Consistency in a marine algal‐grazer interaction over multiple scales

Coralline algae are conspicuous members of many marine assemblages, especially those characterized by intense grazing pressure. We explored whether articulated species, especially Corallina vancouveriensis, depend on grazing invertebrates to both establish and flourish in an exposed rocky intertidal setting, and whether this plant–grazer relationship varied over more than three orders of magnitude (≈100–>300,000 μm). Three experimental manipulations, supplemented by observations on recruitment, demonstrated that (i) C. vancouveriensis failed to recover rapidly from disturbed areas when grazers were experimentally excluded; (ii) recruitment occurred in the presence of grazers; (iii) increasing surface texture of molded surfaces enhanced coralline recruitment more when grazers were present; and (iv) settlement occurred predominately in microtopographical low areas of a molded surface, whereas a competitively superior fleshy red alga tended to recruit to high areas. These results confirm that coralline algal establishment and persistence are enhanced by grazers and reveal that this relationship is consistent over a range of biologically relevant scales.     

Effects of Chiton granosus (Frembly, 1827) and other molluscan grazers on algal succession in wave exposed mid-intertidal rocky shores of central Chile

Molluscan grazers can have important effects on the abundance, colonization rates, and successional pathways of algal assemblages and the entire intertidal community. In general, early successional algae are more readily consumed than corticated algae and kelps, which usually get established later in the community succession. To generalize, however, the effect of different grazers on algal assemblages must be examined on different coasts and under different scenarios. This information could help us understand the mechanisms of ecosystem processes and situations in which general models do not apply. Along the coast of Chile, humans harvest large keyhole limpets, which seem to be the only invertebrate grazers capable of controlling the dominant corticated alga Mazzaella laminarioides, a canopy-forming species that can cover extensive areas of the mid intertidal zone. In this scenario, where large limpets are harvested, the overall effects of the diverse molluscan assemblage of limpets, chitons and snails on algal succession and on corticated algae in particular are not clear. We conducted a 26-month-long experiment to evaluate the effects of molluscan grazers on mid-intertidal algal succession and to isolate the effects of Chiton granosus, the most conspicuous member of the assemblage at these tidal elevations. At sites heavily impacted by humans the molluscan grazer assemblage had strong negative effects on colonization and abundance of green algae such as ulvoids and Blidingia minima. In doing so, the grazer assemblage had a strong negative indirect effect on the establishments of chironomid fly larvae, which were only observed on green algal mats and rarely on bare rock. No significant effects were detected on epilithic microalgae, and effects on sessile invertebrates were highly variable over space and time. C. granosus also had significant negative effects on green algae but did not account for the total grazing pressure exerted by the guild. Limited foraging excursions (ca. 35 cm) from refuges and moderate site (crevice) fidelity in this species may contribute to the patchiness in green algal distribution observed in the field. Nearly 13 months after rock surface were experimentally cleared, M. laminarioides appeared in all experimental plots, but increased over three times faster in enclosures containing C. granosus than in exclosures plots or controls, suggesting that moderate levels of herbivory could actually facilitate the establishment of this alga in the succession and that the green algal cover found in the absence of grazers may delay its establishment.     

Effects of avian grazing on the algal community and small invertebrates in the rocky intertidal zone

In this study of a rocky intertidal habitat in northern Japan, feeding by avian consumers had significant effects on algal assemblages and small herbivorous invertebrates. The effects of the birds on algae were different from those of invertebrate grazers such as urchins and gastropods. The abundance of the dominant algal species decreased during the grazing period, increased again after the grazing period, and indirectly affected algal species richness and evenness. Avian grazing also decreased the density of tube-dwelling amphipods on the dominant alga, but did not change the density of mobile and free-living isopods. These results suggest that avian grazers may act as habitat modifiers rather than exploitative competitors for the small herbivorous crustaceans. Avian herbivores consumed only the upper parts of large algal fronds, apparently reducing the amount of suitable microhabitat for the small herbivorous crustaceans, which are subject to a variety of physical or biological stress. Thus, avian herbivores function as ecosystem engineers, regulating community structure in a manner different to invertebrate herbivores in rocky intertidal habitats.     

