Composition,structure and diversity of cove forest stands in the Great Smoky Mountains: a patch dynamics perspective

Abstract. Cove forests of the Great Smoky Mountains are North American examples of old-growth temperate forest. Ecological attributes of seven stands were studied using one 0.6 - 1.0 ha plot per stand. Stand basal area (39 - 55 m²/ha) and biomass (326 - 471 Mg/ha) were high for temperate deciduous forest. Density ranged from 577 to 1075 stems/ha. All stands had a mixture of deciduous canopy species. Only rarely did a single species comprise more than half of the stand by density, basal area or biomass. Shade-intolerant species were present at low levels (1 - 5 % of total stand density). A wide range of stem diameters was characteristic of most species. However, some species lacked small stems, indicating discontinuous regeneration. Stands tended to have 10 - 20 tree species per ha and at least five species had biomass levels > 10 Mg/ha, indicating high evenness. Canopy gaps covered 10 % of the total area (2 - 21 % by stand). Gaps and conspecific patches of canopy trees > 0.05 ha in size were infrequent. Spatial analyses revealed a variety of patterns among species at inter-tree distances of 1 to 25 m. When all species were combined, juveniles showed aggregation, and adults were often hyperdispersed. Analyses for individual species confirmed that the mosaic of canopy species is influenced by non-random spatial processes. Adults of several species were aggregated at distances > 10 m. Juveniles of all major species exhibited aggregation. Several species exhibited regeneration near conspecific adults. This pattern suggested limited mobility for such species within the shifting mosaic. A diverse patchwork resulted despite the fact that many species did not exhibit segregation of adults and juveniles. Further understanding of patch dynamics and the potential for compositional steady state in cove forests requires long-term study with spatial data.     

Variation of carbon and nitrogen stoichiometry along a chronosequence of natural temperate forest in northeastern China

Aims Carbon (C) and nitrogen (N) stoichiometry contributes to understanding elemental compositions and coupled biogeochemical cycles in ecosystems. However, we know little about the temporal patterns of C:N stoichiometry during forest development. The goal of this study is to explore the temporal patterns of intraspecific and ecosystem components' variations in C:N stoichiometry and the scaling relationships between C and N at different successional stages.Methods Along forest development in a natural temperate forest, northeastern China, four age gradients were categorized into ca. 10-, 30-, 70- and 200-year old, respectively, and three 20 m × 20 m plots were set up for each age class. Leaves, branches, fine roots and fresh litter of seven dominant species as well as mineral soil at depth of 0–10 cm were sampled. A Universal CHN Elemental Analyzer was used to determine the C and N concentrations in all samples.Important findings Intraspecific leaf C, N and C:N ratios remained stable along forest development regardless of tree species; while C, N concentrations and C:N ratios changed significantly either in branches or in fine roots, and they varied with tree species except Populus davidiana (P < 0.05). For ecosystem components, we discovered that leaf C:N ratios remained stable when stand age was below ca. 70 years and dominant tree species were light-demanding pioneers such as Betula platyphylla and Populus davidiana, while increased significantly at the age of ca. 200 years with Pinus koraiensis as the dominant species. C:N ratios in branches and fresh litter did not changed significantly along forest development stages. C concentrations scaled isometrically with respect to N concentrations in mineral soil but not in other ecosystem components. Our results indicate that, leaf has a higher intraspecific C:N stoichiometric stability compared to branch and fine root, whereas for ecosystem components, shifts in species composition mainly affect C:N ratios in leaves rather than other components. This study also demonstrated that C and N remain coupled in mineral soils but not in plant organs or fresh litter during forest development.     

