Food preferences and related behavior of the browsing sea urchin <Emphasis Type="Italic">Tripneustes gratilla</Emphasis> (Linnaeus) and its potential for use as a biological control agent

The short-spined toxopneustid sea urchin Tripneustes gratilla feeds on a wide variety of algal species and on sea grasses. However, the urchin does show preferences when offered a selection of macroalgal species, which it encounters in nature. Preferences among macroalgae were evident in field-collected urchins exposed to pair-wise tests where the variable was either the consumption rate of the algae or observation of which algal species the urchins chose to touch with their lantern teeth. Exposure of lab-housed urchins to one of five species of macroalgae for 5 months did not seem to alter preferences of urchins in three of the exposure groups, but those exposed to Padina sanctae-crucis seemed to show an enhanced preference for this species when offered a choice of the five species of macroalgae at the end of the exposure period, and those exposed to Gracilaria salicornia seemed to avoid the species when offered the choice of the five species. Perhaps more ecologically important than their preferences were two other observations on these urchins: first, when offered only a single species of algae, the urchins on four of five diets ate the same quantity per day. Second, when simultaneously offered the choice among the five macroalgal species, the urchins consumed more macroalgae per day than when offered only one species. These urchins move about a meter a day. They probably encounter food resources in a relatively coarse-grained fashion and have evolved to eat what is available. Because of their limited movements, their habitat overlap with grazing fishes, their acceptance of a wide variety of macroalgae and their preference for macroalgae, these native urchins are thought to have the potential to serve as biological control agents of alien and invasive macroalgae, which have come to dominate some reef zones normally occupied by corals in Hawaii.     

Variability in grazer-mediated defensive responses of green and red macroalgae on the south coast of South Africa

Variabilities in the responses of several South African red and green macroalgae to direct grazing and the responses of one green alga to cues from grazers were tested. We used two feeding experiments: (1) testing the induced responses of three red and one green algae to direct grazing by mesograzers and (2) a multi-treatment experiment, in which the direct and indirect effects of one macrograzer species on the green alga Codium platylobium were assessed. Consumption rates were assessed in feeding assays with intact algal pieces and with agar pellets containing non-polar extracts of the test algae. Defensive responses were induced for intact pieces of Galaxaura diessingiana, but were not induced in pellets, suggesting either morphological defence or chemical defence using polar compounds other than polyphenols. In contrast, exposure to grazing stimulated consumption of Gracilaria capensis and Hypnea spicifera by another grazing species. In the multi-treatment experiment, waterborne cues from both grazing and non-grazing snails induced defensive algal traits in C. platylobium. We suggest that inducible defences among macroalgae are not restricted to brown algae, but that both the responses of algae to grazers and of grazers to the defences of macroalgae are intrinsically variable and complex.     

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

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

Settlement of larval blacklip abalone, Haliotis rubra, in response to green and red macroalgae

Surfaces from the habitat of adult Haliotis rubra were tested as inducers of larval settlement to determine the cues that larvae may respond to in the field. Settlement was high on the green algal species Ulva australis and Ulva compressa (Chlorophyta), the articulated coralline algae Amphiroa anceps and Corallina officinalis, and encrusting coralline algae (Rhodophyta). Biofilmed abiotic surfaces such as rocks, sand and shells did not induce settlement. Ulvella lens was also included as a control. Treatment of U. australis, A. anceps and C. officinalis with antibiotics to reduce bacterial films on the surface did not reduce the settlement response of H. rubra larvae. Similarly, treatment of these species and encrusting coralline algae with germanium dioxide to reduce diatom growth did not significantly reduce larval settlement. These results suggest that macroalgae, particularly green algal species, may play an important role in the recruitment of H. rubra larvae in the field and can be used to induce larval settlement in hatchery culture.     

Waterborne chemical compounds in tropical macroalgae: positive and negative cues for larval settlement

Settlement sites of marine invertebrate larvae are frequently influenced by positive or negative cues, many of which are chemical in nature. Following from the observation that many shallow-water, Hawai'ian marine macroalgae are free of fouling by sessile invertebrates, we predicted that the algae are chemically protected and dependent on either surface-bound or continuously released soluble compounds to deter settling invertebrate larvae. To address the importance of waterborne algal compounds, we experimentally determined whether larvae of two of Hawai'i's dominant hard-surface fouling organisms, the polychaete tube worm Hydroides elegans and the bryozoan Bugula neritina, would settle in the presence of waters conditioned by 12 species of common Hawai'ian macroalgae (representing the Phaeophyta, Chlorophyta, Rhodophyta and Cyanophyta). The results included a full spectrum of biological responses by each larval species to waterborne algal compounds. Larval responses to conditioned water were consistent for each algal species, but the outcomes were not predictable based on the taxonomic relationships of the algae. For example, among the species of Phaeophyta examined, different conditioned waters were: (1) toxic, (2) inhibited settlement, (3) simulated settlement, or (4) had no effect, compared to larvae in control dishes containing filtered seawater. Additionally, larval responses to aged (24 h) conditioned waters could not be predicted from the results of assays run with conditioned waters utilized immediately after preparation. Finally, settlement by larvae of one species did not predict outcomes of tests for the other species. Four of 12 shallow-reef Hawai'ian macroalgae tested released compounds into surrounding waters that immediately killed or inhibited settlement by both H. elegans and B. neritina (toxic: Dictyota sandvicensis; inhibitory: Halimeda discoidea, Sphacelaria tribuloides, Ulva reticulata); the remaining 8 algal species prevented settlement by one of these fouling organisms but for the other had no effect or, in some cases, even stimulated settlement     

Effects of nutrients, herbivory, and depth on the macroalgal community in the rocky sublittoral

