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

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

The interplay of nurse and target plant traits influences magnitude and direction of facilitative interactions under different combinations of stress and disturbance intensities in Andean dry grassland

Aims Facilitation is a key process in vegetation dynamics, driving the response to natural and anthropogenic pressures. In harsh-grazed systems, palatable plants mainly survive when nested under unpalatable tussocks and shrubs. The magnitude and direction of positive interactions are driven by resource availability, extent of herbivory and type of nurse species. We hypothesized that different combinations of disturbance and environmental stress affect community composition in the dry Puna (southern Peruvian Andes) by modifying nurse types and plant interactions in magnitude and specific associations. We investigated whether different combinations of stress and disturbance influence species richness, type and frequency of occurrence of nurse and beneficiary species and magnitude and patterns of plant interactions; whether nurse species influence these interactions and target species change their interactions under different combinations of stress and disturbance and whether plant functional traits differ in the studied communities and influence the pattern of spatial interactions.Methods We selected three plant communities subject to different precipitation and management regimes: in each we laid a number of transects proportional to its extension. Data collected include species presence/absence, type of spatial interactions with nurse species and functional traits. We calculated species richness and rarefaction patterns, described the patterns of plant–plant spatial interactions and investigated the associations between nurse and other species in the three communities using indicator species analysis (ISA). We performed ISA and correlation analysis to investigate whether plant functional traits influenced facilitative interactions.Important findings We found that different combinations of stress and disturbance shaped a complex set of responses, including changes in the nurse species set. Nurse composition influenced magnitude and direction of plant interactions under different stress intensities. Heavy disturbance increased the relative importance of facilitation, even if the overall number of facilitated species decreased. Under equivalent disturbance regimes, increased abiotic stress led to a greater importance of facilitation. Different combinations of stress and disturbance affected the community assemblage also by changing the behaviour of some non-nurse species. Both heavy disturbance and strong stress led to a decrease of trait states; with certain combinations of stress and disturbance, preferential distribution of these states was observed. We also found that plant traits were of key importance in determining facilitative interactions. Some traits were mainly associated with one type of spatial interaction: plant architecture, life cycle and root type influenced the type of interaction between nurses and beneficiaries under different combinations of stress and disturbance. Our results also demonstrate that in plant interaction research the object of observations (species per se, species percentage, etc.) might influence outputs, and to effectively assess the impact of different stress and disturbance intensities on plant interactions it is necessary to work at the community level to consider the whole species pool.     

Original Articles: Differential Influence of Plant Species on Soil Nitrogen Transformations Within Moist Meadow Alpine Tundra

Plant species can influence nitrogen (N) cycling indirectly through the feedbacks of litter quality and quantity on soil N transformation rates. The goal of this research was to focus on small-scale (within-community) variation in soil N cycling associated with two community dominants of the moist meadow alpine tundra. Within this community, the small-scale patchiness of the two most abundant species (Acomastylis rossii and Deschampsia caespitosa) provides natural variation in species cover within a relatively similar microclimate, thus enabling estimation of the effects of plant species on soil N transformation rates. Monthly rates of soil N transformations were dependent on small-scale variation in both soil microclimate and species cover. The relative importance of species cover compared with soil microclimate increased for months 2 and 3 of the 3-month growing season. Growing-season net N mineralization rates were over ten times greater and nitrification rates were four times greater in Deschampsia patches than in Acomastylis patches. Variability in litter quality [carbon:nitrogen (C:N) and phenolic:N], litter quantity (aboveground and fine-root production), and soil quality (C:N) was associated with three principal components. Variability between the species in litter quality and fine-root production explained 31% of the variation in net N mineralization rates and 36% of net nitrification rates. Site variability across the landscape in aboveground production and soil C:N explained 33% of the variation in net N mineralization rates and 21% of net nitrification rates. Within the moist meadow community, the high spatial variability in soil N transformation rates was associated with differences in the dominant species' litter quality and fine-root production. Deschampsia-dominated patches consistently had greater soil N transformation rates than did Acomastylis-dominated patches across the landscape, despite site variability in soil moisture, soil C:N, and aboveground production. Plant species appear to be an important control of soil N transformation in the alpine tundra, and consequently may influence plant community structure and ecosystem function.     

