Fungal succession in the early decomposition process of pine cones on the floor ofPinus densiflora forests

The fungal succession on pine cones on the floor ofPinus densiflora forest was investigated in the early decomposition process (within ca. 30% decrease in dry weight). The fungal flora was examined by both washing and surface-sterilization methods on artificially placed cones and naturally fallen cones. The decomposition rates of artificially placed cones were 0.081–0.082 yr^–1. On withered cones still attached to the tree,Pestalotiopsis spp. were dominant. These fungi also occurred with higher frequencies after cones had lain on the floor and on cones in the L and FH horizons.Xylaria sp. andPhomopsis sp., which seem to colonize the interior of the tissue, occurred with higher frequencies on the cones on the tree, but their occurrence frequencies decreased after cones had lain on the forest floor. Conversely,Mortierella spp. andTrichoderma spp. newly occurred or their occurrence frequencies increased on lying cones. Of these,Trichoderma koningii increased rapidly and showed high occurrence frequencies.Thysanophora penicillioides, which prefers coniferous substrates, showed higher occurrence frequencies in the early stages of lying on the forest floor. On cones lying on the floor, the fungal flora did not significantly change during the investigation period.     

Changes in patch mosaics and vegetation structure of rural forested landscapes under shifting human impacts in South Korea

Landscape change in rural ecosystems is a major global issue because they are an important ecological and socio-cultural resource. The aim of this study was to demonstrate the driving forces behind landscape changes. Changes in the patterns and processes of landscape mosaics and vegetation community in two rural areas (Teokdong-ri, TD and Teokseong-ri, TS) in Korea were studied between 1988 and 2002. Quantitative data of vegetation community, landscape mosaics, and local statistics of the two study areas were analyzed. Agricultural efficiency increased even decreasing household population and the agricultural area. In both areas, as the human dependence on natural resources decreased, the patterns and processes of landscape mosaics and forest structure were changed due to changing human impacts on land use. Owing to the abandonment of forest management for 14 years, the spatial heterogeneity of TD significantly increased more than that of TS. In detrended correspondence analysis ordination, floristic compositions of forest patches of two areas were unevenly located on the ordination axes. Pinus densiflora communities of both areas in 1988 were separately located in 2002 as two directions according to management intensity. The species composition of P. densiflora in one direction of 1988 became similar to those of Quercus forests in 2002. This may be because species composition of P. densiflora has been influenced due to changed habitat environments. Finally, we concluded that the development of forest communities and vegetation succession predicted the spatial pattern of future landscape mosaics and due to the vegetation dynamics of rural ecosystems in Korea.Nomenclature: Miyawaki et al. (1994), Lee (1993), and Lee (1998) for identifying plant species.     

Mycofloral succession onPinus densiflora needles on a moder site

Mycofloral succession on decaying pine needles in aPinus densiflora forest on a moder site was investigated in Sugadaira, Nagano Pref., central Japan. Dead needles on the tree, fallen needles obtained from two recognizable sublayers of the L layer and the upper sub-layer of the F₁ layer in the organic horizon were examined for their fungal flora using both washing and surface sterilization techniques. The major interior colonizer in freshly fallen needles varied with the season:Chaetopsina fulva in summer andSelenosporella curvispora in the other seasons.Thysanophora penicillioides was a remarkable external colonizer of freshly fallen needles in summer, while soil fungi were external colonizers of such needles in the other seasons. A possible successional change of major fungi with the needle decay was suggested. The observed seasonal alternation of the species colonizing freshly fallen needles was discussed in relation to climatic conditions. Contributions from Sugadaira Montane Research Center, No. 152.     

Fungal successions on pine needles fallen at different seasons: The succession of surface colonizers

Field experiments were carried out to investigate influences of seasonal change on the fungal succession occurring on the surface of decaying pine needles at a moder site in Japan. At different seasons, the needles fallen for a short period were collected and marked, then placed on the surface of the O horizon. The needles were removed at intervals and their fungal communities were examined by using a washing technique. Unlike the successions of interior colonizers studied at the same time, those of surface colonizers observed on the fallen needles at four different times are roughly similar to each other.Thysanophora penicillioides was the major first colonizer on the sample needles from the O horizon, andTrichoderma species followed it. In an experiment started in late autumn, three dematiaceous fungi,Chloridium viride var.chlamydosporis, Sporidesmium omahutaense, andChalara sp., commonly occurred and contributed to the darkening of colonized needles. Seasonal variation in climate may have a stronger effect on internal colonizers than external colonizers of needles. Contributions from Sugadaira Montane Research Center, No. 165.     

