Successional theories

Succession is a fundamental concept in ecology because it indicates how species populations, communities, and ecosystems change over time on new substrate or after a disturbance. A mechanistic understanding of succession is needed to predict how ecosystems will respond to land-use change and to design effective ecosystem restoration strategies. Yet, despite a century of conceptual advances a comprehensive successional theory is lacking. Here we provide an overview of 19 successional theories (‘models’) and their key points, group them based on conceptual similarity, explain conceptual development in successional ideas and provide suggestions how to move forward. Four groups of models can be recognised. The first group (patch & plants) focuses on plants at the patch level and consists of three subgroups that originated in the early 20th century. One subgroup focuses on the processes (dispersal, establishment, and performance) that operate sequentially during succession. Another subgroup emphasises individualistic species responses during succession, and how this is driven by species traits. A last subgroup focuses on how vegetation structure and underlying demographic processes change during succession. A second group of models (ecosystems) provides a more holistic view of succession by considering the ecosystem, its biota, interactions, diversity, and ecosystem structure and processes. The third group (landscape) considers a larger spatial scale and includes the effect of the surrounding landscape matrix on succession as the distance to neighbouring vegetation patches determines the potential for seed dispersal, and the quality of the neighbouring patches determines the abundance and composition of seed sources and biotic dispersal vectors. A fourth group (socio-ecological systems) includes the human component by focusing on socio-ecological systems where management practices have long-lasting legacies on successional pathways and where regrowing vegetations deliver a range of ecosystem services to local and global stakeholders. The four groups of models differ in spatial scale (patch, landscape) or organisational level (plant species, ecosystem, socio-ecological system), increase in scale and scope, and reflect the increasingly broader perspective on succession over time. They coincide approximately with four periods that reflect the prevailing view of succession of that time, although all views still coexist. The four successional views are: succession of plants (from 1910 onwards) where succession was seen through the lens of species replacement; succession of communities and ecosystems (from 1965 onwards) when there was a more holistic view of succession; succession in landscapes (from 2000 onwards) when it was realised that the structure and composition of landscapes strongly impact successional pathways, and increased remote-sensing technology allowed for a better quantification of the landscape context; and succession with people (from 2015 onwards) when it was realised that people and societal drivers have strong effects on successional pathways, that ecosystem processes and services are important for human well-being, and that restoration is most successful when it is done by and for local people. Our review suggests that the hierarchical successional framework of Pickett is the best starting point to move forward as this framework already includes several factors, and because it is flexible, enabling application to different systems. The framework focuses mainly on species replacement and could be improved by focusing on succession occurring at different hierarchical scales (population, community, ecosystem, socio-ecological system), and by integrating it with more recent developments and other successional models: by considering different spatial scales (landscape, region), temporal scales (ecosystem processes occurring over centuries, and evolution), and by taking the effects of the surrounding landscape (landscape integrity and composition, the disperser community) and societal factors (previous and current land-use intensity) into account. Such a new, comprehensive framework could be tested using a combination of empirical research, experiments, process-based modelling and novel tools. Applying the framework to seres across broadscale environmental and disturbance gradients allows a better insight into what successional processes matter and under what conditions.     

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.     

Diversity and community biomass depend on dispersal and disturbance in microalgal communities

The evidence for species diversity effects on ecosystem functions is mainly based on studies not explicitly addressing local or regional processes regulating coexistence or the importance of community structure in terms of species evenness. In experimental communities of marine benthic microalgae, we altered the successional stages and thus the strength of local species interactions by manipulating rates of dispersal and disturbance. The treatments altered realized species richness, evenness and community biomass. For species richness, dispersal mattered only at high disturbance rates; when opening new space, dispersal led to maximized richness at intermediate dispersal rates. Evenness, in contrast, decreased with dispersal at low or no disturbance, i.e. at late successional stages. Community biomass showed a non-linear hump-shaped response to increasing dispersal at all disturbance levels. We found a positive correlation between richness and biomass at early succession, and a strong negative correlation between evenness and biomass at late succession. In early succession both community biomass and richness depend directly on dispersal from the regional pool, whereas the late successional pattern shows that if interactions allow the most productive species to become dominant, diverting resources from this species (i.e. higher evenness) reduces production. Our study emphasizes the difference in biodiversity–function relationships over time, as different mechanisms contribute to the regulation of richness and evenness in early and late successional stages.     

