Long-term changes in the woodlands of Clairinsh

Summary

The mixed oak woodland on the island of Clairinsh has been left unmanaged as part of the Loch Lomond National Nature Reserve. Until 1913 the woodlands had been treated as coppice-with-standards. Between 1961 and 1986 changes in the structure and composition of the stand were recorded by means of two transects, within which individual trees and shrubs were charted and measured. Most of the stand remained closed, increasing in basal area, but decreasing in density. However, a limited number of canopy gaps formed, part-icularly as a result of the 1968 hurricane. Of the individuals present in 1961, 30% died by 1986, mortality being particularly high in smaller individuals and in birch. Recruitment was mainly confined to rowan, holly and hazel in the closed woodland, and to birch and rowan in the gaps. The changes are discussed in relation to natural processes and the history of management.     

The last population of the Woodland Brown butterfly (<Emphasis Type="Italic">Lopinga achine</Emphasis>) in the Czech Republic: habitat use,demography and site management

The distribution of Lopinga achine (Lepidoptera Nymphalidae, Satyrinae) in the Czech Republic has declined from thirty grid squares before 1950 to just one extant population, restricted to a single area of deciduous woodland. A review of historical sites shows that this species used to occur in various types of deciduous woodland with a relatively sparse canopy maintained by coppicing and/or grazing. The extant population inhabits mature woodland with a mean canopy cover of 60% (quartiles 50% and 65%), sparse shrubs and a species-rich herb layer containing plant species requiring dry, warm and nutrient-poor conditions. The larval host plants are the fine-leafed sedges, Carex fritschii and C. michelii. In 2006, the total population contained about 10,000 adults but this may be an over-estimate, biased by male behaviour. Measurements of adult mobility, well approximated by an inverse-power function, suggested that all existing colonies are interconnected by dispersal. Continuing existence of the population depends on two conditions; nutrient-poor conditions for a diverse ground flora and a sparse tree canopy. While canopy closure is gradually increasing, the herb layer is threatened by soil enrichment due to the demise of traditional grazing, litter raking and grass mowing in woodlands. Any future management to favour Lopinga achine should include both measures to maintain a sparse canopy and measures to export biomass, such as raking or mowing of ground flora or, preferably, re-establishment of grazing. An erratum to this article can be found at     

Herb layer response to broadleaf tree species with different leaf litter quality and canopy structure in temperate forests

Question: How do broadleaf tree species affect humus characteristics, herb layer composition and species diversity through their leaf litter quality and canopy structure? Location: Mixed broadleaf forests in Brandenburg, NE Germany. Methods: We studied the herb and tree layer composition in 129 undisturbed stands using a 10‐degree cover‐abundance and percentage scale, respectively. The main floristic gradients were extracted by non‐metric multidimensional scaling. Effects of tree species on the herb layer were analysed with partial Spearman rank correlation. We assessed affinities for specific tree species using indicator species analysis. Results: Both beech and oak influenced herb layer composition mainly through their litter quality, which resulted in deep Ol and Of horizons, respectively. The less dense canopy of oak, in contrast to the dense beech canopy, enhanced species diversity in favour of indifferent herb species (species not closely tied to forests). Lime was correlated with a distinct floristic gradient, but a direct effect on the herb layer cannot be proven with the available data. Effects of hornbeam were less pronounced. Conclusions: The relationship between the tree and herb layer must be partly attributed to pH differences. However, tree species effects on humus characteristics and on light flux to the ground were largely responsible as well. The results suggest that tree species can influence herb layer composition and diversity, but the missing correlation with lime and hornbeam raise questions requiring further detailed investigation.     

Long-term change in growth, mortality and regeneration of trees in Denny Wood, an old-growth wood-pasture in the New Forest (UK)

Long-term changes in stand composition and structure were recorded in Denny Wood (New Forest, UK) by means of a permanent transect covering 2 ha. Denny is an ancient, mixed deciduous wood-pasture dominated by beech (Fagus sylvatica), pedunculate oak (Quercus robur) and holly (Ilex aquifolium) whose canopy trees ranged in age from approximately 70 years to over 300 years when the study began in 1956. Individual trees, shrubs and saplings were mapped and measured at irregular intervals until 1996.

