Selective stripping of rowan (Sorbus aucuparia L.) bark by cattle in North-east Scotland

Summary

During winter 1973–4 beef cattle removed bark selectively and extensively from rowan growing in two Aberdeenshire woods. Some bark was also taken from Norway spruce and willow, but damage to all other species, including ash, beech, birch, gean, larch, Scots pine and sycamore, was negligible. In other woodlands utilised by cattle in this area bark-stripping was either less severe or absent. Possible causes for the bark-stripping are discussed. The rowan bark was not distinctive in cover, species of epiphytic lichen or inorganic composition, except perhaps for magnesium.     

Large proportion of wood dependent lichens in boreal pine forest are confined to old hard wood

Intensive forest management has led to a population decline in many species, including those dependent on dead wood. Many lichens are known to depend on dead wood, but their habitat requirements have been little studied. In this study we investigated the habitat requirements of wood dependent lichens on coarse dead wood (diameter >10 cm) of Scots pine Pinus sylvestris in managed boreal forests in central Sweden. Twenty-one wood dependent lichen species were recorded, of which eleven were confined to old (estimated to be >120 years old) and hard dead wood. Almost all of this wood has emanated from kelo trees, i.e. decorticated and resin-impregnated standing pine trees that died long time ago. We found four red-listed species, of which two were exclusive and two highly associated with old and hard wood. Lichen species composition differed significantly among dead wood types (low stumps, snags, logs), wood hardness, wood age and occurrence of fire scars. Snags had higher number of species per dead wood area than logs and low stumps, and old snags had higher number of species per dead wood area than young ones. Since wood from kelo trees harbours a specialized lichen flora, conservation of wood dependent lichens requires management strategies ensuring the future presence of this wood type. Besides preserving available kelo wood, the formation of this substratum should be supported by setting aside P. sylvestris forests and subject these to prescribed burnings as well as to allow wild fires in some of these forests.     

Do moss samples,pollen traps and modern lake sediments all collect pollen in the same way? A comparison from the forest limit area of northernmost Europe

Moss polsters, pollen traps and lake surface sediment samples are commonly used as climate calibration data or as modern analogues for reconstructing vegetation from fossil profiles, but the differences in pollen content between these media have received little attention. This study aims to analyse how the three media differ in reflecting individual vegetation types and spatial differences in vegetation. 119 modern samples (64 moss polsters, 37 lake surface sediment samples and 18 pollen traps from which a collection was made annually) were taken from northern Fennoscandia and the Kola Peninsula as a broad transect crossing the northernmost forest limits of Betula pubescens ssp. czerepanovii (mountain birch), Pinus sylvestris (Scots pine) and Picea abies (Norway spruce). The pollen assemblages from these samples were compared with the surrounding vegetation visually and via PCA (principle components analysis) and cluster analysis. Both comparisons allow a correct distinction between pollen assemblages of arctic/alpine heath, mountain birch dominated areas, and boreal coniferous forests. The differences between the vegetation zones are stronger than the differences between the sampling media. Nevertheless, lake sediment samples from the mountain birch woodland zone tend to overestimate pine and underestimate birch. Pollen traps are biased towards lower tree pollen percentages and higher values of shrubs, herbs and Cyperaceae. This bias is especially strong in traps that have missing years in the data. Irrespective of the vegetation zone, pollen traps tend to have lower Pinus pollen percentages than in the adjacent moss polsters.     

Pattern of Substrate Preferences of Free Living Protists (Myxomycetes) on Decaying Wood

We have studied the influence of the stage of decomposition and acidity of wood, as well as the illumination of the microhabitat on the species composition, abundance, and occurrence of slime molds (Myxomycetes) of the xylobiontic (inhabitants of the wood) substrate complex of forest communities in Siberia (Altai krai, Altai Republic, and Novosibirsk oblast). This work is based on a study of 1777 samples of fruit bodies (sporophores) of myxomycetes. In the analysis of data for communities of Myxomycetes of the xylobiontic substrate complex, we recognize a successional series which clearly correlates with the stage of wood decomposition. The study of the distribution of the species composition of the slime mold on wood of various stages of decomposition shows that the maximum number of species is observed on dead trunks where the wood is of medium density and the bark can easily be separated (W3). The lowest specificity and diversity of species composition is observed on the fallen trunks of trees with very dense wood and dense bark (W1). These dead trees have low water-retaining capacity, tight bark, and almost intact wood containing large amounts of lignin, preventing the invasion of plasmodia and myxamoebae of Myxomycetes into the trunk. A study of the location of slime-mold colonies in relation to the illumination level shows that the greatest number of species is found on the front, side, and bottom parts of dead trunks of woody plants. The least specificity and the least variety of Myxomycetes species are found on apical parts of dead tree trunks facing the sun. The exception are species with large fruit bodies such as Fuligo septica and Reticularia splendens, which are most often observed on the upper parts of the sun facing trunks of woody plants. In the research area, the complex of species typical for wood of coniferous trees is described. It is noted that acidophilic slime molds of the genera Comatricha and Cribraria can be traced in xylobiontic and epiphytic–corticuloid substrate complexes.     