The effects of grazing by gastropods and physical factors on the upper limits of distribution of intertidal macroalgae

Summary The cover of foliose algae is sparse to non-existent above a low-level algal zone on many shores in N.S.W., except in rock-pools. Above this algal zone, encrusting algae, mostly Hildenbrandia prototypus, occupy most of the primary substratum on sheltered shores. Experimental manipulations at midtidal levels were used to test hypotheses about the effects of grazing by molluses and of physical factors during low tide on this pattern of algal community structure.Fences and cages were used to exclude grazers: molluscs grazed under roofs and in open areas. Cages and roofs provided shade, and decreased the harshness of the environment during low tide: fences and open areas had the normal environmental regime.In the absence of grazers, rapid colonization of Ulva and slower colonization by other foliose algae occurred in all experimental areas. The rate of colonization by Ulva sporelings was initially retarded on existing encrusting algae, but after a few months, cover of Ulva equalled that on cleared rock.Most species of algae only grew to maturity inside cages, and remained as a turf of sporelings inside fences. No foliose algae grew to a visible size in open, grazed areas. Grazing thus prevents the establishment of foliose algae above their normal upper limit on the shore, but the effects of physical factors during low tide prevent the growth of algae which become established when grazers are removed. Physical factors thus limit the abundance of foliose algae at mid-tidal levels.The recolonization of cleared areas by Hildenbrandia was not affected by the presence of a turf of sporelings, nor by the shade cast by roofs, but was retarded in cages where mature algae formed a canopy. Even under such a canopy, Hildenbrandia eventually covered as much primary substratum as in open, grazed areas. This encrusting alga is able to escape from the effects of grazing by having a tough thallus, and by its vegetative growth which allows individual plants to cover a lot of substratum, and by the tendency for new individuals to start growing from small cracks and pits in the rock, which are apparently inaccessible to the grazers.Mature foliose algae are removed from the substratum by waves, and many individual plants died during periods of hot weather. Sporelings in a turf were eliminated, after experimental fences were removed, by the combined effects of macroalgal grazers, which invaded the areas, and microalgal grarers which ate the turt from the edges inwards.The results obtained here are discussed with respect to other studies on limits to distribution of intertidal macroalgae, and the role of grazing in the diversity and structure of intertidal algal communities. Some problems of these experimental treatments are also discussed.     

Retention of dead leaves by grasses as a defense against herbivores. A test on the palatable grass Paspalum dilatatum

The vast majority of grass species retain their leaves as they become senescent, inducing an accumulation of dead biomass that may limit plant productivity. In this research, we tested the hypothesis that dead leaf retention by grasses act as a defense against large herbivores. In a natural grassland of the Flooding Pampa, we carried out a factorial experiment with dead-leaf removal and grazing as main factors. Dead leaves were removed from the palatable grass Paspalum dilatatum , and growth was measured either in the absence or in the presence of large grazers. In the absence of grazers, dead-leaf removal promoted plant growth. In the presence of grazers, dead-leaf removal increased consumption by cattle, particularly among the largest plants. As a result, in the presence of grazers, plants with dead leaves removed grew less than intact plants. The results indicate that retaining dead leaves is costly for grass species in the absence of grazers, but, as proposed by our hypothesis, that cost is compensated by a corresponding reduction of grazing consumption.     

Context-dependency in the effects of nutrient loading and consumers on the availability of space in marine rocky environments

Background

Enhanced nutrient loading and depletion of consumer populations interact to alter the structure of aquatic plant communities. Nonetheless, variation between adjacent habitats in the relative strength of bottom-up (i.e. nutrients) versus top-down (i.e. grazing) forces as determinants of community structure across broad spatial scales remains unexplored. We experimentally assessed the importance of grazing pressure and nutrient availability on the development of macroalgal assemblages and the maintenance of unoccupied space in habitats differing in physical conditions (i.e. intertidal versus subtidal), across regions of contrasting productivity (oligotrophic coasts of South Australia versus the more productive coasts of Eastern Australia).