During the 40 years of observations, storms and drought disrupted the stand. Considerable volumes of dead wood accumulated, and canopy gaps extended to 30% of the transect area. Small trees and saplings were severely damaged by ponies and grey squirrels. Regeneration ceased after 1964, due principally to heavy grazing and browsing by deer and ponies. Despite the disturbances, most mortality was due to competitive exclusion within well-stocked parts of the stand.

Historical records from the 17th century onwards demonstrate a long-term change from oak dominance with groups of beech before 1800 to beech dominance in the late 20th century. The stand through which the transect now runs was enclosed in 1870, and this allowed beech to regenerate abundantly, but in the nearby unenclosed part of Denny Wood holly regenerated more abundantly than beech.

The patterns of growth, mortality and regeneration are compared with natural temperate deciduous woodland. The long-term relationship between beech and oak is likely to involve periodic oak regeneration after major disturbances, interspersed with steady increases in the proportion of beech. The implications for managing and monitoring the “Ancient and Ornamental Woods” of the New Forest are considered.     

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.     

Regional differences in floristic change in South Swedish oak forests as related to soil chemistry and land use

Abstract. Changes of the ground layer vegetation between 1983 and 1993 were studied on permanent oak forest plots in Skåne and eastern Småland, two regions of South Sweden rich in oak but with different climatic conditions. 79 of the 123 more common species differed considerably in frequency between Skåne and Småland. The oak forests in Skåne contained more typical forest species, whereas the number of light-demanding grassland species was higher in Småland. Almost half of the more common species showed significant changes in frequency between 1983 and 1993. Several species of clearings and some light-demanding grassland species increased their frequency in Skåne. They were probably favoured by canopy thinning which occurred in 65 out of 95 permanent plots. Only two species decreased in frequency in Skåne, but 14 species showed a decrease in Småland. All were typical grassland species and their decline may be regarded as a long-term response to ceased grazing and tree canopy closure as almost no tree cutting occurred during the study period in Småland. Total species richness increased with management intensity in Skåne. This was due to decreasing species losses and increasing establishment of new species at intense management. A small general increase in species numbers was recorded in Småland. Species richness was more closely related to soil pH in Skåne than in Småland. The floristic variability between the oak forest plots was closely related to soil pH in both Skåne and Småland. We suggest that changes of light conditions as a result of changes in forest management are important for current secondary succession in South Swedish oak forests, whereas the general floristic pattern is to a large extent controlled by soil acidity.     

Seed bank composition and above-ground vegetation in response to grazing in sub-Mediterranean oak forests (NW Greece)

We investigate the persistent soil seed bank composition and its relation to the above-ground flora of grazed and non-grazed sub-Mediterranean deciduous oak forests of NW Greece. Twenty-eight taxa were recorded in the soil seed bank and 83 taxa (70 taxa in plots of seed bank sampling) in the above-ground vegetation. The dominant tree species and many woodland species found in the above-ground vegetation were absent from the soil seed bank. Similarity between the soil seed bank and the above-ground vegetation decreased with grazing, and grazing led to a decrease of species richness in above-ground vegetation and soil seed bank. Beta diversity of vegetation among grazed and among non-grazed plots did not differ, but was significantly higher between grazed and non-grazed areas. Beta diversity of the soil seed bank declined with grazing. When applying classification tree and logistic regression analyses, non-grazed forest sites are clearly differentiated by the presence of Phillyrea latifolia, Euphorbia amygdaloides and Brachypodium sylvaticum. PCA ordination of above-ground species composition reflected a gradient from sites grazed by ruminants to non-grazed sites, but no clear structure was detected in the seed bank.     

