Using patterns of variability to test for multiple community states on rocky intertidal shores

Predictions based on theory of multiple stable states suggest that larger perturbations should lead to more unpredictable patterns of succession. This prediction was tested in the Gulf of Maine using data from 60 intertidal plots of varying size that were experimentally cleared of the rockweed Ascophyllum nodosum and from 14 benchmark sites from throughout the Gulf. Rockweed was removed from the experimental clearings ranging from 1 to 8 m in diameter in 1996 and data collected in 2004 were used to test effects of clearing size and location on divergence and variability in species composition. Benchmark data were collected in 2005, and the 14 sites were from a dataset on 53 sites throughout the Gulf of Maine. The selected sites were randomly chosen from all sites with > 80% canopy cover by A. nodosum and were expected to be similar to uncleared control plots from the experiment. Experimental removal of A. nodosum resulted in clearings at 12 sites within 4 bays. Abundances of gastropods, barnacles, mussels, and fucoid algae and the percentage cover of barnacles, mussels, fucoid algae, bare space, and other species were sampled. CAP and PERMDISP analyses revealed significant differences in multivariate dispersion and variability with both clearing size and location. Variability generally increased with clearing size and location effects were related to the north-south positioning of the sites. Benchmark sites were similar to the experimental control plots but as variable as the largest clearings. Results suggest that succession in larger clearings has been more unpredictable than in small clearings. The pattern of variability in the experimental clearings is consistent with the predictions of multiple stable states. However, the large amount of variation among the benchmark sites was due to mussels and was unexpected. This unexpected variability underscores the importance of sampling benchmark sites as part of experiments.     

Divergent succession and implications for alternative states on rocky intertidal shores

It has been hypothesized that rockweed stands and mussel beds in sheltered bays in the Gulf of Maine, USA, are alternative community states. As a test of this hypothesis, experimental clearings of different sizes were established in stands of the rockweed Ascophyllum nodosum (L.) Le Jolis to determine if successional changes in large clearings developed species assemblages distinctly different from the surrounding A. nodosum stands. Clearings ranging from 1 to 8 m in diameter were created at 12 sites in 4 bays on Swan's Island, Maine, in 1996 to mimic the effects of ice scour, and abundances of gastropods, barnacles, mussels and fucoid algae were monitored until 2002. ANOVAs and MDS showed strong effects of clearing size and divergent successional changes in large clearings. Large clearings were quickly filled in and remained dominated by the alga Fucus vesiculosus L. and the barnacle Semibalanus balanoides (L). There was no evidence for site-specific effects, and Mantel tests showed clearing size was a better predictor of species composition than geographic distances among sites. Results suggest that large pulse disturbances using clearings of 8 m in diameter can initiate divergent successional pathways and have a protracted effect on species composition. Results are also consistent with the hypothesis that mussel beds and rockweed stands in sheltered bays may be alternative community states.     

Survivorship of juvenile barnacles and mussels: spatial dependence and the origin of alternative communities

Mussels, barnacles, and rockweeds often form a distinct mosaic of patches on rocky intertidal shores, and it has been suggested that these communities may represent alternative community states. One way that alternative community states can arise is if early successional events are scale-dependent, but it is not known if juvenile survivorships of mussels and barnacles are, in fact, scale-dependent. Scale-dependence of barnacles (Semibalanus balanoides (L.)) and mussels (Mytilus edulis L.) was tested in the Gulf of Maine, USA. In winter 1997, clearings of 1, 2, 4 and 8 m in diameter and uncleared controls were made in stands of the rockweed Ascophyllum nodosum (L.) Le Jolis at 12 sites spread evenly across four bays on Swan's Island, ME. Summer and fall-winter survivorship of barnacles, which recruited in spring 1997, were estimated by tracking the 1997 cohort until late winter 1998. Survivorship of mussels was estimated from following the fate of transplanted juveniles over 4 days in late August 1997. Both barnacles and mussels showed better survival in 4 and 8 m clearings than in small clearings and controls. There was also significant variation in survivorship among sites. Densities of gastropods in the clearings did not reflect survivorship patterns of barnacles and mussels. Barnacle survivorship increased in fall and winter, and in large clearings was comparable to survivorship in barnacle-dominated habitats. Mussel survivorship was low in all clearing sizes suggesting that mussel beds develop slowly.     