Methodology/Principal findings

In Eastern Australia, grazers were effective in maintaining space free of macroalgae in both intertidal and subtidal habitats, irrespective of nutrient levels. Conversely, in South Australia, grazers could not prevent colonization of space by turf-forming macroalgae in subtidal habitats regardless of nutrients levels, yet in intertidal habitats removal of grazers reduced unoccupied space when nutrients were elevated.

Conclusions/Significance

Assessing the effects of eutrophication in coastal waters requires balancing our understanding between local consumer pressure and background oceanographic conditions that affect productivity. This broader-based understanding may assist in reconciling disproportionately large local-scale variation, a characteristic of ecology, with regional scale processes that are often of greater relevance to policy making and tractability to management.     

Associational resistance in a multiple herbivore system: differential effects of mammal versus insect herbivores

Recent studies have suggested that associational resistance (AR) should be more effective in protecting plants from grazing by mammals than by herbivorous insects. We tested this hypothesis in a multiple-herbivore system by comparing herbivory on, and reproductive success of, fireweed plants growing within meadowsweet stands with fireweeds growing outside meadowsweet stands in abandoned fields in Northern Sweden. We showed that meadowsweet mediates AR against shared mammal grazers, but not against shared insect herbivores. The risk of fireweed plants being browsed by mammals when growing in the absence of meadowsweet was at least double (41?% affected) that of plants growing within meadowsweet patches (17.5?% affected). We also showed that whereas mammal browsing had a negative effect on fireweed performance, the effect of insect herbivory was negligible. Our results support the hypothesis that AR is more effective against grazing mammals than against herbivorous insects.     

Life history of Acrosiphonia (Codiolales, Chlorophyta) in southwestern British Columbia, Canada

This study establishes the phenology of the alternate life history phases of the green alga Acrosiphonia in British Columbia, Canada. Free-living, filamentous plants are seasonal, March-July, with peak percent cover (10%) in April. Plants are fertile immediately after establishment. The unicells, previously identified as Chlorochytrium inclusum and Codiolum petrocelidis, are the sporophyte phase of Acrosiphonia. 'Chlorochytrium,' spherical and 160-280 μm in diameter, colonizes the foliose red alga Mazzaella splendens 1 mo after filamentous Acrosiphonia plants appear. Maximum density (53 'Chlorochytrium' cells/cm(2) of blade) was recorded in May. 'Codiolum,' on the other hand, is stalked (the vesicle measures 150 × 50 μm) and colonizes the red algal crust Petrocelis. Peak density (22?400 'Codiolum' cells/cm(2) of crust) was recorded 2 mo after 'Chlorochytrium' density peaked. The endophytes survive high summer temperatures, which correlate with death of the free-living plants, and overwinter in their hosts. Zoospore release in late winter corresponds to decreased host abundance, suggesting the endophytes have evolved a strategy whereby duration in the host is synchronized with host seasonality. A bet-hedging strategy is proposed for Acrosiphonia's life history: two morphologically different phases have adapted to a seasonally variable environment, with the sporophyte phase capable of colonizing two different hosts.     

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.     

Major consequences of minor damage: impacts of small grazers on fast-growing kelps

Damage by small herbivores can have disproportionately large effects on the fitness of individual plants if damage is concentrated on valuable tissues or on select individuals within a population. In marine systems, the impact of tissue loss on the growth rates of habitat-forming algae is poorly understood. We quantified the grazing damage by an isopod Amphoroidea typa on two species of large kelps, Lessonia spicata and Macrocystis pyrifera, in temperate Chile to test whether non-lethal grazing damage could reduce kelp growth rates and photosynthetic efficiency. For L. spicata, grazing damage was widespread in the field, unevenly distributed on several spatial scales (among individuals and among tissue types) and negatively correlated with blade growth rates. In field experiments, feeding by A. typa reduced the concentration of photosynthetic pigments and led to large reductions (~80 %) in blade growth rates despite limited loss of kelp biomass (0.5 % per day). For M. pyrifera, rates of damage in the field were lower and high densities of grazers were unable to reduce growth rates in field experiments. These results demonstrate that even low per capita grazing rates can result in large reductions in the growth of a kelp, due the spatial clustering of herbivores in the field and the selective removal of photosynthetically active tissues. The impacts of small herbivores on plant performance are thus not easily predicted from consumption rates or abundance in the field, and vary with plant species due to variation in their ability to compensate for damage.     

