Effects of soil organic matter and nitrogen supply on competition between Festuca ovina and Deschampsia flexuosa during inland dune succession

We tested the prediction that the successional replacement of plant species during succession on inland sand dunes results from the effects of an increase in nitrogen mineralization on competitive interactions. The growth and competitive strength of Festuca ovina and Deschampsia flexuosa on soil substrates with different amounts of soil organic matter or nitrogen supply were measured. Small tillers of Festuca ovina and Deschampsia flexuosa were grown in monocultures and 1:1-mixtures on soil columns with undisturbed layers of soil organic matter from different successional age. There was (a) no visible soil organic matter, (b) a thin soil organic layer (0.5 cm) and (c) a thicker soil organic layer (6.0 cm) present on the soil columns. The species were also grown on columns with no visible soil organic matter (bare sand) with two different levels of N fertilization to mimic the increased N mineralization in the older successional stages.In monoculture, Festuca produced more biomass on the substrates with a soil organic layer compared to the unfertilized sand substrate. It also produced more biomass on sand substrates with N fertilization. Deschampsia produced more biomass in treatments with a soil organic layer compared to the bare sand treatments, but did not respond to the ammonium-nitrate addition. In competition, Festuca seemed to be the stronger competitor on the unfertilized sand substrate. Festuca was also the better competitor on the N fertilized sand treatments, while on the treatments with a soil organic layer Deschampsia was the winning species. Our results do not support the hypothesis that an increase in N supply is responsible for the replacement of Festuca by Deschampsia that concur with the accumulation of soil organic matter during succession in inland dunes.     

Carbon and nitrogen in soil and vegetation at sites differing in successional age

We studied vegetation and soil development during primary succession in an inland drift sand area in the Netherlands. We compared five sites at which primary succession had started at different moments in the past, respectively 0, 10, 43 and 121 years ago, and a site at which succession had not yet started. In the three younger sites the vegetation was herbaceous, whereas in the two older sites a pine forest had formed. Forest formation was accompanied by the development of an FH-layer in the soil, an increase in the amount of soil organic matter, and an increase in nitrogen mineralisation rate from 1.9 to 18 g N m^–2 yr^–1. Soil moisture content also increased, whereas pH showed a steady decrease with site age. The vegetation changed from a herbaceous vegetation dominated by mosses and lichens and the grass species Corynephorus canescens and Festuca ovina towards a pine forest with an understorey vegetation dominated by Deschampsia flexuosa and, at the oldest site, with dwarf shrubs Empetrum nigrum and Vaccinium myrtillus. At the same time the total amounts of carbon and nitrogen of the ecosystem increased, with a relatively stronger increase of the carbon pool. The establishment of trees during succession greatly affects the dynamics of the ecosystem, especially its carbon dynamics.     

Pattern analysis in successional communities – An approach for studying shifts in ecological interactions

Abstract. Many ecological studies have addressed issues of vegetation spatial patterns in attempts to understand the processes generating them. We investigated changes in ecological processes during succession via the analysis of shrubs’ spatial patterns in a system of linear sand dunes, an arid ecosystem located in the Negev Desert in Israel during three consecutive years. We hypothesized that spatial patterns change from clustered to regular as succession progresses due to changes in the relative importance of facilitation and competition in this environment. In this ecosystem communities of early successional stages are frequently disturbed by high rates of sand movement, whereas in later successional stages sand stability is high. We mapped in the field individual shrubs on high‐resolution aerial photographs, and converted the digital images to a GIS data set. Using Ripley's K‐function we analysed spatial patterns at three levels: the single‐species level, among species and at the individual level, in three communities characterizing different successional stages. In the early successional communities we found clustered spatial patterns, in comparison with stable habitats where spatial patterns tended to be regular. We argue that these shifts in spatial patterns are indicative of the assumption that in this sand‐dune system ecological interactions change from facilitation to competition as succession progresses. Further, we argue that these interactions operate in different spatial scales at the different successional stages, and that the study of these processes should be conducted at the spatial scales specific to each community.     