Lichens — the biodiversity value of western woodlands

Summary

The Atlantic broad-leaved woodlands of Britain are of international renown for their lichen floras. They are inhabited by 517 lichens, representing 28.3% of the total lichen flora and 73.2% of all British woodland lichens, and they are the main habitat for 165 species. Of these, 31 have a marked southern distribution and do not reach Scotland, whereas 26 species are found in Scotland, but not England or Wales. Their British Red-listed species are outnumbered by the 86 species for which Britain has International Responsibility.

Within the Atlantic broad-leaved woodlands, only 30 lichens show a preponderance for oak. With the exception of some ancient oakwoods in southern England, a high lichen biodiversity is rarely dependent on a dominance of oak in the woodland canopy, more usually it is the result of a long ecological continuity, often a varied tree and shrub composition, a varied canopy density, and good air quality. Consequently, the oak stands within former ‘industrial’ woodlands have a much lower lichen biodiversity compared with woodlands that have a history as ‘pasture woodland’ or, as with some ravine woodlands, have otherwise escaped intensive management.

The life-history of an oak tree is considered in relation to the niches it provides for lichen colonisation with time. Some management scenarios are provided with the enhancement of the lichen interest of former ‘industrial’ oakwoods as an objective.     

Canopy tree mortality depends on the proportion of crown exposed to sunlight,but this effect varies with species' wood density

Understanding what drives changes in tree mortality as well as the covariates influencing trees' response is a research priority to predict forest responses to global change. Here, we combined drone photogrammetry and ground-based data to assess the influence of crown exposure to light (relative to total crown area), growth deviations (relative to conspecifics), tree size, and species' wood density (as a surrogate for light-demanding and shade-tolerant life-history strategies) on the mortality of 984 canopy trees in an Amazon terra firme forest. Trees with lower wood density were less prone to die when their proportion of crown was more exposed to sunlight, but this relationship with relative crown exposure weakened and slightly reversed as wood density increased. Trees growing less than their species average had higher mortality, especially when the species' wood density decreased. The role of wood density in determining the survival of canopy trees under varying light conditions indicates differential responses of light-demanding versus shade-tolerant species. Our results highlight the importance of accounting for life-history strategies, via plant functional types, in vegetation dynamic models aiming to predict forest demography under a rapidly changing climate. Abstract in Spanish is available with online material.     

Regeneration patterns in relation to canopy species composition and site variables in mixed oak forests in the Sierra de Manantlán Biosphere Reserve, Mexico

Regeneration patterns in relation to canopy species composition and site variables were analyzed in mixed oak forests of the Sierra de Manantlán in western Mexico with the aim of establishing an ecological basis for the design of management alternatives. Using ordination (canonical correspondence analysis) and classification (two-way indicator species analysis) methods, five different canopy types and three different seedling associations were revealed according to species composition, all of them dominated by one or more oak species. Red–far red ratio, slope, altitude, topography, canopy type and grazing intensity were the main variables that explained differences in species composition among the seedling associations. Oak seedlings were relatively scarce in the sampling plots, with the lowest frequency values of all species recorded except for those of Quercus crassipes Humb. & Bonpl., and also the lowest density values. The presence of a particular oak seedling species was strongly associated with a particular percentage of canopy openness; Quercus candicans Née, Quercus laurina Humb. & Bonpl. and Quercus rugosa Née were present in the plots with the least-open canopy (6.4%, 2.9 and 6.2%, respectively), while Quercus castanea Née and Quercus crassipes Humb. & Bonpl. were present in the plots with the most-open canopy (13 and 8.1%, respectively). Every oak seedling species was more frequent, although not dependent, on the canopy type where the same oak species dominated. Because of the great heterogeneity in species composition and the physiographical factors of mixed oak forests in the Sierra de Manantlán, we concluded that management alternatives must be prescribed for each ecological situation where the different oak species are growing.     

