Bacterial Succession on the Leaf Surface: A Novel System for Studying Successional Dynamics

Succession is a widely studied process in plant and animal systems, but succession in microbial communities has received relatively little attention despite the ubiquity of microorganisms in natural habitats. One important microbial habitat is the phyllosphere, or leaf surface, which harbors large, diverse populations of bacteria and offers unique opportunities for the study of succession and temporal community assembly patterns. To explore bacterial community successional patterns, we sampled phyllosphere communities on cottonwood (Populus deltoides) trees multiple times across the growing season, from leaf emergence to leaf fall. Bacterial community composition was highly variable throughout the growing season; leaves sampled as little as a week apart were found to harbor significantly different communities, and the temporal variability on a given tree exceeded the variability in community composition between individual trees sampled on a given day. The bacterial communities clearly clustered into early-, mid-, and late-season clusters, with early- and late-season communities being more similar to each other than to the mid-season communities, and these patterns appeared consistent from year to year. Although we observed clear and predictable changes in bacterial community composition during the course of the growing season, changes in phyllosphere bacterial diversity were less predictable. We examined the species–time relationship, a measure of species turnover rate, and found that the relationship was fundamentally similar to that observed in plant and invertebrate communities, just on a shorter time scale. The temporal dynamics we observed suggest that although phyllosphere bacterial communities have high levels of phylogenetic diversity and rapid turnover rates, these communities follow predictable successional patterns from season to season.     

Convergence of community composition during secondary succession on Zokor rodent mounds on the Tibetan Plateau

Aims The community succession theory is much debated in ecology. We studied succession on Zokor rodent mounds on the Tibetan Plateau to address several fundamental questions, among them: (i) During secondary succession, does the community composition converge towards one community state or multiple states depending on the initial colonization? (ii) Do mound communities located in different background communities exhibit different assembly trajectories?Methods In a sub-alpine meadow, we investigated a total of 80 mound communities at several successional stages in three different background communities resulting from different management histories and compared their changes in species composition. The distribution of plant communities over time was analyzed with quantitative classification and ordination methods. The co-occurrence patterns of species were evaluated at each successional stage, and the degree of convergence/divergence among communities was obtained by calculating two beta-diversity indices.Important findings During secondary succession, species richness of mound communities changed over time, and this change was dependent on the background community. Five life-form groups exhibited different dynamic patterns in species richness and plant cover. Community composition and the degree of species co-occurrence between communities increased over time since disturbance. There was much variation in species composition at earlier stages of succession, but communities on older mounds became more similar to each other and to their surrounding vegetation over the course of secondary succession. Post-disturbance succession of Zokor mound communities transitioned from 'multiple alternative states' to 'background-based deterministic community assembly' over time. Tradeoffs between competition and colonization, as well as the characteristics of different life-forms and mass effects within a limited species pool are the mechanisms responsible for convergence of mound communities.     

Species and functional trait re-assembly of ground beetle communities in restored grasslands

Ecosystem restoration provides unique opportunities to study community dynamics under succession and can reveal how consumer communities re-assemble and respond to successional changes. Studying community dynamics from both taxonomic and functional trait perspectives also may provide more robust assessments of restoration progress or success and allow cross-system comparisons. We studied ground beetle (Coleoptera: Carabidae) communities for three years in a restored grassland chronosequence with sites from 0 to 28 years old. We measured traditional community metrics (abundance, richness, Shannon diversity) and functional trait metrics based on species’ body length, wing morphology, activity time, phenology, and diet. Communities had high species richness and abundance in early successional stages, but these declined in later stages to low levels comparable to an adjacent grassland remnant. Species composition also shifted with time, converging with the remnant. Although functional richness, like species richness, declined as succession progressed, functional divergence quickly increased and was maintained over time, suggesting niche differentiation in established communities. Young sites were typified by small, macropterous, phytophagous species, while older sites contained larger species more likely to be flightless and carnivorous. Prescribed burns also affected traits, decreasing prevalence of larger species. This study demonstrates that functionally diverse consumer communities can self-assemble under restoration practices. In a relatively short amount of time both morphological and trophic level diversity are established. However, prescribed fire intended to control non-desirable plants may also shape beetle community functional composition, and restoration managers should consider if plant community benefits of fire outweigh potential declines in consumer function.     

