A note on the dynamics of montane virginPicea yezoensis forest in the Greater Khingan Mts,Northeastern China

The vertical growth ofLarix gmelini in the north of the Greater Khingan Mts., north-eastern China is very fast during the first 50 yr. ThoughPinus sylvestris var.mongolica shows a higher growth rate, it will be replaced by the larch population.Picea jezoensis is the most tolerant tree species and will replace all other trees.The fire cycle is 110–120 yr. Most of the montane areas from 800 to 1000 m a.s.l. are not occupied by spruce forest.There is a sapling bank with large numbers of saplings at ages below 20 yr in the understory in the spruce forest. The sapling bank is the key to maintain the steady state of the spruce forest. The persistent sapling bank in the life history of spruce governs the dynamics of the spruce forest controlled by the intensity and frequency of disturbances.The author is H. Q. Wu     

Consequences of cyclic vegetation management for arthropod survival: Simulation experiments

Many rare arthropod species occupy open grasslands. Mowing or grazing is needed to preserve the habitat for these species. Alternatively the vegetation cover in parts of the managed area can be periodically destroyed by ploughing or rototilling. Such treatment results in a dynamic mosaic of habitat patches in different stages of succession. This mosaic may serve as a habitat for many species. However, the interplay between the frequency of rototilling, the spatial structure of the landscape and life history attributes of the animal species will determine the success of such cyclic management strategies. We used a spatially implicit individual-based metapopulation model to examine optimal spatio-temporal management strategies. The model explicitly incorporates succession, population dynamics and dispersal between habitat patches.Optimal management patterns strongly depend on the species’ properties. Our simulation experiments show that in general the more fertile species with strong intraspecific competition are most robust against the large fluctuations of habitat quality resulting from cyclic management. However, the best management strategies found in our experiments cover the requirements of up to 70% of the species tested. Frequent management of relatively small sub-areas minimizes local fluctiations of patch capacity. Though reducing effective patch capacity by more than 50% these strategies may support the survival of the majority of species.     

Spatial modelling of succession-disturbance dynamics in forest ecosystems: Concepts and examples

Over the last few decades it has become increasingly obvious that disturbance, whether natural or anthropogenic in origin, is ubiquitous in ecosystems. Disturbance-related processes are now considered to be important determinants of the composition, structure and function of ecological systems. However, because disturbance and succession processes occur across a wide range of spatio-temporal scales their empirical investigation is difficult. To counter these difficulties much use has been made of spatial modelling to explore the response of ecological systems to disturbance(s) occurring at spatial scales from the individual to the landscape and above, and temporal scales from minutes to centuries. Here we consider such models by contrasting two alternative motivations for their development and use: prediction and exploration, with a focus on forested ecosystems. We consider the two approaches to be complementary rather than competing. Predictive modelling aims to combine knowledge (understanding and data) with the goal of predicting system dynamics; conversely, exploratory models focus on developing understanding in systems where uncertainty is high. Examples of exploratory modelling include model-based explorations of generic issues of criticality in ecological systems, whereas predictive models tend to be more heavily data-driven (e.g. species distribution models). By considering predictive and exploratory modelling alongside each other, we aim to illustrate the range of methods used to model succession and disturbance dynamics and the challenges involved in the model-building and evaluation processes in this arena.     

Three centuries of fire in montane pine‐oak stands on a temperate forest landscape

Question: What was the role of fire in montane pine‐oak (Pinus‐Quercus) stands under changing human land uses on a temperate forest landscape in eastern North America? Location: Mill Mountain in the central Appalachian Mountains, Virginia, US. Methods: A dendroecological reconstruction of fire history was generated for four stands dominated by xerophytic pine and oak species. The fire chronology began under presettlement conditions following aboriginal depopulation. Subsequent land uses included European settlement, iron mining, logging, and US Forest Service acquisition and fire protection. Results: Fires occurred approximately every 5 years until 1930 without any evidence of a temporal trend in fire frequency. Burning ceased after 1930. Area‐wide fires affecting multiple pine stands were common, occurring at intervals of approximately 16 years. Most living pines became established during the late 1800s and early 1900s. Dead pines indicated that an older cohort established ca. 1730. Most hardwoods were established between the 1920s and 1940s. Conclusions: Except for fire protection, changes in land use had no discernible influence on fire frequency. Lightning ignitions and/or large fire extent may have been important for maintaining frequent burning in the 1700s, while fuel recovery may have constrained fire frequency during later periods. The disturbance regime appears to be characterized by frequent surface fires and occasional severe fires, insect outbreaks or other disturbances followed by pine recruitment episodes. Industrial disturbances appear to have had little influence on the pine stands. The greatest impact of industrial society is fire exclusion, which permitted hardwood establishment.     

