High assemblage persistence in heterogeneous habitats: an experimental test with stream benthic algae

1. The persistence of biological assemblages is positively affected by spatial heterogeneity. This influence may be indirect, through increased species richness. Another possibility is the increased availability of refuges from disturbances, which would prevent local loss. 2. We conducted a field experiment to test the hypothesis that greater roughness (a form of spatial heterogeneity) on the surface of substrata allows higher persistence of assemblages of stream benthic algae and that this relationship does not depend on species richness. Samples were taken on six occasions from smooth and rough artificial substrata used for algal colonisation. We calculated the persistence of assemblages using two analytical approaches: the mean distance to group centroid and the sum of the Euclidean distances between consecutive sampling occasions, both in a multivariate space. We also subsampled the data to take into account differences in species richness between treatments and thus to evaluate the effect of species richness on persistence. 3. Assemblages on rough substrata were more persistent than assemblages on smooth substrata. The effects detected were not due to the greater species richness on rough substrata, since a higher persistence of the assemblages on rough substrata remained after the subsampling procedures. 4. Our results indicate a strong positive relationship between substratum roughness and the persistence of stream benthic algal assemblages. We suggest that this is due to the presence of physical refuges in heterogeneous habitats.     

LINKING BENTHIC ALGAL BIOMASS TO STREAM SUBSTRATUM TOPOGRAPHY1

The physical properties of substrata significantly influence benthic algal development. We explored the relationships among substratum surface texture and orientation with epilithic microphytobenthic biomass accumulation at the whole‐substratum and micrometer scales. Unglazed clay tiles set at three orientations (horizontal, vertical, and 45°), and six substrata of varying surface roughness were deployed in a prairie stream for 3 weeks. Substrata were analyzed for loosely attached, adnate, and total benthic algal biomass as chl a, and confocal laser scanning microscopy was used to measure substrata microtopography (i.e., roughness, microscale slope angles, and three‐dimensional surface area). At the whole‐substratum level, vertical substrata collected significantly (P < 0.05) less algal biomass, averaging 34% and 36% less than horizontal and 45° substrata, respectively. Benthic algal biomass was also significantly less on smoother surfaces; glass averaged 29% less biomass than stream rocks. At the microscale level, benthic algal biomass was the greatest at intermediate values, peaking at a mean roughness of approximately 17 μm, a mean microscale slope of 50°, and a projected/areal surface area ratio of 2:1. The proportion of adnate algae increased with surface roughness (26% and 67% for glass and brick, respectively), suggesting that substratum type changes the efficiency of algal removal by brushing. Individual substrata and microsubstrata characteristics can have a strong effect on benthic algae development and potentially affect reach scale algal variability as mediated by geomorphology.     

How protective are refuges? Quantifying algal protection in rock crevices

1. Refuges can be functionally important if they harbour sufficient organisms during disturbance to augment population recovery. I quantified the protection of stream algae in crevice refuges using the applied, severe disturbance of scrubbing. Scrubbing effectively removed visible surface algae, and algae remaining on stones were considered protected. 2. In a field experiment, substrata with different quantities of crevice (glass bottles; greywacke; schist; pumice) were incubated in a channelised stream. The possible influence of growth conditions was investigated concurrently; half of the substrata were suspended in the water column, the rest were placed on the bottom (providing differential access to grazers and exposing them to different flow conditions). 3. Rougher substrata had greater total algal biomass than smoother substrata; this pattern resulted from more algal biomass in crevices of the rougher substrata. Protection from scrubbing ranged from about 5% of total algal biomass on glass and greywacke to 80% on pumice. In contrast, algal biomass removed by scrubbing was similar among the experimental substrata. Suspended substrata had more chironomid grazers than those on the bottom, and also greater algal biomass, possibly because of high algal concentrations in the chironomid retreats. 4. A survey of stones from three rivers supported the experimental results; namely, rough pumice protected more algae from scrubbing than did smoother greywacke. 5. In a separate experimental assay, there was no difference in algal growth on agar plates with and without added powdered rock substrata, suggesting that crevice characteristics and not substratum chemistry produced the differences in algal assemblages between rough and smooth surfaces. 6. Results indicate that rough stream stones may protect sufficient algae to augment their recovery in streams following disturbance.     