Testing the intermediate disturbance hypothesis: when will there be two peaks of diversity?

Succession after disturbances generates a mosaic of patches in different successional stages. The intermediate disturbance hypothesis predicts that intermediate disturbances lead to the highest diversity of these stages on a regional scale resulting in a hump‐shaped diversity–disturbance curve. We tested this prediction using field data of forest succession and hypothetical succession scenarios in combination with analytical and simulation models. According to our study the main factors shaping the diversity–disturbance curve and the position of the diversity maximum were the transition times between the successional stages, the transition type, neighbourhood effects and the choice of diversity measure. Although many scenarios confirmed the intermediate disturbance hypothesis we found that deviations in the form of two diversity maximums were possible. Such bimodal diversity–disturbance curves occurred when early and late successional stages were separated by one or more long‐lived (compared to the early stages) intermediate successional stages. Although the field data which met these conditions among all those tested were rare (one of six), the consequences of detecting two peaks are fundamental. The impact of disturbances on biodiversity can be complex and deviate from a hump‐shaped curve.     

甘肃兴隆山自然保护区森林演替对鸟类群落结构的影响

运用固定面积法调查了青林三个主要演替阶段森林(阔叶林、混交林和针叶林)的鸟类群落结构。结果表明,混交林中鸟类物种数、鸟类物种多样性和种团多样性最高,阔叶林最低。方差分析和聚类分析均表明,三种演替阶段森林的鸟类群落组成差异显著。所记录到的35种鸟类中有11种差异显著。其中,山雀科和科在阔叶林中占优势,而柳莺亚科在混交林和针叶林中占优势。食性种团中,阔叶林以食虫鸟和食果食虫鸟为主;混交林和针叶林均以食虫鸟、食果食虫鸟、食谷鸟和食果鸟为主。鉴于阔叶林和混交林有向顶级群落(针叶林)演替的趋势,建议采取人为干扰来维持三种演替阶段森林的面积,以便从景观水平上保护鸟类物种多样性。     

Mechanisms of successional dynamics: Consumers and the rise and fall of species dominance

Insight into potential mechanisms of succession following disturbance to an ecological community can be gained by considering processes that contribute to the rise (colonization, interactions with established species) and demise (differential mortality) of specific stages within the successional sequence. Most successional theories focus on the rise to dominance, assuming demise is the result of competition, but other factors can cause differential mortality among species, including physical disturbance, senescence, and consumers. In rocky intertidal communities on the coast of Washington state, USA, gaps in mussel beds exhibit a succession from predator-susceptible to predator-resistant species following disturbance, suggesting that differential consumption by mobile species may play an important role in the demise of early succession species and the eventual dominance of the mussel Mytilus californianus. Experimental manipulation of a dominant species earlier in succession, the blue mussel Mytilus trossulus, demonstrated that this species inhibits the invasion of M. californianus in the absence of predators. Experimental manipulation of predatory snails (Nucella emarginata and Nucella canaliculata), which feed heavily on M. trossulus but not M. californianus, greatly increased the rate at which M. californianus invaded gaps. These results and those of other studies indicate that consumers frequently have important effects on the dynamics of succession in benthic marine systems, and might also play a role in other settings.     