Grazers and Phytoplankton Growth in the Oceans: an Experimental and Evolutionary Perspective

The taxonomic composition of phytoplankton responsible for primary production on continental shelves has changed episodically through Earth history. Geological correlations suggest that major changes in phytoplankton composition correspond in time to changes in grazing and seawater chemistry. Testing hypotheses that arise from these correlations requires experimentation, and so we carried out a series of experiments in which selected phytoplankton species were grown in treatments that differed with respect to the presence or absence of grazers as well as seawater chemistry. Both protistan (Euplotes sp.) and microarthropod (Acartia tonsa) grazers changed the growth dynamics and biochemical composition of the green alga Tetraselmis suecica, the diatom Thalassiosira weissflogii, and the cyanobacterium Synechococcus sp., increasing the specific growth rate and palatability of the eukaryotic algae, while decreasing or leaving unchanged both parameters in the cyanobacteria. Synechococcus (especially) and Thalassiosira produced toxins effective against the copepod, but ciliate growth was unaffected. Acartia induced a 4-6 fold increase of Si cell quota in the diatom, but Euplotes had no similar effect. The differential growth responses of the eukaryotic algae and cyanobacteria to ciliate grazing may help to explain the apparently coeval radiation of eukaryophagic protists and rise of eukaryotes to ecological prominence as primary producers in Neoproterozoic oceans. The experimental results suggest that phytoplankton responses to the later radiation of microarthropod grazers were clade-specific, and included changes in growth dynamics, toxin synthesis, encystment, and (in diatoms) enhanced Si uptake.     

Sediment on rocky intertidal reefs: Effects on early post-settlement stages of habitat-forming seaweeds

Modification of the coastal environment by human activities often leads to an increase in sedimentation of nearshore waters, with potential impacts on benthic marine assemblages. Here we assess the relationships between the levels of sedimentation, wave exposure and benthic organisms on rocky intertidal platforms around the Kaikoura Peninsula in southern New Zealand. We designed and tested five sediment traps to provide a tool for measuring the relative abundance of sediment across sites. Using field- and laboratory-based experiments, we tested hypotheses concerning whether different levels of sedimentation affected algal germling survival and algal zygote attachment, and whether the interactions of grazers and sediments affected germling survival. Levels of sediment and exposure were inversely related across seven sites. The fucoid alga Hormosira banksii characterized the more sedimented wave-sheltered and intermediately wave-exposed sites, with up to 80% cover in the lower mid-tidal zone, while the bull kelp Durvillaea antarctica characterized the three most wave-exposed sites. Grazing molluscs were found across all sites but species abundances varied by sediment and exposure levels. We did two 11-day trials testing the effects of different levels of sediment and different species of molluscan grazers on the survival of 1-week-old Hormosira germlings. Generally, there was no significant treatment effect of grazers, but mortality varied considerably among sites. In particular, one site had very high levels of sediment, which resulted in 100% mortality of germlings across all grazer treatments. Removing sediment at 1-week, 2-week and 4-week intervals made no difference to the survival of Hormosira germlings. In laboratory-based experiments, a light dusting of sediment reduced the percentage of zygotes of Hormosira by 34% and Durvillaea by 71% that attached to primary substratum, and a complete cover of sediment prevented attachment altogether. Overall, the effects of sediments and its interaction with molluscan grazing were highly variable but often large, particularly on the attachment of zygotes to primary substratum.     