Substratum simplification reduces beta diversity of stream algal communities

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Flood disturbance effects on benthic diatom assemblage structure in a semiarid river network

Disturbances such as floods and droughts play a central role in determining the structure of riverine benthic biological assemblages. Extreme disturbances from flash floods are often restricted to part of the river network and the magnitude of the flood disturbance may lessen as floods propagate downstream. The present study aimed to characterize the impact of summer monsoonal floods on the resistance and resilience of the benthic diatom assemblage structure in nine river reaches of increasing drainage size within the Gila River in the southwestern United States. Monsoonal floods had a profound effect on the diatom assemblage in the Gila River, but the effects were not related to drainage size except for the response of algal biomass. During monsoons, algal biomass was effectively reduced in smaller and larger systems, but minor changes were observed in medium systems. Resistance and resilience of the diatom assemblage to floods were related to specific species traits, mainly to growth forms. Tightly adhered, adnate and prostrate species (Achnanthidium spp., Cocconeis spp.) exhibited high resistance to repeated scour disturbance. Loosely attached diatoms, such as Nitzschia spp. and Navicula spp., were most susceptible to drift and scour. However, recovery of the diatom assemblage was very quick indicating a high resilience, especially in terms of biomass and diversity. Regional hydroclimatic models predict greater precipitation variability, which will select for diatoms resilient to bed‐mobilizing disturbances. The results of this study may help anticipate future benthic diatom assemblage patterns in the southwestern United States resulting from a more variable climate.     

Factors influencing algal species assemblages on reef and cobble substrata off Ghana

Subtidal algal assemblages were studied on two substrata, rocky reefs and calcareous cobbles. Although the two occur together at comparable depths, their vegetation differs in species composition and richness, and in patterns of plant size, life form, and longevity. The reef bears a species-rich, patchy cover of small filamentous and crustose forms, with occasional clumps of more robust species. The cobbles support a sparse cover of large leafy and dendritic species in addition to many of the smaller species found on the reef. The floristic separation arises from differential establishment and survival of species under conditions of (1) grazing by fish and urchins (on the reef only), and (2) seasonal physical disturbance during storms leading to the removal of most algae (on the cobbles only). Both substrata show a seasonal floristic cycle, but the trend is more pronounced on cobbles. Species do not depart from randomness in their patterns of co-occurrence on individual cobbles or reef fragments. Interspecific competition appears comparatively unimportant in determining species composition on either substratum.     

The effect of two scales of habitat architecture on benthic grazing in a river

1. This experiment studied the effects of differing levels of the complexity of substratum architecture at two spatial scales on the distribution and abundance of benthic algae and invertebrates, and the strength of the trophic interaction between invertebrate grazers and algae. Some estimates of the effects on invertebrate colonization rates were also made. 2. Four levels of microhabitat architectural complexity were created using artificial substrata (clay tiles) and placed in Mountain River, Tasmania, in two riffle types (bedrock and boulder-cobble) of differing large-scale substratum complexity. After a colonization period, invertebrate grazers were removed from half the tiles to measure the effects of grazing. Invertebrates on the tiles were also counted and identified. At the end of the experiment, algae were removed from the tiles and analysed for chlorophyll a. 3. Invertebrate grazers did not reduce algal biomass during the experiment, and microhabitat-scale architecture influenced algal biomass more strongly than riffle-scale architecture. Highly complex microhabitat architecture increased algal biomass by providing more surface area, but once standardized for surface area, algal biomass decreased as the complexity of microhabitat architecture increased. 4. Microhabitat-scale architecture was also predominant in determining invertebrate density and the identity of the dominant grazer species. In contrast to algal biomass, invertebrate densities and species density increased with the complexity of microhabitat architecture, suggesting that refuges from flow (and possibly predation) were as important to river invertebrates as the distribution of their food source. 5. Riffle-scale architecture had some effect on the colonization of two slow-moving grazer taxa, but, overall, the colonization processes of slow-moving grazers were determined mostly by the complexity of microhabitat-scale architecture.     