Structure,dynamics and disturbance regime of temperate broad-leaved forests in Japan

Abstract. Structure, diversity and dynamics of five Japanese temperate old-growth forests were compared, differing from each other in, i.a., climate, dominant tree type, topography, disturbance regime. The forests were Aya, Ogawa, Kanumazawa Riparian, Kanumazawa upland and Senju. A permanent plot (1–6 ha) was established in each forest and trees were censused several times at intervals of two years. Mean annual recruitment rates and mortality rates in these forests were both in the range of 0.5 to 4.6 %/yr at the community level. Analyses of the structure and dynamics of populations showed that the underlying process was different among the forests. Some forests experienced compositional shifts in their canopies, others had a constant canopy composition but appeared to lack effective regeneration in recent years. The recruitment rate appeared to be strongly affected by competitive undergrowth vegetation such as dwarf bamboo which has been controlled by natural disturbance or human impact. It is likely that the forests with mostly low recruitment rates had a low species diversity. The current variation in structure, diversity and dynamics of the studied forests might have been determined not only by physical conditions (e.g. climate) but also by chance factors (e.g. disturbance, outbreak of deer population).     

Drivers of plant individual-based pollinator visitation network topology in an arid ecosystem

Interactions with pollinators underlie the structure and function of plant communities. Network analysis is a valuable tool for studying plant-pollinator interactions, but these networks are most frequently built by aggregating interactions at the species level. Interactions are between individuals and an advantage of individual-based networks is the ability to integrate inter-individual variation in traits and environmental context within complex ecological networks. We studied the influence of inter-individual variation on pollinator sharing among foundation shrubs and cactus in a desert ecosystem using plant individual-based pollinator visitation networks. We hypothesized that the traits that alter attractiveness of plants to pollinators will also influence an individual plant's role within the visitation network. Foundation plants growing with higher densities of nearby blooming shrubs had higher pollinator visitation rates and had greater access to the conspecific mating pool, suggesting widespread and diffuse pollination facilitation within this community. Further, shrub density influenced the role of betweenness centrality and the effective number of partners (e^H). Floral display size also influenced the effective number of interaction partners but did not directly influence the centrality measures for individual plants or other measures of network structure despite increasing visitation rates. The individual-based visitation networks were significantly modular and module membership was predicted by species identity and pollinator visitation rates. Ecological and individual context mediate the outcome of pollinator-mediated interactions and are fundamental drivers of whole community structure. This study shows that the density of immediate neighbours can influence the overall structure of plant-pollinator interaction networks. Exploring the contribution of intraspecific variation to community interaction networks will improve our understanding of drivers of community-level ecological dynamics.     

Beyond the patch: Disturbance affects species abundances in the surrounding community

The role of disturbance in community ecology has been studied extensively and is thought to free resources and reset successional sequences at the local scale and create heterogeneity at the regional scale. Most studies have investigated effects on either the disturbed patch or on the entire community, but have generally ignored any effect of or on the community surrounding disturbed patches. We used marine fouling communities to examine the effect of a surrounding community on species abundance within a disturbed patch and the effect of a disturbance on species abundance in the surrounding community. We varied both the magnitude and pattern of disturbance on experimental settlement plates. Settlement plates were dominated by a non-native bryozoan, which may have established because of the large amount of initial space available on plates. Percent covers of species within the patch were affected by the surrounding community, confirming previous studies' predictions about edge effects from the surrounding community on dynamics within a patch. Disturbance resulted in lower percent cover in the surrounding community, but there were no differences between magnitudes or spatial patterns of disturbance. Disturbance lowered population growth rates in the surrounding community, possibly by altering the abiotic environment or species interactions. Following disturbance, the recovery of species within a patch may be affected by species in the surrounding community, but the effects of a disturbance can extend beyond the patch and alter abundances in the surrounding community. The dependence of patch dynamics on the surrounding community and the extended effects of disturbance on the surrounding community, suggest an important feedback of disturbance on patch dynamics indirectly via the surrounding community.     