Induced resistance in intertidal macroalgae modifies feeding behaviour of herbivorous snails

Herbivory in terrestrial and marine systems can induce changes in plant chemistry affecting the foraging behaviour of herbivores. A model based on terrestrial plant-herbivore interactions predicts herbivory-induced changes in leaf chemistry to be manifested in (1) increased herbivore mobility, (2) increased feeding dispersal and (3) reduced tissue consumption by herbivores. This study is the first to demonstrate that herbivory-induced changes in the tissue chemistry of the brown seaweed Ascophyllum nodosum elicit the same response in the feeding behaviour of the gastropod Littorina obtusata as predicted for herbivorous insects, providing good evidence for the models validity across different ecosystems. The potential benefit of increased feeding dispersal to terrestrial plants as suggested by the model is the prevention of concentrated damage to apical tissues thereby preserving the plants ability to compete for light; A. nodosum does not conform to these predictions. Increased dispersal of feeding damage on A. nodosum away from primary frond tissues would reduce the likelihood of frond breakage implying a fitness benefit of induced resistance.     

Tracking unstable states: ecosystem dynamics in a changing world

Ecological systems can show complex and sometimes abrupt responses to environmental change, with important implications for their resilience. Theories of alternate stable states have been used to predict regime shifts of ecosystems as equilibrium responses to sufficiently slow environmental change. The actual rate of environmental change is a key factor affecting the response, yet we are still lacking a non-equilibrium theory that explicitly considers the influence of this rate of environmental change. We present a metacommunity model of predator–prey interactions displaying multiple stable states, and we impose an explicit rate of environmental change in habitat quality (carrying capacity) and connectivity (dispersal rate). We study how regime shifts depend on the rate of environmental change and compare the outcome with a stability analysis in the corresponding constant environment. Our results reveal that in a changing environment, the community can track states that are unstable in the constant environment. This tracking can lead to regime shifts, including local extinctions, that are not predicted by alternative stable state theory. In our metacommunity, tracking unstable states also controls the maintenance of spatial heterogeneity and spatial synchrony. Tracking unstable states can also lead to regime shifts that may be reversible or irreversible. Our study extends current regime shift theories to integrate rate-dependent responses to environmental change. It reveals the key role of unstable states for predicting transient dynamics and long-term resilience of ecological systems to climate change.     

The effects of seaweed harvesting on fishes: a response

References Rangeley (1994) has argued that sampling biases, errors in experimental design and low statistical power make the conclusion by Black & Miller (1991), that there was no large impact of an experimental harvest of the seaweedAscophyllum nodosum on fishes, unmerited and misleading. Our study was conducted mainly at two sites close to Lower Argyle, southern Nova Scotia, Canada where harvesting ofAscophyllum nodosum has been carried out continuously for 30 years along-side intense and productive shellfish and finfish fisheries. In southern Nova Scotia, on-going removal ofAscophyllum nodosum with mechanical harvesters is about 17% (Sharpe & Semple 1991); planned annual removal in New Brunswick where Rangeley has worked will be only about 5% (Anon. 1993). The context of our investigation was a small-scale (400 m²) but extreme, experimental removal (100%) of the seaweed. Our focus was on fishes that moved into the intertidal zone as the tide rose in the evening and early morning and were caught in trammel nets, and on the fishes that were retained in a fixed seine that was set at high tide in early morning and the fishes removed at the next low tide. Thus, subject to the constraint on the size of the fishes imposed by the nature of our fishing gear with its stretch mesh of 25 mm, we were concerned with the identity and abundance of fishes that moved in and out of the intertidal zone and their behavioural response to the 400 m² patches of habitat that formed our experimental units. The most striking result was the unexpected, low abundance of fishes caught in our gear.     

