Forest development in canopy gaps in old-growth beech (Nothofagus) forests,New Zealand

Abstract. Tree size and age structures, treefall and canopy gap characteristics, and regeneration responses to treefalls were compared for three stands of old-growth beech (Nothofagus) forest dominated by N. fusca and N. menziesii on the South Island, New Zealand. Treefall gaps (up to 1000 m²) were most often caused by standing trees killed by drought and/or insect attack, or by trees snapped by wind. The causes of gap formation and the size and age distributions of treefall gaps varied between localities because of spatial and temporal differences in the histories of disturbance. At Fergies Bush where drought-related dieback had produced many large gaps with standing dead trees, gaps were generally young. Conversely, at Station Creek, small, old gaps formed by bolesnap dominated the disturbance regime. At Rough Creek, gaps of all ages and sizes were found along with an almost complete fern cover, and abundant shrubs and occasional subcanopy hardwood trees. Although overall patterns of regeneration were unrelated to differences in gap size, the relative abundance of N. fusca and N. menziesii varied between localities according to the seemingly minor differences in forest structure and disturbance history described above. Interpretations of regeneration response to gap parameters, therefore, need to account for differences in disturbance history between sites. Differences in the disturbance history between localities will also influence rates of gap closure, and because closure rates are used to estimate forest turnover times, meaningful comparisons of disturbance regimes for different forest types can only be made if this intersite variability is addressed.     

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     

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     

Succession patterns following soil disturbance in a sagebrush steppe community

Summary A study was begun in 1976 to measure succession patterns following soil disturbance within a sagebrush community in northwestern Colorado. The principal hypothesis was that type of disturbance affects the direction of succession, resulting in different plant communities over time. Successional dynamics were studied through 1988. Four types of soil disturbance resulted in 3 early seral communities: one dominated by grasses, one by annuals, and one intermediate. The annual-dominated communities were opportunistic on these sites, lasting 3–5 years and not determining the direction in which succession proceeded following their replacement. Twelve years after disturbance, 3 communities (one grass-dominated, one shrub-dominated, and one intermediate) occupied the site, the characteristics of which were functions of type of initial soil disturbance. For the period of time covered by this study (12 years), degree of disturbance was found to affect the direction of succession, resulting in different plant communities over time. There were, however, successional characteristics toward the end of the study that suggest that over a longer time period, succession might progress to a single community regardless of type of disturbance.     

Effects of temporal variability of disturbance on the succession in marine fouling communities in northern-central Chile

We investigated the effects of temporal variability in a disturbance regime on fouling communities at two study sites in a northern-central Chilean bay. Fouling assemblages grown on artificial settlement substrata were disturbed by mechanical removal of biomass at different time intervals. Using one single disturbance frequency (10 disturbance events over 5 months) we applied 7 different temporal disturbance treatments: a constant disturbance regime (identical intervals between disturbance events), and 6 variable treatments where both variableness and sequences of intervals between disturbance events were manipulated. Two levels of temporal variableness (low and high, i.e. disturbance events were either dispersed or highly clumped in time) in the disturbance regime were applied by modifying the time intervals between subsequent disturbance events. To investigate the temporal coupling between disturbance events and other ecological processes (e.g. larval supply and recruitment intensity), three different sequences of disturbance intervals were nested in each of the two levels of temporal variableness. Species richness, evenness, total abundance, and structure of communities that experienced the various disturbance regimes were compared at the end of the experiment (15 days after the last disturbance event). Disturbance strongly influenced the community structure and led to a decrease in evenness and total abundance but not species richness. In undisturbed reference communities, the dominant competitor Pyura chilensis (Tunicata) occupied most available space while this species was suppressed in all disturbed treatments. Surprisingly, neither temporal variableness in the disturbance regime nor the sequence of intervals between disturbance events had an effect on community structure. Temporal variability in high disturbance regimes may be of minor importance for fouling communities, because they are dominated by opportunistic species that are adapted to rapidly exploit available space.     

