Preferences of dead wood inhabiting bryophytes for decay stage,log size and habitat types in Hungarian beech forests

Abstract

This study describes the quantitative preferences of epixylic bryophytes for decay stages, log size and habitat type. The two habitats investigated were zonal (situated on plateaux without extreme microclimatic conditions) and ravine-like near-natural beech stands. Preferences of 30 bryophyte species are described using logistic regression modelling. The species pool differed between habitat types. In the ravine-like forests more species are found. Regionally rare epixylic species (mainly liverworts) were limited to these forests. The species richness of individual logs was also higher in the ravine-like than in the zonal forests and the proportion of logs of all sizes colonized by bryophytes was higher in the former. The species were classified into four categories on the basis of their preference for decay stages: strict epixylics, indifferent species, preferential epixylics and epiphytes. The epixylics and preferential epixylics prefer advanced decay stages while the latter extend further into earlier decay stages than strict epixylics. Common epixylic species have a wider tolerance to decay stage than rare ones. Probabilities of species occurrence increase with log size, more strongly in zonal stands than in ravines. A simple successional scheme for bryophyte succession on dead wood is proposed on the basis of the observed species responses.     

Cryptogam communities on fallen logs in the Duke Forest,North Carolina

Abstract. I studied cryptogam (i.e. bryophyte and lichen) communities on fallen logs in the Duke Forest, Durham and Orange Counties, North Carolina, USA, to determine the relationship of log characteristics and microsite to community composition. Species composition and abundance were estimated for 111 randomly selected fallen logs. Interior wood samples were used to identify the tree species. I determined physical and chemical characteristics for each log, and described the log microsite. Canonical Correspondence Analysis (CCA) detected a clear gradient in cryptogam species composition which is correlated with the species of log and the presence of bark. Communities on hardwood bark, hardwood wood, and pine substrates were the most distinct. CCA also revealed that the microsite is not as closely related to species composition as are substrate pH and density. The majority of the cryptogam species encountered on the fallen logs are commonly reported from other substrates in the forest. However, within the habitat type of fallen logs, several species were apparently restricted to certain substrate types.     

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.     

Functional response of lignicolous fungal guilds to bark beetle deforestation

Conifer-dominated forests in the northern hemisphere are prone to large-scale natural disturbances, yet our understanding of their effects beyond changes in species diversity is limited. Bark beetle disturbances provide dead wood for lignicolous fungal guilds and increase insolation but also desiccation. We investigated whether species richness of these guilds increases and functional diversity decreases after bark beetle disturbance, which would promote through habitat filtering the coexistence of species adapted to harsh conditions, i.e. light stress for lichens and substrate desiccation for wood-inhabiting fungi.We sampled epixylic and epiphytic lichens (primary producers) and wood-inhabiting fungi (mainly wood decomposers, some form ectomycorrhizas) in the Bohemian Forest (Long Term Ecological Research – LTER – Site Bavarian Forest National Park), an area in Central Europe most heavily affected by the bark beetle Ips typographus, on undisturbed plots and disturbed plots with spruce (Picea abies) dieback 8 years ago. We analysed species diversity, functional diversity (optimized by phylogeny), and functional compositions.Species richness of lichens but not that of wood-inhabiting fungi was higher on disturbed plots than on undisturbed plots. Community compositions of both guilds differed considerably on disturbed and undisturbed plots. On both types of plots, lichen communities were clustered according to functional diversity, which indicated habitat filtering, and fungal communities were overdispersed, which indicated competition. Disturbance increased the strength of these two patterns only slightly and was significant only for fungi. Single-trait analysis revealed changes in the functional composition; on disturbed plots, lichenous species with larger and more complex growth forms and fungi with large, perennial fruit bodies were favoured. Although the forest canopy changed tremendously because of the bark beetle disturbance, the most important driver of lichen and fungal diversity and mean trait assemblages seemed to be the enrichment of dead wood. The changes in insolation and moisture did not act as habitat filters for either guild. This indicated that the assembly patterns of lichen and fungal communities in coniferous forests are not affected by stand-replacing disturbances in contrast to the predictions for other disturbance regimes.     