Changes of vascular plant diversity along a chronosequence of beech coppice stands,central Apennines,Italy

Abstract

This research sought to understand the patterns of vegetation recovery after disturbances because of coppice management in beech forests. Eighty sampling units from the mountain belt of the Marche region (Apennines, Italy) were collected according to a stratified sampling based on their geological setting (limestone, sandstone), elevation classes and age after last coppicing (to represent a chronosequence, from 1 to 90 years). The expected successional trend of decreasing species richness was confirmed, together with some stabilisation processes for older stand ages. However, more complex diversity patterns were found when total species richness was decomposed into the species richness of five social behaviour types (SBTs), defined based on the species' habitat preference. On both bedrocks, temporal gradients explained the observed diversity changes at the stand level: forest specialists increased whereas non-forest species decreased. A relatively long time after coppicing (40–60 years), the contribution of the beech specialist species doubled, whereas non-forest and species from anthropogenic habitats decreased by about 50%. On sandstone, the contribution of gap species also decreased over a long-period, and the beech stands experienced stronger changes over time. We conclude that the decomposition of total species richness in terms of SBT affords the opportunity to identify temporal references for thresholds which can be used to assess plant diversity status in relation to management schedules and conservation policy decisions.     

间伐对日本黑松海岸林更新的影响

为弄清间伐对海岸林内环境因子 ,进而对海岸林天然更新的影响 ,在黑松海岸林内进行了 4种不同强度的块状间伐试验 .林分间伐后 ,对更新状况和环境因子进行了连续观测 .结果表明 ,间伐可以改善林内光环境、提高土壤的含水量、加强空气流动、促进枯枝落叶的分解等 .4 0年生的黑松海岸林 5 0 %间伐处理后 (密度为 15 0 0株·hm-2 ,林冠开阔度 >30 % )不会对海岸林本身造成风害 ,也不会对防护功能产生较大影响 ,但却能为黑松海岸林天然更新提供良好的生态环境 .试验结果证实了间伐产生的林隙内日本黑松的更新规律 ,提出间伐可作为同龄黑松海岸林向复层、异龄海岸林演化的主要经营措施 .     

Effects of hot ground water on a small swamp-stream in Nova Scotia,Canada

We examined the effects of hot spring water (35–45 °C) on the ecology of a small, intermittant swamp-stream in Nova Scotia, Canada. Temperatures diminished quickly with distance downstream from the hot spring because of abundant inflow of cold ground water (<10 °C), but elevated temperature effects were detectable 130 m downstream. The brook below the hot spring supported a dense mat of the cyanobacterium Oscillatoria (10 m) followed by the green alga Vaucheria taylorii (40 m). Herbaceous vegetation below the algal zones was also altered by the hot water inflow, even where the temperature increase was slight. The structure of the sparse community of benthic invertebrates was sharply different at sites of different stream temperature: only oligochaete worms, ostracods, chironomids, and a single species of snail thrived at the warmest sites; cold downstream sites supported a typical headwater stream community. Mass loss from decomposing leaves of speckled alder was fast at all sites and strongly correlated with water temperature. The changes in community composition and decomposition rate in response to added heat persisted even where the temperature increase was small, indicating a tight coupling of ecosystem structure and function with the physical environment.     

Ecological Risk Assessment of Sediments in Sydney Harbour,Nova Scotia,Canada

Contamination within sediments of Sydney Harbour (once a major industrial port) were evaluated using a multiple lines-of-evidence (LOE) ecological risk assessment (ERA) approach prior to divestiture of the harbor. The multiple LOE approach included: (1) measurement of polycyclic aromatic hydrocarbons (PAHs), polychlorinated biphenyls (PCBs), metals, metalloids, petroleum hydrocarbons(PHCs), and total organic carbon (TOC) concentrations in surface sediments from multiple Sydney Harbour locations; (2) identification and application of sediment quality guidelines (SQGs) from various jurisdictions; (3) comparisons of harbor sediment chemistry against background/reference sediment chemistry; (4) determining number and frequency of exceedances over SQGs; (5) calculating mean probable effect level-quotients (PEL-Qs); (6) PAH forensic source evaluation; (7) review of previous sediment chemistry and biota tissue data; and (8) characterizing benthic habitat at harbor stations. The ERA determined that current sediments exhibited mostly low probability of adverse effects. Furthermore, contaminated sediments exhibiting a high probability of adverse effects were localized to only a few stations within the harbor. Ongoing natural recovery of harbor sediments is likely responsible for attenuating contaminants that historically were higher than those measured in this study and were previously distributed over much wider areas of the harbor. Results suggest that legacy industrial activities and current urban sewage effluents are the major sources of contamination in Sydney Harbour sediments.     