Limpet grazing on a physically stressful Patagonian rocky shore

Many theories of consumer control of communities have come from studies conducted in relatively benign, temperate zone rocky intertidal systems. Here, we examine gastropod grazing and the maintenance of bare space on a dry, wind-swept rocky shore of Patagonia, Argentina. Two limpet species are the primary intertidal grazers. Siphonaria lessoni dominates mid and high intertidal zones, while Nacella magellanica dominates the lower zone. In all zones, limpet densities are positively correlated with bare space and the occurrence of cracks. Tethering experiments revealed that: (1) physical stress sets the upper distribution limit of both limpets, (2) predators, such as oyster catchers, regulate Nacella populations and may restrict them to cracks and vertical surfaces, and (3) desiccation stress appears to drive similar crack distribution patterns of Siphonaria in the upper intertidal. Experimental removal of limpets in each intertidal zone indicated that limpets have: (1) no detectable effect in the high intertidal where physical forces dominate community organization, (2) weak impacts at mid-elevations as grazing only limited the abundance of fleshy algae with physical forces again dominating community structure, and (3) relatively stronger, but still weak impacts in the low zone. These results suggest that grazing impacts on Argentine rocky shores are weak in comparison to the physical stresses (e.g. high winds, low humidity) that largely determine structure in this system. The dominance of physical forcing in this system occurs despite having similar grazer densities to other temperate, but comparatively wet, rocky shorelines (e.g. British Isles) where top-down control is strong.     

Effect of nutrient availability, grazer assemblage and seagrass source population on the interaction between Thalassia testudinum (turtle grass) and its algal epiphytes

Seagrass leaves are often densely covered by epiphytic algae which can suppress seagrass productivity and has been implicated in declines of seagrass meadows worldwide. The net effect of epiphytes on seagrass growth and morphology depends on the independent and interactive effects of a variety of factors, including nutrient availability and the intensity of grazing on epiphytes. Here I report the results of a mesocosm experiment designed to test the effects of nutrient addition and within-functional group variation (grazer species composition and the source population of seagrass) on the strength of the interactions among grazers, epiphytes, and turtle grass (Thalassia testudinum). Turtle grass ramets from two sites in the northern Gulf of Mexico were cleared of epiphytes and transplanted into common-garden mesocosms. Replicate ramets were grown in a split-split plot design with two levels of dissolved nutrients and four different grazer species combinations (Tozeuma carolinense alone, Pagurus maclaughlinae alone, both species together, and no grazers present). As expected, grazers had a significant negative effect on epiphyte biomass/leaf area and a significant positive effect on turtle grass growth in the mesocosms. The two species were more similar in their direct effects on epiphyte biomass than in their indirect effects on turtle grass growth; this may reflect differences in epiphyte community composition under different grazer treatments. The effect of nutrient addition on turtle grass growth depended critically on the intensity of grazing: in the presence of grazers, turtle grass tended to produce a greater biomass of new leaf tissue in the tanks with nutrients added than in the control tanks. However, when grazers were absent, the direction of the effect was reversed, and plants with nutrients added grew less than the control plants. The two source populations of turtle grass differed significantly in epiphyte biomass/leaf area accrued in the mesocosms as well as in the strength of the effect of grazers on turtle grass growth. This suggests that population differentiation in seagrass interactions with epiphytes, as well as spatial and temporal variation in resources and grazer community composition, can greatly effect the role of epiphytes in limiting seagrass productivity.     

Impact of Zooplankton Grazing on the Excystation, Viability, and Infectivity of the Protozoan Pathogens Cryptosporidium parvum and Giardia lamblia