The Red-belted Bracket (Fomitopsis pinicola) colonizes spruce trees early after bark beetle attack and persists

The Red-belted Bracket (Fomitopsis pinicola) is one of the major decomposers of coniferous wood in Europe and can reach high densities after outbreaks of bark beetles. However, factors of dead wood type and decay stage, which determine the growth of reproductive biomass, i.e. basidiomes, remain unclear. In 2013, we surveyed 1280 dead wood objects and vital trees in spruce stands killed by the bark beetle Ips typographus in 2012, 2002, 1992 and in undisturbed stands for the presence, number, mean basidiome size and total volume of basidiomes. Living basidiomes were equally abundant on dead wood 1, 11, and 21 y after bark beetle outbreak, but were lacking on living trees. Our results indicate that F. pinicola is an effective early colonizer of the huge resource pulse of dead wood caused by the outbreak of bark beetles and basidiomes can persist for 21 y.     

Transmission of the Pinewood Nematode, Bursaphelenchus xylophilus,to Slash Pine Trees and Log Bolts by a Cerambycid Beetle,Monochamus titillator, in Florida

Field-collected adults of the southern pine sawyer, Monochamus titillator (F.) (Coleoptera: Cerambycidae), naturally infested with fourth-stage juveniles (dauerlarvae) of the pinewood nematode, Bursaphelenchus xylophilus (Steiner and Buhrer, 1934) Nickle, 1970, were maturation fed on excised shoots of typical slash pine, Pinus elliottii Engelm. var elliottii, for 21 days. During August 1981, a male and female adult beetle were held in a sleeve cage placed on the terminal of a side branch of each of seven replicate, healthy 10-year-old slash pine trees. All seven branch terminals showed evidence of beetle feeding on the bark after 1 week, and pinewood nematodes were present in wood samples taken near these feeding sites. Four of the seven trees showed wilt symptoms in 4-6 weeks and died about 9 weeks after beetle feeding. Pinewood nematodes were recovered from the roots and trunks of the dead trees. Each of seven replicate slash pine log bolts was enclosed in a jar with a pair of the same beetles used in the sleeve cages. After 1 week, wood underlying beetle oviposition sites in the bark of all replicate log bolts was infested with the pinewood nematode.     

<Emphasis Type="Italic">Lecophagus vermicola</Emphasis> sp. nov., a nematophagous hyphomycete with an unusual hunting strategy

Lecophagus vermicola sp. nov. is described and illustrated as a predacious (carnivorous) hyphomycete living in bark fissures of living trees of Platanus and other angiosperm and gymnosperm trees, recorded in Hungary, Luxembourg and France. The fungus captures nematodes unlike other Lecophagus species, which are predators of rotifers and tardigrades. The morphology of the sessile, adhesive knobs differ from all previously described species of the genus which form adhesive pegs. Molecular data confirms that the new species belongs to the Lecophagus clade but without matching existing sequences. The fungus captures victims with adhesive knobs and colonizes its prey with a mycelium of rather broad hyphae on which, again, adhesive knobs are formed which penetrate the cuticule of the victim. Clusters of colonized nematodes form a network utilized to capture more prey. The fungus lives in the xeric, ephemerally aquatic habitat of bark fissures of standing, living or dead, corticated trunks and branches. The genus Haptocara is compared, which has similar adhesive knobs capturing nematodes and similar broad hyphae, but for which no molecular data was available.     