Relationships between ericaceous vegetation and soil nutrient status in a post-fire Kalmia angustifolia-black spruce chronosequence

Post-fire nutrient flushes are an important precursor to secondary succession in fire-driven boreal forest. We studied the magnitude of changes in post-fire soil nutrient status across a chronosequence of ericaceous shrub-dominated boreal forest stands in eastern Newfoundland, Canada. The chronosequence comprised nine stands burned between 1 and 38 years prior to the study. These sites have resisted tree reestablishment following forest fire-induced mortality of black spruce and a concomitant increase in dominance of the ericaceous dwarf shrub Kalmia angustifolia L. Our objectives were: (1) to identify the factors driving soil nutrient status in these post-fire stands dominated by ericaceous plants, and (2) to test hypotheses that specific relationships exist among environmental factors, dominant vegetation and indicators of soil nutrient status. Macronutrients such as NH₄⁺, total organic N and mineral soil P concentrations showed non-linear declines with time since fire. These parameters were also negatively associated with cover of ericaceous plants. Potential phytotoxins such as total phenolics and aluminium concentrations increased with increasing cover of K. angustifolia. Variability in net ammonification, total P and total phenolic acids in organic soils were strongly related to ericaceous dominance even when the effect of time since fire was partialled out using regression analysis. These findings suggest a strong capacity for ericaceous vegetation to have top-down effects on soil chemical property particularly in the organic horizon with the increase in its post-fire dominance.     

Markov models in the study of post-fire succession in heathland communities

Data on the presence of a number of vegetation states (defined in terms of species dominance in areas of 10×10 cm) and transition probabilities were derived from permanent quadrats in a number of recently burned heath stands. Data were taken from a species-rich community, a species-poor type and a high-level Calluna-Eriophorum bog. Simple Markovian models were constructed using these data, and the model predictions were compared with known or expected trends. Models for species-rich heath yielded poor simulations of expected trends since matrices derived from data for the first years after fire did not contain sufficient information on transitions to states important later in the developmental sequence. Model results for the simpler species-poor and bog communities were more satisfactory and simulated expected trends. In these types all species recovered quickly after fire and less rearrangement of species abundances took place. Maximum likelihood statistics carried out on the transition matrices produced inconclusive results for the species-rich and species-poor types, but indicated that the data from the Calluna-Eriophorum bog approximated a first-order time-homogeneous Markov chain. It was concluded that Markov models lack predictive ability except in relatively simple systems, but that they may be useful in illustrating variations in short-term community dynamics.Nomenclature follows Tutin et al. (1964–80) for vascular plants and Smith (1978) for mosses.I am grateful to Prof. C. H. Gimingham for help and advice during this study, which was carried out during tenure of a Natural Environment Research Council studentship. Thanks are also due to Dr M. B. Usher and an anonymous referee for their comments on the draft.     

Forest responses to the large-scale east coast fires in Korea

The east coast forest fires of April 2000 were Koreas largest recorded fires. This, along with the fact that they took place in the region most frequently affected by fire, attracted a great deal of attention. Due to the variations in wind, topography and pre-fire forest stands, a heterogeneous landscape mosaic of burn severity was created across the region. It turned out to be an excellent opportunity to study various landscape-scale impacts of fires on forest dynamics. Therefore, we investigated stands in the 23794ha of burned forest region, in terms of burn severity, vegetation regeneration and forested landscape change as a measure of community stability. Using the geographic information system technique, we analyzed the differential severity and post-fire recovery of pre-fire forest types of different stand age both at stand and species level. Analysis showed that pre-fire vegetation was composed of mainly pine (Pinus densiflora) stands that occupied 70% of the whole forested area, while pine-hardwood and hardwood stands occupied only 28% and 3%, respectively. In addition, two-thirds of all stands were less than 30-years-old. Pine stands were the most severely burned, while conversely pine-hardwood and hardwood stands were less vulnerable. This implied that pine forests had fire-prone characteristics. Vegetation recovery went the opposite way; that is, the regenerating vegetation cover was 71% at pre-fire hardwood stands, and 65% and 53% at pine-hardwood and pine stands, respectively. However, these recovery rates were strikingly fast, considering that investigation took place about 3months after the fires. Fire did not initiate successional processes, but tended to accelerate the predicted successional changes by releasing pre-fire understory species that survived the fires and regenerated by sprouting. The dominant pre-fire tree species (P. densiflora) was susceptible to fire and not resilient enough to reestablish in competition with oak species. Contrary to pines, the abilities of oak species, mainly Quercus mongolica and Q. variabilis, to survive fires and to resprout vigorously made them dominant at most post-fire stands. These shifts in species abundance caused drastic changes to the landscape: from pine-dominated to oak-dominated stands without any notable change in species composition. The patterns in forest regeneration that we observed in Korea may be representative of forest responses to any long-term repeated disturbances, including fire.     