Species richness of tropical wood-inhabiting macrofungi provides support for species-energy theory

A study was undertaken at the El Verde Field Station in Puerto Rico to determine the effect of energy available from newly dead trees on the species richness of macrofungal communities that inhabit them. It is hypothesized that there is a positive relationship between available energy and species richness. Energy was measured using the volume of the dead trees and the wood density of living trees of the same species. One hundred ninety-four logs of known tree species were surveyed 1 y for fruiting bodies of macrofungi at monthly intervals. For individual logs, log volume had a significant positive effect on macrofungal species richness. Younger logs had significantly higher species richness than older logs, and those with less apparent decay had more species than those with more decay. When logs were grouped by tree species, total wood volume and density of live wood had a significant positive effect and average log diameter had a negative effect on total species richness and abundance of the wood-inhabiting macrofungi. Macrofungal richness and abundance constantly increased with initial wood density; there was no evidence for a unimodal relationship. These results support the proposed relationship between species richness and energy.     

Sequence of bryophytes and lichens in relation to substrate variables of decaying coniferous wood in Northern Sweden

Söderström, L. 1988. Sequence of bryophytes and lichens in relation to substrate variables of decaying coniferous wood in Northern Sweden. - Nord. J. Bot. 8: 89–97. Copenhagen. ISSN 0107–055X.
The species sequence on decaying logs during the decay period has been investigated in a forest in Northern Sweden. The species present can be divided, by means of a weighted average technique, into four groups, facultative epiphytes, early and late epixylics, and ground flora species.
Species sequence on several variables reflecting the wood decay, was measured for 25 species. Wood texture (degree of surface erosion) was the most important variable in separating species. Logs high above ground had very few species, probably due to low moisture content. Log diameter was important and several species did not occur, or only rarely occurred, on small logs. Small logs are more rapidly overgrown by ground flora species and very few reach late decay stages before being overgrown. Larger logs also have greater waterholding capacity which facilitates growth of many species sensitive to drought.     

Bryophytes and decaying wood– a comparison between managed and natural forest

A comparative study was made of one managed forest and one natural forest regarding the supply of decaying wood on the ground and the occurrence of bryophytes on the wood, especially epixylic specialists.
The total substrate surface of wood is larger in the primeval forest. The larger quantity of decaying wood increases the probability of wood occurring in ail decay stages. Logs of Populus tremula occurred only in the natural forest. An important proportion of the wood substrate in the natural forest consisted of logs of large diameter class; these logs were missing in the managed forest. Fiftyfour bryophyte species were found on decaying wood. They were separated into four different groups. Facultative epiphytes first colonized decaying wood, followed by epixylic specialists and finally by competitive epigeics. Opportunistic generalists showed a more irregular pattern of occurrence. The bryophyte flora on decaying wood was most species rich in the natural forest. Sixteen epixylic specialists occurred here, several considered to be threatened by forest management, while only 5 of these species occurred in the managed forest.
The higher frequency of epixylic specialists in the natural forest is related to the greater frequency of favourable habitats, specifically: logs of large diameter, decayed wood, Populus tremula and (to a lesser extent) Picea abies.     

Surface Covering of Downed Logs: Drivers of a Neglected Process in Dead Wood Ecology

Many species use coarse woody debris (CWD) and are disadvantaged by the forestry-induced loss of this resource. A neglected process affecting CWD is the covering of the surfaces of downed logs caused by sinking into the ground (increasing soil contact, mostly covering the underside of the log), and dense overgrowth by ground vegetation. Such cover is likely to profoundly influence the quality and accessibility of CWD for wood-inhabiting organisms, but the factors affecting covering are largely unknown. In a five-year experiment we determined predictors of covering rate of fresh logs in boreal forests and clear-cuts. Logs with branches were little covered because they had low longitudinal ground contact. For branchless logs, longitudinal ground contact was most strongly related to estimated peat depth (positive relation). The strongest predictor for total cover of branchless logs was longitudinal ground contact. To evaluate the effect on cover of factors other than longitudinal ground contact, we separately analyzed data from only those log sections that were in contact with the ground. Four factors were prominent predictors of percentage cover of such log sections: estimated peat depth, canopy shade (both increasing cover), potential solar radiation calculated from slope and slope aspect, and diameter of the log (both reducing cover). Peat increased cover directly through its low resistance, which allowed logs to sink and soil contact to increase. High moisture and low temperatures in pole-ward facing slopes and under a canopy favor peat formation through lowered decomposition and enhanced growth of peat-forming mosses, which also proved to rapidly overgrow logs. We found that in some boreal forests, peat and fast-growing mosses can rapidly cover logs lying on the ground. When actively introducing CWD for conservation purposes, we recommend that such rapid covering is avoided, thereby most likely improving the CWD''s longevity as habitat for many species.     

