Connell and Slatyer's models of succession in the biodiversity era

Understanding how species interactions drive succession is a key issue in ecology. In this study we show the utility of combining the concepts and methodologies developed within the biodiversity-ecosystem functioning research program with J. H. Connell and R. O. Slatyer's classic framework to understand succession in assemblages where multiple interactions between early and late colonists may include both inhibitory and facilitative effects. We assessed the net effect of multiple species interactions on successional changes by manipulating the richness, composition, and abundance of early colonists in a low-shore assemblage of algae and invertebrates of the northwestern Mediterranean. Results revealed how concomitant changes in species richness and abundance can strongly alter the net effect of inhibitory vs. facilitative interactions on succession. Increasing richness of early colonists inhibited succession, but only under high levels of initial abundance, probably reflecting the formation of a highly intricate matrix that prevented further colonization. In contrast, increasing initial abundance of early colonists tended to facilitate succession under low richness. Thus, changes in abundance of early colonists mediated the effects of richness on succession.     

Geographic variation in resource dominance-discovery in Brazilian ant communities

A predictive framework for the ecology of species invasions requires that we learn what limits successful invaders in their native range. The red imported fire ant (Solenopsis invicta) is invasive in the United States, Puerto Rico, Australia, New Zealand, and China. Solenopsis invicta appears to be a superior competitor in its introduced range, where it can cause the local extirpation of native species, but little is known about its competitive ability in its native range in South America. Here we examine the competitive ability of S. invicta for food resources in three widely separated Brazilian ant communities. Each of these communities contains 20-40 ant species, 8-10 of which were common and frequently interacted with S. invicta. S. invicta at all three sites was attacked by several species-specific phorid parasitoids, and at one site, two other species were attacked by their own specialized parasitoids. We examined interactions in these local communities for evidence that trade-offs among ant species between resource dominance and resource discovery, and between resource dominance and parasitoid vulnerability facilitate local coexistence. The trade-off between resource dominance and resource discovery was strong and significant only at Santa Genebra, where parasitoids had no effect on the outcome of confrontations at resources. At Bonito, parasitoids significantly reduced the ability of S. invicta, which was the top-ranked behavioral dominant, from defending and usurping food resources from subordinate species. In the Pantanal, S. invicta ranked behind three other ant species in a linear hierarchy of behavioral dominance, and lost the majority of its interactions with a fourth more subordinate species, Paratrechina fulva, another invasive species. Parasitoids of S. invicta were uncommon in the Pantanal, and did not affect its low position in the hierarchy relative to the other two sites. Parasitoids, however, did affect the ability of Linepithema angulatum, the top-ranked behavioral dominant in this community, from defending and usurping resources from behavioral subordinates. These results indicate that both interspecific competition and trait-mediated indirect effects of phorid parasitoids affect the ecological success of the red imported fire ant in its native range, but that the relative importance of these factors varies geographically.     

Neighborhood analyses of canopy tree competition along environmental gradients in New England forests.

We use permanent-plot data from the USDA Forest Service's Forest Inventory and Analysis (FIA) program for an analysis of the effects of competition on tree growth along environmental gradients for the 14 most abundant tree species in forests of northern New England, USA. Our analysis estimates actual growth for each individual tree of a given species as a function of average potential diameter growth modified by three sets of scalars that quantify the effects on growth of (1) initial target tree size (dbh), (2) local environmental conditions, and (3) crowding by neighboring trees. Potential growth of seven of the 14 species varied along at least one of the two environmental axes identified by an ordination of relative abundance of species in plots. The relative abundances of a number of species were significantly displaced from sites where they showed maximum potential growth. In all of these cases, abundance was displaced to the more resource-poor end of the environmental gradient (either low fertility or low moisture). The pattern was most pronounced among early successional species, whereas late-successional species reached their greatest abundance on sites where they also showed the highest growth in the absence of competition. The analysis also provides empirical estimates of the strength of intraspecific and interspecific competitive effects of neighbors. For all but one of the species, our results led us to reject the hypothesis that all species of competitors have equivalent effects on a target species. Most of the individual pairwise interactions were strongly asymmetric. There was a clear competitive hierarchy among the four most shade-tolerant species, and a separate competitive hierarchy among the shade-intolerant species. Our results suggest that timber yield following selective logging will vary dramatically depending on the configuration of the residual canopy, because of interspecific variation in the magnitude of both the competitive effects of different species of neighbors and the competitive responses of different species of target trees to neighbors. The matrix of competition coefficients suggests that there may be clear benefits in managing for specific mixtures of species within local neighborhoods within stands.     