Changes in Mediterranean plant succession: old-fields revisited

Abstract. Old-field plots used for a study of succession in Mediterranean France were revisited after 12–14 yr. Our aims were: (1) to verify if predicted patterns of species richness, turnover and composition are confirmed; (2) to compare the development in disturbed plots with that in undisturbed ones; (3) to discuss the impact of management changes. In undisturbed plots species richness and turnover decreased with successional age. Floristic composition changed in a way consistent with the predicted successional development in most plots. Therophytes decreased and phanerophytes increased; anemochorous species decreased and endozoochorous species increased, as expected. In plots disturbed since the first analysis richness decreased with successional age, but generally remained higher than in undisturbed plots. Floristic composition, species turnover and an increase in therophytes indicated changes towards younger successional stages. Thus, disturbance changed succession but not much. This is probably linked with the regeneration abilities typical of mediterranean species, e.g. resprouting. At the landscape scale, richness did not change and species turnover was low. The plots studied were situated in two distinct locations. One had not been disturbed between the two observation periods, while the other is a mosaic of undisturbed and disturbed sites. Observations fitted predictions much more closely at the undisturbed location. We conclude that permanent plot studies are powerful in identifying successional trends and can also provide additional insights into the effects of disturbance some of the mechanisms underlying the dynamics of diversity.     

Directionality,convergence, and rate of change during early succession in the Inland Pampa,Argentina

Abstract. We tested the following hypotheses forthe first five years of a grassland succession: (a) community changes are mainly directional and related to time after disturbance ratherthanto environmental fluctuations; (b) rates of succession decrease over time, and (c) plant communities in different plots converge on a similar composition within five years of succession. We tested those hypotheses using a Principal Components Analysis applied to data from four successional plots established in successive y ears in a large cropland in the Inland Pampa, Argentina. Community changes were correlated to the age of the plots, and unrelated to rainfall variability, a major environmental variable in grasslands. Successional rates were constant over the five years, probably because of the continued dominance of different annuals; we conclude that successional rates depend on the life history of the dominant species rather than on any emergent community property. We found no evidence of convergence ordivergence; we concluded that the results of successional studies may depend on the temporal and spatial scale of observation.     

Impacts of climate variability on tree demography in second growth tropical forests: the importance of regional context for predicting successional trajectories

Naturally regenerating and restored second growth forests account for over 70% of tropical forest cover and provide key ecosystem services. Understanding climate change impacts on successional trajectories of these ecosystems is critical for developing effective large‐scale forest landscape restoration (FLR) programs. Differences in environmental conditions, species composition, dynamics, and landscape context from old growth forests may exacerbate climate impacts on second growth stands. We compile data from 112 studies on the effects of natural climate variability, including warming, droughts, fires, and cyclonic storms, on demography and dynamics of second growth forest trees and identify variation in forest responses across biomes, regions, and landscapes. Across studies, drought decreases tree growth, survival, and recruitment, particularly during early succession, but the effects of temperature remain unexplored. Shifts in the frequency and severity of disturbance alter successional trajectories and increase the extent of second growth forests. Vulnerability to climate extremes is generally inversely related to long‐term exposure, which varies with historical climate and biogeography. The majority of studies, however, have been conducted in the Neotropics hindering generalization. Effects of fire and cyclonic storms often lead to positive feedbacks, increasing vulnerability to climate extremes and subsequent disturbance. Fragmentation increases forests’ vulnerability to fires, wind, and drought, while land use and other human activities influence the frequency and intensity of fire, potentially retarding succession. Comparative studies of climate effects on tropical forest succession across biogeographic regions are required to forecast the response of tropical forest landscapes to future climates and to implement effective FLR policies and programs in these landscapes.     

Succession-driven changes in soil respiration following fire in black spruce stands of interior Alaska

Boreal forests are highly susceptible to wildfire, and post-fire changes in soil temperature and moisture have the potential to transform large areas of the landscape from a net sink to a net source of carbon (C). Understanding the ecological controls that regulate these disturbance effects is critical to developing models of ecosystem response to changes in fire frequency and severity. This paper combines laboratory and field measurements along a chronosequence of burned black spruce stands into regression analyses and models that assess relationships between moss succession, soil microclimate, decomposition, and C source-sink dynamics. Results indicate that post-fire changes in temperature and substrate quality increased decomposition in humic materials by a factor of 3.0 to 4.0 in the first 7 years after fire. Bryophyte species exhibited a distinct successional pattern in the first five decades after fire that corresponded to decreased soil temperature and increased C accumulation in organic soils. Potential rates of C exchange in mosses were greatest in early successional species and declined as the stand matured. Residual sources of CO₂ (those not attributed to moss respiration or humic decomposition) increased as a function of stand age, reflecting increased contributions from roots as the stand recovered from disturbance. Together, the field measurements, laboratory experiments, and models provide strong evidence that interactions between moss and plant succession, soil temperature, and soil moisture largely regulate C source-sink dynamics from black spruce systems in the first century following fire disturbance.     