The effect of a liquid seaweed extract from Ascophyllum nodosum (Fucales,Phaeophyta) on the two-spotted red spider mite Tetranychus urticae

This paper presents evidence from an initial trial in favor of claims that the liquid seaweed extract of Ascophyllum nodosum, marketed in the UK as Maxicrop, can reduce numbers of the two-spotted red spider mite Tetranychus urticae on crop plants. Seven applications of Maxicrop Triple over 20 days significantly suppressed the population build-up of two-spotted red spider mites on treated strawberry plants (Fragaria sp. var. Redgauntlet) raised under high polythene tunnels as compared to untreated controls.     

Detection of alternative stable states in marine communities

During the last 20 years there have been several attempts to test the theory of alternative stable states using marine systems. The results have been mixed, and there have been conflicting interpretations of the outcomes. In an attempt to resolve some of the problems, the theory and evidence for alternative stable states are reviewed. There are several different views of what is meant by alternative stable states, and so the key elements of the theory of alternative stable states are set in the context of marine systems. Appropriate experimental designs for detecting alternative states are discussed, and recent experimental studies in coral reefs, rocky intertidal systems, soft-sediment beds, and subtidal systems are briefly reviewed. Suggestions for improving experimental designs are proposed and unresolved issues are highlighted.     

Oscillating trophic control induces community reorganization in a marine ecosystem

Understanding how climate regulates trophic control may help to elucidate the causes of transitions between alternate ecosystem states following climate regime shifts. We used a 34-year time series of the abundance of Pacific cod ( Gadus macrocephalus ) and five prey species to show that the nature of trophic control in a North Pacific ecosystem depends on climate state. Rapid warming in the 1970s caused an oscillation between bottom–up and top–down control. This shift to top–down control apparently contributed to the transition from an initial, prey-rich ecosystem state to the final, prey-poor state. However, top–down control could not be detected in the final state without reference to the initial state and transition period. Complete understanding of trophic control in ecosystems capable of transitions between alternate states may therefore require observations spanning more than one state.     

Effects of harvesting onAscophyllum nodosum (L.) Le Jol. (Fucales,Phaeophyta): a demographic approach

Although populations ofAscophyllum nodosum are harvested commercially, little is known about the effects on demographic vital rates (growth, reproduction, survival). This study examines the effects of harvesting season and harvesting intensity on growth, reproduction and mortality of intact fronds in four size classes and in fronds truncated by the harvest. Knowledge of size-specific vital rates was used to evaluate the response of the population to harvesting.Harvesting season and harvesting intensity did not exert a significant effect on growth. Growth in plots not subject to harvesting was less than in harvested plots. No major differences in growth, reproduction and survival between intact and severed fronds emerged. The number of fronds attaining reproduction was enhanced by increased harvesting intensity and by cutting in summer. Harvesting did not seem to induce breakage, and breakage appeared higher in the uncut plots. Most harvesting treatments did not influence survivorship and survivorship was similar among all size classes. Growth rates were inversely related to sizes of fronds.Assessment of variation across size classes yielded more accurate estimates of growth rates than those of previously used methods. Accurate size class specific-growth rates will be a useful criterion when regulating intervals between harvests. Furthermore, assessment of size-specific vital rates allows identification of the frond size classes most relevant to the preservation of resources. Because of their fast growth rates and abundance, fronds in class 1, and, to a lesser extent, class 2, are responsible for most of the population regrowth after harvest. In contrast, classes 3 and 4 contribute little to recovery. This finding provides a strong basis for a harvesting strategy that targets the largest fronds.Author for correspondence     

Reduction of nitrate leaching from soil treated with an Ascophyllum nodosum based soil conditioning agent

A commercially available soil conditioning agent prepared from a brown marine alga, Ascophyllum nodosum, was tested for its ability to effect leaching rates of nitrate from two soil types. The use of the material led to a significant (p < 0.05) reduction of nitrate leaching from a soil composed of sand and compost (50:50). This revised version was published online in August 2006 with corrections to the Cover Date.     