Very little is known about the ability of the zooplankton grazer Daphnia pulicaria to reduce populations of Giardia lamblia cysts and Cryptosporidium parvum oocysts in surface waters. The potential for D. pulicaria to act as a biological filter of C. parvum and G. lamblia was tested under three grazing pressures (one, two, or four D. pulicaria grazers per 66 ml). (Oo)cysts (1 × 10⁴ per 66 ml) were added to each grazing bottle along with the algal food Selenastrum capricornutum (6.6 × 10⁴ cells per 66 ml) to stimulate normal grazing. Bottles were rotated (2 rpm) to prevent settling of (oo)cysts and algae for 24 h (a light:dark cycle of 16 h:8 h) at 20°C. The impact of D. pulicaria grazing on (oo)cysts was assessed by (i) (oo)cyst clearance rates, (ii) (oo)cyst viability, (iii) (oo)cyst excystation, and (iv) oocyst infectivity in cell culture. Two D. pulicaria grazers significantly decreased the total number of C. parvum oocysts by 52% and G. lamblia cysts by 44%. Furthermore, two D. pulicaria grazers significantly decreased C. parvum excystation and infectivity by 5% and 87%, respectively. Two D. pulicaria grazers significantly decreased the viability of G. lamblia cysts by 52%, but analysis of G. lamblia excystation was confounded by observed mechanical disruption of the cysts after grazing. No mechanical disruption of the C. parvum oocysts was observed, presumably due to their smaller size. The data provide strong evidence that zooplankton grazers have the potential to substantially decrease the population of infectious C. parvum and G. lamblia in freshwater ecosystems.     

Mutualism between the territorial intertidal limpet Patella longicosta and the crustose alga Ralfsia verrucosa

Mutualistic relations between plants and animals are well documented on land but have received less attention in marine systems. This study examined the relationship between the territorial intertidal limpet Patella longicosta and the crustose brown alga Ralfsia verrucosa. Adult Patella are found exclusively in association with Ralfsia, on which they feed, while Ralfsia occurs primarily, but not exclusively, in Patella territories. Ralfsia benefits directly from both the presence and the territorial behaviour of Patella. Algal productivity was assessed by measuring oxygen evolution and utilization in situ and deriving photosynthesis/irradiance curves. Productivity was increased by about 30% by the presence of Patella in both summer (P _max of grazed algae 0.0098; ungrazed algae 0.0063 mg C · cm^-2 · h^-1) and winter (P _max grazed algae 0.0081; ungrazed algae 0.0053 mg^-2 · C · h^-1). Algal growth rates were not significantly increased by the application of limpet mucus in the laboratory. We did not examine nutrient regeneration by the limpet, but the increase in photosynthetic rate may depend on the limpet's grazing pattern which creates secondary sites for growth. Ralfsia also benefited from the territorial behaviour of Patella. The effects of different grazing regimes were investigated in different seasons by removing territorial limpets and either excluding all limpets using copper-based antifouling paint, or allowing access to non-territorial limpets (mostly P. oculus) using partial paint barriers. Exclusion of all limpets resulted in rapid overgrowth of Ralfsia plants by the foliose green alga Ulva sp.. Where non-territorial limpets had access to the plants overgrowth was reduced but Ralfsia plants were entirely removed by destructive grazing. Non-territorial grazers removed 90% of Ralfsia plants within 4 weeks in summer and 60% in winter. In control treatments P. longicosta prevented overgrowth by Ulva and actively excluded vagrant grazers, preventing overgrazing. Based on these findings, the association between the limpet and alga can be regarded as a nonobligate mutualism.     

Physical Organization of Two Reservoirs in Serbia as a Crucial Factor in Development of Their Hydrophilic Flora: a Comparative Study

Marine invertebrate herbivores occasionally prefer particular tissues of a given seaweed depending on tissue age. For the intertidal alga Mazzaella parksii (=M. cornucopiae), however, important tissue differences result from abiotic stress: distal tissues of many fronds become bleached under strong irradiance and desiccation during spring and summer low tides. Snails of the genus Littorina (periwinkles) are significant grazers of M. parksii. Through a multiple-choice feeding-preference experiment, we found that periwinkles actively consume bleached tissues, but almost no photosynthetic tissues. This suggests that bleached tissues of M. parksii may support Littorina populations to a good extent during spring and summer. In addition, since photosynthetic tissues are basal, the impact of periwinkles on frond mortality might be lower than if they preferred photosynthetic tissues. Photosynthetic tissues are actively consumed by periwinkles when they are the only food choice, as shown by other researchers. Therefore, retaining bleached tissues for some time might have evolved in M. parksii to divert grazing pressure away from vital tissues.     