Phylogenetic turnover during subtropical forest succession across environmental and phylogenetic scales

Although spatial and temporal patterns of phylogenetic community structure during succession are inherently interlinked and assembly processes vary with environmental and phylogenetic scales, successional studies of community assembly have yet to integrate spatial and temporal components of community structure, while accounting for scaling issues. To gain insight into the processes that generate biodiversity after disturbance, we combine analyses of spatial and temporal phylogenetic turnover across phylogenetic scales, accounting for covariation with environmental differences. We compared phylogenetic turnover, at the species‐ and individual‐level, within and between five successional stages, representing woody plant communities in a subtropical forest chronosequence. We decomposed turnover at different phylogenetic depths and assessed its covariation with between‐plot abiotic differences. Phylogenetic turnover between stages was low relative to species turnover and was not explained by abiotic differences. However, within the late‐successional stages, there was high presence‐/absence‐based turnover (clustering) that occurred deep in the phylogeny and covaried with environmental differentiation. Our results support a deterministic model of community assembly where (i) phylogenetic composition is constrained through successional time, but (ii) toward late succession, species sorting into preferred habitats according to niche traits that are conserved deep in phylogeny, becomes increasingly important.     

How predictable are reptile responses to wildfire?

Natural disturbances are key processes in the vast majority of ecosystems and a range of ecological theories have been developed in an attempt to predict biotic responses to them. However, empirical support for these theories has been inconsistent and considerable additional work remains to be done to better understand the response of biodiversity to natural disturbance. We tested predictions from the intermediate disturbance hypothesis and the habitat accommodation model of succession for reptile responses to fire history and a single major fire event. We focused our work on a broad range of vegetation types spanning sedgeland to temperate rainforest located within a national park in south‐eastern Australia. We found no significant relationships between reptile species richness and the number of fires over the past 35 years, the time since the last fire, or the severity of a major fire in 2003. Thus, we found no strong evidence to support the intermediate disturbance hypothesis. A correspondence analysis of reptile assemblages revealed a gradient in species responses to fire history. However, we found limited evidence for an ordered succession of reptiles. Nor could the responses of individual species be readily predicted from life history attributes. Thus, our findings were generally not consistent with predictions from the habitat accommodation model of succession. A possible explanation for the absence of a predictable sequence of recovery following disturbance might be the rapidity of post‐fire recovery of many components of native vegetation cover that were found to be important for reptiles (e.g. the extent of grass cover). This would have limited the time for early successional conditions to prevail and limited opportunities for species associated with such conditions. We found that most reptile species responses were much more strongly linked to vegetation type than fire variables, emphasizing a need to understand relationships with vegetation before being able to understand possible fire effects (if and where they exist). We found the disturbance concepts we examined were limited in their ability to accurately predict reptile responses to past fire history or the impacts of a single major fire in 2003. Practical management might be best guided not by disturbance theory, but by carefully setting objectives to meet conservation goals for particular individual species of reptiles.     