Investigating the role of impoundment and forest structure on the resistance and resilience of mangrove forests to hurricanes

The present study evaluates how a prolonged artificial flooding regime in impounded mangrove forests influences the regeneration after medium-sized forest destruction created by two hurricanes in 2004 in the Indian River Lagoon, Florida, USA. Disturbance patterns, forest structure, and regeneration were investigated. We found disturbed areas, characterized by uprooted and snapped trees, and intact forests in close proximity. The canopy turnover was greater in forests in higher succession stages with median tree diameter at breast height of 7.6 ± 5.7 cm compared to the intact forest with 3.7 ± 1.2 cm. Larger trees with lower densities were more susceptible to hurricane damage. We observed that regeneration of the open patches was dominated by the flood-tolerant species Rhizophora mangle (89.9%) instead of the faster-growing pioneer species Laguncularia racemosa (7.0%). Some of the disturbed areas created by the hurricanes were not recolonized. The regeneration rate of the disturbed areas expressed by vegetation closure >50% differed between almost zero to 100% in three different impoundments. We concluded that the artificial flooding regime imposed by impoundment is the predominant selective force in the successive process according to the species composition. However, we were not able to detect the driving factors that prevented mangrove establishment in disturbed areas.     

Size distribution and expansion of canopy gaps in a northern Appalachian spruce-fir forest

Canopy gap area/age distributions and growth mechanisms were examined in a virgin subalpine forest in the White Mountains, New Hampshire, USA. The gap area distribution was negative exponential in form. Whithin gap tree ages varied widely in response to stepwise gap expansion caused by windthrow of peripheral trees or death of standing mature Picea rubens at gap edges. As a consequence, the density of small gaps may have been underestimated and the density of large gaps overestimated. The estimates of canopy turnover time, 303 yr, and of patch birth rate on an area basis, 3.3×10^-3 ha new patches/ha land area/yr, were not affected by the gap expansion phenomenon. However, any estimate of patch birth rate as numbers of new patches formed per year would have been too low. Because of increasingly widespread Picea death, the patch area/age distribution of this forest may not currently be in steady-state.     

The latitude-elevation relationship for spruce-fir forest and treeline along the Appalachian mountain chain

Spruce-fir forests extend along the Appalachian Mountains of eastern North America from 35° to 49° N. This montane vegetation differs from boreal spruce-fir forest in that it is dominated by Picea rubens, has a higher vascular species richness, has wind, rather than fire, dominated dynamics, and has a mean annual temperature above 2 °C. Using field reconnaissance, remote sensing, and literature review we described and modeled the latitude-elevation relationship for Appalachian spruce-fir. The elevation of the sprucefir/deciduous forest ecotone decreases from 1,680 m at 35° N to 150 m at 49° N, while the elevation of treeline (spruce-fir/tundra ecotone) decreases from 1,480 m at 44° N to 550 m at 55° N. Linear regressions gave latitude-elevation relationships of –100 m/l^o Latitude for the spruce-fir/deciduous forest ecotone and –83 m/l^o Latitude for treeline. These values compare to literature reports of –54 to –230 m/l^o Latitude and are most similar to values reported from eastern Asia. The latitude-elevation relationship for eman July temperature ( –94 to –121 m/l^o Latitude) was more similar to the slopes of these ecotones than is the slope for mean annual temperature ( –170 to –220 m/l^o Latitude). The spruce-fir/deciduous forest ecotone was correlated with a mean July temperature of approximately 17 °C. Treeline was correlated with a mean July temperature of approximately 13 °C.     