Epiphytes on the rocky intertidal red alga RhodomelaLarix (Turner) C. Agardh: Negative effects on the host and food for herbivores?

The diversity of epiflora and fauna associated with a dominant turf-forming alga was examined in intertidal communities on the central Oregon coast. Epiphytes associated with the red alga, Rhodomelalarix (Turner) C. Agardh, were examined by surveying intertidal areas for the presence of epiphytes, and by following changes in epiphyte cover in marked quadrats of R. larix. The alga is host for at least 17 species of sessile plants and animals. To determine the role of some of the larger epiphytes in the community, Rhodomela plants were marked and monitored and herbivore feeding was examined. Data suggest that epiphytes decrease the growth rate of their host, increase the probability of axis breakage and decrease reproductive output. Epiphytes provide food for littorine snails and gammarid amphipods that live in the beds of the plant. Amphipods were found to decrease epiphyte cover on R. larix in laboratory tanks, suggesting that these herbivores may have beneficial effects on the host plant.     

Grazing of the invasive alga Codium fragile ssp. tomentosoides by the common periwinkle Littorina littorea: Effects of thallus size, age and condition

We examined the potential of herbivory by the common periwinkle Littorina littorea to limit recruitment and vegetative re-growth of the invasive green alga Codium fragile ssp. tomentosoides in a series of manipulative field experiments in tidepools on a wave-exposed rocky shore in Nova Scotia, Canada. Snails were excluded or included from circular plots (14 to 20 cm diameter) with cages to compare growth and survival of C. fragile against procedural (partial cages) or natural (uncaged) control plots. Our results show that L. littorea may restrict growth and survival of C. fragile by grazing new recruits (< 2 cm thallus length), fronds of adult thalli that are bleached and necrotic, and residual holdfasts (< 2 mm thickness) of detached thalli (artificially severed to mimic wave dislodgement). Once recruits grow beyond a critical size (~ 3 cm), or damaged tissues or holdfasts regenerate, grazing of C. fragile by L. littorea appears to be limited and ineffectual. Our experimental results corroborate correlative evidence from previous studies that herbivory by L. littorea limits the abundance of C. fragile in tidepools on the Atlantic coast of Nova Scotia, particularly pools in the high intertidal zone where these snails are abundant. Lower on the shore, littorinid grazing and physical stressors that render algae more vulnerable to grazers (e.g. UV radiation and freezing) are less intensive, and probably have less of a regulatory effect on populations of C. fragile.     

Interactions of plant stress and herbivory: intraspecific variation in the susceptibility of a palatable versus an unpalatable seaweed to sea urchin grazing

Summary Interactions among environmental stresses, plant defensive characteristics, and plant nutrient status may significantly affect an alga's susceptibility to herbivores. Following desiccation, the palatable seaweed Gracilaria tikvahiae was less susceptible to grazing by the sea urchin Arbacia punctulata while the unpalatable alga Padina gymnospora became more susceptible. Increased grazing on desiccated Padina appeared to result from a loss of chemical defenses following desiccation. Palatable plants treated with organic extracts from desiccated Padina plants were consumed at more than twice the rate of plants treated with extracts from undesiccated plants. Increased susceptibility of Padina did not correlate with changes in protein content of the alga; reduced grazing on desiccated Gracilaria was associated with a decrease in protein content. When Padina was grazed by Arbacia or mechanically damaged to mimic urchin grazing, its susceptibility to Arbacia decreased within 1 to 5 days. These results demonstrate that history of physical or biological stress may affect a plant's susceptibility to herbivory. We hypothesize that urchins cue primarily on attractiveness features (e.g. nutrient content) of highpreference algae and deterrent features (e.g. chemical defenses) of low-preference algae. Stresses may therefore increase, decrease, or not affect a plant's susceptibility to herbivory depending upon the primary feeding cues used by the herbivore, the defensive mechanisms used by the plant, and the way these are altered by various environmental stresses.