Effects of anthropogenic habitat disturbance and Giardia duodenalis infection on a sentinel species' gut bacteria

Habitat disturbance, a common consequence of anthropogenic land use practices, creates human–animal interfaces where humans, wildlife, and domestic species can interact. These altered habitats can influence host–microbe dynamics, leading to potential downstream effects on host physiology and health. Here, we explored the effect of ecological overlap with humans and domestic species and infection with the protozoan parasite Giardia duodenalis on the bacteria of black and gold howler monkeys (Alouatta caraya), a key sentinel species, in northeastern Argentina. Fecal samples were screened for Giardia duodenalis infection using a nested PCR reaction, and the gut bacterial community was characterized using 16S rRNA gene amplicon sequencing. Habitat type was correlated with variation in A. caraya gut bacterial community composition but did not affect gut bacterial diversity. Giardia presence did not have a universal effect on A. caraya gut bacteria across habitats, perhaps due to the high infection prevalence across all habitats. However, some bacterial taxa were found to vary with Giardia infection. While A. caraya's behavioral plasticity and dietary flexibility allow them to exploit a range of habitat conditions, habitats are generally becoming more anthropogenically disturbed and, thus, less hospitable. Alterations in gut bacterial community dynamics are one possible indicator of negative health outcomes for A. caraya in these environments, since changes in host–microbe relationships due to stressors from habitat disturbance may lead to negative repercussions for host health. These dynamics are likely relevant for understanding organism responses to environmental change in other mammals.     

Drought impacts and recovery are driven by local variation in species turnover

In forest systems, drought acts as a disturbance that can generate impacts on community structure and composition at multiple scales. This study focused on a 2-month drought event within an early successional forest system to determine the mechanism(s) of community response to, and recovery from, drought. Drought induced a 28% decline in neighborhood species richness and cover as a result of decreased colonization and increased extinction rates. Following drought, neighborhood richness quickly recovered via increased colonization rates while extinction rates were unaltered. Drought had little long-term effect on neighborhood structure (species richness and cover) and generated only subtle changes in neighborhood composition. Ruderal (annual and biennial) species were more likely to change (increase or decrease) in cover and frequency than the more stress tolerant perennial and woody species. However, population dynamics appeared to be generally driven by stochastic species turnover among fields and not by uniform shifts in species performance across the site. Although drought impacts and recovery appeared predictable at the neighborhood scale, population responses to drought within the site were rather unpredictable. Our findings suggest that stochastic fine-scale processes can generate predictable course-scale dynamics within a disturbed system. The scale-specific mechanisms of community change presented here should be explored in other systems to determine the extent of their generality in driving disturbance impacts on communities.     

Turnover of plants on small islets of the eastern Aegean Sea within two decades

Aim To estimate species turnover of plants on 32 small islands within a 20‐year period and to assess possible changes in community composition and properties, such as species richness and factors affecting it, nestedness, species co‐occurrence and overall community similarity. Additionally, to assess the possible effects of grazing, gull colonies and fire on turnover values. Location Thirty‐two islets in the eastern Aegean Sea (Greece). Methods Complete sampling of plants was performed in 1974 and in 1990–94 (mostly in 1994, which was used as the reference year). Species turnover rates were estimated using both per island and per species approaches. Multiple regression was used to evaluate factors affecting species richness. Chi‐square tests were applied to compare community composition among sampling periods. The effects of various factors on turnover rates and species richness were examined using one‐way anova and ancova . Mann–Whitney tests were applied in order to check for differences between frequencies of occurrence of extinct, immigrant and persisting species. Community nestedness was calculated using bitmatnest and the C‐score index for co‐occurrence was estimated using EcoSim7. Species similarities among islands in each of the 1974 and 1994 data sets were assessed using Jaccard’s index and the two similarity matrices were compared using a Mantel test. Results Of 391 species recorded on the islets, 334 were present in 1974, 301 in 1994 and 244 were common to both these periods. Species richness in the 1974 and 1994 data sets was significantly correlated with elevation and area, but not with distance from the nearest large island. Richness was positively affected by grazing, but not by fire or gull colonies. The slopes of species–area and species–elevation regressions were almost identical in 1974 and 1994. Mean relative turnover was 2.06 (species per islet) and 3.26 (islets per species). Turnover was not correlated with area, elevation or distance from the nearest large island. Nestedness and co‐occurrence levels were very similar. Tables of islet by islet floral similarity (Jaccard’s index) did not differ between the 1974 and 1994 data sets. Main conclusions The turnover rates found are among the highest recorded for plants; at the same time the islet communities exhibit notable stability in overall properties. Our results provide evidence for rapid shifts in species number that may nonetheless be considered as equilibrial dynamics, as these islets are able to respond rapidly to environmental change and disturbance. Human activities, notably the application of grazing, have a significant complicating effect on community dynamics, enhancing observed turnover rates.     