Epiphytic algae on eelgrass at Roscoff,France

Algae growing as epiphytes on eelgrass, Zostera marina L., at Roscoff were investigated during one year by examining the species richness and abundance. The abundance of the algae, expressed as cover values, appeared to be influenced by the characteristics of the substrate, especially the ephemeral status and growth rate of the leaves.Colonization of the seagrass leaves occurred primarily on tips and edges, subsequently spreading over the faces to the base. The pioneers appeared to be the dominant, persistent taxa, of which Audouinella virgatula (Harv.) Dixon, Fosliella lejolisii (Rosan.) Howe, Myrionema magnusii (Sauv.) Lois. and the diatoms were most prominent. The oldest parts of fully grown leaves showed the most diverse epiphytic vegetation. Though there was a form of succession, a climax community was not reached.On examining the spatial distribution of the epiphytes, a group of characteristic taxa was seen to be accompanied by a second group, whose variation in occurrence seemed to be correlated with the height in the littoral.     

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.     

Reproductive development and maternal effects in the pine weevil Hylobius abietis

1. In a laboratory study of maturation feeding of female pine weevil Hylobius abietis on current and 1‐year‐old stem bark of transplants of Scots and Corsican pine, Norway and Sitka spruce, Douglas fir, and hybrid larch, the length of the pre‐oviposition period was influenced by the species on which weevils fed. The shortest pre‐oviposition period was on hybrid larch (11.8 days) and the longest on Douglas fir (15.5 days). 2. The species on which weevils fed also affected fecundity but there was evidence of a species–year interaction. Over a period of 36 days, most eggs were laid by weevils feeding on current stem of Norway spruce and Corsican and Scots pine and fewest on current stem of Sitka spruce. 3. Significant maternal effects on egg size were observed both in relation to female size and conifer species. The largest eggs were laid on Corsican pine and the smallest on Douglas fir, with no evidence of a trade‐off between number of eggs laid and their size. 4. There was a positive relationship between egg and larval size and between larval size and survival on logs of four conifer species. Residual resistance mechanisms in the bark of recently cut stumps and larval competition are discussed briefly in relation to the importance of the observed maternal effects on weevil population dynamics.     

Effects of introduced ungulates on forest understory communities in northern Patagonia are modified by timing and severity of stand mortality

Natural disturbances such as fires, windstorms, floods, and herbivory often act on plant communities, affecting their structure and the abundance and composition of their species. Most research has focused on the effects of single disturbances on plant communities whereas the synergistic effects of several disturbances have received less attention. In this study, we evaluated how timing and severity of tree mortality modified plant use by introduced deer and early post-mortality successional trajectories in northern Patagonian conifer forests. We sampled understory composition and deer use in Austrocedrus chilensis (ciprés de la cordillera) forest stands undergoing varying timing and severity of forest mortality as reconstructed using dendroecological techniques. In addition, we evaluated the effect of fallen logs on plant composition and deer use of plants by monitoring areas of massive dieback where fallen logs had been removed for fire hazard reduction, and nearby control areas not subjected to such removal. Stepwise regression analyses showed that history and severity of tree mortality strongly influence plant composition and deer use of plants. For deer use (with pellet counts and browsing index as response variables), results showed a positive relationship with degree of stand mortality and a negative relationship with cover of fallen logs. Similarly, cover of unpalatable shrub species was explained by canopy mortality history, whereas cover of palatable shrub species was positively associated with severity of canopy mortality. In areas where fallen logs had been removed, pellet counts were six times higher than those in control areas. Though total shrub species cover was similar between log removal and control areas, proportion of unpalatable shrubs increased in areas where fallen logs had been removed. In conclusion, deer use of plants was strongly limited by tall fallen logs, allowing palatable species to establish and grow. Fallen log removal accelerated deer entrance and changed understory composition toward more browse-resistant and unpalatable species. These results underscore the importance of considering the dynamics (timing, severity, and extent) of fallen woody debris influencing understory herbivory and post-disturbance succession. In addition, experimental results underpin the importance of maintaining snags and large woody debris in disturbed landscapes where salvage logging is a routine procedure.     