Models,mechanisms and pathways of succession

The study of succession has been hampered by the lack of a general theory. This is illustrated by confusion over basic concepts and inadequacy of certain models. This review clarifies the basic ideas of pathway, mechanism, and model in succession. Second, in order to prevent inappropriate narrowness in successional studies, we analyze the mechanistic adequacy of the most widely cited models of succession, those of Connell and Slatyer. This analysis shows that models involving a single pathway or a dominant mechanism cannot be treated as alternative, testable hypotheses. Our review shows much more mechanistic richness than allowed by these widely cited models of succession. Classification of the mechanisms of specific replacement, called for by existing models, is problematic and less valuable than the search for the actual mechanisms of particular seres. For example, the “tolerance” mechanism of succession has at least two contrasting meanings and is unlikely to be disentangled from the “inhibition” mechanism in field experiments. However, the understanding of particular species replacements through experiment and knowledge of the conditions of a particular sere and species life histories is a reasonable and desirable goal. Finally, we suggest the need for a broad mechanistic concept of succession. Thus, based on classical causes of succession that have survived recent scrutiny, we erect a framework of successional mechanisms. This framework aims at comprehensiveness, and specific mechanisms are nested within more general causes. As a result of its breadth and hierarchical structure, the framework performs two important functions: First, it provides a context for studies at specific sites and, second, is a scheme for formulating general and testable hypotheses. The review of specific successional mechanisms and the general mechanistic framework can together guide future work on succession, and may foment the development of a broad theory.     

Primary succession of Bistorta vivipara (L.) Delabre (Polygonaceae) root‐associated fungi mirrors plant succession in two glacial chronosequences

Glacier chronosequences are important sites for primary succession studies and have yielded well‐defined primary succession models for plants that identify environmental resistance as an important determinant of the successional trajectory. Whether plant‐associated fungal communities follow those same successional trajectories and also respond to environmental resistance is an open question. In this study, 454 amplicon pyrosequencing was used to compare the root‐associated fungal communities of the ectomycorrhizal (ECM) herb Bistorta vivipara along two primary succession gradients with different environmental resistance (alpine versus arctic) and different successional trajectories in the vascular plant communities (directional replacement versus directional non‐replacement). At both sites, the root‐associated fungal communities were dominated by ECM basidiomycetes and community composition shifted with increasing time since deglaciation. However, the fungal community's successional trajectory mirrored the pattern observed in the surrounding plant community at both sites: the alpine site displayed a directional‐replacement successional trajectory, and the arctic site displayed a directional‐non‐replacement successional trajectory. This suggests that, like in plant communities, environmental resistance is key in determining succession patterns in root‐associated fungi. The need for further replicated study, including in other host species, is emphasized.     

Understanding ecological community succession: Causal models and theories,a review

Critical review of explanations for patterns of natural succession suggests a strong, common basis for theoretical understanding, but also suggests that several well known models are incomplete as explanations of succession. A universal, general cause for succession is unlikely, since numerous aspects of historical and environmental circumstances will impinge on the process in a unique manner. However, after disturbance, occupation of a site by any species causes changes in the conditions at the site. Sorting of species may result, since different species are adapted to different regions of environmental gradients. Such sorting can generate several patterns of species abundance in time, but commonly results in sequential replacements of species adapted to the varying conditions. This may be due to constraints on species' strategies, or life history traits, placed by the limited resources available to the organism. These constraints often result in inverse correlation between traits which confer success during early and late stages of succession. Facilitatory or inhibitory effects of species on each other are best understood in terms of these life history interactions, perhaps as restrictions on, or as moderation of, these processes.Strong support for the importance of correlations in life history traits stems from comparisons of simulated succession with and without these correlations. These simulations are reviewed in some detail, and followed by brief reviews of other prominent models for succession. Several aspects of the confusion and controversies in the successional literature are then discussed, with a view to a more optimistic synthesis and direction for successional ecology.     

Implications from the Buell‐Small Succession Study for vegetation restoration

Abstract. Succession is relevant to restoration because managers have to prevent, enhance or replace natural vegetation dynamics. Features of a permanent plot study of post‐agricultural succession in central New Jersey, USA, illustrate important implications of vegetation dynamics for restoration. In the past, such implications had to be drawn from chronosequences and coarse resolution studies, neither of which exposes the local contingencies relevant to site specific restoration. However, the fine scale and continuous nature of the current study reveal that succession is highly contingent on historical and local spatial heterogeneity. For example, the absence of one generally expected dominant stage, the demise of shrubs without replacement by later successional tree species, and the long and multimodal persistence of individual species suggest that neither the relay floristic or initial floristic models of succession is adequate to guide restoration. At the local scale, volleys of species appear through the succession, and reflect spatial contingencies such as neighboring vegetation and edge relationships, and patchy behavior of different functional groups. The role of introduced species and of plant consumers are additional sources of local contingency. These local and time‐specific behaviors in the vegetation are the patterns that restorationists must understand either to choose appropriate reference states, to prevent unexpected local dynamics, or to design interventions that are appropriate to the specific site of interests.     