Effects of small-scale disturbances of canopy and grazing on intertidal assemblages on the Swedish west coast

The effects of small-scale disturbances (80×30-cm plots) of canopy and grazers on intertidal assemblages were investigated in this 4-year experiment on sheltered rocky shores on the Swedish west coast. Canopy disturbances due to ice scouring were mimicked by removal of adult plants of the seaweed Ascophyllum nodosum (L.) Le Joli. Density of the main epilithic grazing gastropods, Littorina spp., was lowered by exclosure and handpicking. Based on earlier experiments in other areas, the general hypothesis was that canopy removal and grazer exclosure, alone or in combination, should increase the recruitment of A. nodosum or other fucoid juveniles, and change the structure of the understorey assemblage.There was an effect of canopy removal on the development of this assemblage, lasting for more than 31 months. Both increased and decreased abundances of species were found as short-term effects, but there was also a longer-term effect with increased abundance. Grazer exclosure was only effective in combination with canopy removal, causing a short-term increase in ephemeral green algae. Short-term effects of canopy removal were also the increase in recruitment of Semibalanus balanoides (Linnaeus) and the decrease of the red alga Hildenbrandia rubra (Sommerfelt) Meneghini. Fast recruitment and growth of fucoid species (Fucus serratus L. and F. vesiculosus L.) restored the canopy and conditions of the understorey within 18 months. Thus, the canopy removal changed the physical conditions for the understorey, making it possible for other species to coexist in this community. Surprisingly, no effect of canopy removal or grazer exclusion was found on the recruitment of juvenile A. nodosum, neither by canopy removal nor grazer exclosure. The lack of such effects might be due to the early mortality caused by other grazers (small, mobile crustaceans), or to the low density of periwinkles on these shores. However, despite the patchy and generally low recruitment of A. nodosum juveniles, observations suggested that the cover of A. nodosum in manipulated patches would return to initial levels, either by recruitment or regrowth of small holdfasts and from growth of edge plants.     

Enhanced leaf chlorophyll levels in plants treated with seaweed extract

Application to the soil of an aqueous alkaline extract ofAscophyllum nodosum resulted in higher concentrations of chlorophyll in the leaves of treated plants in comparison to control plants treated with an equivalent volume of water. Positive results were obtained with all species tested (tomato, dwarf French bean, wheat, barley, maize). When the seaweed extract was applied as a foliar spray, similar effects on leaf chlorophyll contents were obtained, except in the case of dwarf French bean plants, for which no significant difference was recorded between test and control plants. When the betaines present in the seaweed extract were applied as a mixture in the same concentrations as those in the diluted seaweed extract (-aminobutyric acid betaine 0.96 mg L^–1, -aminovaleric acid betaine 0.43 mg L^–1, glycinebetaine 0.34 mg L^–1), very similar leaf chlorophyll levels were recorded for the seaweed extract and betaine treated plants. This suggests strongly that the enhanced leaf chlorophyll content of plants treated with seaweed extract is dependent on the betaines present.     