Separating forest continuity from tree age effects on plant diversity in the ground and epiphyte vegetation of a Central European mountain spruce forest

Forest continuity has been identified as an important factor influencing the structure and diversity of forest vegetation. Primary forests with centuries of continuity are usually more diverse than young secondary forests as forest are colonized only slowly and because the former are richer in old tree individuals. In the present study, performed in unmanaged high-elevation spruce forests of the Harz Mountains, Germany, we had the unique opportunity to separate the effects of forest continuity and tree age on plant diversity. We compared an old-growth spruce forest with century-long habitat continuity with an adjacent secondary spruce forest, which had naturally established on a former bog after 1796 when peat exploitation halted. Comparative analysis of the ground and epiphyte vegetation showed that the plant diversity of the old-growth forest was not higher than that of the secondary forest with a similar tree age of >200 years. Our results suggest that a period of >200 years was sufficient for the secondary forest to be colonized by the whole regional species pool of herbaceous and cryptogam forest plants and epiphytes. Therefore, it is likely that habitat structure, including the presence of old and decaying trees, was more important for determining plant diversity than the independent effect of forest continuity. Our results are probably not transferrable to spruce forests younger than 200 years and highly fragmented woodlands with long distances between new stands and old-growth forests that serve as diaspore sources. In addition, our results might be not transferable to remote areas without notable air pollution, as the epiphyte vegetation of the study area was influenced by SO₂ pollution in the second half of the 20th century.     

Woody vegetation structure of xeric forest stands under different edaphic site conditions and disturbance histories in the Biosphere Reserve 'Parque Costero del Sur', Argentina

We studied stand structures and soil properties in an old-growth forest and two 30-yr-old second-growth stands. In the old-growth forest, the total density and basal area average 1566 trees > 1.25 m height ha^-1 and 46.73 m² ha^-1. The dominant trees are Scutia buxifolia and Celtis tala. Using multivariate techniques we distinguished three stands: PS1, dominated by S. buxifolia, is 1000 m far from the river. Its soil is shallow and loamy. PS2 and PS3, co-dominated by S. buxifolia and C. tala, are over 1200 m distant from the river. There the soil is deeper and has thicker texture and higher phosphorus and calcium concentrations than the near-the-river forest soil. Scutia buxifolia shows reverse J-shaped size-distributions and has morphological features of stress-tolerant species. Celtis tala shows oscillating decay size-curves that suggest recruitment pulses related to small gaps and it has morphological features of competitive species. Celtis tala was selectively cut in the past in the second-growth stands SNRD and SRD. The stand SNRD, 1000 m far from the river, is dominated by S. buxifolia. The second species is Schinus polygamus which presents the bell-shaped size-structure of the pioneer species. SNRD does not differ from its old-growth counterpart PS1 in total tree density, basal area and tree branching. The stand SRD, over 1200 m distant from the river, is co-dominated by S. buxifolia and by C. tala trees regenerated from stumps. SRD does not differ from its old-growth counterparts PS2 and PS3 in total tree density and basal area. As to tree branching, it does not differ from PS3, but differs from PS2. All the stands are being invaded by the exotic tree Ligustrum lucidum.The differences between the old-growth stands seem to be related to the gradients of soil texture and nutrient concentrations raising edaphic stress towards the river. In SNRD, the stress, the stress-tolerance of S. buxifolia, and the aptitude of S. polygamus to recruit in disturbed habitats seem to have prevented the post-logging recruitment of C. tala. In SRD, C. tala regenerated possibly due to a better competitive performance in a more favorable site. Under protection the second-growth stands recovered the old-growth quantitative features. We recommend the restoration of the qualitative features and the control of L. lucidum.     