Gradients in compositional variation on lahars,Mount St. Helens,Washington, USA

How isolation affects primary succession remains unresolved. Our hypotheses are: 1) the rate of succession is slowed within 50 m from sources and 2) dominance increases more rapidly if the growing season is longer. We sampled lahar vegetation near conifer forests on Mount St. Helens 23 years after the eruption using transects (Muddy) and grids (Butte Camp, BC). Transects were compared to isolated plots of the same age, while grids were compared to vegetation that was 400 years older. Cover declined with distance and with elevation on the Muddy due to denser seed rain near forests and shorter growing seasons at higher elevation. On BC-1, next to a forest, richness measures increased with distance, while cover decreased with elevation. On BC-2, more exposed than BC-1, mean richness and cover declined with elevation, but increased towards seed sources. Equitability increased with distance and with elevation on more stressful transects and on BC-1. Percent similarity (PS), a measure of floristic heterogeneity, declined with elevation on Muddy-SW. BC-1 PS decreased with both distance and elevation. Data from grids collected since 1987 showed that both sites became more homogeneous through time. Directional changes on BC-1 were greater than on BC-2, while annual variation of DCA scores declined only on BC-2. These differences are related to conifer dominance patterns. Succession is influenced by isolation, which controls the seed rain, and by stress, which controls establishment, the rate of biomass accumulation, and the spread of immigrants. Although community development is governed by environmental factors, this study shows that the effects of dispersal that result from distance factors can persist and may be a source of unexplained variation in mature communities.     

Phylogenetic and functional dissimilarity does not increase during temporal heathland succession

Succession has been a focal point of ecological research for over a century, but thus far has been poorly explored through the lens of modern phylogenetic and trait-based approaches to community assembly. The vast majority of studies conducted to date have comprised static analyses where communities are observed at a single snapshot in time. Long-term datasets present a vantage point to compare established and emerging theoretical predictions on the phylogenetic and functional trajectory of communities through succession. We investigated within, and between, community measures of phylogenetic and functional diversity in a fire-prone heathland along a 21 year time series. Contrary to widely held expectations that increased competition through succession should inhibit the coexistence of species with high niche overlap, plots became more phylogenetically and functionally clustered with time since fire. There were significant directional shifts in individual traits through time indicating deterministic successional processes associated with changing abiotic and/or biotic conditions. However, relative to the observed temporal rate of taxonomic turnover, both phylogenetic and functional turnover were comparatively low, suggesting a degree of functional redundancy among close relatives. These results contribute to an emerging body of evidence indicating that limits to the similarity of coexisting species are rarely observed at fine spatial scales.     

Successional Patterns of Microfungi in Fallen Leaves of Ficus pleurocarpa (Moraceae) in an Australian Tropical Rain Forest1

Successional patterns of microfungi on decaying leaves of Ficus pleurocarpa were assessed as part of a study to enumerate microfungi in tropical rain forest leaf litter. Leaves degraded into fragments over a period of 3 mo. Two methods were applied, a direct observational method and a particle filtration protocol. Using a direct method, 104 species were observed, while 53 sporulating taxa and 100 sterile morphotaxa were isolated by particle filtration. Overall patterns of succession were confirmed by both methods, but the relative abundance of species detected differed between the two methods. Nonmetric Multidimensional Scaling identified at least four successional stages and suggested that microfungal communities increased in similarity with advancing decay. Data collected by the direct method indicated a slow but steady decline of diversity with advancing decay, whereas an increase in diversity was detected by particle filtration. Synecological succession studies provide a useful tool to identify patterns and generate hypotheses. Understanding the underlying causes of successional patterns will require further autecological studies.     

Intraspecific variation in plant size,secondary plant compounds,herbivory and parasitoid assemblages during secondary succession

