Branching morphology as an indicator of environmental disturbance: Testing the vegetative fragmentation of Acanthophora spicifera and the turf morphology of Laurencia papillosa

A new method for interpreting branching morphology is introduced and demonstrated to be a practical means of assessing ecological events. The “turf” configuration of Laurencia papillosa (Forssk.) Grev. and the vegetative fragmentation of Acanthophora spicifera (Vahl) Børg. are examined on a fringing reef platform at Galeta Point, Panama. Predictions of branching morphology are tested by comparing plants in the fore-reef, that are exposed to intense wave action and potentially large numbers of herbivorous fish, with those in the back-reef, where wave action and grazing are minimal. Differences in plant size, number of branches and length of branch segments, together with the number and location of missing branches, are quantified. It is shown that in some instances branching morphology can be used to test ecological events.There is little difference in the branching structure of Laurencia papillosa in the foreand back-reef. The number and the length of branch segments were similar and few branches were lost to wave action or herbivory. Some grazing of determinate branches occurred on the reef flat, but this had no apparent effect upon branching morphology. Plants in the fore-reef, however, were significantly smaller than those in the back-reef. It follows that fore-reef plants are best characterized as dwarf plants as opposed to a “turf”.The branching morphology of Acanthophora spicifera is altered by wave action in the fore-reef. Plants were shorter and had proportionally fewer branches in the fore-reef than in the back-reef. There was, however, no difference in the length of branch segments between the two areas. Numerous branching scars, resulting from the loss of indeterminate branches, were found. In the fore-reef, the smaller-sized plants of A. spicifera are best explained by vegetative fragmentation.     

Responses to nutrient enrichment,wave action and disturbance in rocky shore communities

A high degree of resistance against nutrient enrichment has previously been demonstrated for macroalgal-dominated rocky shore communities in the presence of moderate to large amounts of macroinvertebrate grazers. To experimentally examine, under controlled conditions, the possible roles for this resistance of two other factors, i.e. disturbance (presence/absence of the macroalgal canopy itself) and wave action, the canopy algae and associated algal and animal assemblages were removed by scraping from approximately one third of the area of eight littoral mesocosms, subjected to two different wave action regimes. After this, excessive nutrients were added to four mesocosms with the factor nutrients fully crossed with the factor wave action with two replicate mesocosm basins of each nutrient/wave treatment combination. Disturbance was added to the design as a within-basin factor thus making up a split-plot experiment. The abundance of grazers was allowed to vary freely and under the influence of the treatments. After 11 summer weeks, there were significant differences in community structure between nutrient enrichment levels for both algal and animal assemblages when examined by multivariate statistical techniques. Univariate analyses confirmed a significantly stimulated colonisation by green algae, mainly Ulva lactuca, in both disturbed (scraped) and undisturbed areas of nutrient-enriched mesocosms. In un-enriched mesocosms, the green algae were absent from undisturbed areas and rare in disturbed areas, where mainly brown Ectocarpus spp. and red algae had settled. Among the macrofauna, the total abundance of grazers was stimulated in nutrient-enriched mesocosms with individuals of the amphipod genus Gammarus and the isopod genus Jaera being especially numerous. With regard to wave action, no significant differences occurred in community structure, although there were indications of significant nutrient × wave effects for both the amount of exported red algae and the amount of accumulated brown algae. The study shows that eutrophication-related community shifts on rocky shores may occur very rapidly, regardless of the level of wave-energetic stress and the abundance of grazers, if the nutrient concentrations are high and the colonisation and growth of opportunistic algae are facilitated by disturbance such as (naturally or anthropogenically driven) canopy gap forming processes.     