The importance of defining focal assemblages when evaluating amphibian and reptile responses to land use

Habitat loss and degradation are primary threats to amphibians and reptiles, but the relative effects of common land uses on assemblages and the mechanisms that underlie faunal responses are poorly studied. We reviewed the effects of four prevalent types of habitat alteration (urbanization, agriculture, livestock grazing, and silviculture) on amphibian and reptile species richness and abundance by summarizing reported responses in the literature and by estimating effect sizes across studies for species richness in each land‐use type. We then used a multinomial model to classify species as natural habitat specialists, generalists, and disturbed habitat specialists and examined variation in effect sizes for each land‐use type according to habitat specialization categories. There were mixed conclusions from individual studies, some reporting negative, neutral, or positive effects of land use on species richness and total abundance. A large proportion of studies reported species‐specific effects of individual species abundance. However, in our analysis of effect sizes, we found a general trend of negative effects of land use on species richness. We also demonstrate that habitat associations of common species and species turnover can explain variation in the effect of land use on herpetofauna. Our review highlights the pervasive negative effects of common land uses on amphibians and reptiles, the importance of identifying groups vulnerable to land‐use change (e.g., forest‐associated species) in conservation studies, and the potential influence of disturbance‐associated species on whole assemblage analyses.     

Shifts in southern Wisconsin forest canopy and understory richness, composition, and heterogeneity

We resurveyed the under- and overstory species composition of 94 upland forest stands in southern Wisconsin in 2002-2004 to assess shifts in canopy and understory richness, composition, and heterogeneity relative to the original surveys in 1949-1950. The canopy has shifted from mostly oaks (Quercus spp.) toward more mesic and shade-tolerant trees (primarily Acer spp.). Oak-dominated early-successional stands and those on coarse, nutrient-poor soils changed the most in canopy composition. Understories at most sites (80%) lost native species, with mean species density declining 25% at the 1-m2 scale and 23.1% at the 20-m2 scale. Woody species have increased 15% relative to herbaceous species in the understory despite declining in absolute abundance. Initial canopy composition, particularly the abundance of red oaks (Quercus rubra and Q. velutina), predicted understory changes better than the changes observed in the overstory. Overall rates of native species loss were greater in later-successional stands, a pattern driven by differential immigration rather than differential extirpation. However, understory species initially found in early-successional habitats declined the most, particularly remnant savanna taxa with narrow or thick leaves. These losses have yet to be offset by compensating increases in native shade-adapted species. Exotic species have proliferated in prevalence (from 13 to 76 stands) and relative abundance (from 1.2% to 8.4%), but these increases appear unrelated to the declines in native species richness and heterogeneity observed. Although canopy succession has clearly influenced shifts in understory composition and diversity, the magnitude of native species declines and failure to recruit more shade-adapted species suggest that other factors now act to limit the richness, heterogeneity, and composition of these communities.     

Functional identity and functional structure change through succession in a rocky intertidal marine herbivore assemblage

Despite the great interest in characterizing the functional structure and resilience of functional groups in natural communities, few studies have examined in which way the roles and relationships of coexisting species change during community succession, a fundamental and natural process that follows the release of new resources in terrestrial and aquatic ecosystems. Variation in algal traits that characterize different phases and stages of community succession on rocky shores are likely to influence the magnitude, direction of effects, and the level of redundancy and complementarity in the diverse assemblage of herbivores. Two separate field experiments were conducted to quantify per capita and population effects and the functional relationship (i.e., redundancy or complementarity) of four herbivore species found in central Chile during early and late algal succession. The first experiment examined grazer effects on the colonization and establishment of early-succession algal species. The second experiment examined effects on the late-successional, dominant corticated alga Mazzaella laminarioides. Complementary laboratory experiments with all species and under natural environmental conditions allowed us to further characterize the collective effects of these species. We found that, during early community succession, all herbivore species had similar effects on the ephemeral algae, ulvoids, but only during the phase of colonization. Once these algae were established, only a subset of the species was able to control their abundance. During late succession, only the keyhole limpet Fissurella crassa could control corticated Mazzaella. The functional relationships among these species changed dramatically from redundant effects on ephemeral algae during early colonization, to a more complementary role on established early-successional algae, to a dominant (i.e., keystone) effect on late succession. This study highlights that functional relationship within consumer assemblages can vary at different phases and times of community succession. Differentiation in herbivore roles emphasizes the need to evaluate consumer's impacts through different times of community succession, and through experimental manipulations to make even broad predictions about the resilience or vulnerability of diverse intertidal assemblages to human disturbances.     