Lack of tree layer control on herb layer characteristics in a subtropical forest,China

Question: Knowledge of the interaction between understorey herb and overstorey tree layer diversity is mostly restricted to temperate forests. How do tree layer diversity and environmental variables affect herb layer attributes in subtropical forests and do these relationships change in the course of succession? Do abundance and diversity of woody saplings within the herb layer shift during succession? Location: Subtropical broad‐leaved forests in southeast China (29°8′18″‐29°17′29″N, 118°2′14″118°11′12″E). Methods: A full inventory of the herb layer including all plants below 1‐m height was done in 27 plots (10 × 10 m) from five successional stages (<20, <40, <60, <80 and ≥80 yr). We quantified the contribution of different life forms (herbaceous, woody and climber species) to herb layer diversity and productivity and analysed effects of environmental variables and tree layer diversity on these attributes. Results: Herb layer composition followed a successional gradient, as revealed by non‐metric multidimensional scaling (NMDS), but diversity was not correlated to the successional gradient. There was no correlation of diversity across layers. Herb layer productivity was neither affected by tree layer diversity nor by herb layer diversity. Although abundance of woody species in the herb layer decreased significantly during succession, woody species contributed extraordinarily to herb layer species diversity in all successional stages. All environmental factors considered had little impact on herb layer attributes. Conclusions: The subtropical forest investigated displays an immense richness of woody species in the herb layer while herbaceous species are less prominent. Species composition of the herb layer shows a clear successional pattern, however, the presence or absence of certain species appears to be random.     

Effects of landslides on the mountain vegetation of Flores Island,Azores

Question: What are the consequences of frequently occurring landslides on vegetation dynamics, floristic and structural diversity? Location: 39°27′N; 31°13′W – Morro Alto, Flores Island, Azores, Portugal. Methods: Six comparable landslides were selected. Plots were placed at the top, slope and toe of landslides. Data on floristic composition and biovolume, demography and size structure of the dominant tree species (Juniperus brevifolia) were collected. Hierarchical agglomerative clustering and Principal Component Analysis were used in order to identify succession stages and compare succession pathways and vegetation recovery in different parts of the landslides. Results: Four stages of primary succession on substrates formed by landslides were identified: pioneer (Festuca‐Sphagnum grassland), assembly (Juniperus‐Festuca‐Sphagnum open scrub), building (Juniperus‐Sphagnum scrub) and mature (Juniperus‐Sphagnum woodland). Concerning J. brevifolia populations, the succession pathways are independent of location on the landslide. However, at the floristic level, there are some differences, mainly in the pioneer stage at the toes of landslides. Better abiotic conditions, resulting in a higher succession rate, are probably responsible for a faster vegetation recovery on landslide toes. Conclusion: Landslides trigger succession processes that enable a massive regeneration of the dominant tree species and existence of species not present in mature forests. They are also responsible for the simultaneous occurrence of vegetation of different structures. Overall, landslides increase the floristic and structural diversity of the vegetation, consequently increasing landscape heterogeneity.     

毛乌素沙地油蒿群落的循环演替

流动沙地→半流动沙地白沙蒿群落→半固定沙地油蒿＋白沙蒿群落→固定沙地油蒿群落→固定沙地油蒿＋本氏针茅＋苔藓群落→地带性的本氏针茅草原及其迅速沙化的植被发展过程是毛乌素沙地植被自发演替的基本过程。油蒿具有耐沙埋、抗风蚀、耐土壤贫瘠等特性,是该地区最主要的优良固沙植物和重要牧草。半固定、固定沙地池蒿群落在毛乌素沙地的生态环境保护和农牧业生产方面发挥着极其重要的作用,维持其稳定十分重要。然而,其沙化的现     

Temporal development of vegetation and geomorphology in a man-made beach-dune system by natural processes

Twenty-four yrs of primary succession in a man-made beach-dune system at the Baltic coast of Denmark, built of calcareous, marine sand and the dune planted with Ammophila arenaria , was studied by qualitative observations, quantitative records in permanent plots, levelling and soil analysis within a selected area of 0.75 ha. The aim of the study was to discuss whether human initiated and influenced beach and dune dynamics mimick natural beach and dune processes, and to discuss the relationship between succession and zonation. During the study period, which started in 1979, the man-made dune stabilized rather fast. In the stabilized dune, a sand-pararendzina with a thin A-horizon developed. The beach expanded by accretion of less calcareous, marine sand. On the beach new dunes, 3 m high, successively developed. The number of species in the study area increased from 16 to 55, accompanied by species and life form dynamics, characteristic for primary succession on sandy coasts. A gradual change in species composition of the permanent plots, which appeared by a DCA analysis, could by using TWINSPAN be structured into four groups or plant communities, which reflect succession as well as zonation. Two groups represent the vegetation of sandy beaches with annual dicotyledon species, and mobile dunes dominated by rhizomatous geophytes, especially A. arenaria. The two other groups represent stabilized, calcareous dune, dominated by hemicryptophytes, specially Festuca rubra , and less stabilized dune with F. rubra and the invasive alien Rosa rugosa. It was concluded, that the main trends in the geomorphological and vegetational development of the man-made beach-dune system is similar to the development in natural dunes. In the future, further accretion and seaward dune formation may be expected, but is it also expected, that larger parts of the area gradually will be covered by low scrubs of R. rugosa.     