Large grazers modify effects of aboveground-belowground interactions on small-scale plant community composition

Aboveground and belowground organisms influence plant community composition by local interactions, and their scale of impact may vary from millimeters belowground to kilometers aboveground. However, it still poorly understood how large grazers that select their forage on large spatial scales interact with small-scale aboveground–belowground interactions on plant community heterogeneity. Here, we investigate how cattle (Bos taurus) modify the effects of interactions between yellow meadow ants (Lasius flavus) and European brown hares (Lepus europaeus) on the formation of small-scale heterogeneity in vegetation composition. In the absence of cattle, hares selectively foraged on ant mounds, while under combined grazing by hares and cattle, vertebrate grazing pressure was similar on and off mounds. Ant mounds that were grazed by only hares had a different plant community composition compared to their surroundings: the cover of the grazing-intolerant grass Elytrigia atherica was reduced on ant mounds, whereas the relative cover of the more grazing-tolerant and palatable grass Festuca rubra was enhanced. Combined grazing by hares and cattle, resulted in homogenization of plant community composition on and off ant mounds, with high overall cover of F. rubra. We conclude that hares can respond to local ant–soil–vegetation interactions, because they are small, selective herbivores that make their foraging decisions on a local scale. This results in small-scale plant patches on mounds of yellow meadow ants. In the presence of cattle, which are less selective aboveground herbivores, local plant community patterns triggered by small-scale aboveground–belowground interactions can disappear. Therefore, cattle modify the consequences of aboveground–belowground interactions for small-scale plant community composition.     

Dances with fire: Tracking metapopulation dynamics of<Emphasis Type="Italic">polygonella basiramia</Emphasis> in Florida scrub (USA)

The regional persistence of species subject to local population colonization and extinction necessarily depends on how landscape features and disturbance affect metapopulation dynamics. Here, we characterize the metapopulation structure and short-term dynamics ofPolygonella basiramia. This rare, short-lived perennial herb is endemic to Florida scrublands and lacks a seed bank. Fires create the open sand gaps within a shrub matrix that support this species but also kill established plants. Thus, persistence depends on frequent colonization of unoccupied gaps. We are monitoring population dynamics within and among 1204 gaps distributed among 19 shrub patches. Considerable subpopulation turnover is evident at the gap level with rates of gap extinction exceeding rates of colonization in the first year. Whether declines in overall abundance continue is likely to depend on patterns of disturbance and regional stochasticity in this dynamic landscape.Polygonella is more likely to occupy larger and less isolated gaps, demonstrating that landscape features and disturbance strongly affect metapopulation dynamics. BecausePolygonella basiramia displays characteristics, occupancy patterns, and turnover dynamics consistent with metapopulation theory, it represents a model system for studying plant metapopulations.     

