Markov Chain Analysis of Succession in a Rocky Subtidal Community

Abstract

We present a Markov chain model of succession in a rocky subtidal community based on a long-term (1986)(1987)(1988)(1989)(1990)(1991)(1992)(1993)(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 The dynamics of an ecological community are often described by changes in species composition over time. We use the term "succession" to refer to these changes. Succession is no longer viewed as a deterministic development toward a unique stable climax community (Connell and Slatyer 1977). Instead, it is widely recognized that disturbance, dispersal, colonization, and species interactions produce patterns and variability on a range of temporal and spatial scales.Markov chains were introduced as models of succession by Waggoner and Stephens (1970) and Horn (1975). These models imagine the landscape as a large (usually infinite) set of patches or points in space. The state of a point is given by the list of species that occupy it. In one class of models, this list may include multiple coexisting populations; such models are called patch-occupancy models (e.g., Caswell andCohen 1991a, 1991b). In another class of models, points are occupied by single individuals rather than populations. Such models have been applied to forests (Waggoner and Stephens 1970;Horn 1975;Runkle 1981;Masaki et al. 1992), plant communities (Isagi and Nakagoshi 1990; Aaviksoo 1995), insect assemblages (Usher 1979), coral reefs (Tanner et al. 1994(Tanner et al. , 1996, and rocky intertidal communities (Wootton 2001b(Wootton , 2001c.Successional dynamics are modeled by defining the pro...

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