Ants accelerate succession from mountain grassland towards spruce forest

Question: What is the role of mound‐building ants (Lasius flavus) in successional changes of a grassland ecosystem towards a spruce forest? Location: Slovenské Rudohorie Mountains, Slovakia; ca. 950 m a.s.l. near the Obrubovanec point (1020 m a.s.l.; 48°41′N, 19°39′E). Methods: Both chronosequence data along a successional gradient and temporal data from long‐term permanent plots were collected on ants, spruce establishment, and vegetation structure, together with additional data on spruce growth. Results: There are more spruce seedlings on ant mounds (4.72 m^?2) than in the surrounding vegetation (0.81 m^?2). Spruce seedlings grow faster on these mounds compared to surrounding areas. The first colonization wave of seedlings was rapid and probably occurred when grazing prevailed over mowing. Ant colony presence, mound volume, and plant species composition change along the successional gradient. Mounds become bigger when partly shaded but shrink in closed forest, when ant colonies disappear. Shade‐tolerant acidophylic species replace grassland plants both on the mounds and in surrounding areas. Conclusions: The massive occurrence of Lasius flavus anthills contributes to a runaway feedback process that accelerates succession towards forest. The effect of ants as ecosystem engineers is scale‐dependent: although they stabilize the system at the scale of an individual mound, they may destabilize the whole grassland system over a longer time scale if combined with changes in mowing regime.     

Long-term vegetation changes in stands of Pinus sylvestris in southern Finland

Abstract. Understorey vegetation dynamics in Pinus sylvestris L. stands were examined over 30 yr in southern Finland. Data were collected from 112 permanent sample plots twice, first in 1950–1956 and again in 1983 - 1986. Between the two surveys 52 stands were clear-felled and in the 60 remaining old stands thinnings of various magnitude were conducted. In the mature, thinned stands species presence remained virtually unchanged. With the exception of a few species, including Vaccinium vitis-idaea, Hylocomium splendens, and Dicranum polysetum, the changes in species cover were also slight. In the clear-felled plots the changes in species presence were minor but the cover of many species changed strongly. The average cover of Vaccinium myrtillus and Hylocomium splendens dropped to near zero and that of Vaccinium vitisidaea and Pleurozium schreberi decreased 2 - 4 times on all sites. The cover of Calluna vulgaris approximately doubled, and Cladina rangiferina and Cladina arbuscula also increased after canopy removal. In addition to intrinsic successional change, a special interest in this study was to investigate vegetation dynamics in relation to forest land classification methods in which understorey vegetation is used as part of the classification criteria. The results support the Cajanderian site classification approach, in the case of boreal Scots pine stands, that sites can be classified on the basis of understorey vegetation at different successional phases. However, because of the strong successional and site-to-site variation in species cover, it is suggested that site classification be based primarily on species presence     

Primary succession of Bistorta vivipara (L.) Delabre (Polygonaceae) root‐associated fungi mirrors plant succession in two glacial chronosequences

Glacier chronosequences are important sites for primary succession studies and have yielded well‐defined primary succession models for plants that identify environmental resistance as an important determinant of the successional trajectory. Whether plant‐associated fungal communities follow those same successional trajectories and also respond to environmental resistance is an open question. In this study, 454 amplicon pyrosequencing was used to compare the root‐associated fungal communities of the ectomycorrhizal (ECM) herb Bistorta vivipara along two primary succession gradients with different environmental resistance (alpine versus arctic) and different successional trajectories in the vascular plant communities (directional replacement versus directional non‐replacement). At both sites, the root‐associated fungal communities were dominated by ECM basidiomycetes and community composition shifted with increasing time since deglaciation. However, the fungal community's successional trajectory mirrored the pattern observed in the surrounding plant community at both sites: the alpine site displayed a directional‐replacement successional trajectory, and the arctic site displayed a directional‐non‐replacement successional trajectory. This suggests that, like in plant communities, environmental resistance is key in determining succession patterns in root‐associated fungi. The need for further replicated study, including in other host species, is emphasized.     