Precipitation and elephants,not fire,shape tree community composition in Serengeti National Park,Tanzania

Savannas are spatially diverse, variable and are susceptible to high rates of disturbance from fire and herbivory. There is significant interest in woody cover dynamics in relation to disturbance regimes. Less effort has been devoted to understand processes that drive tree community composition. In this study, tree species composition data collected at the landscape scale in the Serengeti were used to identify key environmental factors driving variation in species composition. A system of 38 plots clustered within 10 sites spanning the mean annual precipitation (MAP) gradient was used to assess the relative role of bottom‐up (precipitation, soil nutrients and soil texture) vs. top‐down factors (fire and elephant herbivory) on tree community composition. We developed candidate models relating tree species composition (based on multivariate community analysis) to different combinations of plot‐level environmental covariates. Results suggest that tree community composition is largely driven by MAP and is associated with elephant population density. Strikingly, we found no evidence that fire influences species compositional turnover. In a second analysis, we used structural equation model (SEM) to explore the possible direction of association between elephant density and tree species composition. We compared a model that included elephant effects on composition to one that included community composition effects on elephant density. Results suggest that variation in elephant population density across space is more likely to drive tree community composition and not vice versa. We propose that precipitation and herbivory, rather than fire, determine tree species composition in Serengeti Acacia tree community.     

Rates of change in tree communities of secondary Neotropical forests following major disturbances

Rates of change in tree communities following major disturbances are determined by a complex set of interactions between local site factors, landscape history and structure, regional species pools and species life histories. Our analysis focuses on vegetation change following abandonment of agricultural fields or pastures, as this is the most extensive form of major disturbance in Neotropical forests. We consider five tree community attributes: stem density, basal area, species density, species richness and species composition. We describe two case studies, in northeastern Costa Rica and Chiapas, Mexico, where both chronosequence and annual tree dynamics studies are being applied. These case studies show that the rates of change in tree communities often deviate from chronosequence trends. With respect to tree species composition, sites of different ages differ more than a single site followed over time through the same age range. Dynamic changes in basal area within stands, on the other hand, generally followed chronosequence trends. Basal area accumulation was more linked with tree growth rates than with net changes in tree density due to recruitment and mortality. Stem turnover rates were poor predictors of species turnover rates, particularly at longer time-intervals. Effects of the surrounding landscape on tree community dynamics within individual plots are poorly understood, but are likely to be important determinants of species accumulation rates and relative abundance patterns.     

Spatial patterns in benthic invertebrate community structure: products of habitat stability or are they habitat specific?

1. The effects of habitat stability on benthic invertebrate community structure were examined at eleven sites (ten streams and a wind-swept lake shore) with similar physicochemical characteristics but differing stability. 2. Habitat characteristics were assessed to place the study sites within the framework of the disturbance-productivity-diversity model, the intermediate disturbance hypothesis and a variant of the habitat templet model to examine their predictions with respect to community structure. 3. Many of the most common invertebrate species were present at all the study site, although their relative abundance and density differed markedly between sites. Thus, while stability did not appear to affect colonization of the study sites by these taxa, it did affect their relative success. 4. Communities at unstable sites were very similar and shared a number of taxa such as Deleatidium, Austrosimulium and several species of chironomid, presumably well-adapted to surviving and recolonizing after flood events. 5. Communities at the stable sites differed markedly, both from each other and the group of unstable sites. The characteristic fauna at each of the stable sites seemed to be a result of the site's intrinsic character and possibly biotic interactions. 6. Although stability was a pervading influence on community structure, acting as a bottleneck to the development of a site-specific suite of taxa, none of the above models could adequately explain the observed patterns.     

Selective gopher disturbance influences plant species effects on nitrogen cycling

Plant species effects on ecosystem processes are mediated by traits such as litter quality and exudation. These same traits also influence the activity and distribution of animals that play key roles in regulating ecosystem dynamics. We planted monocultures of eight plant species commonly found in California grasslands to investigate the relative importance of plant species direct effects on nitrogen cycling, versus their indirect effects mediated by plant interactions with gophers. Plant species differed in their litter C:N ratio, which closely related to species effects on rates of net mineralization and nitrification in undisturbed soil. However, the effect of selective gopher disturbance on N cycling greatly altered these species effects.
Plant species differed in their effects on the type and timing of gopher disturbance. Small feeding holes were formed in late spring in plots containing species with high tissue quality. These feeding holes minimally disturbed the soil and did not alter N cycling rates over the short term. Large gopher mounds were formed in the winter and early spring, primarily in plots containing the grass, Aegilops triuncialis , and to a lesser extent in plots containing Avena barbata . These large mounds significantly disturbed the soil and greatly increased net nitrification rates, but had no consistent effects on net N mineralization. In undisturbed soil, Aegilops had the highest litter C:N ratio and one of the lowest rates of net nitrification. However, gophers preferentially built large mounds in Aegilops plots. Once the effects of gopher burrowing were considered, Aegilops had one of the highest rates of net nitrification, indicating that the indirect effects of plant species on N cycling can be more important than the direct effects alone. This experiment indicates that it is vital to consider interactions between plants and other organisms in order to predict the ecosystem effects of plant communities.     