Competition among eucalyptus trees depends on genetic variation and resource supply

Genetic variation and environmental heterogeneity fundamentally shape the interactions between plants of the same species. According to the resource partitioning hypothesis, competition between neighbors intensifies as their similarity increases. Such competition may change in response to increasing supplies of limiting resources. We tested the resource partitioning hypothesis in stands of genetically identical (clone-origin) and genetically diverse (seed-origin) Eucalyptus trees with different water and nutrient supplies, using individual-based tree growth models. We found that genetic variation greatly reduced competitive interactions between neighboring trees, supporting the resource partitioning hypothesis. The importance of genetic variation for Eucalyptus growth patterns depended strongly on local stand structure and focal tree size. This suggests that spatial and temporal variation in the strength of species interactions leads to reversals in the growth rank of seed-origin and clone-origin trees. This study is one of the first to experimentally test the resource partitioning hypothesis for intergenotypic vs. intragenotypic interactions in trees. We provide evidence that variation at the level of genes, and not just species, is functionally important for driving individual and community-level processes in forested ecosystems.     

Competition,disturbance and local diversity patterns of substratum-bound clonal organisms: A simulation

We have conducted a series of simulations in order to compare the relative diversification effects of disturbance and intransitive (i.e., A > B, B > C but C > A) competition on a hypothetical set of clonal organisms. We found large significant differences between these two effects only when disturbance levels were low or absent. We suggest that natural assemblages should be examined for the frequency of intransitive interactions along disturbance gradients in order to assess the significance of these competitive relationships. We also argue that intransitive competition ought to be incorporated into nonequilibrium disturbance models; diversification effects of disturbance and intransitive competition need not be mutually exclusive.     

Biodiversity and Ecosystem Function

In at least some circumstances, biodiversity affects various ecosystem functions and the ways in which ecosystems respond to disturbance. Because these interactions occur at many spatial and temporal scales and throughout all levels of biological organization, it is difficult to decide where to focus attention on interactions between biodiversity and ecosystem function. The loci for initial attention is important for setting research priorities to understand these interactions further, for organizing known information to instruct the development of natural resource policies, and for identifying biodiversity conservation priorities. Holling (1992) argues that ecosystem behavior can be understood from a few dominating ecological processes that structure the ecosystem. In the temporal dimension, these key structuring processes dictate a few dominant temporal frequencies that drive other processes. Thus, the most effective strategy for studying interactions between biodiversity and ecosystem function is to focus on the key structuring processes at intermediate scales of space and time. Thereafter, other ecological conditions signify situations in which the interactions between biodiversity and ecosystem function are particularly strong: early to midsuccessional status, low soil fertility, intermediate levels of disturbance, biotic interactions only where there is collaborative indication of importance, invading species that differ significantly from native species in resource acquisition or utilization, and ecotones.     

Effects of Postfire Salvage Logging on Deadwood‐Associated Beetles

Abstract: In Canada and the United States pressure to recoup financial costs of wildfire by harvesting burned timber is increasing, despite insufficient understanding of the ecological consequences of postfire salvage logging. We compared the species richness and composition of deadwood‐associated beetle assemblages among undisturbed, recently burned, logged, and salvage‐logged, boreal, mixed‐wood stands. Species richness was lowest in salvage‐logged stands, largely due to a negative effect of harvesting on the occurrence of wood‐ and bark‐boring species. In comparison with undisturbed stands, the combination of wildfire and logging in salvage‐logged stands had a greater effect on species composition than either disturbance alone. Strong differences in species composition among stand treatments were linked to differences in quantity and quality (e.g., decay stage) of coarse woody debris. We found that the effects of wildfire and logging on deadwood‐associated beetles were synergistic, such that the effects of postfire salvage logging could not be predicted reliably on the basis of data on either disturbance alone. Thus, increases in salvage logging of burned forests may have serious negative consequences for deadwood‐associated beetles and their ecological functions in early postfire successional forests.     