Depth distribution and composition of seed banks under different tree layers in a managed temperate forest ecosystem

In the present work we examined the composition and distribution across three soil layers of the buried soil seed bank under three different overstory types (Fagus sylvatica, Quercus robur, Pinus sylvestris) and in logging areas in a 4383-ha forest in central Belgium. The objectives were: (1) to investigate whether species composition and species richness of soil seed banks are affected by different forest stands; (2) to examine how abundant are habitat-specific forest species in seed banks under different planted tree layers. The study was carried out in stands which are replicated, managed in the same way (even-aged high forest), and growing on the same soil type with the same land-use history. In the investigated area, the seed bank did show significant differences under oak, beech, pine and in logging areas, respectively in terms of size, composition and depth occurrence. All species and layers taken together, the seed bank size ranked as follows: oakwood > beechwood > logging area > pinewood. The same pattern was found for forest species. Seed numbers of Betula pendula, Calluna vulgaris, Dryopteris dilatata and Rubus fruticosus were significantly higher under the beech canopy. Carex remota, Impatiens parviflora and Lotus sp. showed a significantly denser seed bank in logging areas, while Digitalis purpurea seeds were significantly more abundant in soils under the oak canopy. The fact that the seed bank of an originally homogeneous forest varies under different planted stands highlights that a long period of canopy conversion can affect the composition and depth of buried seeds.     

TEMPORAL AND SPATIAL PATTERNS OF SPECIFIC LEAF WEIGHT IN SUCCESSIONAL NORTHERN HARDWOOD TREE SPECIES

Temporal and spatial patterns of specific leaf weight (SLW, g/m²) were determined for deciduous hardwood tree species in natural habitats in northern lower Michigan to evaluate the utility of SLW as an index of leaf photosynthetic capacity. No significant diurnal changes in SLW were found. Specific leaf weight decreased and then increased during leaf expansion in the spring. Most species, especially those located in the understory, then had relatively constant SLW for most of the growing season, followed by a decline in SLW during autumn. Specific leaf weight decreased exponentially down through the canopy with increasing cumulative leaf area index. Red oak (Quercus rubra), paper birch (Betula papyrifera), bigtooth aspen (Populus grandidentata), red maple (Acer rubrum), sugar maple (A. saccharum), and beech (Fagus grandifolia) generally had successively lower SLW, for leaves at any one level in the canopy. On a given site, comparisons between years and comparisons of leaves growing within 35 cm of each other showed that differences in SLW among species were not due solely to microenvironmental effects on SLW. Bigtooth aspen, red oak, and red maple on lower-fertility sites had lower SLW than the same species on higher-fertility sites. Maximum CO₂ exchange rate, measured at light-saturation in ambient CO₂ and leaf temperatures of 20 to 25 C, increased with SLW. Photosynthetic capacities of species ranked by SLW in a shaded habitat suggest that red oak, red maple, sugar maple, and beech are successively better adapted to shady conditions.     

The important role of scattered trees on the herbaceous diversity of a grazed Mediterranean dehesa

Scattered trees are considered keystone structures and play an important role in Mediterranean sylvopastoral systems. Such systems are associated with high biodiversity and provide important natural resources and ecosystem services. In this study, we measured the contribution of scattered trees and different grazing management (cattle, sheep and wildlife only) to the diversity of the grassland sward in a dehesa (open holm oak woodland) located in Central Spain. We analyzed alpha and beta diversity through measurement of species richness, Shannon-Wiener, and Whittaker indices, respectively; and the floristic composition of the herb layer using subplots within two adjacent plots (trees present vs. trees absent) under three different grazing management regimes, including wildlife only, during a year. We found a 20–30% increment in the alpha diversity of wooded plots, compared to those without trees, regardless of grazing management. All beta indices calculated showed more than 60% species turnover. Wooded plots were occupied by different herbaceous species in different heterogeneous microsites (under the canopy, in the ecotone or on open land) created by the trees. Livestock grazing modified species composition (e.g. more nitrophilous species) compared to wildlife only plots. In addition to all their other benefits, trees are important to maintaining grassland diversity in Mediterranean dehesas.     