During secondary succession on abandoned agricultural fields the diversity and abundance of insect communities often increases, whereas the performance and nutritional quality of early successional plants often declines. As the diversity and abundance of insects on a single plant are determined by characteristics of the environment as well as of the host plant, it is difficult to predict how insects associated with a single plant species will change during succession. We examined how plant characteristics of the early successional plant species ragwort (Jacobaea vulgaris), and the herbivores and parasitoids associated with these plants change during secondary succession. In ten grasslands that differed in time since abandonment (3–26 years), we measured the size and primary and secondary chemistry of individual ragwort plants. For each plant we also recorded the presence of herbivores in flowers, leaves and stems, and reared parasitoids from these plant parts. Ragwort plants were significantly larger but had lower nitrogen concentrations in recently abandoned sites than in older sites. Pyrrolizidine alkaloid (PA) composition varied among plants within sites but also differed significantly among sites. However, there was no relationship between the age of a site and PA composition. Even though plant size decreased with time since abandonment, the abundance of stem-boring insects and parasitoids emerging from stems significantly increased with site age. The proportion of plants with flower and leaf herbivory and the number of parasitoids emerging from flowers and leaves was not related to site age. Parasitoid diversity significantly increased with site age. The results of our study show that ragwort and insect characteristics both change during secondary succession, but that insect herbivore and parasitoid abundances are not directly related to plant size or nutritional quality.     

Microbial leaf degraders in boreal streams: bringing together stochastic and deterministic regulators of community composition

1. Leaves that fall into the water represent a new habitat for microorganisms to colonise in streams, providing an opportunity to study colonisation and the subsequent regulation of community structure. We explored community composition of bacteria and fungi on decomposing alder leaves in nine streams in central Sweden, and describe their relationship with environmental variables. Succession of the microbial community was studied in one of the streams for 118 days. Microbial community composition was examined by denaturing gradient gel electrophoresis on replicate samples of leaves from each stream. 2. During succession in one stream, maximum taxon richness was reached after 34 days for bacteria and 20 days for fungi respectively. Replicate samples within this stream differed between each other earlier in colonisation, while subsequently such variation among replicate communities was low and remained stable for several weeks. Replicate samples taken from all the nine streams after 34 days of succession showed striking similarities in microbial communities within‐streams, although communities differed more strongly between streams. 3. Canonical analysis of microbial communities and environmental variables revealed that water chemistry had a significant influence on community composition. This influence was superimposed on a statistical relationship between the properties of stream catchments and microbial community composition. 4. The catchment regulates microbial communities in two different ways. It harbours the species pool from which the in‐stream microbial community is drawn and it governs stream chemistry and the composition of organic substrates that further shape the communities. We suggest that there is a random element to colonisation early in succession, whereas other factors such as species interactions, stream chemistry and organic substrate properties, result in a more deterministic regulation of communities during later stages.     

Stability and succession of the rhizosphere microbiota depends upon plant type and soil composition

We examined succession of the rhizosphere microbiota of three model plants (Arabidopsis, Medicago and Brachypodium) in compost and sand and three crops (Brassica, Pisum and Triticum) in compost alone. We used serial inoculation of 24 independent replicate microcosms over three plant generations for each plant/soil combination. Stochastic variation between replicates was surprisingly weak and by the third generation, replicate microcosms for each plant had communities that were very similar to each other but different to those of other plants or unplanted soil. Microbiota diversity remained high in compost, but declined drastically in sand, with bacterial opportunists and putative autotrophs becoming dominant. These dramatic differences indicate that many microbes cannot thrive on plant exudates alone and presumably also require carbon sources and/or nutrients from soil. Arabidopsis had the weakest influence on its microbiota and in compost replicate microcosms converged on three alternative community compositions rather than a single distinctive community. Organisms selected in rhizospheres can have positive or negative effects. Two abundant bacteria are shown to promote plant growth, but in Brassica the pathogen Olpidium brassicae came to dominate the fungal community. So plants exert strong selection on the rhizosphere microbiota but soil composition is critical to its stability. microbial succession/ plant–microbe interactions/rhizosphere microbiota/selection.     

Non-Native Plants Disrupt Dual Promotion of Native Alpha and Beta Diversity

Non-native species can alter patterns of species diversity at multiple spatial scales, but the processes that underlie multi-scale effects remain unclear. Here we show that non-native species reduce native diversity at multiple scales through simultaneous disruption of two processes of native community assembly: species immigration, which enhances alpha diversity, and community divergence, which enhances beta diversity. Community divergence refers to the process in which local communities diverge over time in species composition because the history of species immigration and, consequently, the way species affect one another within communities are variable among communities. Continuous experimental removal of species over four years of floodplain succession revealed that, when non-native species were excluded, stochastic variation in the timing of a dominant native species’ arrival allowed local communities to diverge, thereby enhancing beta diversity, without compromising promotion of alpha diversity by species immigration. In contrast, when non-native species were allowed to enter experimental plots, they not only reduced native alpha diversity by limiting immigration, but also diminished the dominant native species’ role in enhancing native beta diversity. Our results highlight the importance of community assembly and succession for understanding multi-scale effects of non-native species.     