Alien marine fishes deplete algal biomass in the Eastern Mediterranean

One of the most degraded states of the Mediterranean rocky infralittoral ecosystem is a barren composed solely of bare rock and patches of crustose coralline algae. Barrens are typically created by the grazing action of large sea urchin populations. In 2008 we observed extensive areas almost devoid of erect algae, where sea urchins were rare, on the Mediterranean coast of Turkey. To determine the origin of those urchin-less 'barrens', we conducted a fish exclusion experiment. We found that, in the absence of fish grazing, a well-developed algal assemblage grew within three months. Underwater fish censuses and observations suggest that two alien herbivorous fish from the Red Sea (Siganus luridus and S. rivulatus) are responsible for the creation and maintenance of these benthic communities with extremely low biomass. The shift from well-developed native algal assemblages to 'barrens' implies a dramatic decline in biogenic habitat complexity, biodiversity and biomass. A targeted Siganus fishery could help restore the macroalgal beds of the rocky infralittoral on the Turkish coast.     

Factors involved in herbivore food preference

Herbivore food preference can be measured either in terms of attractiveness or of edibility: these quantities are not necessarily correlated. The isopod Idotea baltica (Pallas) is attracted to large, tough, branched algae (perennials) while the amphipod Ampithoe valida (Smith) is attracted to softer, filamentous or bladed algae (ephemerals). Attractiveness for Ampithoe is related to the nutritive value of the algae; this is not so for Idotea, which responds more to algal morphology and availability. Idotea's mobility, possible susceptibility to fish predation, and preference for moderate wave exposures may select for a primary response to algae as habitat rather than as food source.     

The relative impacts of grazing by caribbean coral reef fishes and Diadema: effects of habitat and surge

The effects of grazing on fleshy algal turfs by large herbivorous fishes and by the black spiny urchin, Diadema antillarum (Philippi), were investigated in three shallow (<3 m), moderately exposed, reef flat habitats in the San Blas Islands on the Caribbean coast of Panama. Grazing by Diadema was found to be a primary determinant of the biomass of fleshy algae in this shallow reef environment, and its impact was exposure-dependent. Feeding activity of Diadema was inhibited during the protracted periods of heavy wave action that characterize the dry season when strong north winds (24–27 kph) are typical. In the shallowest, most exposed elkhorn bluff habitats urchin grazing was minimal even during calm periods due to the effects of waves generated by unpredictable, light winds. In a relatively protected backreef habitat, water movement was inadequate to discourage grazing significantly, even during the turbulent dry season. As a result, algal biomass (decalcified dry weight) was typically three-fold greater in the elkhorn bluff habitat than on the ackreef. In habitats of intermediate exposure (elkhorn fields) algal biomass was reduced during calm periods, but increased to the level characteristic of the elkhorn bluff habitat during protracted rough periods that inhibited grazing by Diadema.A caging experiment in the backreef habitat demonstrated a relatively minor effect of grazing by herbivorous fishes in comparison to that of grazing by Diadema. Observations on the distributions and abundances of herbivorous fishes also indicated a relatively minor role for these grazers as determinants of algal biomass in the more exposed habitats.     

Responses to nutrient enrichment,wave action and disturbance in rocky shore communities

A high degree of resistance against nutrient enrichment has previously been demonstrated for macroalgal-dominated rocky shore communities in the presence of moderate to large amounts of macroinvertebrate grazers. To experimentally examine, under controlled conditions, the possible roles for this resistance of two other factors, i.e. disturbance (presence/absence of the macroalgal canopy itself) and wave action, the canopy algae and associated algal and animal assemblages were removed by scraping from approximately one third of the area of eight littoral mesocosms, subjected to two different wave action regimes. After this, excessive nutrients were added to four mesocosms with the factor nutrients fully crossed with the factor wave action with two replicate mesocosm basins of each nutrient/wave treatment combination. Disturbance was added to the design as a within-basin factor thus making up a split-plot experiment. The abundance of grazers was allowed to vary freely and under the influence of the treatments. After 11 summer weeks, there were significant differences in community structure between nutrient enrichment levels for both algal and animal assemblages when examined by multivariate statistical techniques. Univariate analyses confirmed a significantly stimulated colonisation by green algae, mainly Ulva lactuca, in both disturbed (scraped) and undisturbed areas of nutrient-enriched mesocosms. In un-enriched mesocosms, the green algae were absent from undisturbed areas and rare in disturbed areas, where mainly brown Ectocarpus spp. and red algae had settled. Among the macrofauna, the total abundance of grazers was stimulated in nutrient-enriched mesocosms with individuals of the amphipod genus Gammarus and the isopod genus Jaera being especially numerous. With regard to wave action, no significant differences occurred in community structure, although there were indications of significant nutrient × wave effects for both the amount of exported red algae and the amount of accumulated brown algae. The study shows that eutrophication-related community shifts on rocky shores may occur very rapidly, regardless of the level of wave-energetic stress and the abundance of grazers, if the nutrient concentrations are high and the colonisation and growth of opportunistic algae are facilitated by disturbance such as (naturally or anthropogenically driven) canopy gap forming processes.     