Growth strategies of main trees and forest architecture of aFagus-Magnolia forest in Florida, USA

Growth strategies of six species of trees are compared and used to analyze forest architecture. They included the overstory speciesFagus grandifolia, Magnolia grandiflora, Pinus glabra andLiquidambar styraciflua, and the understory speciesOstrya virginiana andIlex opaca. The six species were abundant in Woodyard Hammock, an old-growth forest in northern Florida, USA. Height, stem diameter, crown projection and radial growth were measured in trees between 5 and 35 m tall. Three different, but non-exclusive, growth strategies were identified for the tree species: survival (Fagus grandifolia, Magnolia grandiflora, Ilex opaca), occupy open space (Fagus grandifolia, Ostrya virginiana, Ilex opaca), and reach above competitors (Liquidambar styraciflua, Pinus glabra). In two transects (0.42 ha) and one quadrat (1 ha), heights of dominant trees were used to distinguish different phases of forest development, which were mapped. In the quadrat, juvenile canopy trees in the undergrowth were mapped. The combination of presence of different developmental phases, presence of juveniles in the undergrowth, growth strategies of main tree species, and disturbance regime was used to assess forest development in the near future.Fagus grandifolia is predicted to become the main dominant species, but the frequent hurricanes open the forest canopy and provide opportunities for understory species (Ostrya virginiana andIlex opaca), and light-demanding overstory species (Liquidambar styraciflua andPinus glabra).     

Wind disturbance in remnant forest stands along the prairie-forest ecotone,Minnesota, USA

Strong winds are an important disturbance agent in northern Minnesota forests. On June 19, 1994, strong winds (>160 km h^(-1)) associated with a tornado damaged forested areas within the Rydell National Wildlife Refuge, situated in Polk County Minnesota along the prairie-forest boundary. Field sampling was conducted immediately following the storm to quantify the type and extent of damage in four different community types, and to project future composition based on the nature of the storm damage and current understory characteristics, including the impact of overbrowsing by deer. Basal area in six sampled remnant forest stands was reduced by 33.5%, although the damage was heterogeneous; basal area in one stand was reduced by 68.1%. The overall effect of the storm was the removal of early- successional species (primarily Populus tremuloides) in larger size classes. Trees situated at stand edges were not more susceptible to snapping or uprooting than interior trees. Projections of future stand composition indicate that wind disturbance, unlike other agents of disturbance such as fire, may accelerate succession on the Refuge, such that early-successional stands will assume a later-successional character, while Acer-Tilia stands should maintain their late-successional character. Overbrowsing and preferential foraging by deer may significantly alter stand recovery patterns.     

Vegetation responses to burning in a rain forest in Borneo

During the 1997/98 ENSO (El Niño Southern Oscillation) event more than 5 million ha of East Kalimantan, Indonesia burned. Here we quantify the initial stages of regeneration (19982001), both in forest that burned and in unburned controls. Sapling and seedling density and species richness remained significantly lower in burned than in unburned forest and community composition remained substantially different between both forest types throughout the sampling period. The only pronounced edge effect was a significantly higher density of seedlings in the interior of unburned forest. Sapling density increased and seedling density declined in both unburned and burned forest during the four-year study period. In the unburned forest, sapling and seedling species richness remained stable, but sapling species richness declined significantly with time in the burned forest. The pioneer community in the burned forest was, furthermore, characterised by higher growth and recruitment than in the unburned forest but mortality did not differ between both forest types. Differences in environment (burned versus unburned: 2965% of variation explained) and the distance between sample sites (1323% of variation explained) explained substantial amounts of variation in sapling and seedling community similarity. Similarity was, however, only marginally (< 1% explained) related to the edge position and temporal variation (difference among sample events). Our results, four years after the initial burn, indicate that burned forest still differed greatly from unburned forest in terms of density, species richness and community composition. There was also no clear trend of a return to pre-disturbance conditions, which indicates that the burned forest may remain in a severely degraded state for a prolonged period of time.     

Scale and resolution of forest structural pattern

An individual tree-based forest succession model was modified to simulate a forest stand as a grid of contiguous 0.01-ha cells. We simulated a 9 ha stand for 750 years and sampled the stand at 50 yr intervals, outputting structural variables for each grid cell. Principal components analysis was used to depict temporal patterns in forest structure as observed in 0.01 ha samples (individual grid cells). We then resampled the grid using square aggregates of 4 to 100 grid cells as quadrats. Principal component scores recalculated for the aggregates, using the original (0.01 ha scale) scoring matrix, depict the effects of obervational scale on perceived patterns in forest structure. Larger quadrats reduce the apparent variation in forest structure and decrease the apparent rate of structural dynamics. Results support a scale-dependent conceptualization of forest systems by illustrating the qualitative difference in forest dynamics as viewed at the scale of individual gap elements as compared to the larger scale steady state mosaic. The aggregation exercise emphasizes the relationship between these two observational scales and serves as a general framework for understanding scaling relationships in ecological phenomena.     