Effects of Selective Logging on the Butterflies of a Bornean Rainforest

Abstract: Selective logging has been the main cause of disturbance to tropical forests in Southeast Asia, so the extent to which biodiversity is maintained in selectively logged forest is of prime conservation importance. We compared the butterfly assemblages of Bornean primary rainforest to those of rainforest selectively logged 6 years previously. We sampled by means of replicated transects stratified into riverine and ridge forests and we included roads in the logged forest. There was a three-fold variation in species richness and abundance over the 8-month sampling period. More species and individuals were observed in the logged forest, although between-replicate variability was high. Rarefied species richness was positively correlated with canopy openness within the range of disturbance levels encountered at our forest sites. Within families, there was no significant difference in the number of species between primary and logged forest. There was a significant difference in the relative abundance of species, but this was due largely to the abundance of one or two species. Community ordination separated the sites along a gradient of disturbance and revealed strong differences between riverine and ridge-forest butterfly assemblages in primary forest that were obscured in logged forest. There was no evidence that logging has resulted in a change in the composition of the butterfly assemblages from species with a local distribution to more widespread species. We conclude that at a logged forest site in close proximity to primary forest, low intensities of logging do not necessarily reduce the species richness or abundance of butterflies, although assemblage composition is changed.     

Density-dependent effects of ants on selection for bumble bee pollination in Polemonium viscosum

Mutualisms are commonly exploited by cheater species that usurp rewards without providing reciprocal benefits. Yet most studies of selection between mutualist partners ignore interactions with third species and consequently overlook the impact of cheaters on evolution in the mutualism. Here, we explicitly investigate how the abundance of nectar-thieving ants (cheaters) influences selection in a pollination mutualism between bumble bees and the alpine skypilot, Polemonium viscosum. As suggested in past work with this species, bumble bees accounted for most of the seed production (78% +/- 6% [mean +/- SE]) in our high tundra study population and, in the absence of ants, exerted strong selection for large flowers. We tested for indirect effects of ant abundance on seed set through bumble bee pollination services (pollen delivery and pollen export) and a direct effect through flower damage. Ants reduced seed set per flower by 20% via flower damage. As ant density increased within experimental patches, the rate of flower damage rose, but pollen delivery and export did not vary significantly, showing that indirect effects of increased cheater abundance on pollinator service are negligible in this system. To address how ants affect selection for plant participation in the pollination mutualism we tested the impact of ant abundance on selection for bumble bee-mediated pollination. Results show that the impact of ants on fitness (seed set) accruing under bumble bee pollination is density dependent in P. viscosum. Selection for bumble bee pollination declined with increasing ant abundance in experimental patches, as predicted if cheaters constrain fitness returns of mutualist partner services. We also examined how ant abundance influences selection on flower size, a key component of plant investment in bumble bee pollination. We predicted that direct effects of ants would constrain bumble bee selection for large flowers. However, selection on flower size was significantly positive over a wide range of ant abundance (20-80% of plants visited by ants daily). Although high cheater abundance reduces the fitness returns of bumble bee pollination, it does not completely eliminate selection for bumble bee attraction in P. viscosum.     

Biotic Impoverishment and Homogenization in Unfragmented Forest Understory Communities

Abstract:  Ecological change is often hard to document because of a lack of reliable baseline data. Several recent then-versus-now surveys of temperate forest and grassland communities demonstrate losses of local plant species, but most are based on data from a single site. We resurveyed understory communities in 62 upland forest stands in northern Wisconsin (U.S.A.) for which quantitative baseline data exist from 50 years ago. These stands are within a largely unfragmented region but vary in species composition and successional stage. We collected data on changes in (1) total and native species richness, (2) the ratio of exotic to native species, (3) the relative abundance of habitat generalists, and (4) community similarity among sites. We also compared how these rates of change varied over time. Over the past 50 years, native species density declined an average of 18.5% at the 20-m² scale, whereas the ratio of exotic species to native species increased at 80% of all sites. Habitat generalists increased, and habitat specialists declined, accounting in part for an 8.7% rise in average similarity in species composition among sites. Most of these changes cannot be related to succession, habitat loss, or invasion by exotic species. Areas without deer hunting showed the greatest declines in native species density, with parks and research natural areas faring no better than unprotected stands. Animal-pollinated and animal-dispersed species also declined, particularly at unhunted sites. These results demonstrate the power of quantitative multistand data for assessing ecological change and identify overabundant deer as a key driver of community change. Because maintaining forest habitats alone fails to preserve plant diversity at local scales, local biotic simplification seems likely to continue in the region unless active efforts are taken to protect diversity.     