On the Distribution of <Emphasis Type="Italic">Pleurozium schreberi</Emphasis> (Bryophyta,Hylocomiaceae) in the East European Plain and Eastern Fennoscandia

Pleurozium schreberi is one of the most common moss species in the forest area. It is dominant in the moss layer of blueberry and wood sorrel forests. It can occur in small quantities in almost all types of forests (even in bogs). It is also a typical component of the moss layer in tundra. The article considers the distribution of Pleurozium schreberi in the East European Plain and Eastern Fennoscandia. On the basis of literature sources on the occurrence of the species in different regions (according to point data), a model map of species distribution using the kriging-method has been created. The overlaying of the model map on the maps of spatial distribution of climatic parameters and vegetation zones in this area has revealed that the biogeographical preferences of the species. P. schreberi is characterized by its highest distribution in the forest zone. It often occurs here and represents a phytocenotically active species. The occurrence of Pleurozium schreberi dramatically decreases in the transition from the forest to the steppe zone, where it is a rather rare species, growing exclusively in pine and birch pegs. This species disappears in the open steppe. From the steppe zone to the south, the occurrence of Pleurozium schreberi gradually decreases with increase in summer temperatures and decrease in precipitation and with forest disappearance. In the north, where the species is highly active, its range abruptly ends on the coast of the Arctic Ocean. This pattern of distribution of Pleurozium schreberi is associated both with cenotic preferences and with climate: it becomes rare in regions with summer temperatures higher than +23°C and annual precipitation of less than 400 mm.     

Biology of xylophilous larvae of the rhagionid flies <Emphasis Type="Italic">Chrysopilus dives</Emphasis> and <Emphasis Type="Italic">Chrysopilus nigrifacies</Emphasis> (Diptera,Rhagionidae)

The larval morphology and biology of two species of rhagionid flies (Diptera, Rhagionidae) from the Russian Far East are described for the first time. Larvae of Chrysopilus dives Loew develop in the tunnels of xylobiont insects in the trunks of ash trees. Larvae of Ch. nigrifacies Nagatomi were found under the bark of the elm, Sakhalin cork tree, birch, and fir. Morphological characteristics of larvae of Chrysopilus are discussed.     

The utilization of dead wood resources by woodpeckers in Britain

Dead wood is important for woodpeckers, providing foraging, roost and nest-sites. In this paper, data from long-term studies of woodpeckers and dead wood in oakwoods in southern England are used to examine the dead wood requirements of the three British resident woodpecker species. Both Great Dendrocopos major and Lesser Spotted Woodpeckers Dendrocopos minor select dead trees for nest-sites although the former is able to nest in living trees too. On the other hand a smaller fraction of Lesser Spotted Woodpecker nests are in living trees. Green Woodpecker Picus viridis shows no selection for dead nesting trees. Hence the smallest woodpecker species appears to be most dependent on dead and decaying trees for nest-sites. Great and Lesser Spotted Woodpeckers show no preference for foraging on dead trees although they both make use of dead branches on living trees. Lesser Spotted Woodpeckers forage on smaller branches higher in the tree than Great Spotted Woodpeckers. There has been a trend for increasing dead wood resources in the study woods with both dead wood on the ground and standing dead trees (snags) increasing in the last 20 years. The levels of dead wood are shown to be the result of continual processes of creation and decay. Around 0.5% of oak Quercus spp., Ash Fraxinus excelsior and Hornbeam Carpinus betulus and 3.4% of the birch Betula spp. trees die each year in the woods resulting in a continuity of new dead snags and fallen trees. There is a high turnover of standing dead snags of oak and birch with 95% and 80% annual survival, respectively. Snags are only suitable for nesting Great Spotted Woodpeckers for a few years after their creation. It is suggested that these stand and dead wood dynamics are likely to provide habitats more favourable for the Great Spotted than the Lesser Spotted Woodpecker.     

Spatial and temporal patterns of rowan (<Emphasis Type="Italic">Sorbus aucuparia</Emphasis> L.) regeneration in West Carpathian subalpine spruce forest

Non-random seed shadows are commonly seen in plant species whose seeds are dispersed by animals, in particular by birds. The behaviour of birds can influence the spatial pattern of seed dispersal and, consequently, the entire regeneration process of fleshy-fruited trees. This study examined regeneration patterns in a fleshy-fruited tree species, rowan (Sorbus aucuparia L.), growing in West Carpathian subalpine spruce forests, focussing on two problems: the temporal relationship between rowan regeneration and gap formation, and the spatial relationship between rowan regeneration and stand structure. It was found that rowan seedlings and saplings were recruited in advance of gap formation. Establishment of new rowan individuals in gaps was infrequent, but gaps enhanced their regeneration nearby under spruce canopy, where they occurred densely in a narrow belt about 15 m wide. Inside spruce stands, the highest density of young rowans was directly under crowns, especially near trunk bases. Few rowan saplings were found growing under mature rowan trees. The presence of a rowan seedling and sapling bank determines whether rowans fill spruce stand gaps. Dense rowan groves can develop mainly in extensive but slowly expanding gaps.     