Stream geomorphology regulates the effects on periphyton of ecosystem engineering and nutrient enrichment by Pacific salmon

1. Pacific salmon (Oncorhynchus spp.) returning to streams deliver substantial quantities of nutrients (nitrogen and phosphorus) that may stimulate primary production. Salmon can also affect the phytobenthos negatively via physical disturbance during nest excavation, a process that may counteract the positive effects of salmon‐derived nutrients on benthic algae. The ability of salmon to disturb benthic habitats may be a function of substratum particle size, and therefore, the geomorphology of streams could determine the net effect of salmon on benthic communities. 2. Based on surveys of 17 streams in southwest Alaska before the salmon run and during peak salmon density, we identified size thresholds for the disturbance of substratum particles by salmon and classified particles as vulnerable (<60 mm B‐axis), invulnerable (>110 mm) or transitional (61–110 mm). At the scale of individual rocks, algal biomass on vulnerable substrata decreased at peak spawning (relative to values before the run) as a power function of salmon density; transitional and invulnerable substrata showed no quantifiable pattern. However, invulnerable substrata in streams with more than 0.11 salmon m^?2 showed net algal accrual, or relatively smaller declines in algal biomass, than vulnerable substrata, indicating that large rocks provide refuge for benthic algae from salmon disturbance. 3. We expected that streams with proportionally larger rocks would respond positively to salmon at the whole‐stream scale, after accounting for the relative abundance of rocks of different sizes within streams. Invulnerable rocks made up only 0–12% of the total substratum particle size distribution in salmon‐bearing streams, however, and algal accrual on invulnerable substrata did not outweigh the strong disturbance effects on the more spatially extensive vulnerable substrata. The change in whole‐stream benthic algal biomass among streams was negatively related to salmon density. 4. Stable isotopes of nitrogen (δ¹⁵N) were used to track nutrients from salmon into benthic biota. Periphyton δ¹⁵N on rocks of all size classes was higher at peak salmon spawning than before the salmon run, indicating the uptake of salmon‐derived nitrogen. Peak δ¹⁵N values were positively related to salmon abundance and followed a two‐isotope mixing relationship. The per cent of N from salmon in periphyton was also related to salmon density and was best explained by a saturating relationship. Spring δ¹⁵N was unrelated to salmon returns in the previous year, suggesting little annual carryover of salmon nutrients.     

Local disturbance history affects patchiness of benthic river algae

1. Recent research has shown that high‐flow events in streams leave a small‐scale mosaic of bed patches that have experienced scouring, sediment deposition (fill), or remained stable. Few studies have investigated if this ‘local disturbance history’ contributes to the patchy distribution of benthic organisms in streams and rivers. 2. In the present research, we demonstrate that local disturbance history in a mid‐sized river can have both short‐ and long‐term effects on epilithic algae. Chains buried vertically in the substratum of the river bed (236 in a 800‐m reach) indicated that two floods (return periods ≤1 year) caused a mosaic of bed patches with different disturbance histories. Once after the first and twice after the second flood, we sampled epilithic algae (mainly diatoms) in replicate patches that had been scoured, filled, or remained stable during the respective event. Algal biomass and cell density per substratum area were determined. 3. Three months after the first flood, algal biomass, total diatom density, diatom taxon richness, and densities of six of nine most common taxa were highest in fill patches. Six days after the second flood, biomass was highest in stable patches, indicating a refugium function of these patches. The refugium patches consisted of average‐sized stones, in contrast to previous studies of flood refugia for benthic algae in which these refugia were always large and/or immobile substrata. Four weeks after the second flood, diatoms tended to be most abundant in scour patches. With one exception, these differences between patch types could not be attributed to differences in local near‐bed current velocity or water depth. 4. The effects of disturbance history were more complex than a simple refugium function of stable patches because algal patterns changed with time since the last disturbance, possibly depending on the successional state of the algal mats.     

Periphyton distribution and abundance on substrata of different size along a gradient of stream trophy de Montréal

In stream, substrata of different size present different degree ofstability, current, erosion, and deposition to colonizingorganisms. In this study, we tested the importance of substratumsize ranging from sand to small boulders for periphytondistribution and abundance. Because trophy strongly affect streamorganisms, we sampled at nine sites chosen to represent the rangeof nutrients typical of Eastern Ontario and Western Québec. Alarge part of the variability in algal biomass (as chlorophyll)among sites was explained by trophy (as seston or totalphosphorus). However, there was also an effect of substratum size.Cobbles had the highest biomass, and gravel the lowest; sand andboulders were intermediate. Assemblages on different substrata weredifferent in taxonomy and life forms. Cyanobacterial colonies andmotile diatoms dominated the finer substrata while adnate andfilamentous algae were more developed on the larger ones.Consequently periphyton on fine sediments was more loosely attachedthan on rocks. Average algal size was not related to substratumsize but increased significantly with trophy confirmingobservations in benthic and planktonic assemblages inlakes.     