Fish community reassembly after a coral mass mortality: higher trophic groups are subject to increased rates of extinction

Since Gleason and Clements, our understanding of community dynamics has been influenced by theories emphasising either dispersal or niche assembly as central to community structuring. Determining the relative importance of these processes in structuring real‐world communities remains a challenge. We tracked reef fish community reassembly after a catastrophic coral mortality in a relatively unfished archipelago. We revisited the stochastic model underlying MacArthur and Wilson's Island Biogeography Theory, with a simple extension to account for trophic identity. Colonisation and extinction rates calculated from decadal presence‐absence data based on (1) species neutrality, (2) trophic identity and (3) site‐specificity were used to model post‐disturbance reassembly, and compared with empirical observations. Results indicate that species neutrality holds within trophic guilds, and trophic identity significantly increases overall model performance. Strikingly, extinction rates increased clearly with trophic position, indicating that fish communities may be inherently susceptible to trophic downgrading even without targeted fishing of top predators.     

Reduced aboveground tree growth associated with higher arbuscular mycorrhizal fungal diversity in tropical forest restoration

Establishing diverse mycorrhizal fungal communities is considered important for forest recovery, yet mycorrhizae may have complex effects on tree growth depending on the composition of fungal species present. In an effort to understand the role of mycorrhizal fungi community in forest restoration in southern Costa Rica, we sampled the arbuscular mycorrhizal fungal (AMF) community across eight sites that were planted with the same species (Inga edulis, Erythrina poeppigiana, Terminalia amazonia, and Vochysia guatemalensis) but varied twofold to fourfold in overall tree growth rates. The AMF community was measured in multiple ways: as percent colonization of host tree roots, by DNA isolation of the fungal species associated with the roots, and through spore density, volume, and identity in both the wet and dry seasons. Consistent with prior tropical restoration research, the majority of fungal species belonged to the genus Glomus and genus Acaulospora, accounting for more than half of the species and relative abundance found on trees roots and over 95% of spore density across all sites. Greater AMF diversity correlated with lower soil organic matter, carbon, and nitrogen concentrations and longer durations of prior pasture use across sites. Contrary to previous literature findings, AMF species diversity and spore densities were inversely related to tree growth, which may have arisen from trees facultatively increasing their associations with AMF in lower soil fertility sites. Changes to AMF community composition also may have led to variation in disturbance susceptibility, host tree nutrient acquisition, and tree growth. These results highlight the potential importance of fungal–tree–soil interactions in forest recovery and suggest that fungal community dynamics could have important implications for tree growth in disturbed soils.     

The interaction of disturbances and small mammal community dynamics in a lowland forest in Belize

1. Three floods (July 2000, August 2002, September 2003) and a hurricane (October 2001) that occurred in a lowland forest in the southern Maya Mountains of Belize presented an opportunity to evaluate the influence of these disturbances on the structure of a small mammal assemblage. 2. Four terrestrial and four primarily scansorial/arboreal species were trapped July 2000-March 2005 in six grids over 14 irregularly spaced trapping periods. 3. Community dynamics were characterized more by changes in species composition than changes in diversity. The dynamics were driven by species-specific variation in abundance, with changes in composition generally, but not exclusively, due to the occurrence or disappearance of species at low abundance. Despite the disturbances, species richness remained relatively constant. Evenness within the assemblage was consistently low, primarily as a result of dominance by one species, Heteromys desmarestianus. 4. Effects of flooding on community structure were direct but relatively brief (< 1 year), and varied with the duration and intensity of flooding. Effects from the hurricane were indirect but long-lasting and strongly related to severely reduced food resources. 5. This study suggests that long-term dynamics in the structure of many animal communities in the tropics often results from interactions between direct and indirect effects of disturbance. It also suggests that community resistance will depend on variation in disturbance type and regime, but resilience will be determined by the life-history characteristics of each species.     