Effects of “Reduced” and “Business-As-Usual” CO2 Emission Scenarios on the Algal Territories of the Damselfish Pomacentrus wardi (Pomacentridae)

Turf algae are a very important component of coral reefs, featuring high growth and turnover rates, whilst covering large areas of substrate. As food for many organisms, turf algae have an important role in the ecosystem. Farming damselfish can modify the species composition and productivity of such algal assemblages, while defending them against intruders. Like all organisms however, turf algae and damselfishes have the potential to be affected by future changes in seawater (SW) temperature and pCO₂. In this study, algal assemblages, in the presence and absence of farming Pomacentrus wardi were exposed to two combinations of SW temperature and pCO₂ levels projected for the austral spring of 2100 (the B1 “reduced” and the A1FI “business-as-usual” CO₂ emission scenarios) at Heron Island (GBR, Australia). These assemblages were dominated by the presence of red algae and non-epiphytic cyanobacteria, i.e. cyanobacteria that grow attached to the substrate rather than on filamentous algae. The endpoint algal composition was mostly controlled by the presence/absence of farming damselfish, despite a large variability found between the algal assemblages of individual fish. Different scenarios appeared to be responsible for a mild, species specific change in community composition, observable in some brown and green algae, but only in the absence of farming fish. Farming fish appeared unaffected by the conditions to which they were exposed. Algal biomass reductions were found under “reduced” CO₂ emission, but not “business-as-usual” scenarios. This suggests that action taken to limit CO₂ emissions may, if the majority of algae behave similarly across all seasons, reduce the potential for phase shifts that lead to algal dominated communities. At the same time the availability of food resources to damselfish and other herbivores would be smaller under “reduced” emission scenarios.     

Consumptive and non-consumptive effects of an invasive marine predator on native coral-reef herbivores

Invasive predators typically have larger effects on native prey populations than native predators, yet the potential roles of their consumptive versus non-consumptive effects (CEs vs. NCEs) in structuring invaded systems remains unclear. Invasive lionfish (Pterois volitans) may have ecosystem-level effects by altering native fish grazing on benthic algae that could otherwise displace corals. Lionfish could reduce grazing by decreasing the abundance of herbivorous fishes (CEs), and/or the predation risk posed by lionfish could alter grazing behavior of fishes (NCEs). To test for these CEs, we manipulated lionfish densities on large reefs in The Bahamas and surveyed fish populations throughout June 2009–2011. In July 2011, NCEs of lionfish were measured by observing fish grazing behavior on algal-covered substrata placed in microhabitats varying in lionfish presence at different spatial scales, and quantifying any resulting algal loss. Lionfish reduced small herbivorous fish density by the end of the 2010 summer recruitment season. Grazing by small and large fishes was reduced on high-lionfish-density reefs, and small fish grazing further decreased when in the immediate presence of lionfish within-reefs. Lionfish had a negative indirect effect on algal loss, with 66–80 % less algae removed from substrata in high-lionfish-density reefs. Parrotfishes were likely driving the response of herbivorous fishes to both CEs and NCEs of lionfish. These results demonstrate the importance of considering NCEs in addition to CEs of invasive predators when assessing the effects of invasions.     