Effect of treefall gaps on the patchiness and species richness of Neotropical ant assemblages

Natural formation of treefall gaps plays an integral role in the ecological and evolutionary dynamics of many tropical forests, affecting the spatiotemporal distribution of plants and the animals that interact with them. This study examines the impact of treefall gaps on the spatial and temporal patchiness of ant assemblages in a moist lowland forest in Panama. Using pitfall traps and honey baits, we compared ant assemblages in five 1 to 2-year-old treefall gaps (ca 100 m²) and five adjacent plots (ca 100 m²) in undisturbed forest understory at three different times of year (late wet season, late dry season, and early wet season). We found little evidence that ant assemblages respond dramatically to the formation of treefall gaps and the differences in habitat qualities they produce. Ant abundance, species richness, species composition, and rates of resource discovery did not differ between gaps and forest understory. However, we did find significant differences in numerical abundance related to forest stratum (ground vs vegetation) and resource type in pitfall traps (oil-cockroach vs honey), and significant differences in ant species richness and rates of resource discovery across seasons. While habitat effects by themselves were never statistically significant, habitat and seasonal differences in species richness interacted significantly to produce complex, season-dependent differences among gap and forest habitats. These results suggest that the formation of natural treefall gaps has less of an effect on Neotropical ant assemblages compared to other groups of organisms (e.g., plants, birds) or other causes of patchiness (e.g., ant mosaics, moisture availability, army ant predation). The results of our study also have important implications for the underlying causes of habitat differences in the distribution of ant-defended plants. Received: 3 February 1998 / Accepted: 7 April 1998     

Patch dynamics of coral reef macroalgae under chronic and acute disturbance

The patch dynamics (colonisation rate, growth rate, and extinction rate) are quantified for two dominant species of macroalgae on a Caribbean forereef in Belize: Lobophora variegata (Lamouroux) and Dictyota pulchella (Hörnig and Schnetter). Measurements were taken on time scales of days, weeks, months, and years during which three hurricanes occurred. All patches were followed on naturally occurring ramets of dead Montastraea annularis. The first hurricane (Mitch) caused massive coral mortality and liberated space for algal colonisation. The cover of Lobophora increased throughout the study and herbivores did not appear to limit its cover within a 4 year time frame. In contrast, the cover of D. pulchella fluctuated greatly and showed no net increase, despite an increase in parrotfish biomass and settlement space. Variation in the overall percent cover of an alga is not indicative of the underlying patch dynamics. The steady rise in the cover of Lobophora took place despite a high turnover of patches (12–60% of patches per year). The patch dynamics of Dictyota were slower (7–20%), but a greater patch density and threefold higher lateral growth rate led to greater fluctuations in total cover. The dynamics of algal patches are size-specific such that larger patches are less likely to become extinct during hurricanes.     

The significance of spatial and temporal patterns of algal mat deposition in structuring salt marsh vegetation

Question: Are there hot spots of algal mat deposition in space and time at the marsh scale and, if so, how does this affect the coexistence of a dominant (Spartina anglica) and gap dependent (Salicornia europaea) species? Location: The Rattekaai salt marsh in the Scheldt estuary in the southwestern Netherlands (NW Europe). Methods: Mat cover and the abundance of the gap dependent species Salicornia europaea were monitored at the scale of a marsh. The effects of mat cover on the vegetation structure were studied by applying three mat removal treatments over three growing seasons. Results: The low marsh border was found to be a hot spot of algal mat deposition during the growing season, which had a correlated spatial pattern between two successive years at a 20 m X 20 m scale. The combination of duration, timing and repetition of mat cover determined growth inhibition of the competitive dominant Spartina anglica, and thereby the abundance of subordinates such as Salicornia europaea. Mat cover reduced the storage of carbon reserves in Spartina and our results imply that repetition of non‐lethal mat cover can lead to ‘gap creation’. Gaps gave only temporary habitat to less dominant species since Spartina quickly re‐invaded them. The gap dependent annual Salicornia was most abundant at intermediate levels of disturbance measured as a function of both space and time. Conclusions In addition to disturbance level, the spatial and temporal distribution of disturbance are important in creating and maintaining habitat for gap dependent species. Relatively small disturbances will have a large effect on diversity if the spatial and temporal distribution of the disturbances leads to ‘disturbance hot spots’.     