Towards the analysis of vegetation succession

Attention is drawn to canonical analysis as a plausible model for analyzing vegetation succession. An assessment of the opportunities afforded by canonical analysis for this purpose is then made by reference to two applications of the method. The applications deal with characteristics of hydroseral processes and with the dynamic status of an area of lowland tropical rain forest. On the basis of these and other studies the conclusion is drawn that canonical analysis could contribute usefully in efforts to place the study of dynamic ecosystem processes on a more analytic footing.This work was supported by a Natural Sciences and Engineering Research Council award to L. Orlóci. It is a pleasure to acknowledge my indebtedness both to Dr. Orlóci for his support and encouragement and to the Department of Plant Sciences at the University of Western Ontario for hospitality and help. Appreciation is expressed also to Dr. John Ogden, Research School of Pacific Studies, The Australian National University, Canberra for kindly placing his rain forest data at my disposal and for his invaluable help in interpreting the results of analyses of these data.     

Spatial variation of seedling distribution in an east Mediterranean pine woodland at the beginning of post-fire succession

Most of the area in pine woodlands is occupied by perennial seeders that regenerate from seeds in the first winter after the fire and by annuals. Control of the germination in the regenerating vegetation after wildfire is therefore a primary ecological component of the post-fire succession in this ecosystem. The aim of the study presented here was to determine the distribution of Pinus, Cistus and other plants seeds around burned Pinus halepensis trees, and to measure the conditions related to seed germination in the upper soil layers in the same locations. The study was carried out in a 50-year old planted Pinus halepensis woodland that was burned down by a wildfire in July 1995. The variation of seedbank density was determined by collecting samples under the canopies of burned trees and in a nearby open area. Pine seedbank density decreased and that of Cistus and annuals increased with increasing distance from the burned trunks. Most pine seeds were present in the ash layer while those of the other plants were in the soil. In situ germination experiments showed that seedling density decreased with distance from the burned trunks while the proportion of pines in the seedling population increased. This was a result of seedbank variation and germination inhibition by the high pH conditions caused by the ash. The establishment of sparse pine seedling under the dead tree canopies insured their rapid development without interference by other plants and played a key role in the regeneration and stability of the pine woodland community. The concomitant mass germination of the perennial seeders in the rest of the area prevented invasion by annuals.     

北京地区油松人工林不同演替类型空间结构对林下植被及土壤的影响

以北京地区油松(Pinus tabuliformis)人工林不同演替类型林分为研究对象,研究油松纯林、油松-栓皮栎(Quercus variabilis)混交林和栓皮栎纯林三种不同演替类型林分的空间结构、林下植被和土壤水分的变化规律及其相互作用关系。结果表明：(1)林分水平及垂直空间结构、草本层物种多样性、更新幼树生长、土壤持水和透气性能等指标在三种不同演替类型林分间差异显著(P<0.05),林分空间结构参数中的角尺度、林层指数和开敞度显著影响了各类型林分的灌草多样性,混交度、林层指数和大小比数显著影响了更新幼树的生长,混交度和林层指数显著影响了土壤水分的变化(P<0.05)。(2)松栎混交林灌草生物量、天然更新幼树的生长以及土壤水分物理状况均好于纯林,并主要受林分混交度和林层指数的共同作用。(3)各演替类型林分内均存在栓皮栎更新幼树,混交林栓皮栎更新幼树数量最多、长势最好,对林地资源的竞争最为激烈。因此,可以通过调整林分空间结构实现种间关系及林地资源的调控,以充分发挥森林生态系统的各项功能与价值。     