Succession of secondary forests affected by pine wilt disease in western Japan followed on vegetation maps

Abstract. Succession, changes in the distribution pattern of forest vegetation, and Pinus forest survival following pine wilt disease were clarified based on phytosociological analysis and vegetation maps. Survival of Pinus forests was restricted to the early successional stages, which were located on ridges and the upper part of slopes. Subsequent to pine wilt disease, the succession progressed from early to late substages of Pinus forest, mixed deciduous and evergreen Quercus, to evergreen Quercus forest. Succession occurs in abandoned pine forests which apparently are in a bad state and are vulnerable to attacks by pine wilt disease.     

Monitoring succession from space: A case study from the North Carolina Piedmont

Question: Is the successional transition from pine to hardwood, which has been inferred from chronosequence plots in previous studies, validated through a time line of satellite imagery? Location: Durham, North Carolina, USA. Methods: We examined successional trends in a time‐series of winter‐summer pairs of Thematic Mapper imagery from 1986 to 2000. We calculated the normalized difference of vegetation index (NDVI) for winter and summer, as well as the difference between summer and winter NDVI (i.e., summer increment NDVI). A set of approximately 50 forest stands of known age and phenology were used to interpret patterns in winter and summer increment NDVI over successional time, and a continuum was found to exist between pine‐dominance and hardwood‐dominance. We fitted a series of linear regressions that modeled the change in winter and summer increment NDVI as a function of initial winter and summer increment NDVI, and additional explanatory variables. Results: All regressions were highly significant (P < 0.0001, R²= ca. 0.3). Predicted dynamics are in accord with successional theory, with pixels moving from evergreen dominance to deciduous dominance along a line of fairly constant summer NDVI. A large disturbance event that occurred over the course of this study, Hurricane Fran, appeared to slow rates of succession in the short term (1–3 years), but increase the rate of conversion to hardwoods over longer time spans. Conclusions: We conclude that temporal sequences of remote sensing images provide an excellent opportunity for broad‐scale monitoring of successional processes, and that continuous metrics of that change are essential to accurate monitoring.     

内蒙古锡盟白音锡勒牧场沙地植被的空间分布格局与演替

1985—1986年期间,作者调查了白音锡勒牧场沙地植被。本文具体分析沙地植被的空间分布和动态演替。1．沙地西段的植被呈疏林灌丛草原景观,沙地东段的植被则呈密林灌丛草原景观。由西向东随着海拔高度的上升形成一个植被生态系列。植被PCA分析的结果表明沙地基质稳定性梯度和沙地环境水分梯度是控制沙地植物群落空间分布格局的两个主要环境因素。2．描述了沙地原生植被演替的各个阶段。沙地植被演替阶段与沙地固定过程相一致,群落稳定性与沙地地貌稳定性相关。由于地形分异引起立地水热条件组合变化,植被演替系列分化为阴坡系列和阳坡系列,前者向森林化方向发展,后者则沿草原化方向发展。本文还就沙地植被资源的开发、利用和保护等方面提出6项建议。     

Mobile dunes and eroding salt marshes

The paper deals with general outlines of salt marsh and dune vegetation in the Ellenbogen and Listland area on Sylt (Schleswig-Holstein, FRG). The composition of current salt marsh vegetation is considered to be mainly the result of a long-lasting process of tidal inundation, grazing, and a permanent influence of groundwater seepage from the surrounding dunes. The lower salt marsh communities have shown constancy for 67 years, due to the effect of heavy grazing. The mid-upper salt marsh communities demonstrated a succession from aPuccinellia maritima-dominated community of the lower marsh to aJuncus gerardii-dominated community of the mid-upper salt marsh, which may be due to the transport of sand — over a short time — on the surface of the marsh. The area covered by plant communities of annuals below Mean High Water (MHW) seemed to diminish. Salt marsh soils, especially of the mid-upper marsh, indicate sandy layers resulting from sand drift of the dunes. Dry and wet successional series of the dunes in the Listland/Ellenbogen area both show grassy stages shifting to dwarf shrubs as final stages. White primary dunes can only be found on the accreting shoreline of the Ellenbogen, which is also grazed by sheep; vegetation cover therefore remains dominated by grasses, mosses and lichens. Three mobile dunes (as the most prominent features of this landscape) have been left unaffected by seeding and planting by local authorities. Grazing is considered to be an inadequate tool in nature conservation as long as natural processes are to prevail in the landscape as major determinants.     

