and could be divided into three phases: (1) Young stands (up to 40 years)acted as a net source for C of 6-10 mol C m-2 year-1 because the previous generation CWD pool originating from stand-replacing crown fires decayed much faster than biomass increased. During this period the C pool in the unburned lichen type chronosequence decreased from 807 to 480 mol C m-2. (2) Middle aged stands (40-100 years) being in a stage of maximum biomass accumulation were a net sink of 8-10 mol C m-2 year-1. (3)Maturestands (100 to > 350 years) continued to sequester C at a lower rate (0.8-2.5mol C m-2 year-1). Differences in the rates of C sequestration during the two later phases could be explained by the complex interaction between surface fire regime and site type. Recurrent surface fires resulted in enhanced mortality and regularly redistributed C from the living to the CWD pool thereby lowering the rate of C sequestration. Site quality determined the potential to recover from disturbance by fire events. Differences in site type did not correlate with soil and total ecosystem N pool size. However, the N status of needles as well as the N pool of physiologically active tissue was highest in the stands of the Vaccinium type. The woody C pool (biomass + CWD) was sensitive to differences in surface fire regime and site type. It was lowest in the heavily burned lichen type chronosequence (297 ± 108 mol C m-2), intermediate in the unburned and moderately burned lichen type chronosequence (571 ± 179 mol C m-2) and highest in the moderately burned Vaccinium type chronosequence (810 ± 334 mol C m-2). In contrast, the total soil C pool (organic plus mineral layer down to a depth of 25 cm) was independent of stand age, surface fire regimeand site type and fluctuated around a value of 250 mol C m-2. The organic layer C pool oscillated in response to recurring surface fires and its C pool was dependent on time since fire increasing at a rate of about 1.5 mol C m-2 year-during the first 40 years and then reaching a plateau of 170 mol C m-2. The total ecosystem N pool was 7.4 ± 1.5 mol N m-2 on average of which only 25 % were stored in biomass or coarse woody debris. Total ecosystem N was independent of stand age, surface fire regime and site type. No correlation was found between total ecosystem C and N pools. Average total ecosystem C:N ratio was 114 ± 35 mol C mol N-1. A conceptual model illustrating how changes in the regime of stand-replacing crown fires and recurrent surface fires and changes in site quality interact in determining the long-term C balance in Siberian Scots pine forests is presented.