Fungal succession on pine needles in Germany

The mycofloral succession on the needles ofPinus sylvestris was investigated in Tübingen, southwest Germany. Dead needles attached to the branches (D-type), those caught on branches (C-type) and three types of fallen needles, i.e., freshly fallen (L-type), slightly discolored (OL-type) and almost black needles (F-type) were examined for their fungal flora. Common primary saprophytes were rich on the dead needles on the tree, and on the L-type needles. They were replaced by successive species that contained the well-known species preferring pine needles as their substratum, such asVerticicladium trifidum orSympodiella acicola. Their ecological niches in pine leaf litter and their distribution patterns from a biogeographical viewpoint were discussed.     

The dynamics of a succulent karoo vegetation

Strong inference techniques were used in a preliminary study of the structure and dynamics of the vegetation the Prince Albert region of the Great Karoo, South Africa. Seedling emergence studies indicate that open areas in the interstices of the existent vegetation are colonized by mound-building species of the Mesembryanthemaceae. Later, these species serve as sites of establishment for seedlings of several species of woody shrubs. Eventually, the woody shrubs replace the mound-building mesems through interspecific competition. The woody shrubs persist in the community until they reach senescence and die or are removed through overgrazing. Superimposed on this dynamic pattern is a further temporal pattern involving a combination of disturbance and subsequent soil changes. Much of this effect is caused by fossorial animals, which are associated with large circular (10–20 m diameter) mounds. Burrowing by these animals changes the soil characteristics and establishes an additional sequence of vegetation succession in which the mounds serve as refuges from which non-palatable species, such as Pteronia pallens, can invade the rangeland. Later, as the mounds are abandoned and disturbance effects cease, more palatable species, such as P. empetrifolia, will replace P. pallens, overcoming finally the effects of the adjacent mounds on the surrounding vegetation.     

Quantitative analysis of a stand of Pinus densiflora undergoing succession to Quercus mongolica ssp. crispula: 1. A 31-year record of growth and population dynamics of the canopy trees

We describe the population dynamics, growth of dominant species and plant biomass production in an ecosystem undergoing succession from Pinus densiflora dominance to Quercus mongolica ssp. crispula dominance. The diameter at breast height (DBH) was recorded for a period of 31 years. As DBH increased, the tree density decreased. We estimated the net biomass production of the canopy trees, the biomass of the understory and the average amount of litter. These results suggest that the biomass production of an ecosystem depends on its species composition. In order to estimate future ecosystem productivity, it is necessary to predict temporal changes in species composition.     

Chains of opportunity: A Markov model for acquisition of reusable resources

Summary A vacancy chain is a unique type of resource acquisition process composed of an interconnected series of events in which the gaining of a particular resource unit by one individual depends directly on prior acquisition events by other individuals. Taken from the sociological literature, vacancy chains may also describe the distribution of many types of animal resources such as burrows, dwellings and shelters. Using data on hermit crabs, we present a Markov model simulating a vacancy chain process, and test the model against field data. Our results show that a simple Markov model adequately describes shell acquisition in hermit crabs, and that models combining shell size and crude estimates of quality fit the data extremely well. We illustrate in detail how to generate vacancy chain models from ecological data, how to determine the number and size of organisms gaining new resource units from resource introductions of specific sizes, and how to statistically evaluate the accuracy of Markov models. Not recognizing the presence of a vacancy chain system may lead to serious errors in estimating resource dynamics and therefore in demographic and competition models based on these dynamics. Finally, we suggest some ways in which vacancy chain models can aid studies of competition, population dynamics, life histories, and conservation in species using this type of resource acquisition process.     

Temporal and vertical changes in the humus form profile during a primary succession of Pinus sylvestris

The development of the humus form profile during a primary succession of Pinus sylvestris has been studied along chronosequences on dunes and in blow-outs. Attention was given to vertical variation within the humus form and how this changes during profile development. The mor-type ectorganic profile features marked vertical gradients of several soil attributes, while its constituting horizons show no or only small changes of chemical properties during succession. These changes in particular involve increasing calcium and nitrogen concentrations in the organic matter. After an initial high rate of organic matter accumulation in the successive organic horizons, these rates are strongly reduced, suggesting the attainment of a dynamic equilibrium within the time span of the chronosequences on dunes and blow-outs. Blow-outs differ from dunes in the sense that they have a lower amount of organic matter and a higher F/H ratio. This different ratio likely relates to microclimatic conditions less conducive to decomposition.An attempt is made to explain the vertical trends in terms of processes affecting the characteristics of the organic horizons. Main conclusions are that the development of the ectorganic profile results from a combined effect of decay dynamics, rhizosphere processes and atmospheric deposition, which cannot be unentangled quantitatively with the data available. Furthermore, the distinction between F and H horizons has morphological rather than chemical or ecological relevance, as major vertical changes occur within the F horizon.     