Effects of nitrogen limitation on species replacement dynamics during early secondary succession on a semiarid sagebrush site

Summary A soil nitrogen (N) availability gradient was induced on a disturbed sagebrush site in northwestern Colorado by fertilizing with nitrogen (high available N), applying sucrose (low available N), and applying neither nitrogen nor sucrose (control). Species composition was studied for 3 years. At the end of the study, N concentration of aboveground tissue of 3 major species was determined. The rate of species replacement was most rapid on plots receiving the sucrose treatment and was slowest on plots receiving the N treatment. Early-seral dominats had greater tissue N concentrations when availability of the resource was high but lower tissue N concentrations when available soil N became limited. Midseral dominants displayed the opposite pattern. These results suggest that the supply of available soil N, and therefore the dynamics of N incorporation in perennial plant tissue, is a primary mechanism in controlling the rate of secondary succession within this semiarid ecosystem.     

Interactions between Limonium vulgare and Plantago maritima in the Plantagini-Limonietum on the Boschplaat,Terschelling, The Netherlands

Limonium vulgare and Plantago maritima are growing together over large areas and in varying proportions on the Boschplaat salt marsh. The question is how both species interact and which environmental factors play a role in those interactions.There are indications of different measures of niche differentiation in the Plantagini-Limonietum association related to differences in flooding frequency and substrate. Interactions between both species are described according to three models (related to de Wit's competition model). Mechanisms that may cause niche differentiation are: differences in growth rhythm, in salt tolerance, in rooting depth, in preference to types of substrate, in morphology, in mycorrhizae and diseases.On a small scale (spatial heterogeneity in the marsh and variation in subsequent seasons) Limonium can achieve a bigger portion in the vegetation by occupying open sites faster than Plantago. On a larger scale (the whole marsh, decades) the impossibility of Plantago to germinate and rejuvenate after sedimentation of silt and salinisation of the soil is the determining factor causing an overall decrease of this species (compared with Limonium) and increase of Limonium.The life cycle of both species, criteria for scale in space and time, and the question whether the Plantagini-Limonietum should be considered a syntaxon, are discussed.Nomenclature follows Heukels-van Ooststroom (1977).I like to thank Dr. P. Ketner and Dr. C. J. T. Spitters for their accompaniment and their ideas, so that I was aware of the wood (Boschplaat) for the trees. I also like to thank Drs. A. J. M. Roozen and Prof. Dr. E. van der Maarel for their discussions about this subject on many occasions. Finally I would like to thank Dr. W. G. Beeftink for his ideas and support and Ron Don, Wim van Hooff, Sjef Jansen, Alex Jonker, Annemiek Demon, Wim Heijligers, Marlie Becks and Frans Wilms for providing the basic concept (the Plantagini-Limonietum) and for their help.     

Scale- and intensity-dependent disturbance determines the magnitude of opportunistic response

Opportunistic, fugitive or pioneer species are species that posses life-history characteristics that allow them to respond quickly to disturbances. Their abundance during the early stages of succession is central to ecological models of benthic soft-sediment succession and these species can play important roles in affecting subsequent successional trajectories. Nevertheless, numerous studies have demonstrated seemingly random patterns of opportunistic responses following disturbance, questioning the generality of currently accepted successional models. In this paper we provide examples from two case studies and argue that the spatial scale or magnitude of disturbance is key to the development of opportunistic responses, and that the scale of disturbance may be particularly important in determining (a) the levels of resources made available and (b) the magnitude of release from competitive interactions, which permit opportunists to flourish.     

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.     

Differential effects of lichens,mosses and grasses on respiration and nitrogen mineralization in soils of the New Jersey Pinelands

In the New Jersey Pinelands, severely disturbed areas often do not undergo a rapid succession to forest; rather, a patchy cover of lichens, mosses and grasses persists for decades. We hypothesized that these plant covers affect soil microbial processes in different ways, and that these effects may alter the successional dynamics of the patches. We predicted that the moss and grass covers stimulate soil microbial activity, whereas lichens inhibit it, which may in turn inhibit succession. We collected soil cores from beneath each type of cover plus bare soil within two types of highly disturbed areas—sites subjected to hot wildfires, and areas mined for sand. Organic matter (OM) content, soil respiration and potential N mineralization were measured in the cores. Soils under mosses were similar to those under grasses; they accumulated more OM and produced more mineral N, predominantly in the form of ammonium, than either the bare soils or the soils beneath lichens. Mineralization under lichens, like that of the bare soils but unlike the soils beneath mosses or grasses, was dominated by net nitrification. These patterns were reproduced in experimentally transplanted moss and lichen mats. Mosses appear to create high-nutrient microsites via high rates of OM accumulation and production of ammonium, whereas lichens maintain low-nutrient patches similar to bare soil via low OM accumulation rates and production of mineral N predominantly in the mobile nitrate form. These differences in soil properties may explain the lack of vascular plant invasion in lichen mats, in contrast to the moss-dominated areas.     