Differential effects of host plant hybridization on herbivore community structure and grazing pressure on forest canopies

Interspecific hybridization in plants is known to have ecological effects on associated organisms. We examined the differences in insect herbivore community structure and grazing pressure on tree canopy leaves among natural hybrids and their parental oak species. We measured leaf traits, herbivore community structure, and grazing pressure on leaves of two oak species, Quercus crispula and Q. dentata, and their hybrids. The concentration of nitrogen in canopy leaves was greater in hybrids and in Q. dentata than in Q. crispula. The concentration of total phenolics was lower in hybrids than in Q. crispula. The concentration of condensed tannin was greater in hybrids than in Q. crispula. Relative herbivore abundance and species richness were greater on oak hybrids than on either parental species; herbivore species diversity and composition on hybrids were close to those on Q. crispula. Herbivore grazing pressure was lower on hybrids and Q. dentata than on Q. crispula. There was a negative correlation between herbivore grazing pressure and leaf nitrogen, suggesting that interspecific variation among oak taxa in herbivore pressure may be explained by leaf nitrogen; variation in herbivore community structure among oak taxa is likely to be controlled by polygenic leaf traits. Differing responses of (1) herbivore community structure and (2) herbivore grazing pressure to host plant hybridization may play important roles in regulating herbivore biodiversity in cool‐temperate forest canopies.     

Patterns and drivers of lichen species composition in a NW-European lowland deciduous woodland complex

We surveyed lichens in an extensive area of lowland deciduous oak and beech dominated forest under wood pasture management within the New Forest National Park, southern England. This provided the basis for an investigation of factors affecting the species density and composition of epiphytic lichen communities. Fifteen 1 hectare plots were established in the same sites as a parallel invertebrate survey, of which nine were in old growth forest and six in oak plantations of between c. 150 and 300 years old. In each plot 12 trees were identified for sampling and lichens were sampled on four aspects of each trunk. Results at the plot level showed that species density was significantly higher in the old growth woodland plots from those in plantations. Plot age had a significant effect on species on oak but this was not significant for lichen communities on beech. The species density of lichens associated with Trentepohliaceae photobionts on oaks showed a linear positive relationship with increasing plot age while the species density of species with other coccal green Chlorophyta decreased with age of the plot. A major part of the lichenised fungi with Trentepohliaceae photobionts includes indicator and notable species used in conservation evaluation and this component is most affected by fragmentation and isolation of forest sites. The results emphasise the importance of long term connectivity and ecological continuity in this extensive mosaic of lowland deciduous forest.     

Cattle exclosure and vegetation dynamics in an ancient,Irish wet oakwood

The effects of a 10-year period of cattle exclosure on the ground flora and tree regeneration of an Irish ancient lowland wet oakwood (Corylo-Fraxinetum deschampsietosum) community are assessed. The approach was to record changes in a quadrat sample located in a National Nature Reserve and to model change by multivariate statistical analysis using Principal Components Analysis and Redundancy Analysis ordination. Exclosure gave significant increases in the abundance of regenerating ash Fraxinus excelsior and holly Ilex aquifolium. Non-native tree species establishment also occurred. In the ground flora, there were significant decreases in the abundance of ruderal species and grasses and significant increases in graze-resistant and graze-sensitive species usually restricted to broadleaf woodland habitats. Changes were similar under low and high canopy sites, but greater under a high canopy. In Irish lowland wet oakwood, managed historically to favour oak and currently grazed by cattle, exclosure promotes the succession of ash to the canopy and holly to the understorey. With time, this is likely to lead to a shift from oak dominance. In a conservation context this, together with an increased risk of non-native tree species establishment, can be ecologically damaging, particularly in small sites without natural dynamics, as is the case in the hedged agricultural landscapes of western Europe. We conclude that site-based conservation objectives should be prioritised when implementing generic, landscape-scale management strategies, such as rotational exclosure.     