Hierarchical structure of juvenile hybrid aspen xylem revealed using X-ray scattering and microtomography

X-ray scattering and microtomography (μCT) are useful techniques to reveal the structure of wood at the nano- and micrometer scales. The nanostructure of xylem in greenhouse-grown 2.5- to 3.5-month-old Populus tremula L.?×?tremuloides Michx. trees was characterized using wide-angle X-ray scattering (WAXS), and the cellular structure was investigated using μCT. For comparison, the nanostructure of wood in 2-year-old silver birch, Norway spruce and Scots pine saplings was determined. Based on the μCT results, the lengths of fiber lumina of the hybrid aspen saplings were shorter than any previous results on the lengths of wood fibers. The mean microfibril angles of the hybrid aspen saplings were significantly lower (8°–14°) than those of the birch, spruce and pine saplings (27°–35°) implicating that cellulose microfibrils were oriented nearly parallel to the cell axis in the young hybrid aspen saplings. Hybrid aspen saplings were found to contain tension wood based on the histochemical analysis and μCT images. However, typical tension wood properties, i.e. larger crystallite width and higher crystallinity than in normal wood, were detected only in a few hybrid aspen samples, while in most of the hybrid aspen saplings, the crystallite widths (3.0?±?0.1?nm) and the crystallinities (30?±?5?%) corresponded to those of normal wood. The deformations of cellulose crystallites were determined using WAXS in situ upon dehydration of the never-dried samples. In all the species studied, the cellulose unit cell dimension decreased and disorder of cellulose chains increased parallel to the chains upon drying. Also, the transverse disorder of chains increased in birch, spruce and pine, while no changes were detected in this direction in hybrid aspen. The crystallite widths and drying deformation results might indicate that the gelatinous layer has not fully developed in the young hybrid aspen saplings.     

The Influence of Location,Tree Age and Forest Habitat Type on Basic Fuel Properties of the Wood of the Silver Birch (<Emphasis Type="Italic">Betula pendula</Emphasis> Roth.) in Poland

The most extensive study to be carried out in Poland, and one of only a few worldwide, regarding the influence of location, tree age and forest habitat type on the basic fuel properties of silver birch (Betula pendula Roth.) wood was conducted in 12 Forestry Districts of the Polish State Forests. The field study included trees in three age groups of approximately 30, 50 and 70 years. The research was carried out in a fresh broadleaved forest (FBF) habitat type, where in Poland silver birch stands predominate in respect of coverage area and merchantable volume. Additionally, for five selected Forestry Districts, a comparative study was conducted in a fresh mixed broadleaved forest (FMBF), the second most important habitat of this tree species. A total of 306 test trees were examined. For every sample, calorific value and contents of ash, carbon, hydrogen, nitrogen, sulphur and chlorine were determined. The results indicated a statistically significant influence of location on the calorific value (p?=?0.0001) and on the contents of ash (p?<?0.0001), carbon (p?<?0.0001), hydrogen (p?<?0.0001), nitrogen (p?<?0.0001) and chlorine (p?<?0.0001) in the analysed wood. Moreover, statistically significant differences were observed between values of ash content (p?=?0.046) and of calorific value (p?=?0.0026) depending on the forest habitat type. Tree age was found to have no significant influence on the calorific value of silver birch wood.     

Tree species and position matter: the role of pests for survival of other insects

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The fungal matrices of <Emphasis Type="Italic">Ophiostoma ips</Emphasis> hinder movement of the biocontrol nematode agent, <Emphasis Type="Italic">Deladenus siricidicola</Emphasis>, disrupting management of the woodwasp, <Emphasis Type="Italic">Sirex noctilio</Emphasis>