Top-down control of natural phyto- and bacterioplankton prey communities by Daphnia magna and by the natural zooplankton community of the hypertrophic Lake Blankaart

Biomanipulation, through the reduction of fish abundance resulting in an increase of large filter feeders and a stronger top-down control on algae, is commonly used as a lake restoration tool in eutrophic lakes. However, cyanobacteria, often found in eutrophic ponds, can influence the grazing capacity of filter feeding zooplankton. We performed grazing experiments in hypertrophic Lake Blankaart during two consecutive summers (1998, with and 1999, without cyanobacteria) to elucidate the influence of cyanobacteria on the grazing pressure of zooplankton communities. We compared the grazing pressure of the natural macrozooplankton community (mainly small to medium-sized cladocerans and copepods) with that of large Daphnia magna on the natural bacterioplankton and phytoplankton prey communities. Our results showed that in the absence of cyanobacteria, Daphnia magna grazing pressure on bacteria was higher compared to the grazing pressure of the natural zooplankton community. However, Daphnia grazing rates on phytoplankton were not significantly different compared to the grazing rates of the natural zooplankton community. When cyanobacteria were abundant, grazing pressure of Daphnia magnaseemed to be inhibited, and the grazing pressure on bacteria and phytoplankton was similar to that of the natural macrozooplankton community. Our results suggest that biomanipulation may not always result in a more effective top-down control of the algal biomass.     

In situ tests on inducible defenses in Dictyota kunthii and Macrocystis integrifolia (Phaeophyceae) from the Chilean coast

Numerous experimental studies have reported inducible defenses in macroalgae, but most of them have been conducted in laboratory environments where algae were maintained detached from the substratum and in artificial flow regimes. The results of those experiments might not reflect the natural situation, which can only be studied in situ. We examined whether the brown macroalgae Dictyota kunthii (C. Agardh) Greville and Macrocystis integrifolia (Bory) show inducible defenses following exposure to different grazing levels (direct, water-borne cues from nearby grazed conspecifics, presence of a non-grazing herbivore and natural grazing) in field experiments, striving to maintain natural conditions as much as possible. We measured palatability of algae after exposure to different grazing levels by using live algae and agar-based food containing non-polar extracts. M. integrifolia showed no induction of defenses (at least not of non-polar compounds), suggesting constitutive defenses, absence of defenses (tolerance) or use of another strategy to avoid herbivory. These results are similar to those from previous laboratory experiments. In D. kunthii, defense was induced after two weeks of direct grazing by amphipods under field conditions. Water-borne cues from nearby grazed conspecifics, presence of a non-grazing herbivore and natural grazing did not induce defenses. Induction of defense in response to direct grazing agrees with results from a previous laboratory study, but while indirect cues induced defenses in the laboratory, there was no measurable induced defense in the field. Probably chemical cues from grazers are diluted quickly in the field, not reaching concentrations that cause induction of defenses. This might be the reason why in some algae induction by direct grazing is a more important defensive strategy than induction by water-borne cues. The results from our study also suggest that laboratory experiments showing induced defenses in response to grazed neighbours or mere grazer presence need to be interpreted with caution.     

Echinoid grazing as a structuring force in coral communities: Whole reef manipulations