Effects of habitat-conditioning by the damselfish Stegastes nigricans (Lacepède) on the community structure of benthic algae

We compared the community structure of benthic algae inside and outside pomacentrid damselfish (Stegastes nigricans) territories in a moat at Sesoko Island, Okinawa, Japan. S. nigricans maintained “algae farms” that were dominated by the filamentous rhodophyte, Womersleyella setacea. Species richness and biomass were higher inside damselfish territories than outside, while species diversity and evenness were higher outside. Detrended correspondence analysis (DCA) based on species composition showed that the dominance of W. setacea was maintained throughout the year in all samples collected from inside damselfish territories. The observed strong dominance of filamentous rhodophytes was consistent with the findings of most studies on damselfish territories worldwide. However, the dominance of a single species of alga and low species diversity inside the territories was in contrast to the findings of previous studies, in which the reduction of grazing pressure caused intermediate disturbance and enhanced algal species diversity. This discrepancy in algal species diversity inside the damselfish territories seems to have been caused by unique characteristics of the alga and the fish. W. setacea traps sediment, which reduces the availability of firm substrata for attachment and inhibits the recruitment of some algae. Moreover, S. nigricans “weeds” indigestible calcareous and thicker algae. The algal assemblage outside damselfish territories varied among samples, and included mat-forming cyanophytes (Calothrix aeruginosa and Calothrix codicola), a prostrate laminar phaeophyte (Padina sp.), thin and small-scaled algae (Cladophora sp. and Feldmannia indica), finely branched filamentous rhodophytes (Taenioma perpusillum and Herposiphonia obscura), and a coarsely branched rhodophyte (Gelidiopsis variabilis).

We placed artificial slate plates inside and outside damselfish territories, and showed that the W. setacea inside territories gradually increased in biomass, reaching the same levels of biomass and dominance as W. setacea on natural substrata. Outside the territories, the algal assemblage underwent succession from early colonizers, i.e., thin and small-scaled algae, to grazing-resistant algae such as mat-forming cyanophytes and prostrate laminar Padina sp. Under heavy grazing, the flora outside the territories was composed of early colonizers, grazing-resistant algae, and scattered erect algae that had probably escaped grazing by chance.

Our findings suggest that sediment trapped by the turf of W. setacea inhibited recruitment of some algae, and that moderate cropping and selective weeding by S. nigricans excluded grazing-resistant algae and prevented early colonizers and competitively superior algae from out-competing W. setacea. Consequently, low species diversity and a high-biomass “farm” suitable for harvesting was maintained.     

Adhesion of algal cells to surfaces

This paper reports the cell–substratum interactions of planktonic (Chlorella vulgaris) and benthic (Botryococcus sudeticus) freshwater green algae with hydrophilic (glass) and hydrophobic (indium tin oxide) substrata to determine the critical parameters controlling the adhesion of algal cells to surfaces. The surface properties of the algae and substrata were quantified by measuring contact angle, electrophoretic mobility, and streaming potential. Using these data, the cell–substratum interactions were modeled using thermodynamic, DLVO, and XDLVO approaches. Finally, the rate of attachment and the strength of adhesion of the algal cells were quantified using a parallel-plate flow chamber. The results indicated that (1) acid–base interactions played a critical role in the adhesion of algae, (2) the hydrophobic alga attached at a higher density and with a higher strength of adhesion on both substrata, and (3) the XDLVO model was the most accurate in predicting the density of cells and their strength of adhesion. These results can be used to select substrata to promote/inhibit the adhesion of algal cells to surfaces.     