Small-scale plant dynamics in temperate Themeda triandra grasslands of southeastern Australia

Abstract. Vegetation subjected to two long-term burning regimes (annual or biennial burning) was studied in permanent plots, at two spatial scales: 0.01 m² and 1 m², to determine the small-scale dynamics of plants in temperate Themeda triandra grasslands of southeastern Australia. Species turnover rates were estimated by presence/absence data while species mobility was assessed using cumulative frequency data. While mean species richness did not fluctuate greatly between years, the vegetation was internally dynamic rather than static. Cumulative species richness increased by 50% at both spatial scales and sites over the 4-yr study period. However, few species became cumulatively frequent (i.e. occurred in 80% of plots in the first and/or subsequent years), suggesting that cumulative species richness increases were due to small- or local-scale movements of plants, rather than ‘shifting clouds’ of species moving across the entire site. The vegetation's dynamics did not differ greatly at sites subject to different (frequent) fire intervals. Species turnover and mobility were individualistic, but the dynamics of many species was greater at the smaller spatial scale: 31–48% of the species present at both spatial scales at the two sites had higher turnover rates at the 0.01 m² scale. Similarly, some ‘non-mobile’ species at the 1-m² scale (i.e. ‘constant’ or ‘local’ mobility types), were more mobile at the smaller-scale. Turnover rate and mobility type were strongly associated with life form in some cases, particularly at the annually-burnt site. In general, therophytes (and to a lesser degree, geophytes) were positively associated with high turnover and mobility in most years, while hemicryptophytes were negatively associated with high turnover in many instances. Hemicryptophytes included many species with a range of mobility types and hence, few significant associations between mobility and this life form were found. The previously unrecognized internal dynamics of this community under ‘stable’ management regimes contributes to species coexistence by allowing plants with different dynamics properties to persist in a spatially and temporally unpredictable manner. Frequent burning is presumably the important component driving much of the non-directional, small-scale dynamics because it regularly destroys individual plants and aerial plant parts and creates opportunities for seedling regeneration, whilst permitting the vegetative persistence and spread of established plants in non-light-limited microsites.     

Influence of temporal heterogeneity in nitrogen supply on competitive interactions in a desert shrub community

Soil nutrients in arid systems are supplied to plants in brief pulses following precipitation inputs. While these resource dynamics have been well documented, little is known about how this temporal heterogeneity influences competitive interactions. We examined the impacts of the temporal pattern of N supply on competitive intensity and ability in an N-limited desert shrub community. At our field site, the three codominant shrubs, Atriplex confertifolia, A. parryi, and Sarcobatus vermiculatus, differ in seasonal growth patterns, with A. confertifolia and S. vermiculatus achieving higher growth rates earlier in the growing season than A. parryi. We predicted that these timing differences in maximum growth rate may interact with temporal variation in N supply to alter competitive abilities over time. Seedlings of the two Atriplex species were planted either individually in field plots or as target plants surrounded by neighbor seedlings. After one year of establishment, the same amount of ¹⁵N was applied to plots either as early spring pulses, mid spring pulses or continuously through the second growing season. Competitive effects were observed under continuous and pulsed N supply. Averaged across all target–neighbor treatments, competitive intensity was ∼1.8-fold greater when N was pulsed compared to when N was supplied continuously, but overall, the outcome of competitive interactions was not influenced by N pulse timing. While the timing of resource supply did not differentially influence the competitive abilities of coexisting species in this system, the temporal pattern of resource supply did alter the intensity of competitive interactions among species. While additional studies in other systems are needed to evaluate the generality of these results, this study suggests that competitive intensity may not necessarily be a direct function of productivity or resource availability as traditionally assumed. Instead, the intensity of competitive interactions in resource-poor systems may depend upon the temporal pattern of resource supply.