To examine the effects of echinoid grazing on shallow water coral community structure, the entire Diadema antillarum Philippi population (> 3000 individuals) was eliminated from a patch reef in Discovery Bay, Jamaica W.I. in January 1974. All remaining regular echinoids, notably including Echinometra viridis A. Agassiz (> 7000 individuals), were removed from one-half of the same reef 6 months later.Where Echinometra was present, its grazing was highly patchy, even at densities of up to 50/m². On the other hand, the effects of grazing by Diadema were more uniformly distributed at equivalent or lower densities.Percent-cover of adult corals was enhanced in the presence of Echinometra viridis, however, it was greatly reduced in the absence of all echinoid grazing as a result of massive overgrowth by algae. Success of coral recruitment (density of coral spat) increased in the absence of Diadema due to the alleviation of biological disturbance imposed by this urchin's feeding activities. The presence of Echinometra viridis, even at the high densities naturally present on the reef, did not deter coral settlement.Coral diversity was highest in the presence of all echinoids, intermediate in the presence of E. viridis (without Diadema), and lowest in the absence of all echinoids. This was due to the disproportionately successful recruitment and resultant high relative abundance of Agaricia at low echinoid densities. Favia, Millepora, Helioseris and Acropora all exhibited universally low recruitment levels. Predation by Diadema and Echinometra is responsible for damaging larger coral colonies, particularly Porites.As echinoid species composition and density was altered, the species composition of benthic algae and associated invertebrates shifted. In this way, the array of species competing for space with coral spat shifted as did their competitive relationships. Major epifaunal competitors included spirorbid polychaetes, various foraminifera (particularly Gypsina), and sponges. Algal competitors included filamentous chlorophytes, corallines (e.g. Jania), and numerous other red algae; the species composition of algal competitors shifted through time. Competitive success in corals varied as echinoid density changed and was species-specific with respect to coral genus. Interspecific competition between coral spat (i.e., interspecific aggression via extracoelenteric digestion) was negligible under all treatments.The presence of Echinometra viridis in high densities (in the absence of Diadema) created conditions where growth and fusion in Agaricia spat were optimized. Under these conditions, fusion allows a young coral colony to attain a large enough size to survive damage incurred from grazing or from competitive overgrowth, decreasing the probability of whole-colony mortality during the early stages of development.     

In vitro experimental assessment of the grazing pressure of two gastropods on Zostera marina L. ephiphytic algae

This study experimentally assessed the impact of gastropod grazing on epiphyton biomass and taxonomic composition on leaves of the seagrass Zostera marina L. along the coasts of Brittany, France. Image analysis was used to measure the same section of a leaf before and after grazing and assess grazing pressure. It was found that: (1) both grazers, Gibbula umbilicalis and Jujubinus striatus, were highly efficient in reducing epiphytic cover; (2) J. striatus showed a higher specific grazing rate than G. umbilicalis at an equivalent animal biomass; (3) J. striatus increased its activity when epiphytic biomass increased. Analysis of the variability of epiphytic cover along the leaves showed considerable heterogeneity from the basal part of the external leaves of a shoot to the apex. The development of long filamentous algae (mainly ectocarpales) in spring and summer and their concentration at the apex intensified shading impact on the seagrass bed. The grazing of both gastropod species was also concentrated at the apex, thereby enhancing their role in moderating the shading effect.     

Impact of hard-bottom substrata on the small-scale distribution of fish and decapods in shallow subtidal temperate waters

The micro-scale spatial distribution patterns of a demersal fish and decapod crustacean assemblage were assessed in a hard-bottom kelp environment in the southern North Sea. Using quadrats along line transects, we assessed the in situ fish and crustacean abundance in relation to substratum types (rock, cobbles and large pebbles) and the density of algae. Six fish and four crustacean species were abundant, with Ctenolabrus rupestris clearly dominating the fish community and Galathea squamifera dominating the crustacean community. Differences in the substratum types had an even stronger effect on the micro-scale distribution than the density of the dominating algae species. Kelp had a negative effect on the fish abundances, with significantly lower average densities in kelp beds compared with adjacent open areas. Averaged over all of the substrata, the most attractive substratum for the fish was large pebbles. In contrast, crustaceans did not show a specific substratum affinity. The results clearly indicate that, similar to other complex systems, significant micro-scale species–habitat associations occur in northern hard-bottom environments. However, because of the frequently harsh environmental conditions, these habitats are mainly sampled from ships with sampling gear, and the resulting data cannot be used to resolve small-scale species–habitat associations. A detailed substratum classification and community assessment, often only possible using SCUBA diving, is therefore important to reach a better understanding of the functional relationships between species and their environment in northern temperate waters, knowledge that is very important with respect to the increasing environmental pressure caused by global climate change.     