Pathways for algal recolonization in seasonally‐flowing streams

1. In semi‐arid climates, seasonally‐flowing streams provide most of the water required for human use, but knowledge of how water extraction affects ecological processes is limited. Predicted alterations in stream flows associated with the impacts of climate change further emphasize the need to understand these processes. Benthic algae are an important base for stream food webs, but we have little knowledge of how algae survive dry periods or respond to altered flow regimes. 2. We sampled 19 streams within the Grampians National Park, south‐eastern Australia and included four components: a survey of different drought refuges (e.g. permanent pools, dry biofilm on stones and dry leaf packs) and associated algal taxa; a survey of algal regrowth on stones after flows recommenced to determine which refuges contributed to regrowth; reciprocal transplant experiments to determine the relative importance of algal drift and regrowth from dry biofilm in recolonization; direct measurement of algal drift to determine taxonomic composition in relation to benthic assemblage composition. 3. Algae showed little specificity for drought refuges but did depend on them; no species were found that were not present in at least one of the perennial pool, dry biofilm or leaf pack refuges. Perennial pools were most closely correlated with the composition of algal assemblages once flows resumed, but the loss or gain of perennial pools that might arise from stream regulation is unlikely to affect the composition of algal regrowth. However, regulated streams were associated with strong increases in algal density in dry biofilm, including increased densities of Cyanobacteria. 4. A model for algal recolonization in seasonally‐flowing streams identified three pathways for algal recolonization (drift‐dependent, dry biofilm‐dependent and contributions from both), depending on whether streams are diatom‐dominated or dominated by filamentous algae. The model predicted the effects of changes to stream flow regimes on benthic algal recolonization and provides a basis for hypotheses testable in streams elsewhere.     

Scale-dependant control of motile epifaunal community structure along a coral reef fishing gradient

Large-scale fishing is mostly conducted using towed gears that reduce the biomass and diversity of benthic invertebrates. However, it is impossible to differentiate between the physical disturbance effect of towed gears from the effect of fish predator removal upon benthic invertebrate communities. Here we explore the impact of fish removal alone on the community structure of small motile coral reef invertebrates (epifauna) along a subsistence fishing intensity gradient in the Lau group, Fiji. We deployed settlement plates at three areas in each of six fishing grounds and examined the density and class richness of the motile epifaunal communities and the associated algal communities in relation to the structure of fish and benthic communities. Motile epifaunal density was unrelated to fishing intensity. However, at smaller inter-area scale (0.5-10 km) motile epifaunal density was negatively related to plate algal biomass, whereas at the larger inter-fishing-ground scale (4-180 km) motile epifaunal density was positively related to the rugosity (substrate complexity) of the surrounding benthos. The class richness and diversity (Margalef's d) of motile epifaunal communities were negatively related to fishing intensity, but unrelated to grazing intensity, rugosity or algal biomass at either scale. Benthic community structure varied significantly with fishing intensity; hard-coral cover was lower and turf-algal cover was higher at high fishing pressure. The variation in benthic community structure was associated with variation in fish community structure, which in turn varied with fishing intensity. Motile epifaunal community structure upon plates was linked to the structure of the surrounding benthic community, but was not directly linked to the plate algal community. We suggest the decline in richness of the motile epifauna community along the fishing gradient is attributable to either to exploiter-mediated coexistence or the reduction in ‘habitat quality’ of the surrounding benthos. At the large spatial scale substrate complexity is the key determinant of motile epifaunal density, suggesting predation by fishes plays an important structuring role at this scale. Assuming that rugosity is inversely related to predation risk then this study represents the first evidence for spatial-dependence on the top-down (predation) vs. bottom-up (algal biomass) control of community structure. We argue fisheries exploitation, in the absence of a physical disturbance can negatively influence motile epifaunal community structure at large spatial scales.     

Effects of intensity and seasonal timing of disturbances on a rocky intertidal benthic community on the southern coast of Korea

The effects of intensity and timing of disturbances on recovery of marine benthic organisms were investigated on a rocky intertidal shore in Gwangyang Bay, Korea. We hypothesized that the recovery pattern of the benthic community structure would be affected by disturbance intensity and season. Twenty-eight permanent plots were set up, with disturbance intensity (cleared plots and sterile plots) and seasonal disturbance (fall 1999 and spring 2000) incorporated into the experimental design. To monitor natural seasonal variation in benthic community abundances, we established seven permanent unmanipulated plots. Turf-forming algae were observed in the unmanipulated plots throughout the experimental period, whereas green algae and invertebrate presence varied with season. In the disturbance-intensity experiment, turf-forming and green algae were dominant in cleared plots. The highest coverage of sessile organisms was observed in sterile plots, which exhibited the highest species richness because of their relatively low macroalgal coverage. Seasonal effects of disturbance were an important factor in the recovery pattern of benthic organisms under high disturbance intensity. Coverage of green algae was higher in sterile spring plots than in sterile fall plots; this result was attributed to low spatial competition, as the disturbances occurred just before green algal blooms. On the other hand, the abundances of barnacles and bivalves were highest on sterile fall plots, as these organisms were suppressed by green algal blooms in other periods. These results indicate that the effects of disturbance intensity on benthic community recovery patterns can be influenced by season of disturbance.     