Field study of beech (t Fagus sylvatica L.) and melojo oak (t Quercus pyrenaica Willd) leaf litter decomposition in the centre of the Iberian Peninsula

The effect of the canopy on leaf decomposition of beech (t Fagus sylvatica) and melojo oak (t Quercus pyrenaica) was studied during a period of 660 days in a mountain forest of central Spain; response of leaves to leaching was also studied to determine the effects of rainfall after leaf fall. Beech leaves lost 5.8% of their weight by leaching, whereas melojo oak leaves lost 13%. Under both types of canopy, beech leaves showed almost no difference in their decomposition patterns, with decay constants of 0.31 and 0.32 respectively. Melojo oak leaves showed quite a different behaviour under both canopies; decay constant was 0.47 under t Quercus pyrenaica and 0.77 under beech canopy. Total immobilization of nitrogen was less in the melojo oak forest. Effects of summer dryness were sharper in the melojo oak forest, where decomposition stopped during the summer. This delay in the decomposition might have been due to the lower canopy density in the melojo oak forest. The decomposition patterns of the leaves of both species, under the canopy of the other species, suggests what might happen in mixed stands. Thus, the presence of melojo oaks in beech forest would increase decay and decrease nitrogen immobilization. The presence of beech trees in melojo oak forests would improve microclimatic conditions and increase decay, whereas beech litter on the soil would immobilizate more nitrogen.     

Tree size and herbivory determine below-canopy grass quality and species composition in savannahs

Large single-standing trees are rapidly declining in savannahs, ecosystems supporting a high diversity of large herbivorous mammals. Savannah trees are important as they support both a unique flora and fauna. The herbaceous layer in particular responds to the structural and functional properties of a tree. As shrubland expands stem thickening occurs and large trees are replaced by smaller trees. Here we examine whether small trees are as effective in providing advantages for grasses growing beneath their crowns as large trees are. The role of herbivory in this positive tree-grass interaction is also investigated. We assessed soil and grass nutrient content, structural properties, and herbaceous species composition beneath trees of three size classes and under two grazing regimes in a South African savannah. We found that grass leaf content (N and P) beneath the crowns of particularly large (ca. 3.5 m) and very large trees (ca. 9 m) was as much as 40% greater than the same grass species not growing under a tree canopy, whereas nutrient contents of grasses did not differ beneath small trees (<2.3 m). Moderate herbivory enhanced these effects slightly. Grass species composition differed beneath and beyond the tree canopy but not between tree size classes. As large trees significantly improve the grass nutrient quality for grazers in contrast to smaller trees, the decline of the former should be halted. The presence of trees further increases grass species diversity and patchiness by favouring shade-tolerant species. Both grazing wildlife and livestock will benefit from the presence of large trees because of their structural and functional importance for savannahs.     

Plant and arthropod communities in young oak stands: are they determined by site history?

Effect of site history on forest plant and insect communities was studied by comparing afforestations on former agricultural land with reafforestations on ancient woodland sites. Vascular plants, mosses, true bugs, lacewings and saproxylic beetles were surveyed at 18 young broadleaved forest sites dominated by oak (Quercus robur), established between 1986 and 1994 in three different growth regions in Bavaria, Germany. Two strata, near ground level and the canopy, were sampled. Compared to woodland reafforestations greater species density and abundance of plants and true bugs were observed in field layer of afforestation sites. Proportion of forest species among plants and true bugs was however significantly lower in afforestations than on ancient woodland sites. In the canopy, zoo-phytophagous true bugs were significantly better represented in afforestations and zoophagous true bugs in reafforestations. Saproxylic beetles, especially inhabitants of old dead wood, were species-poor in afforestations. Results indicate that site history affects both producer and consumer communities in multiple ways, even 20 years after afforestation of former agricultural land. However, afforestations adjacent to existing forest stands can be regarded as valuable to nature conservation in effectively extending forest habitats. Investment in such afforestation therefore represents more than just an agricultural subsidy. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.