Latitudinal variation in intertidal algal community structure: the influence of grazing and vegetative propagation

Summary The hypothesis that sea urchin grazing and interactions with turf-forming red algae prevent large brown algae from forming an extensive canopy in the low intertidal zone of southern California was tested with field experiments at two study sites. Experimental removal of sea urchins resulted in rapid algal recruitment. Crustose coralline algae which typically dominate the substratum in areas with dense urchin populations were quickly overgrown by several species of short-lived green, brown and red algae. The removal of urchins also significantly increased the recruitment of two long-lived species of large brown algae (Egregia laevigata and Cystoseira osmundacea at one study site and E. laevigata and Halidrys dioica at the other). The experimental plots at both sites were eventually dominated by perennial red algae.A two-factorial experiment demonstrated that sea urchin grazing and preemption of space by red algae in areas where urchins are less abundant are responsible for the rarity of large brown algae in the low intertidal of southern California. The three dominant perennial red algae, Gigartina canaliculata, Laurencia pacifica and Gastroclonium coulteri, recruit seasonally from settled spores but can rapidly fill open space with vigorous vegetative growth throughout the year. These species encroach laterally into space created by the deaths of large brown algae or by other disturbances. Once extensive turfs of these red algae are established further invasion is inhibited. This interaction of algae which proliferate vegetatively with algae which recruit only from settled spores is analogous to those which occur between solitary and colonial marine invertebrates and between solitary and cloning terrestrial plants.It is suggested that a north-south gradient in the abundance of vegetatively propagating species, in grazing intensity and in the frequency of space-clearing disturbances, may account for latitudinal variation in intertidal algal community structure along the Pacific coast of North America.     

Organismal defenses versus environmentally mediated protection from herbivores: Unraveling the puzzling case of Desmarestia viridis (Phaeophyta)

The role of anti-herbivore organismal defenses in algae-herbivore interaction is frequently investigated without taking into account the potential role of environmental factors in mediating the interaction. Here we reexamine the interaction between the highly acidic, brown alga Desmarestia viridis and the green sea urchin, Strongylocentrotus droebachiensis, by incorporating a previously overlooked facet, the effect of changes in the wave environment on the ability of the urchin to establish contact with the alga. Factorial experiments in a wave tank (presence versus absence of waves; real versus mimic algae) showed that the aggregation of urchins on D. viridis was more than 2-fold greater in the absence than in the presence of waves. Similar numbers of urchins made contact with natural and mimic D. viridis plants, both with and without waves, indicating that any external release of chemicals (acid) from the alga had no perceptible repulsive effect on the urchin. The ability of the urchins to climb onto D. viridis increased markedly when urchin density attained a critical level. These results were consistent with field observations that urchins readily attack D. viridis under conditions of low wave action but do not under conditions of moderate wave action. We conclude (1) that the chemical makeup of D. viridis alone is neither necessary nor sufficient to limit contacts by the urchins and that (2) wave action is a major factor explaining the survival of D. viridis on urchin barrens, because waves limit the movements of the urchins towards the alga. We recommend that studies addressing marine algal defenses against herbivores be more comprehensive and examine interactions between algal traits, the physical environment, and the abundance and behavioral repertoire of herbivores.     

The depletion of algal beds by geese: a predictive model and test

The short-term impact of herbivores on plant productivity has been well studied. Demonstrating long-term effects of grazing on plant populations is much more difficult, but knowledge of such long-term effects is vital in understanding plant-herbivore interactions. We address this issue in a relatively simple plant herbivore system involving the dark-bellied brent goose Branta bernicla bernicla and two marine macroalgae, Enteromorpha spp. and Ulva lactuca, on which the geese graze. In 3 years of sampling, goose grazing was responsible for depleting between 23% and 60% of the algae in the autumn, while wave damage caused between 15% and 70% depletion. The degree of depletion in autumn had no effect on the biomass of algae present the following summer, suggesting no long-term consequences of grazing for the population dynamics of the algae. A model simulating the change in mean algal biomass over the autumn and winter, incorporating changes due to depletion by geese, wave action and productivity, successfully described the date at which geese abandoned the algal bed in six different years. These years varied in numbers of geese, strength and timing of storms and initial biomass available. The most important factor determining the date of abandonment of the algal bed was a tradeoff between the timing of storms and the numbers of geese. When storms occur early, most depletion is due to wave action and geese must abandon the bed early, regardless of the numbers grazing there. As the algae become depleted, the geese switch to feeding on saltmarsh, pastures or arable crops. The rate of depletion of algae affects the timing of this movement, and the model presented here enables the switch to be predicted.     