The gothic earthflow revisited: a chronosequence examination of colonization on a subalpine earthflow

I examine the process of plant community colonization through a chronosequence study of permanent plots spanning 45 years on a 70-year-old subalpine earthflow in south-western Colorado, USA.While local diversity and average density of vegetation remained stable throughout the study, the overall diversity increased through the early years of the study with little change over the past 20 years. A gradual shift in species composition occurred through time with species with good dispersal and stress tolerance abilities colonizing the site initially followed by more generalist species.After over 70 years the disturbed communities remained distinct from adjacent relatively undisturbed areas in diversity, density, and species composition. Soil translocation experiments suggest that it is the severe microclimate of the earthflow which is limiting further colonization of the site.     

Revegetation following soil disturbance in a California meadow: the role of propagule supply

Revegetation following a disturbance event initially should be constrained by the abundance and types of propagules available at the disturbed site. I tested this idea by conducting two experiments in which I created artificial soil disturbances by excavating or burying pre-existing grassland vegetation. In the first experiment, I varied disturbance intensity (depth), to investigate the consequences for revegetation when numbers of surviving propagules (dormant seeds and bulbs) were altered. In the second experiment, I varied the timing of disturbance, to investigate the consequences when disturbed sites experienced differing exposures to seasonal patterns of clonal growth and seed dispersal. I sampled these experiments from 1991 to 1993, and have interpreted their results using measurements of the seed bank, the bulb bank, and the seed rain. In the first (depth) experiment, bulbs declined in abundance with burial depth and were scarcer in deeper excavations. In contrast, numbers of annual graminoids initially showed no trends with respect to disturbance depth. These results reflect the depth distributions of the seed and bulb banks. Since bulbs occur deeply in the soil, progressively deeper disturbances left fewer survivors. Similarly, perennial graminoids could grow through the shallowest burials. In contrast, since the annual-graminoid-dominated seed bank is concentrated near the soil surface, disturbance depth mattered less to these species: any disturbance removing the surface layer was equally destructive. In the second (timing) experiment, more annual graminoids initially occurred in older plots. This result reflects seasonal patterns of seed production: plots exposed to more of the annual-graminoid-dominated seed rain supported higher densities of annual graminoids as a result. In subsequent years, the vegetation of most plots in both experiments was increasingly dominated by annual graminoids, again as a consequence of their great abundance in the seed rain. These results indicate that interactions between soil disturbances and sources of propagules play an important role in controlling early stages of succession in newly created gaps. They also suggest that disturbance may play different roles in communities characterized by species with different reproductive strategies. Understanding sources of colonists will improve our ability to predict the effects of disturbance.     

Disturbance frequency and community structure in a twenty-five year intervention study

Models of community regulation commonly incorporate gradients of disturbance inversely related to the role of biotic interactions in regulating intermediate trophic levels. Higher trophic-level organisms are predicted to be more strongly limited by intermediate levels of disturbance than are the organisms they consume. We used a manipulation of the frequency of hydrological disturbance in an intervention analysis to examine its effects on small-fish communities in the Everglades, USA. From 1978 to 2002, we monitored fishes at one long-hydroperiod (average 350 days) and at one short-hydroperiod (average 259 days; monitoring started here in 1985) site. At a third site, managers intervened in 1985 to diminish the frequency and duration of marsh drying. By the late 1990s, the successional dynamics of density and relative abundance at the intervention site converged on those of the long-hydroperiod site. Community change was manifested over 3 to 5 years following a dry-down if a site remained inundated; the number of days since the most recent drying event and length of the preceding dry period were useful for predicting population dynamics. Community dissimilarity was positively correlated with the time since last dry. Community dynamics resulted from change in the relative abundance of three groups of species linked by life-history responses to drought. Drought frequency and intensity covaried in response to hydrological manipulation at the landscape scale; community-level successional dynamics converged on a relatively small range of species compositions when drought return-time extended beyond 4 years. The density of small fishes increased with diminution of drought frequency, consistent with disturbance-limited community structure; less-frequent drying than experienced in this study (i.e., longer return times) yields predator-dominated regulation of small-fish communities in some parts of the Everglades. Electronic Supplementary Material Supplementary material is available for this article at