Current reclamation practices after oil and gas development do not speed up succession or plant community recovery in big sagebrush ecosystems in Wyoming

Reclamation is an application of treatment(s) following disturbance to promote succession and accelerate the return of target conditions. Previous studies have framed reclamation in the context of succession by studying its effectiveness in reestablishing late‐successional plant communities. Reestablishment of plant communities is especially important and challenging in drylands such as shrub steppe ecosystems where succession proceeds slowly. These ecosystems face threats from climate change, invasive species, altered fire regimes, and land‐use change, as well as fossil‐fuel extraction and associated disturbance. As such, the need for effective reclamation after this type of energy development is great. However, past research regarding this type of reclamation has focused on mining rather than oil and gas development. To better understand the effect of reclamation on rates of succession in dryland shrub steppe ecosystems, we sampled oil and gas wellpads and adjacent undisturbed big sagebrush plant communities in Wyoming, U.S.A., and quantified the extent of recovery for forbs, grasses, and shrubs on reclaimed and unreclaimed wellpads relative to undisturbed plant communities. Reclamation increased the recovery rate for early‐successional types, including combined forbs and grasses and perennial grasses, but did not affect recovery rate of late‐successional types, particularly big sagebrush and perennial forbs. Rather, subsequent analyses showed that recovery of late‐successional types was affected by soil texture and time since wellpad abandonment. This is consistent with studies in other ecosystems where reclamation has been implemented, suggesting that reclamation may not help reestablish late‐successional plant communities more quickly than they would reestablish naturally.     

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.     

Spontaneous succession in Central‐European man‐made habitats: What information can be used in restoration practice?

Abstract. The results of previous studies concerning spontaneous vegetation succession in various man‐made or disturbed habitats in Central Europe are discussed in relation to restoration ecology. An attempt is made to answer the main questions which are important to restoration programs. These concern the rate and direction of succession; participation of target species and communities, woody species, ruderals and aliens; arresting or diverting succession; changes in species diversity; variability of successional pathways and the role of abiotic environmental conditions. The great potential for using successional theory in restoration programs is emphasized.     

Markov chain analysis of succession in a rocky subtidal community

We present a Markov chain model of succession in a rocky subtidal community based on a long-term (1986-1994) study of subtidal invertebrates (14 species) at Ammen Rock Pinnacle in the Gulf of Maine. The model describes successional processes (disturbance, colonization, species persistence, and replacement), the equilibrium (stationary) community, and the rate of convergence. We described successional dynamics by species turnover rates, recurrence times, and the entropy of the transition matrix. We used perturbation analysis to quantify the response of diversity to successional rates and species removals. The equilibrium community was dominated by an encrusting sponge (Hymedesmia) and a bryozoan (Crisia eburnea). The equilibrium structure explained 98% of the variance in observed species frequencies. Dominant species have low probabilities of disturbance and high rates of colonization and persistence. On average, species turn over every 3.4 years. Recurrence times varied among species (7-268 years); rare species had the longest recurrence times. The community converged to equilibrium quickly (9.5 years), as measured by Dobrushin's coefficient of ergodicity. The largest changes in evenness would result from removal of the dominant sponge Hymedesmia. Subdominant species appear to increase evenness by slowing the dominance of Hymedesmia. Comparison of the subtidal community with intertidal and coral reef communities revealed that disturbance rates are an order of magnitude higher in coral reef than in rocky intertidal and subtidal communities. Colonization rates and turnover times, however, are lowest and longest in coral reefs, highest and shortest in intertidal communities, and intermediate in subtidal communities.