New aspects of grassland recovery in old‐fields revealed by trait‐based analyses of perennial‐crop‐mediated succession

Classical old‐field succession studies focused on vegetation changes after the abandonment of annual croplands or on succession after the elimination of cultivated crops. Perennial‐crop‐mediated succession, where fields are initially covered by perennial crops, reveals alternative aspects of old‐field succession theories. We tested the validity of classical theories of old‐field succession for perennial‐crop‐mediated succession. We formulated the following hypotheses: (1) functional diversity increases with increasing field age; (2) resource acquisition versus conservation trade‐off shifts toward conservation at community level during the succession; (3) the importance of spatial and temporal seed dispersal decreases during the succession; and (4) competitiveness and stress‐tolerance increases and ruderality decreases at community level during the succession. We studied functional diversity, trait distributions and plant strategies in differently aged old‐fields using chronosequence method. We found increasing functional richness and functional divergence, but also unchanged or decreasing functional evenness. We detected a shift from resource acquisition to resource conservation strategy of communities during the succession. The role of spatial and temporal seed dispersal was found to be important not only at the initial but also at latter successional stages. We found an increasing stress‐tolerance and a decreasing ruderality during succession, while the competitiveness remained unchanged at the community level. Despite the markedly different starting conditions, we found that classical and perennial‐crop‐mediated old‐field successions have some similarities regarding the changes of functional diversity, resource acquisition versus conservation trade‐off, and seed dispersal strategies. However, we revealed also the subsequent differences. The competitive character of communities remained stable during the succession; hence, the initial stages of perennial‐crop‐mediated succession can be similar to the middle stages of classical old‐field succession. Moreover, the occupied functional niche space and differentiation were larger in the older stages, but resources were not effectively utilized within this space, suggesting that the stabilization of the vegetation requires more time.     

Explaining productivity-diversity relationships in plants

Relationships between productivity and diversity in plant communities have been widely documented. Unimodal productivity-diversity relationships are most common along natural productivity gradients, and fertilization generally reduces diversity. Five distinct hypotheses invoke changes in competition to explain why diversity should decline from intermediate to high productivity. Because experiments measuring the effects of competition on diversity are rare, four of the five hypotheses have not been directly tested, but each hypothesis makes unique predictions that allow for indirect tests. The indirect evidence is often conflicting, and while none of the hypotheses can be rejected, only the dynamic equilibrium hypothesis is consistently supported. A new hypothesis, however, is supported by indirect evidence and may help to explain the variation in the shape of productivity-diversity relationships, as well as the most common patterns. Diversity may be high in environments that promote size symmetric competition, where soil resources limit growth and are homogeneously distributed within the soil volume explored by individual plants. Conversely, diversity may be low in environments that promote size asymmetric competition, where light is limiting, or where soil resources are limiting and are patchily distributed within rooting zones.     

Community disassembly under global change: Evidence in favor of the stress‐dominance hypothesis

Ecological theory suggests that communities are not random combinations of species but rather the results of community assembly processes filtering and sorting species that are able to coexist together. To date, such processes (i.e., assembly rules) have been inferred from observed spatial patterns of biodiversity combined with null model approaches, but relatively few attempts have been made to assess how these processes may be changing through time. Specifically, in the context of the ongoing biodiversity crisis and global change, understanding how processes shaping communities may be changing and identifying the potential drivers underlying these changes become increasingly critical. Here, we used time series of 460 French freshwater fish communities and assessed both functional and phylogenetic diversity patterns to determine the relative importance of two key assembly rules (i.e., habitat filtering and limiting similarity) in shaping these communities over the last two decades. We aimed to (a) describe the temporal changes in both functional and phylogenetic diversity patterns, (b) determine to what extent temporal changes in processes inferred through the use of standardized diversity indices were congruent, and (c) test the relationships between the dynamics of assembly rules and both climatic and biotic drivers. Our results revealed that habitat filtering, although already largely predominant over limiting similarity, became more widespread over time. We also highlighted that phylogenetic and trait‐based approaches offered complementary information about temporal changes in assembly rules. Finally, we found that increased environmental harshness over the study period (especially higher seasonality of temperature) led to an increase in habitat filtering and that biological invasions increased functional redundancy within communities. Overall, these findings underlie the need to develop temporal perspectives in community assembly studies, as understanding ongoing temporal changes could provide a better vision about the way communities could respond to future global changes.     