No bottom-up effects of food addition on predators in a tropical forest

Several studies in temperate forests have demonstrated effects of litter addition on decomposers and predators. However, adding litter does not allow separating the effects of food availability and habitat space. We investigated the response of decomposers and predators to increased food resources and space in forests of the southern Mata Atlântica of Brazil. In two forest ecosystems representing an early successional stage of secondary forests and old-growth forest, we added nutrient-rich organic material, artificial litter of no nutritional value, or a combination of both to the soil surface of 120 plots to separate the effects of habitat space and food on soil food webs. We sampled litter- and soil-dwelling arthropods after three months using pitfall traps, soil sample extraction, and sticky traps just above the soil. Adding artificial litter had no positive effect on any of the 17 analyzed arthropod groups. Combining all sampled arthropods the effect was even significantly negative. Adding food had a positive effect on the abundance of decomposers, but not predators. We found no interactions between added artificial litter and added organic material. Our results suggest that the soil fauna in tropical forests is food limited. The lack of a bottom-up effect on predators suggests that they are not predominantly regulated by the abundance of epigeic prey but rather by competition or predation.     

Rapid expansion of lichen woodlands within the closed-crown boreal forest zone over the last 50 years caused by stand disturbances in eastern Canada

Aim Our two main goals are first to evaluate the resilience of the boreal forest according to latitude across the closed‐crown forest zone using the post‐disturbance distribution and cover of lichen woodlands and closed‐crown forests as a metric, and second to identify the disturbance factors responsible for the regeneration and degradation of the closed‐crown forest according to latitude since the 1950s. Location The study area extends between 70°00′ and 72°00′ W and throughout the closed‐crown forest zone, from its southern limit near 47°30′ N to its northern limit at the contact with the lichen woodland zone at around 52°40′ N. Methods Recent (1972–2002) and old (1954–1956) aerial photos were used to map the distribution of lichen woodlands across the closed‐crown forest zone. Forest disturbances such as fire, spruce budworm (Choristoneura fumiferana (Clemens)) outbreak, and logging were recorded on each set of aerial photos. Each lichen woodland and stand disturbance was validated by air‐borne surveys and digitized using GIS software. Results Over the last 50 years, the area occupied by lichen woodlands has increased according to latitude; that is, 9% of the area that was occupied by closed‐crown forests has shifted to lichen woodlands. Although logging activities have been concentrated in the same areas during the last 50 years, the area covered by logging has increased significantly. Outbreaks by the spruce budworm occurred predominantly in the southern (47°30′ N to 48°30′ N) and central (48°53′ N to 50°42′ N) parts of the study area, where balsam fir stands are extensive. In the northern part of the study area (51°–52°40′ N), extensive fires affected the distribution and cover of closed‐crown forests and lichen woodlands. Main conclusions Over the last 50 years, the area occupied by closed‐crown forests has decreased dramatically, and the ecological conditions that allow closed‐crown forests to establish and develop are currently less prevalent. Fire is by far the main disturbance, reducing the ability of natural closed‐crown forests to self‐regenerate whatever the latitude. Given the current biogeographical shift from dense to open forests, the northern part of the closed‐crown forest zone is in a process of dramatic change towards the dominance of northern woodlands.     