Interactions among invaders: community and ecosystem effects of multiple invasive species in an experimental aquatic system

With ecosystems increasingly supporting multiple invasive species, interactions among invaders could magnify or ameliorate the undesired consequences for native communities and ecosystems. We evaluated the individual and combined effects of rusty crayfish (Orconectes rusticus) and Chinese mystery snails [Bellamya (=Cipangopaludina) chinensis] on native snail communities (Physa, Helisoma and Lymnaea sp.) and ecosystem attributes (algal chlorophyll a and nutrient concentrations). Both invaders are widespread in the USA and commonly co-occur within northern temperate lakes, underscoring the importance of understanding their singular and joint effects. An outdoor mesocosm experiment revealed that while the two invaders had only weakly negative effects upon one another, both negatively affected the abundance and biomass of native snails, and their combined presence drove one native species to extinction and reduced a second by >95%. Owing to its larger size and thicker shell, adult Bellamya were protected from crayfish attack relative to native species (especially Physa and Lymnaea), suggesting the co-occurrence of these invaders in nature could have elevated consequences for native communities. The per capita impacts of Orconectes (a snail predator) on native snails were substantially greater than those of Bellamya (a snail competitor). Crayfish predation also had a cascading effect by reducing native snail biomass, leading to increased periphyton growth. Bellamya, in contrast, reduced periphyton biomass, likely causing a reduction in growth by native lymnaeid snails. Bellamya also increased water column N:P ratio, possibly because of a low P excretion rate relative to native snail species. Together, these findings highlight the importance of understanding interactions among invasive species, which can have significant community- and ecosystem-level effects.     

Experimental confirmation of aggressive mimicry by a coral reef fish

A number of potential mimetic relationships between coral reef fishes have been described, but the underlying mechanisms are poorly understood. Similarities in colour between species have often been attributed to aggressive mimicry (where predators resemble models in order to deceive prey), however this has not been tested. The fang blenny, Plagiotremus rhinorhynchos is a specialized predator that feeds on tissues of other fishes. Some individuals appear to mimic the harmless cleaner wrasse Labroides dimidiatus in order to deceive fish visiting cleaning stations, thereby increasing access to food. In this study, the ecological relationship between the mimic and model was examined at Kimbe Bay (Papua New Guinea) and the hypothesis that colour similarities represent facultative aggressive mimicry was experimentally evaluated. Some juveniles exhibited a striking resemblance to the juvenile colouration of the cleaner wrasse, but only when in close proximity to the wrasse and only when similar in size. As predicted for mimics, P. rhinorhynchos co-occurred with L. dimidiatus, but was rare relative to the model. Among site comparisons showed that the abundance of mimetic type blennies was positively correlated with the abundance of juvenile cleaner wrasses. Approximately 50% of all P. rhinorhynchos were found 1 m from the nearest L. dimidiatus, a distance significantly shorter than expected if they were not associated. A cleaner wrasse removal experiment was carried out to test whether the colour displayed by the blenny and its foraging success were contingent upon the presence of a model. In all cases, removal of the model prompted a rapid colour change to a general non-mimetic colouration in P. rhinorhynchos. Removal of L. dimidiatus also resulted in a ~20% reduction in the average foraging success of the blenny compared to controls, supporting the hypothesis that the blenny is a facultative aggressive mimic of the cleaner wrasse.     

The effects of seaweed harvesting on fishes: a critique

Synopsis Black & Miller (1991) concluded that there was no large impact of an experimental harvest of rockweed,Ascophyllum nodosum, on fishes. A critique of their study demonstrates that this conclusion cannot be substantiated by their data because of sampling biases, errors in experimental design and low statistical power. Further, evidence is presented which supports the hypothesis that rockweed may provide an important nursery habitat for juvenile fishes.     

Quantitative estimation of betaines in commercial seaweed extracts using overpressured layer chromatography

An off-line, overpressured layer chromatographic procedure has been developed for the evaluation of betaines in commercial seaweed extracts using Dragendorff's reagent for the detection of the compounds and densitometry for their quantitative estimation. Using continuous development and a low sample volume, the amounts of -aminobutyric acid betaine and -aminovaleric acid betaine can be estimated, but a larger sample volume is necessary for the estimation of glycinebetaine and minor betaines.The critical steps of this analytical method are the visualization and the quantitative evaluation of the spots produced. Temperature plays a major role in the resolution, sensitivity and precision after derivatization with Dragendorff's reagent; this has led to the adoption of standardized conditions.Author for correspondence     