Stand dynamics of a yellow-poplar (Liriodendron tulipifera L.) forest in the Appalachian Mountains, Virginia, USA

Yellow-poplar (Liriodendron tulipifera L.) is a large, rapidly growing, shade-intolerant tree species common after disturbances on moist sites in the Appalachian Mountains. The species is typically scattered throughout old-growth mesophytic forests, where periodic gap formation creates conditions favorable for yellow-poplar establishment and growth. On abandoned agricultural fields, however, it is common for nearly monospecific forests of yellow-poplar to develop.This study examines stand dynamics of a yellow-poplar forest in western Virginia, USA that was established on agricultural fields abandoned in the late 1940s. Increment cores were collected from yellow-poplar trees growing on exposed ridgetops and in a more sheltered hollow. Tree-ring data show that the forest is even-aged. Tree establishment began about 5 years earlier on the ridgetops than in the hollow. Major ice storms disturbed the forest in 1978 and 1994, with two separate events in 1994. Ice storms disturb forests by depositing heavy loads of freezing rain on trees, breaking or uprooting them. The dendroecological data collected for this study provide little support for the hypothesis that ice storm disturbance promotes the establishment of new yellow-poplar cohorts. However, the data show that radial growth of some trees increased after ice storm disturbance, a pattern that reflects the increased availability of light following disturbance. Radial growth declined in some other trees as a consequence of severe injury during the storms.Radial growth responses following the 1978 ice storm were stronger on the ridgetops than in the hollow, suggesting that tree damage was more severe on the higher, more exposed sites. Growth responses were relatively mild following the storms of 1994, and did not exhibit pronounced topographic variations.     

Patch choice and population size

The distribution of animals between feeding patches has been the subject of considerable theoretical and empirical investigation. When all animals are equal and fitness is well represented by intake rate, the ideal free distribution requires the animals to be distributed in such a way as to equalize intake rate in each feeding patch. We refer to this as the equal rates policy. This approach ignores the effect of stochasticity in the food supply on starvation. It also ignores predation. An alternative approach is based on the assumption that each animal tries to minimize its death rate. An optimal policy now involves making decisions about which patch to use on the basis of the current level of energy reserves. We investigate a simple model of population dynamics in which over-winter mortality is either derived from animals adopting the equal rates policy or the optimal state-dependent policy to decide between two feeding patches. We show that the state-dependent policy results in a larger equilibrium population size than the equal rates policy. This difference can be considerable when the foraging environment is very stochastic. Furthermore, the state-dependent policy may result in a viable equilibrium population when the equal rates policy does not. The equilibrium under the state-dependent policy may be less stable than that under the equal rates policy. We identify conditions under which the state-dependent policy results in approximately equal intake rates on the two feeding patches. Levels of mortality as a result of predation are investigated. We show that, under some circumstances, the proportion of mortality that is due to predation may decrease as the predation pressure increases.     

Recovery of forest understories buried by tephra from Mount St. Helens

To determine the effects of tephra (volcanic aerial ejecta) on forest understory plants, six sites were chosen along a tephra depth gradient (23 to 150 mm) northeast of Mount St. Helens, USA. All sites were in old forests beyond the limits of direct blast damage from the volcanic eruption. At each site, 150 one m² plots were permanently marked; all tephra was removed from 50 of these in 1980. Cover and density of plant species were recorded during 1980, 1981, and 1982.Tephra 23 mm deep had almost no effect on cover and density of vascular plants, and reduced bryophyte cover for only two years. Tephra 45 mm deep destroyed almost all bryophytes. Although damaged by 45 mm tephra, deciduous herbs recovered by 1982, but some evergreen species did not. Tephra 75 mm deep reduced herb cover in 1982 to 32% and density to 26% of that in cleared plots. At two sites with an average tephra depth of 150 mm, almost all herbs were eliminated except in microsites where tephra was thin, but shrub abundance was greatly reduced only where snow had been present during tephra deposition. Almost all cover was contributed by plants established previous to the eruption; seedling cover never exceeded 0.2%. Refugia with thin tephra, resulting from erosion, were vital to the survival of many species, especially bryophytes.Nomenclature of vascular plants follows Hitchcock & Cronquist (1973); moss nomenclature follows Lawton (1971).We thank T. Hill, C. Halpern and B. Smith for field assistance. The USDA Forest Service, and especially J. F. Franklin, facilitated entry into the restricted area around Mount St. Helens. This word was supported by the National Science Foundation, USDA Science and Education Administration, and the U.S. Forest Service Pacific Northwest Forest and Range Experiment Station.