The influence of resprouting forest canopy species on richness in Southern Cape forests, South Africa

We investigated the relationship between species richness and numbers and types of individuals and species present in forests with different physiognomies in the southern Cape Province, South Africa. Data were collected from three different ‘plot’ types: 400 m², canopy‐scaled (plot length is directly proportional to canopy height) and per 100 individuals closest to a point. Plots were designed to control for the effect of scale on local richness. Canopy species richness was inversely proportional to the abundance of resprouting species. The strength of the relationship between the abundance of resprouters and canopy species richness increased progressively from the 400 m² plots to the canopy‐scaled plots and finally to the plots of 100 individuals. Resprouter abundance decreased, while canopy species richness increased, with increasing canopy height. Resprouters are able to retain their in situ position in the forests for longer periods of time than do reseeders. This reduces individual and species turnover, thus reducing species richness in resprouter‐dominated forests.     

Comparison of Plankton-Water Chemistry Relationships in Three Acid Stressed Lakes in Nova Scotia,Canada

A field program was conducted in 1980 and 1981 on three lakes in Kejimkujik National Park, Nova Scotia, to study plankton-water chemistry relationships in a region of potential acid precipitation stress. Among lake and between year comparisons were made. The three study sites represent two different types of lakes. Kejimkujik Lake (pH 4.8) and Pebbleloggitch Lake (pH 4.5) are both high-colored, dystrophic acid lakes. Beaverskin Lake is a clear, oligotrophic lake which is not as acidic (pH 5.4). The phytoplankton of Kejimkujik Lake was dominated by diatoms, while Pebbleloggitch Lake was dominated by chlorophytes and chrysophytes, and Beaverskin Lake was dominated by cyanophytes. Kejimkujik Lake had the highest total algal cell volume per liter, and Pebbleloggitch Lake the lowest. Rotifer populations composed the majority of the zooplankton communities, while the crustacean zooplankton were dominated by the acid-tolerant copepod Diaptomus minutus. Kejimkujik Lake had the lowest zooplankton biomass and Pebbleloggitch Lake the highest zooplankton biomass per m³. A pattern of highest algal density in Beaverskin Lake, but highest zooplankton density in Pebbleloggitch Lake emerged in both years. This is likely an effect of abundant, but unsuitable food (blue-green algae) in Beaverskin Lake and important detrital food resources in Pebbleloggitch Lake in both years. Multiple regression analysis of water chemistry variables with plankton species produced many significant effects, but failed to show clear patterns. Cause and effect relationships in aquatic ecosystems are poorly delineated by such techniques.     

Effects of a clearcut on the net rates of nitrification and N mineralization in a northern hardwood forest,Catskill Mountains,New York,USA

The Catskill Mountains of southeastern New York receive among the highest rates of atmospheric nitrogen (N) deposition in eastern North America, and ecosystems in the region may be sensitive to human disturbances that affect the N cycle. We studied the effects of a clearcut in a northern hardwood forest within a 24-ha Catskill watershed on the net rates of N mineralization and nitrification in soil plots during 6 years (1994–1999) that encompassed 3-year pre- and post-harvesting periods. Despite stream NO₃^– concentrations that increased by more than 1400 mol l^–1 within 5 months after the clearcut, and three measures of NO₃^– availability in soil that increased 6- to 8-fold during the 1^st year after harvest, the net rates of N mineralization and nitrification as measured by in situ incubation in the soil remained unchanged. The net N-mineralization rate in O-horizon soil was 1– 2 mg N kg^–1 day^–1 and the net nitrification rate was about 1 mg N kg^–1 day^–1, and rates in B-horizon soil were only one-fifth to one-tenth those of the O-horizon. These rates were obtained in single 625 m² plots in the clearcut watershed and reference area, and were confirmed by rate measurements at 6 plots in 1999 that showed little difference in N-mineralization and nitrification rates between the treatment and reference areas. Soil temperature increased 1 ± 0.8 °C in a clearcut study plot relative to a reference plot during the post-harvest period, and soil moisture in the clearcut plot was indistinguishable from that in the reference plot. These results are contrary to the initial hypothesis that the clearcut would cause net rates of these N-cycling processes to increase sharply. The in situ incubation method used in this study isolated the samples from ambient roots and thereby prevented plant N uptake; therefore, the increases in stream NO₃^– concentrations and export following harvest largely reflect diminished uptake. Changes in temperature and moisture after the clearcut were insufficient to measurably affect the net rates of N mineralization and nitrification in the absence of plant uptake. Soil acidification resulting from the harvest may have acted in part to inhibit the rates of these processes. The US Governments right to retain a non-exclusive, royalty-free license in and to any copyright is acknowledged.     