Deladenus (=?Beddingia) siricidicola (Tylenchida: Neotylenchidae) is the most effective biocontrol agent used against the invasive wood wasp, Sirex noctilio (Fabricius) (Hymenoptera: Siricidae). The nematodes feed and reproduce on the wood-inhabiting fungus, Amylostereum areolatum (Chaillet ex Fr.) Boidin (Russulales: Amylostereaceae) and parasitise larvae of S. noctilio. In the nematode biocontrol program, the nematodes are inoculated into herbicide-weakened ‘trap trees’. Recent declines in nematode parasitism of S. noctilio in Australia have coincided with an increased incidence of an exotic bark beetle, Ips grandicollis (Eichhoff) (Coleoptera: Curculionidae), attacking trap trees and vectoring a wood-inhabiting fungus, Ophiostoma ips (Rumbold) Nannfelt (Ophiostomatales: Ophiostomataceae), which may inhibit migration of the nematode within the tree to the detriment of S. noctilio biocontrol. Several in vitro and in vivo experiments were conducted to investigate the effect of fungal interactions on the ability of D. siricidicola to locate and reproduce on A. areolatum. Deladenus siricidicola showed preference to A. areolatum in the presence and absence of O. ips, but the presence of O. ips negatively affected the choice response and the number of eggs laid by the nematodes. Deladenus siricidicola was unable to survive and reproduce on O. ips. Results give a clearer understanding of the choice response of D. siricidicola in I. grandicollis infested trees, explaining the disruptive impact of bark beetles on biocontrol of S. noctilio, an effect that could extend from Australia to other important pine growing countries.     

Biotic resistance and the spatiotemporal distribution of an invading woodwasp, <Emphasis Type="Italic">Sirex noctilio</Emphasis>

Quantifying the strength of interactions among introduced and native species across space and time is critical in understanding biological invasions as they can attenuate or amplify the magnitude of impact. The European woodwasp, Sirex noctilio F., a global threat to pines, is a recent invader to North America. It attacks and kills primarily Pinus resinosa and Pinus sylvestris, and encounters a diverse assemblage of potential competitors for this resource. We quantified spatial colonization patterns of this woodwasp and resident competitors including scolytine bark beetles, woodboring cerambycid and buprestid beetles, and the fungal root rot diseases Armillaria and Heterobasidion across an 80 year old pine plantation over 4 years. All xylophages were spatially aggregated, but only on a localized scale of 15–20 m. Colonizers occurred non-randomly within trees, with S. noctilio negatively or neutrally associated with all other colonizing agents, whereas all other insect and root rot colonizers were mostly positively co-associated. An autologistic regression with spatially-weighted variables indicated the probability of a dead tree exhibiting symptoms of S. noctilio attack was positively associated with tree density, host species (P. sylvestris), and density of S. noctilio-attacked trees from the current and previous year. Interspecific stand patterns were weaker; probability of attack was negatively associated only with root rot pathogens. Across spatial scales, there were mixed (woodborers) and neutral (bark beetles) associations between S. noctilio and other co-colonizing insects. Our results suggest that competitive interactions with resident species may be contributing to the limited success of S. noctilio in North America, but are unlikely to be driving it by themselves.     

Regenerative and age dynamic of spruce population in pine plantations under conditions typical for spruce development

We have studied the natural regeneration of spruce (Picea abies L.) under a canopy of pine (Pinus sylvestris L.) on loamy soils. Spruce survival and growth depend on the duration of the regeneration period from the creation of the plantation and the local conditions formed as a result of uneven thinning of pine and spruce canopy. The formation of spruce population is mainly determined by trees that regenerated upon intensive thinning of 20- to 40-year-old pine trees. Spruce regeneration may be enhanced by timely cleaning cutting in pine plantations. The first one, done at the age of 15–20 years, favors pine growth and spruce regeneration. At the normal reproduction of spruce population under the canopy of 80-year-old pine plantations, the second spruce layer is formed. Trunk reserve in this layer is 20–25% of the reserves of the first layer. After its formation, the light regime in the forest depends greatly on the space volume occupied by spruce crowns. Their percentage is especially high at the relative height equal to 0.4–0.7 of the mean spruce height in the second layer. Smaller spruce trees may exist for a long time period, but their development is slowed down and they die at the undergrowth stage. When the plantation is 150 years old, the reserves of spruce trees regenerated under the canopy of pine comprise one-third of the total reserves of the plantation. If the growing conditions are favorable for spruce (C₃), the stability and productivity of pine-spruce plantations exceed those of the pure spruce plantations. The reasonability of natural spruce regeneration for the creation of pine-spruce plantations under C₃ conditions should be substantiated with the consideration of their designation, ages of cuttings, and the possibility of plantation creation and sanitary cuttings according to the valid regulations.     

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.