Comparison of the structure of populations of Ascophyllum nodosum (Fucales,Phaeophyta) at sites with different harvesting histories

Changes in the structure of the Ascophyllum nodosum population at Pubnico, southwest Nova Scotia, Canada, at an experimental site subjected to mechanical harvest and at two control sites never subjected to mechanical harvest were monitored from 1991 to 1994. A bimodal population structure measured in terms of plant length was characteristic of all these sites before the experiment. The population structure of the experimental site became unimodal immediately after experimental harvest by machine and remained unimodal for the subsequent two years. However, a bimodal population structure began to appear in the third year. A bimodal population structure remained evident at the control site with bedrock as the substratum but was less evident at the other control site where the substratum is made up of boulders and cobbles. Movement of loose rocks with rockweeds still attached may have contributed to the less distinct modal structure of this control site. Other sites with different harvesting histories monitored in the summer of 1992 showed some interesting patterns. A unimodal population structure was evident in Argyle Sound and Pubnico Point South and at Charlesville, which had been harvested one and two years before, respectively. A bimodal population structure was more evident at Frenchman's Point, which had been harvested three years prior. The rate of change from a unimodal to a bimodal population structure may depend on the intensity of harvest. Extensive canopy removal in intensively harvested areas may be conducive to an influx of recruits and to regeneration from the holdfast. Hence, plant length modal structure may be a useful measure of the relative state of recovery of a harvested population.     

Drift and upstream movement of invertebrates in a springbrook community ecosystem

Structure, drift, and upstream movement of populations of benthic macroinvertebrates, in particular Synurella dentata Hubricht and Lirceus fontinalis Raf., were examined within a temperate spring ecosystem. Chemical and physical aspects of the springbrook were also investigated and life histories of the gammarids and asellids noted.Chemically and physically the spring proved both constant and predictable, much more so than other lotic systems.Species diversity was low from November through February and increased in March, April, and May. Equitability followed the same trends as species diversity. Both indices were most affected by large fluctuations in the populations of aquatic insects.Significant changes in the numbers of amphipods, isopods, and total macroinvertebrates was evident over a seven month period. Males were present in the isopod population year-round, but only from November to January in the amphipod population. Breeding by the isopods occurred throughout the year and peaked during winter. Amphipods copulated only in the late fall and early winter.Significant diel peaks in the amphipod, isopod, and total invertebrate drift negatively correlated with light intensity levels. Amphipods and isopods did not exhibit any preferential upstream movement during either the day or night; however, total macroinvertebrate upstream movement was greater at night. The total number of invertebrates moving upstream were lower than values reported from other lotic environments.     

Changes in the population structure of Ascophyllum nodosum (L.) Le Jolis due to mechanical harvesting

The use of a Norwegian suction cutter to harvest populations of the brown alga Ascophyllum nodosum (L.) Le Jolis in southwestern Nova Scotia started in 1985. The impact of this type of mechanical harvest on the algal population structure was evaluated. Changes in the length and density of individual plants (clumps) within 0.25 m^–2 quadrats, as well as the length of individual shoots within clumps were monitored before and after an experimental harvest. The mechanical harvest cut mainly the longer plants, thus changed the initial bimodal size structure of the population to unimodal. There was a 20 to 36% plant mortality, reducing the plant density from 92.6 to 73.6 individuals m^–2. Pre-harvest size distribution of the tagged shoots was skewed to the right and became more normal after the harvest. Tagged shoots in the harvested quadrats suffered a 42% mortality as compared to 11% of those in the control quadrats. An understanding of the impact of the mechanical harvesting on the harvested population is essential in the design of a management strategy. Sources of variation in the impact of mechanical harvest include the tide level at time of harvest, length of time the machine operated in one site, skill of the machine operator, and sharpness of the machine cutting blades.