Fire and regeneration: the role of seed banks in the dynamics of northern heathlands

Questions: How do species composition and abundance of soil seed bank and standing vegetation vary over the course of a post‐fire succession in northern heathlands? What is the role of seed banks – do they act as a refuge for early successional species or can they simply be seen as a spillover from the extant local vegetation? Location: Coastal Calluna heathlands, Western Norway. Methods: We analysed vegetation and seed bank along a 24‐year post‐fire chronosequence. Patterns in community composition, similarity and abundances were tested using multivariate analyses, Sørensen's index of similarity, vegetation cover (%) and seedling counts. Results: The total diversity of vegetation and seed bank were 60 and 54 vascular plant taxa, respectively, with 39 shared species, resulting in 68% similarity overall. Over 24 years, the heathland community progressed from open newly burned ground via species rich graminoid‐ and herb‐dominated vegetation to mature Calluna heath. Post‐fire succession was not reflected in the seed bank. The 10 most abundant species constituted 98% of the germinated seeds. The most abundant were Calluna vulgaris (49%; 12 018 seeds m^?2) and Erica tetralix (34%; 8 414 seeds m^?2). Calluna showed significantly higher germination the first 2 years following fire. Conclusions: Vegetation species richness, ranging from 23 to 46 species yr^?1, showed a unimodal pattern over the post‐fire succession. In contrast, the seed bank species richness, ranging from 21 to 31 species yr^?1, showed no trend. This suggests that the seed bank act as a refuge; providing a constant source of recruits for species that colonise newly burned areas. The traditional management regime has not depleted or destroyed the seed banks and continued management is needed to ensure sustainability of northern heathlands.     

Vegetation succession on the dunes near Oostvoorne,The Netherlands,since 1934, interpreted from air photographs and vegetation maps

The vegetation succession on the dunes near Oostvoorne, The Netherlands has been followed by means of a novel combination of repeated large-scale vegetation mapping and air photograph interpretation. Vegetation units have been discerned on the formation level because these could be distinguished fairly easily on the photographs and because the rates of change are appropriate to the time interval chosen. Nineteen formations were distinguished. Five 1:6250 maps were constructed, reflecting the formation pattern in 1934, 1943, 1959, 1972 and 1980. An overlay with 2736 grid points at 25 m field distance was used to quantify changes in the formation pattern.The results suggest a pronounced multiple pathway succession with nevertheless three principal trajectories of succession from pioneer to woodland vegetation. The outer dunes, which have originated since 1910, are distinct in successional characteristics from the inner dunes, which already existed but were released from heavy grazing pressure in 1910. The rate of change in the outer dunes was high in the beginning and is slowly decreasing eversince. In the inner dunes it went the other way around. Through visual extrapolation the likely formation patterns in 1910 and in 2000 were estimated.Transition frequencies proved highly variable for most formations. Moreover, strong spatial dependence was found. Limitations in the use of Markov models in cases of long-term succession in heterogeneous environments are discussed.Nomenclature follows the same sources as in van der Maarel et al. (1984).Field work carried out 1980–1981 when the authors were at the Division of Geobotany, University of Nijmegen. We thank Jos Rijntjes, Nijmegen, for his cooperation in the field. Vegetation maps were prepared and calculations performed at Uppsala. We thank the Foundation Het Zuid-Hollands Landschap, Rotterdam, for providing facilities and a grant for fieldwork as well as additional means to reproduce the vegetation maps. Two reviewers gave useful comments.     