Disturbance,coral reef communities,and changing ecological paradigms

We examine changing ecological theory regarding the role of disturbance in natural communities and relate past and emerging paradigms to coral reefs. We explore the elements of this theory, including patterns (diversity, distribution, and abundance) and processes (competition, succession, and disturbance), using currently evolving notions concerning matters of scale (temporal and spatial), local versus regional species richness, and the equilibrium versus nonequilibrium controversy. We conclude that any attempt to categorize coral reef communities with respect to disturbance regimes will depend on the question being asked and the desired level of resolution: local assemblage versus regional species pool, successional versus geological time, and on the taxonomic and tropic affinities of species included in the study. As with many communities in nature, coral reefs will prove to be mosaics of species assemblages with equilibrial and nonequilibrial dynamics.     

The effects of species pool,dispersal and competition on the diversity–productivity relationship

Aim The diversity–productivity relationship is a controversial issue in ecology. Diversity is sometimes seen to increase with productivity but a unimodal relationship has often been reported. Competitive exclusion was cited initially to account for the decrease of diversity at high productivity. Subsequently, the roles of evolutionary history (species pool size) and dispersal rate have been acknowledged. We explore how the effects of species pool, dispersal and competition combine to produce different diversity–productivity relationships. Methods We use a series of simulations with a spatially explicit, individual‐based model. Following empirical expectations, we used four scenarios to characterize species pool size along the productivity gradient (uniformly low and high, linear increase and unimodal). Similarly, the dispersal rate varied along the productivity gradient (uniformly low and high, and unimodal). We considered both neutral communities and communities with competitive exclusion. Results and main conclusions Our model predicts that competitive interactions will result in unimodal diversity–productivity relationships. The model often predicts unimodal patterns in neutral communities as well, although the decline in richness at high productivity is less than in competing communities. A positive diversity–productivity relationship is simulated for neutral communities when the species pool size increases with productivity and the dispersal rate is high. This scenario is probably more widespread in nature than the others since positive diversity–productivity relationships have been observed more frequently than previously expected, especially in the tropics and for woody species. Our simulated effects of species pool, dispersal and competition on diversity patterns can be linked to empirical observations to uncover mechanisms behind the diversity–productivity relationship.     

Functional Roles Affect Diversity-Succession Relationships for Boreal Beetles

Species diversity commonly increases with succession and this relationship is an important justification for conserving large areas of old-growth habitats. However, species with different ecological roles respond differently to succession. We examined the relationship between a range of diversity measures and time since disturbance for boreal forest beetles collected over a 285 year forest chronosequence. We compared responses of “functional” groups related to threat status, dependence on dead wood habitats, diet and the type of trap in which they were collected (indicative of the breadth of ecologies of species). We examined fits of commonly used rank-abundance models for each age class and traditional and derived diversity indices. Rank abundance distributions were closest to the Zipf-Mandelbrot distribution, suggesting little role for competition in structuring most assemblages. Diversity measures for most functional groups increased with succession, but differences in slopes were common. Evenness declined with succession; more so for red-listed species than common species. Saproxylic species increased in diversity with succession while non-saproxylic species did not. Slopes for fungivores were steeper than other diet groups, while detritivores were not strongly affected by succession. Species trapped using emergence traps (log specialists) responded more weakly to succession than those trapped using flight intercept traps (representing a broader set of ecologies). Species associated with microhabitats that accumulate with succession (fungi and dead wood) thus showed the strongest diversity responses to succession. These clear differences between functional group responses to forest succession should be considered in planning landscapes for optimum conservation value, particularly functional resilience.     