Effects of Forest Fragmentation and Habitat Degradation on West African Leaf-Litter Frogs

Abstract:  Habitat degradation alters the dynamics and composition of anuran assemblages in tropical forests. The effects of forest fragmentation on the composition of anuran assemblages are so far poorly known. We studied the joint influence of forest fragmentation and degradation on leaf-litter frogs. We specifically asked whether the processes structuring leaf-litter anuran assemblages in fragmented forests are the same as those in continuous forests. We analyzed anuran assemblages with respect to habitat characteristics, including fragmentation and degradation parameters. In comparison with continuous forests, species richness and diversity were lower and assemblage composition was altered in forest fragments. These changes seemed to be mainly caused by habitat degradation rather than forest fragmentation. Availability of aquatic sites for breeding, vegetation structure (including those variables indicating degradation), and leaf-litter cover had the most influence on the presence of single species. The comparatively small impact of fragmentation on anurans might be due to the location of the study area; it still possessed large tracts of continuous forest. These forest blocks may stabilize the regional rainforest climate and thus weaken the effects of fragmentation .     

Species richness and stand stability in conifer forests of the Sierra Nevada

Theoretical and empirical studies have long suggested that stability and complexity are intimately related, but evidence from long-lived systems at large scales is lacking. Stability can either be driven by complex species interactions, or it can be driven by the presence/absence and abundance of a species best able to perform a specific ecosystem function. We use 64 years of stand productivity measures in forest systems composed of four dominant conifer tree species to contrast the effect of species richness and abundance on three stability measures. To perform this contrast, we measured the annual growth increments of > 900 trees in mixed and pure forest stands to test three hypotheses: increased species richness will (1) decrease stand variance, (2) increase stand resistance to drought events, and (3) increase stand resilience to drought events. In each case, the alternate hypothesis was that species richness had no effect, but that species composition and abundance within a stand drove variance, resistance, and resilience. In pure stands, the four species demonstrated significant differences in productivity, and in their resistance and resilience to drought events. The two pine species were the most drought resistant and resilient, whereas mountain hemlock was the least resistant and resilient, and red fir was intermediate. For community measures we found a moderately significant (P = 0.08) increase in the community coefficient of variation and a significant (P = 0.03) increase in resilience with increased species richness, but no significant relationship between species richness and community resistance, though the variance in community resistance to drought decreased with species richness. Community resistance to drought was significantly (P = 0.001) correlated to the relative abundance of lodgepole pine, the most resistant species. We propose that resistance is driven by competition for a single limiting resource, with negative diversity effects. In contrast resilience measures the capacity of communities to partition resources in the absence of a single limiting resource, demonstrating positive diversity effects.     

Crab herbivory regulates plant facilitative and competitive processes in Argentinean marshes

Interactions among plants have been hypothesized to be context dependent, shifting between facilitative and competitive in response to variation in physical and biological stresses. This hypothesis has been supported by studies of the importance of positive and negative interactions along abiotic stress gradients (e.g., salinity, desiccation), but few studies have tested how variation in biotic stresses can mediate the nature and strength of plant interactions. We examined the hypothesis that herbivory regulates the strength of competitive and facilitative interactions during succession in Argentinean marshes dominated by Spartina densiflora and Sarcocornia perennis. Spartina densiflora is preferred by the dominant herbivore in the system, the crab Chasmagnathus granulatus. We experimentally manipulated crab herbivory, plant structure, and shade, and we found that, when herbivory was low in the spring and summer, competitive interactions between plants were dominant, but in the fall, when herbivory was highest, facilitative interactions dominated, and Spartina densiflora survival was completely dependent upon association with Sarcocornia perennis. Moreover, experimental removal of Sarcocornia perennis across recently disturbed tidal flats revealed that, while Sarcocornia perennis positively affected small Spartina densiflora patches by decreasing herbivory, as patch size increases and they can withstand the impact of herbivory, competitive interactions predominated and Spartina densiflora ultimately outcompeted Sarcocornia perennis. These results show that herbivory can mediate the balance between facilitative and competitive processes in vascular plant communities and that the strength of consumer regulation of interactions can vary seasonally and with patch size.     