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.     

Changes in the seasonal rhythm of two forest communities during secondary succession

Changes in the seasonal rhythm of two plant phytocoenoses in a submountain beech forest during secondary succession were studied. Investigations were done on four monitoring plots with different stand density over the period of four successive years. The rhythm of the associations Dentario bulbiferae-Fagetum and Carici pilosae-Fagetum reflects the course of succession processes running six years after the human impact (cutting) in the ecosystem. Results of the phenological observations of the understorey species with the focus on the changes in flowering and colour spectrum allowed to make the comparisons between both associations in connection with different phyto-climatic conditions and in dependence on time. The most conspicuous changes in the seasonal rhythm and structure of the examined associations were found in conditions of the former clear-cut, currently in succession phase. A clear decrease (56%) in number of taxons with the dominance > 1% in one association towards the end of the 4-year study period was detected here. Simultaneously, a decrease in the number of flowering species was observed, while the relative rate of species being in the vegetative stage increased considerably (from 6 to 67%) over the growing season. The course of flowering of both of the associations missed discernible trends and peaks as well as colour spectra were partially changed during four monitored successive years on the formerly unstocked area.     

Cycling of soil carbon in a Japanese red pine forest I. Before a clear-felling

Cycling of soil carbon was measured synthetically and quantitatively throughout a year in two Japanese red pine forest stands on mid- and foot-slopes at Mt. Takao, Hiroshima Prefecture, west Japan. There was no distinct difference of soil temperature along the slopes, but the soil water content was higher on the foot-slope than on the midslope. The carbon flow (litterfall, soil respiration, etc.) rates were larger on the foot-slope than on the mid-slope, but there was no significant difference of the accumulation of soil carbon (A₀ layer or human in mineral soil) between the areas. The results of the analysis of soil carbon cycling based on a compartment model show that the relative decomposition rate of A₀ layer and humaus in mineral soil increased 1.4–1.5 fold from the mid- to the foot-slopes, corresponding to the soil moisture condition. The relative decomposition rate of A₀ layer was, however, about one-third of that in a evergreen oak forest. This fact suggests that the great resistance of needle litter to decomposition is one of the main limiting factors of the cycling of soil carbon and prevents the fertilization of mineral soil in the pine forest, which was also proven by the simulation of dynamics of soil carbon cycling.     

Role of vegetation in determining carbon sequestration along ecological succession in the southeastern United States

Vegetation plays a central role in controlling terrestrial carbon (C) exchange, but quantifying its impacts on C cycling on time scales of ecological succession is hindered by a lack of long‐term observations. The net ecosystem exchange of carbon (NEE) was measured for several years in adjacent ecosystems that represent distinct phases of ecological succession in the southeastern USA. The experiment was designed to isolate the role of vegetation – apart from climate and soils – in controlling biosphere–atmosphere fluxes of CO₂ and water vapor. NEE was near zero over 5 years at an early successional old‐field ecosystem (OF). However, mean annual NEE was nearly equal, approximately ?450 g C m^?2 yr^?1, at an early successional planted pine forest (PP) and a late successional hardwood forest (HW) due to the sensitivity of the former to drought and ice storm damage. We hypothesize that these observations can be explained by the relationships between gross ecosystem productivity (GEP), ecosystem respiration (RE) and canopy conductance, and long‐term shifts in ecosystem physiology in response to climate to maintain near‐constant ecosystem‐level water‐use efficiency (EWUE). Data support our hypotheses, but future research should examine if GEP and RE are causally related or merely controlled by similar drivers. At successional time scales, GEP and RE observations generally followed predictions from E. P. Odum's ‘Strategy of Ecosystem Development’, with the surprising exception that the relationship between GEP and RE resulted in large NEE at the late successional HW. A practical consequence of this research suggests that plantation forestry may confer no net benefit over the conservation of mature forests for C sequestration.