Effect of larval fish and nutrient enrichment on plankton dynamics in experimental ponds

The hypotheses that larval fish density may potentially affect phytoplankton abundance through regulating zooplankton community structure, and that fish effect may also depend on nutrient levels were tested experimentally in ponds with three densities of larval walleye, Stizostedion vitreum (0, 25, and 50 fish m^–3), and two fertilizer types (inorganic vs organic fertilizer). A significant negative relationship between larval fish density and large zooplankton abundance was observed despite fertilizer types. Larval walleye significantly reduced the abundances of Daphnia, Bosmina, and Diaptomus but enhanced the abundance of various rotifer species (Brachionus, Polyarthra, and Keratella). When fish predation was excluded, Daphnia became dominant, but Daphnia grazing did not significantly suppress blue-green algae. Clearly, larval fish can be an important regulator for zooplankton community. Algal composition and abundance were affected more by fertilizer type than by fish density. Inorganic fertilizer with a high N:P ratio (20:1) enhanced blue-green algal blooms, while organic fertilizer with a lower N:P ratio (10:1) suppressed the abundance of blue-green algae. This result may be attributed to the high density of blue-green algae at the beginning of the experiment and the fertilizer type. Our data suggest that continuous release of nutrients from suspended organic fertilizer at a low rate may discourage the development of blue-green algae. Nutrient inputs at a low N:P ratio do not necessarily result in the dominance of blue-green algae.     

Measurement of rates of grazing of the ostracod Cyprinotus carolinensis on blue-green algae

Measurements were made of the rates of grazing of the ostracod Cyprinotus carolinensis fed ¹⁴C-labelled filamentous blue-green algae (cyanobacteria). The grazing rate was a linear function of food concentration at densities below 1.1 µg dry weight of algae · ml^-1 and independent of concentration at densities above 11.5 µg algae · ml^-1. Starvation affected grazing rates significantly, but light vs. dark feeding, animal density, and the volume of feeding container did not. Grazing on Nostoc sp. was a linear function of ostracod size. Respiration of C. carolinensis and the blue-green algae was not a significant factor in the tests of grazing.     

Benthic development on large-scale engineered reefs: A comparison of communities among breakwaters of different age and natural reefs

Breakwaters represent large-scale engineered artificial reefs that can develop diverse and abundant communities and are likely to play an increasing role in marine ecosystems as human populations grow in coastal urban areas. Information on how these communities develop and if and when these communities begin to resemble those on natural hard-bottom habitat is essential for marine management, but is not well understood. In this study, benthic communities on six breakwaters ranging from 1 to 31 years of age were compared to provide an understanding of patterns of community development on engineered coastal defenses, and these were compared to communities on natural reefs to gain an understanding of how communities develop on artificial structures relative to those in natural habitats. Multivariate analyses indicated that benthic communities on breakwaters became more similar to natural reefs with increasing age, but that communities on even the most mature (31 years) breakwater were distinct from those on natural reefs (ANOSIM p < 0.001). Generally, breakwaters ≤5.5 years had higher abundance of turf algae, sponges, bivalves, and bare pavement, while more mature (≥25 years) breakwaters were dominated by corals. Coral cover on 25 and 31 years old breakwaters (46% and 56%, respectively) was significantly higher than on natural reefs (37%; HSD test p < 0.05 and p < 0.001, respectively). These results indicate that breakwaters develop benthic communities that continue to change over periods exceeding 31 years, and that although they become more similar to communities on natural reefs with increasing age, these communities remain distinct.     

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.