Dependence on Fire in Wood-living Insects: An Experiment with Burned and Unburned Spruce and Birch Logs

Several boreal wood-living insect species breed exclusively in recently burned forest. However, the reason for this dependence on fire is largely unknown. Here wood-living insects and other arthropods were sampled from burned and unburned logs of birch and spruce in a burned forest, together with unburned logs at a clearing and in an uncut forest, during two years of succession after tree death. Burned spruce logs hosted fewer beetles than unburned logs. Notably, bark-beetles and their associated fauna, responded negatively to fire-scorching of the logs while arthropods that feed on ascomycete fungi responded positively. Fire-scorched logs more often had visible ascomycete fungi, and lost their bark faster than unburned logs. However, despite this obvious effect of fire-scorching of the logs, the species composition in burned and unburned logs at the burned site was more similar than in unburned logs at the three different sites. A larger diversity of beetles, when measured with rarefaction, was found for fire-scorched logs. When sites were compared, birch logs had the most diverse fauna at the burned site and spruce logs in the uncut forest. Pyrophilous insect species were almost exclusively confined to the burned forest, but occurred in both burned and unburned logs. These species may be divided into two groups: (1) mycophagous species that need burned substrate per se because ascomycete fungi are favoured by burning, and (2) phloem-feeders and predators that are favoured by some habitat characteristic of recently burned forest rather than of burned wood.     

Conifer seedling recruitment in a southeastern Canadian boreal forest: the importance of substrate

Abstract. In order to explain conifer species recruitment in Canada's southeastern boreal forest, we characterized conifer regeneration microsites and determined how these microsites vary in abundance during succession. Microsite abundance was evaluated in deciduous, mixed and coniferous stands along a 234-yr postfire chronosequence. Conifers were most often found in relatively well-illuminated microsites, devoid of litter, especially broad-leaf litter, and with a reduced cover of lower vegetation (< 50 cm tall). Although associated with moss-rich forest floor substrates, Abies balsamea was the most ubiquitously distributed species. Picea glauca and especially Thuja occidentalis seedlings were frequently found on rotten logs. Light measurements did not show differences among seedling species nor between stand types. The percentage cover of broad-leaf litter decreased significantly during succession. Also, rotten logs covered with moss occupied a significantly larger area in the mid-successionnal stands than in early successional deciduous or late successional coniferous stands. The results suggest that the presence of specific forest floor substrate types is a factor explaining low conifer recruitment under deciduous stands, conifer codominance in the mid-successional stage, and delayed Thuja recolonization after fire. Results also suggest that some facilitation mechanism is responsible for the observed directional succession.     

The herb and dwarf shrubs colonization of decaying logs in subalpine forest in the Polish Tatra Mountains

This is a study of the colonization pattern of herbs and dwarf shrubs on rotten logs in subalpine spruce forests (Plagiothecio Piceetum) in the Tatra Mountains. On four study plots (total area 1.43 ha.) all dead logs were measured and the decomposition stage was estimated using the 8-degree scale. For each log the cover of all vascular species, bryophytes and lichens was determined according to the methods of classical phytosociology. Constancy and an index of coverage were calculated for all vascular species growing on logs. The total volume of logs was relatively high (93 m³ ha^–1) and constituted 22% of the volume of living trees. Logs and stumps covered 411 m² ha^–1. These values are similar to those known from natural spruce forest from Carpathians and Scandinavia. The 8 stages of decomposition were equally represented, which indicates a constant supply of dead wood to the forest floor over time. The colonization of dead wood starts with lichens, followed by bryophytes and finally herbs and tree saplings. The first vascular plant colonists of dead logs appear at decay stage nr. 3 at least 20 years after tree death. The most suitable condition for most of the herb species corresponds to decay stage nr. 6 ca. 50 years after tree death. The herb cover is distinctively dominated by Vaccinium myrtillus. Simultaneously with herb species, tree seedlings colonize the logs. Constancy and abundance of Norway spruce saplings increases with advanced decomposition. It seems that the herb cover of logs does not hinder the regeneration of spruce.     

Effects of Epixylic Vegetation Removal on the Dynamics of the Microbial Community Composition in Decaying Logs in an Alpine Forest

Ecosystems - Epixylic vegetation may be important in dead wood decay by altering the microenvironment and, thereby, microbial communities in logs. However, the interaction between epixylic...     