Effects of disturbance intensity and frequency on early old‐field succession

Abstract. Early old‐field succession provides a model system for examining vegetation response to disturbance frequency and intensity within a manageable time scale. Disturbance frequency and intensity can interact with colonization and competition to influence relative abundance of earlier and later successional species and determine, respectively, how often and how far succession can be reset. We tested the joint effects of disturbance frequency and intensity on vegetation response (species richness, abundance, canopy structure) during the first six years of succession by clipping the dominant species (D) or all species (T) in spring and fall of each year (S), once per year in summer (Y1), each two years in summer (Y2), or each four years in summer (Y4). Vegetation response reflected disturbance effects on expansion of a later monospecific dominant perennial herb, Solidago altissima, and persistence of the early, richer flora of annuals. A more abundant and taller top Solidago canopy developed on plots clipped each 2 yr or less frequently. Plots clipped yearly or seasonally were richer, but had less abundant, shorter, and differently stratified canopy. Disturbance mediated the relative abundance of early and later successional species; however, frequency and intensity effects were not completely congruent. Persistence of a richer early successional flora increased through the most frequent disturbance (S), and was magnified by disturbance intensity. Disturbance as extreme as clipping all vegetation twice yearly did not cause a drop in species richness, but maintained the early successional community over the first six years of succession. We conclude that clipping disturbance influenced the rate of succession, but the early community could rebound through the range of disturbance frequency and intensity tested.     

Soil seed bank and driftline composition along a successional gradient on a temperate salt marsh

Abstract. This study focuses on the relationship between vegetation succession and soil seed bank composition on the Schiermonnikoog (The Netherlands) salt marsh over 100 yr. The importance of driftline material in seed dispersal and the relationship with succession is also investigated. The results indicate that the majority of species have a transient or short‐term seed persistent bank. Seeds of most species are able to float over the salt marsh and become concentrated in the driftline higher up the marsh. After plants have established a seed bank forms, which disappears when vegetation is replaced by later‐successional species. Exceptions are Spergularia mar‐itima, which is still present in the seed bank of late successional stages, and Juncus gerardi and Glaux maritima, which appear in the seed bank of early successional stages, but are absent in the vegetation. Based on the results of this study constraints and possibilities for salt‐marsh restoration by de‐embankment are discussed.     

Population dynamics of Echinops gmelinii Turcz. at different successional stages of biological soil crusts in a temperate desert in China

• The effects of biological soil crusts (BSC) on vascular plant growth can be positive, neutral or negative, and little information is available on the impacts of different BSC successional stages on vascular plant population dynamics.
• We analysed seedling emergence, survival, plant growth and reproduction in response to different BSC successional stages (i.e. habitats: bare soil, cyanobacteria, lichen and moss crusts) in natural populations of Echinops gmelinii Turcz. in the Tengger Desert of northwest China. The winter annual E. gmelinii is a dominant pioneer herb after sand stabilisation.
• During the early stages of BSC succession, the studied populations of E. gmelinii were characterised by high density, plant growth and fecundity. As the BSC succession proceeded beyond moss crusts, the fecundity decreased sharply, which limited seedling recruitment. Differences in seedling survival among the successional stages were not evident, indicating that BSC have little effect on survival in arid desert regions. Moreover, E. gmelinii biomass allocation exhibited low plasticity, and only reproductive allocation was sensitive to the various habitats. Our results further suggest that the negative effects of BSC succession on population dynamics are primarily driven by increasing topsoil water‐holding capacity and decreasing rain water infiltration into deeper soil.
• We conclude that BSC succession drives population dynamics of E. gmelinii, primarily via its effect on soil moisture. The primary cause for E. gmelinii population decline during the moss‐dominated stage of BSC succession is decreased fecundity of individual plants, with declining seed mass possibly reducing the success of seedling establishment.

     

Changes in two Minnesota forests during 14 years following catastrophic windthrow

Abstract. We measured tree damage and mortality following a catastrophic windthrow in permanent plots in an oak forest and a pine forest in central Minnesota. We monitored changes in forest structure and composition over the next 14 years. Prior to the storm, the oak forest was dominated by Quercus ellipsoidalis, and the pine forest by Pinus strobus. The immediate impacts of the storm were to differentially damage and kill large, early‐successional hardwoods and pines. Subsequent recovery was characterized by the growth of late‐successional hardwoods. In both forests the disturbance acted to accelerate succession. Ordination of tree species composition confirmed the trend of accelerated succession, and suggested a convergence of composition between the two forests.