Clonal structure of Elytrigia atherica along different successional stages of a salt marsh

Elytrigia atherica is a tall clonal grass species typical of higher salt marshes, but is gradually invading to the lower marshes. At young successional stages of a salt marsh, E. atherica is found sparsely dispersed in small groups of ramets. These patches increase in size and ramet density over time, eventually forming extensive swards as succession proceeds. This study investigates the change in the clonal diversity of E. atherica stands during colonization as a result of its reproductive strategy. Clonal diversities of differently sized patches of E. atherica were investigated on two lower salt-marsh sites of different age, 25 years and 35 years, respectively. Microsatellite fingerprint patterns were used to determine genet identities and to estimate relatedness and genetic differentiation between the sites, between patches within sites and within patches. The majority of the patches on both sites contained more than one genet. On the older site, the clonal diversity was higher than on the younger site. However, the clonal diversity tended to decrease with increasing patch size. Low genetic differentiation was found between the two sites, indicating habitat differentiation, whereas differentiation between patches within sites was high. It is reasoned that different environmental conditions could have resulted in different clonal structures: On an older marsh, the increase of successful seedling recruitment, due to more suitable environmental conditions, leads to an increase in clonal diversity. Over time, with increasing ramet density, intraspecific competition is likely to increase, resulting in a decrease of clonal diversity.     

Short-term rates of parasite evolution predict the evolution of host diversity

Coevolution with parasites has been implicated as an important factor driving the evolution of host diversity. Studies to date have focussed on gross effects of parasites: how host diversity differs in the presence vs. absence of parasites. But parasite-imposed selection is likely to show rapid variation through time. It is unclear whether short-term fluctuations in the strength of parasite-imposed selection tend to affect host diversity, because increases in host diversity are likely to be constrained by both the supply of genetic variation and ecological processes. We followed replicate populations of coevolving, initially isogenic, bacteria and phages through time, measuring host diversity (with respect to bacterial colony morphologies), host density and rates of parasite evolution. Both host density and time-lagged rates of parasite evolution were good independent predictors of the magnitude of bacterial within- and between-population diversities. Rapid parasite evolution and low host density decreased host within-population diversity, but increased between-population diversity. This study demonstrates that short-term changes in the rate of parasite evolution can predictably drive patterns of host diversity.     

Geographical variation and population structure in the White-rumped Sandpiper Calidris fuscicollis as shown by morphology, mitochondrial DNA and carbon isotope ratios

Several plant-herbivore hypotheses are based on the assumption that plants cannot simultaneously allocate resources to growth and defence. We studied seasonal patterns in allocation to growth and putatively defensive compounds by monitoring several chemical and physical traits in the leaves of mountain birch from early June (budburst) to late September (leaf senescence). We found significant seasonal changes in all measured characteristics, both in terms of concentrations (mg g^-1) and amounts (mg leaf^-1). Changes were very rapid in the spring, slow in the middle of the season, and there was another period of fast changes in the senescing leaves. Co-occurring changes in physical leaf traits and concentrations of several compounds indicated a seasonal decline in foliage suitability for herbivores. Concentrations of protein and free amino acids declined through the growing season whereas individual sugars showed variable seasonal patterns. The seasonal trends of phenolic groups differed drastically: concentrations of soluble proanthocyanidins increased through the season, whereas cell wall-bound proanthocyanidins, gallotannins and flavonoid glycosides declined after an initial increase in young leaves. We failed to find proof that the seasonal accumulation of phenolics would have been seriously compromised by leaf or shoot growth, as assumed by the growth/differentiation balance hypothesis and the protein competition model hypothesis. On the contrary, there was a steady increase in the total amount of phenolics per leaf even during the most active leaf growth.     

Marine invertebrates in an algal succession. I. Variations in abundance and diversity with succession

This is a study of the effects of successional algal changes on the community of small invertebrate animals living in the algae at a marine intertidal boulder field. At each of 14 sampling times over a 2-yr period, replicate samples were taken from all stages of succession. By sampling all successional stages simultaneously at each date over an extended period, we could analyze spatial and temporal variations within and among stages. Species richness, diversity, and abundance of the whole invertebrate community increased from early to middle successional stages, while remaining similar from middle to late stages. Evenness remained about constant throughout succession. The increases occur primarily through the addition of species to the community. Some taxonomic groups in this study showed different patterns. Unlike patterns seen in birds, insects, and spiders associated with terrestrial plant succession, species replacements are rare, and there are few early and middle successional specialists. Trophic structure changed markedly during succession. Spatial variation did not change with succession but temporal variation declined. Many of these results are different from the predictions of equilibrium-based theory. Succession in this marine animal community is very different from that in terrestrial animal groups, probably because of the higher rate of exchange of matter in this open coast system.