Habitat structure,resources and diversity: the separate effects of surface roughness and macroalgae on stream invertebrates

Habitat structure has pervasive effects on community composition and diversity, with physically complex habitats often containing more species than physically simple ones. What factors or mechanism drive this pattern is little understood, but a complicating problem is that different sources of habitat structure can be confounded in both surveys and experiments. In this study, we carried out an experiment in which two sources of habitat structure, attached macroalgae and substrate surface texture, were separately manipulated to discern their joint and separate effects upon the diversity and composition of colonizing macroinvertebrates in a stony, upland stream. Because stream algae vary markedly in abundance in both space and time, we also sampled the epilithon of stream stones at two spatial scales on eight dates over 2 years to gain some preliminary data on how stream algae vary between individual substrata over time. Experimental substrata had either a smooth (siltstones, sandstones, crystal-poor felsic volcanics, plain paving bricks) or rough (granodiorites, crystal-rich felsic volcanics, sand-blasted paving bricks) surface. We allowed these substrata to be colonized naturally by macroalgae, mostly the filamentous red alga Audouinella hermannii. Half of each of the rough and smooth substrata were selected at random and the macroalgae gently sheared off. All substrata were defaunated with a household insecticide with little field persistence, set out randomly through the study riffle, and invertebrates allowed to colonize them for 14 days. Some substrata were sampled immediately to check the efficacy of faunal and algal removals, which proved to be successful. Experimental results showed that both surface texture and macroalgae increase species richness independently of each other. Surface texture had no effect on densities, while macroalgae increased colonization densities, but rarefaction showed that both sources of habitat structure increased species richness above values expected simply on the basis of the numbers of colonists. However, reference stones with high macroalgal cover had the same species richness as those with low cover, suggesting that the effects of macroalgae on species richness are transient relative to those associated with surface texture. Epilithon samples taken at different times suggest that the magnitude of spatial variation in plant growth alters with time. If plants generally recolonize rough surfaces more quickly than smooth, then the effects of habitat structure on macroinvertebrates ought to be strongest after major disturbances during growing seasons of plants. Received: 1 September 1999 / Accepted: 10 January 2000     

Effects of territorial damselfish on an algal-dominated coastal coral reef

Territorial damselfish are important herbivores on coral reefs because they can occupy a large proportion of the substratum and modify the benthic community to promote the cover of food algae. However, on coastal coral reefs damselfish occupy habitats that are often dominated by unpalatable macroalgae. The aim of this study was to examine whether damselfish can maintain distinctive algal assemblages on a coastal reef that is seasonally dominated by Sargassum (Magnetic Island, Great Barrier Reef). Here, three abundant species (Pomacentrus tripunctatus, P. wardi and Stegastes apicalis) occupied up to 60% of the reef substrata. All three species promoted the abundance of food algae in their territories. The magnitudes of the effects varied among reef zones, but patterns were relatively stable over time. Damselfish appear to readily co-exist with large unpalatable macroalgae as they can use it as a substratum for promoting the growth of palatable epiphytes. Damselfish territories represent patches of increased epiphyte load on macroalgae, decreased sediment cover, and enhanced cover of palatable algal turf.     

The responses of a community to disturbance: The importance of successional age and species' life histories