Recovery of Endemic Dragonflies after Removal of Invasive Alien Trees

Abstract:  Because dragonflies are very sensitive to alien trees , we assessed their response to large-scale restoration of riparian corridors. We compared three types of disturbance regime—alien invaded , cleared of alien vegetation , and natural vegetation (control)—and recorded data on 22 environmental variables. The most significant variables in determining dragonfly assemblages were percentage of bank cover and tree canopy cover , which indicates the importance of vegetation architecture for these dragonflies. This finding suggests that it is important to restore appropriate marginal vegetation and sunlight conditions. Recovery of dragonfly assemblages after the clearing of alien trees was substantial. Species richness and abundance at restored sites matched those at control sites. Dragonfly assemblage patterns reflected vegetation succession. Thus , initially eurytopic , widespread species were the main beneficiaries of the removal of alien trees , and stenotopic , endemic species appeared after indigenous vegetation recovered over time. Important indicator species were the two national endemics ( Allocnemis leucosticta and Pseudagrion furcigerum ) , which , along with vegetation type , can be used to monitor return of overall integrity of riparian ecology and to make management decisions. Endemic species as a whole responded positively to restoration , which suggests that indigenous vegetation recovery has major benefits for irreplaceable and widespread generalist species .     

How well do sunken vessels approximate fish assemblages on coral reefs? Conservation implications of vessel-reef deployments

The amount of artificial habitat (termed ??artificial reef??, AR) in marine systems is rapidly increasing, yet the effect of most types of AR on reef communities remains unknown. We examined the role of well-established vessel-reefs in structuring coral reef fish assemblages by comparing assemblages on 7 World War II wrecks (>65?years old) to those on interspersed coral patch reefs of comparable size in a tropical lagoon. Fish abundance, species richness, diversity and feeding guild structure on wrecks were similar to natural reefs; however, species composition differed between the two reef types (R?=?0.189?C0.341, average dissimilarity: 67.3?C68.8?%). Despite being more species-rich and diverse, fish assemblages on larger wrecks were less similar to assemblages on their adjacent natural reefs than smaller wrecks. Wrecks may also have affected fish abundance on adjacent natural reefs, with reefs adjacent to larger wrecks supporting higher abundances than reefs adjacent to smaller wrecks. Our results indicate that increases in vessel-reef habitat may not greatly affect reef fish assemblage parameters, but may affect the relative abundances of particular species.     

Temperature-associated population diversity in salmon confers benefits to mobile consumers

Habitat heterogeneity can generate intraspecific diversity through local adaptation of populations. While it is becoming increasingly clear that population diversity can increase stability in species abundance, less is known about how population diversity can benefit consumers that can integrate across population diversity in their prey. Here we demonstrate cascading effects of thermal heterogeneity on trout-salmon interactions in streams where rainbow trout rely heavily on the seasonal availability of anadromous salmon eggs. Water temperature in an Alaskan stream varied spatially from 5 degrees C to 17.5 degrees C, and spawning sockeye salmon showed population differentiation associated with this thermal heterogeneity. Individuals that spawned early in cool regions of the 5 km long stream were genetically differentiated from those spawning in warmer regions later in the season. Sockeye salmon spawning generates a pulsed resource subsidy that supports the majority of seasonal growth in stream-dwelling rainbow trout. The spatial and temporal structuring of sockeye salmon spawn timing in our focal stream extended the duration of the pulsed subsidy compared to a thermally homogeneous stream with a single population of salmon. Further, rainbow trout adopted movement strategies that exploited the multiple pulses of egg subsidies in the thermally heterogeneous stream. Fish that moved to track the resource pulse grew at rates about 2.5 times higher than those that remained stationary or trout in the reference stream with a single seasonal pulse of eggs. Our results demonstrate that habitat heterogeneity can have important effects on the population diversity of dominant species, and in turn, influence their value to species that prey upon them. Therefore, habitat homogenization may have farther-reaching ecological effects than previously considered.     

Community succession following massive ice-scour on a rocky intertidal shore: recruitment,competition and predation during early,primary succession

The importance of species interactions and recruitment variability was examined during the first year and a half of primary succession (1988–1989) on an exposed rocky seashore near Halifax, Nova Scotia. Previous work suggested that emergent rock on these shores is normally dominated by fucoid rockweeds because predatory whelks control the sessile animal competitors, mussels and barnacles, and because herbivorous littorinids control ephemeral algal competitors. Abundances of all species except seasonal ephemeral algae were very small throughout this experiment and we found no significant effects of carnivory, herbivory, plant-animal competition or plant-plant competition. A slight facilitation of Fucus recruitment is attributed to a thin mat of ephemeral, blue-green algae. Very few other studies have directly manipulated intertidal ephemeral algae. As primary succession may be very rare in this assemblage, these results may be specific to these circumstances, but they highlight the varying importance of species interactions with variable recruitment. In particular, it appears that variations in recruitment success may be important to community structure, even when recruitment is not limited by propagule supply. The scale of the study also provides insight into successional processes occurring after the recent, extensive ice-scour of exposed seashores in this region.