Fungal wood decomposer activities influence community structures of myxomycetes and bryophytes on coarse woody debris

Dead wood is an important habitat for forest organisms, and wood decay fungi are the principal agents determining the dead wood properties that influence the communities of organisms inhabiting dead wood. In this study, we investigated the effects of wood decomposer fungi on the communities of myxomycetes and bryophytes inhabiting decayed logs. On 196 pine logs, 72 species of fungi, 34 species and seven varieties of myxomycetes, and 16 species of bryophytes were identified. Although white rot was the dominant decay type in sapwood and heartwood, brown and soft rots were also prevalent, particularly in sapwood. Moreover, white rot and soft rot were positively and brown rot negatively correlated with wood pH. Ordination analyses clearly showed a succession of cryptogam species during log decomposition and showed significant correlations of communities with the pH, water content, and decay type of wood. These analyses indicate that fungal wood decomposer activities strongly influence the cryptogam communities on dead wood.     

Predicting decay and ground vegetation development in <Emphasis Type="Italic">Picea abies</Emphasis> snag stands

In a Picea abies (L.) Karst. (Norway spruce) mountain forest on the Gandberg site in the northern Swiss Alps, trees were killed by bark beetles in 1992–1997. A combination of field studies and dynamic modelling was used to project snag decay and future ground vegetation succession in these steep, unharvested stands. In permanent plots, ground vegetation cover and natural tree regeneration have been monitored annually since 1994. To obtain additional information on the abundance of snags, logs, boulders and other microsite types in these stands, the relative frequency of the microsite types was quantified along four strip transects on the montane and subalpine elevational levels. A dynamic model of snag decay and ground vegetation development was constructed (modified matrix model approach). Based on field data and literature values, the model was parameterised and initialised separately for the montane and the subalpine level. For model validation, microsite types were quantified in 2001 with the line-intercept method on both elevational levels. Starting from the conditions in the stands before the bark beetle attacks, it was possible to project short-term succession and to accurately simulate the decay and ground vegetation patterns eight years after tree die-back. Long-term simulations suggest that on the montane level, raspberries (Rubus idaeus L.) will be replaced by Picea abies, while on the subalpine level ferns will dominate for a long time.     

Species diversity in plant communities on ultramafic soils in relation to pine afforestation

Abstract. The influence of pine afforestation on the species diversity of plant communities on ultramafic substrate was investigated in an area of Tuscany, central Italy, by means of species-area relationships, plant unit area, the Gini coefficient and the pattern of the index of Jaccard in relation to plot size. The species-area relationship was found to best fit the semilogarithmic model. Contrary to the available data for temperate ecosystems, tree canopy cover was found to increase the α-diversity of the understorey vegetation and its cover. The nutrient input due to the pine canopy caused an increase in the abundance of the grass Festuca inops — which was already present in the community — and subsequently the spread of several grassland species leading to a reduction in dominance concentration and to the formation of a species-rich grassland. The pine cover caused a decrease in floristic resemblance between plots, especially when the canopy cover is scattered. The increase in species richness found under the pine canopy, where the metal content in the soil is higher, suggests that potentially toxic metals are not the most limiting factor in Tuscan ultramafic soils. The typical poorness of vegetation on ultramafic soils should be first of all related to hydrological and nutritional stresses.     

The importance of substrate type, shading and scorching for the attractiveness of dead wood to saproxylic beetles

Modern forestry management has reduced the amount of dead wood in forest ecosystems and this has become a serious threat to flora and fauna. Efforts are therefore being made to reverse this trend but one problem is that we still lack detailed knowledge regarding the substrate requirements of many saproxylic species. In a field experiment, conducted in three forest types (forest reserve, mature managed forest and clear-cut), we evaluated the value, from a conservation perspective, of different substrate types (logs, snags and tops) of Norway Spruce, Picea abies, and if the quality of spruce logs as saproxylic habitats can be improved by simple log treatments (scorching and shading). We collected 9982 individuals representing 262 saproxylic beetle species in window traps. Both substrate type and, to a lesser extent, log treatment had a significant effect on the abundance and species richness of saproxylic beetles attracted to the different dead wood substrates. However, more importantly, the composition of the beetle assemblages differed significantly between both substrates and log treatments. Snags, logs and tops all attracted significantly different beetle assemblages and scorched logs differed from untreated control logs. Sixteen red listed species were trapped, with the highest number (11 species) being found on scorched logs. We found strong evidence that some species preferred a specific substrate type, mainly logs, in some cases treated logs (scorched or shaded), but not snags, the substrate commonly provided for conservation purposes on e.g. clear-cuts. This stresses the importance of conducting forestry in such a way that a multitude of both forest habitats and dead wood substrates are available continuously in the forest landscape to maintain biodiversity.