Spatial structure and diversity of woody plants and ectomycorrhizal fungus sporocarps in a natural subtropical forest

Spatial patterns of ectomycorrhizal fungi, ectomycorrhizal plants, and non-ectomycorrhizal plants were investigated in a natural subtropical forest using second-order analysis. The results of spatial pattern analysis showed that the degree of clumping of woody plants and ectomycorrhizal sporocarps were correlated. There was a significantly positive correlation of relative aggregation indices between ectomycorrhizal fungi and both non-ectomycorrhizal trees and ectomycorrhizal saplings. Correlations between percentage of ectomycorrhizal trees and sporocarp occurrence of ectomycorrhizal fungi and between diversities of woody plants and ectomycorrhizal fungi were distance-dependent or scale-related. A significantly high percentage of ectomycorrhizal trees was found only at relatively short distance from ectomycorrhizal fungal sporocarps, and significantly positive correlation of the diversity between woody plants and ectomycorrhizal fungi was found only at relative long distance, which implied that ectomycorrhizal sporocarps prefer ectomycorrhizal-tree-dominant micro-sites at near distances and at relatively large scales, diverse ectomycorrhizal sporocarps could be found in woodlands with high diversity of woody plants. Important factors affecting the spatial distribution, occurrence, and diversity of ectomycorrhizal fungi include spatial pattern of ectomycorrhizal plants and non-ectomycorrhizal plants, percentage of ectomycorrhizal plants, and plant diversity in a natural forest.     

Disturbance and regeneration dynamics of a mixed Korean pine dominated forest on Changbai Mountain,North-Eastern China

We used dendrochronological methods to study disturbance history of a mixed Korean pine (Pinus koraiensis Siebold et Zuccarini) dominated forest on the northern slope of Changbai Mountain, North Eastern China, over 1770–2000. Frequent small-scale canopy gaps and infrequent medium-scale canopy disturbances dominated natural disturbance regime in the forest, which did not experience stand-replacing disturbances over the studied period. Percentages of growth releases in subcanopy trees were below 6% in most decades, suggesting that disturbances initiating these releases were of low intensity. Strong winds were likely cause of moderate disturbance events. Two episodes with increased disturbance rates (19% and 13%) were dated to the 1920s and 1980s, timing of the 1980s event was consistent with a hurricane occurred in 1986 on the western slope of the Changbai Mountain. Age structure and growth release analyses revealed species-specific regeneration strategies of canopy dominants. Shade-intolerant Olga bay larch (Larix olgensis Henry) recruited mainly before the 1860s. Recruitment of moderately shade-tolerant P. koraiensis occurred as several regeneration waves (1820s, 1850s, 1870–1880s, 1930s, and 1990–2000s) of moderate intensity. Shade-tolerant Jezo spruce (Picea jezoensis Carr. var. komarovii (V. Vassil.) Cheng et L.K.Fu) and Manchurian fir (Abies nephrolepis (Trautv.) Maxim.) regenerated continuously over the last 220 and 130 years, respectively. Enhanced recruitment of P. koraiensis, P. jezoensis, and A. nephrolepis was observed during the 1930s and 1990s, coinciding with increased growth release frequency in the 1920s and 1980s, and suggesting disturbance events of moderate intensity. Our results indicate that the current disturbance regime of the mixed Korean pine dominated forest maintains coexistence of light-demanding and shade-tolerant species and that change in wind climate may be particularly important for future forest composition.