Summary The responses of different successional stages of a temperate intertidal algal community to disturbance were investigated with a field experiment. The experiment was conducted in a low intertidal boulder field in southern California. In this habitat, the top surfaces of boulders are covered with algae. The composition of the assemblage on any particular boulder depends on the length of time since it was last overturned by wave action. When a boulder is overturned, the algae on what was formerly the top surface, are killed in whole or part by a combination of sea urchin grazing, anoxia, light levels below compensation intensity, and mechanical damage caused by crushing or abrasion. The length of time that a boulder remains overturned and the local abundance of sea urchins determines the intensity of the disturbance. When the boulder is righted, recolonization begins either by vegetative regrowth of survivors and/or by spores from outside.Using a three-factorial design, this natural form of disturbance was experimentally mimicked and the responses of three different successional stages of the algal community monitored. Boulders in each successional category were overturned for periods of 17, 27 and 54 days in areas with and without sea urchins, then righted. Two aspects of community response to perturbation were evaluated. These were (1) the assemblage's ability to resist change and (2) its ability, if altered, to adjust to some semblance of its original state. The resistance of each assemblage and of its component species to change was measured by the percent decrease in algal cover and by the decline in percent similarity of the community to its original composition. The recovery rate of each assemblage and of the cover lost by each species during the first 35 days following a disturbance was measured by the rate of increase in percent similarity to the original composition and the percent reestablishment of lost cover.The experimental evidence demonstrates that the successional stages of the producer level of an intertidal algal community differ significantly in their responses to disturbance. Early successional communities suffer more damage from a given level of perturbation but recover more quickly than either middle or late successional communities. Damage to any particular assemblage of algae, irrespective of successional age, is more extensive and recovery slower, the longer the boulder is overturned and/or sea urchins are present. Several thresholds in these responses were also identified.Differences in community responses and non-linearities in these responses were attributable to the life history characteristics of the component species rather than emergent properties of the assemblage. These characteristics have evolved in response to a variety of recurrent natural disturbances. This interpretation is in agreement with recent critical reevaluations of the trends and mechanisms of successional change in natural communities.     

Experimental manipulation of shell color and morphology of the limpets Lottia asmi (Middendorff) and Lottia digitalis (Rathke) (Mollusca: Patellogastropoda)

We experimentally show that substratum can determine an individual patellogastropod's color and morphology. Lottia asmi (Middendorff), previously considered a high-domed, black stenotopic species on trochid gastropods, is similar to other northeastern Pacific limpet species with low profile and tessellated color patterns, when it is translocated to rock substrata. Lottia digitalis (Rathke), a highly variable species with at least two ecological variants that have been considered to be genetically determined, changes between these two forms when translocated from stalked barnacles to rocks. Color changes because the ingested pigments (algal, periostracal, etc.) and other materials (e.g., calcium carbonate) vary between substrata; morphological changes occur as responses to the topographical complexity and constraints of substrata. The geographical distribution of ecological variants of polytypic limpet species, and the ability to achieve cryptic coloration, may be determined by the geographic distributions of algae on which the limpets feed rather than by intrinsic characteristics of the limpets.     

Effect of fire on benthic algal assemblage structure and recolonization in intermittent streams

Abstract: Dry biofilm on rocks and other substrata forms an important drought refuge for benthic algae in intermittent streams following the cessation of flow. This dry biofilm is potentially susceptible to disturbance from bushfires, including direct burning and/or scorching and damage from radiant heat, particularly when streams are dry. Therefore, damage to dry biofilms by fire has the potential to influence algal recolonization and assemblage structure in intermittent streams following commencement of flow. The influence of fire on benthic algal assemblages and recolonization was examined in intermittent streams of the Grampians National Park, Victoria, Australia, using a field survey and manipulative field experiment. The field survey compared assemblages in two intermittent streams within a recently burnt area (within 5 months of the fire) with two intermittent streams within an unburnt area. The two burnt streams were still flowing during the fire so most biofilms were not likely to be directly exposed to flames. Considerable site‐to‐site and stream‐to‐stream variation was detected during the field survey, which may have obscured potential differences attributable to indirect effects of the fire. The manipulative field experiment occurred in two intermittent streams and consisted of five treatments chosen to replicate various characteristics of bushfires that may influence dry biofilms: dry biofilm exposed directly to fire; dry biofilm exposed to radiant heat; dry biofilm exposed to ash; and two procedural controls. After exposure to the different treatments, rocks were replaced in the streams and algae were sampled 7 days after flow commenced. Differences occurred across treatments, but treatment differences were inconsistent across the two streams. For example, direct exposure to fire reduced the abundance of recolonizing algae and altered assemblage structure in both streams, while radiant heat had an effect on assemblage structure in one stream only. The manipulative field experiment is likely to have represented the intensity of a small bushfire only. Nonetheless, significant differences across treatments were detected, so these experimental results suggest that fire can damage dry biofilms, and hence, influence algal recolonization and assemblage structure in intermittent streams.