We studied the interacting roles of nutrient availability and herbivory in determining the macroalgal community in a rocky littoral environment. We conducted a factorial field experiment where we manipulated nutrient levels and herbivory at two sublittoral depths and measured macroalgal colonization and the following young assemblage during the growing season. At the community level, grazing reduced algal colonization, though the effect varied with depth and its interaction with nutrient availability varied in time. In shallow water, the total density of macroalgae increased in response to nutrient enrichment, but the ability of grazers to reduce macroalgal density also increased with the nutrient enrichment, and thus, the community could not escape from the top-down control. In deep water, the algal density was lower, except in July when nutrient enrichment caused a very dense algal growth. Grazing at the greater depth, though effective, was generally of smaller magnitude, and in July it could not limit algal recruitment and growth. Species richness peaked at the intermediate nutrient level in deep but not in shallow water during most of the growing season. Grazing had no effect on diversity of the algal community at either depth and only a minor effect on species richness at the greater depth. Opportunistic and ephemeral algae benefited from the nutrient enrichment but were also grazed to very low densities. Slowly growing and/or perennial species colonized poorly in the nutrient enriched treatments, and depending on the species, either suffered or indirectly benefited from herbivory. For all species, effects of nutrients on colonization depended on depth; usually both nutrient and herbivory effects were more pronounced at the shallow depth. We conclude that grazers are able to reduce macroalgae over a large range of nutrient availabilities, up to 12-fold nutrient enrichment in the current experiment, and that the sublittoral depth gradient generates variation in the algal community control exerted by both herbivory and nutrient availability. Thus temporal and spatial variability in both top-down and bottom-up control and in their interaction, especially along the depth gradient, may be crucially important for producer diversity and for the successional dynamic in a rocky sublittoral environment.     

Resource utilization for decorating in three intertidal majid crabs (Brachyura: Majidae)

 For three spider crabs (Tiarinia cornigera, Micippaplatipes and Pugettia quadridens quadridens), patterns of algal utilization for decorating were compared with the dynamics of algae on an intertidal rocky shore reef where the crabs co-occurred. T. cornigera and P. quadridens quadridens were most abundant from autumn to spring when the dominant algae (Boodleacoacta, Sargassum hemiphyllum, S. thunbergii and Corallina pilulifera) occurred in high coverage, while M. platipes was most abundant from spring to autumn. Monthly change of algae used for decorating was not correlated with algae growing in the crab habitat for T. cornigera, but for M. platipes, it was positively correlated for two algal species, and for P. quadridens quadridens, negatively correlated for one algal species. Each species of the spider crabs used some algal species preferentially for decoration. Decorating preference experiments conducted in the laboratory showed that M. platipes and P. quadridens quadridens exhibited similar preference to their algal utilization in the field, whereas for T. cornigera, algal preference in the experiment differed from utilization in the field. Comparisons between materials used for decoration and gut contents revealed that T. cornigera and M. platipes used algal species differently for decorating and feeding, while P. quadridens quadridens used the same algal species for both decorating and feeding. Different tactics for camouflage are discussed in terms of algal utilizations by the three majid species. Received: 25 September 1999 / Accepted: 22 June 2000     

Effects of micro- and mesograzers on intertidal macroalgal recruitment

This study examined the effects of a guild of micrograzing harpacticoid copepods (dominated by two species of Paradactylopodia sp. nov. and one species of Scutellidium sp. nov.) and a mesograzing periwinkle, Afrolittorina praetermissa, on the early recruitment of intertidal macroalgae on a wave-exposed, rocky shore. This is the first study, to our knowledge, to examine the effects of micrograzers (<500 μm) on intertidal macroalgal recruitment. Data showed that microscopic harpacticoid copepods altered the assemblages and reduced the densities of several macroalgal taxa, while A. praetermissa changed the assemblages and reduced both the density and number of macroalgal taxa. Recruitment of encrusting coralline algae was actually higher in copepod inclusions than exclusions, suggesting that copepods may be beneficial to the recruitment of this algal group. These results contribute to the understanding of grazing as a factor causing high mortality of algal recruits, but also highlight the need for more studies that examine the effects of micro- and mesograzers on the distribution and abundance of macroalgae.     

Herbivory by the Caribbean king crab on coral patch reefs

Caribbean coral reefs are increasingly dominated by macroalgae instead of corals due to several factors, including the decline of herbivores. Yet, virtually unknown is the role of crustacean macrograzers on coral reef macroalgae. We examined the effect of grazing by the Caribbean king crab (Mithrax spinosissimus) on coral patch reef algal communities in the Florida Keys, Florida (USA), by: (1) measuring crab selectivity and consumption of macroalgae, (2) estimating crab density, and (3) comparing the effect of crab herbivory to that of fishes. Mithrax prefers fleshy macroalgae, but it also consumes relatively unpalatable calcareous algae. Per capita grazing rates by Mithrax exceed those of most herbivorous fish, but Mithrax often occurs at low densities on reefs and its foraging activities are reduced in predator-rich environments. Therefore, the effects of grazing by Mithrax tend to be localized and when at low density contribute primarily to spatial heterogeneity in coral reef macroalgal communities.     

The role of the amphipod <Emphasis Type="Italic">Gammarus locusta</Emphasis> as a grazer on macroalgae in Swedish seagrass meadows

Mesograzers are thought to play a critical role in seagrass beds by preventing overgrowth of ephemeral algae. On the Swedish west coast, eelgrass Zostera marina has decreased in recent decades as a result of eutrophication and increased growth of macroalgal mats (mainly filamentous Ulva spp. and Ectocarpales), with no indication of grazer control of the algae. The aim of this study was to investigate the ability of the amphipod Gammarus locusta to control algal blooms during nutrient-enriched and ambient conditions, using a combination of laboratory, field and model studies. Laboratory experiments demonstrated that juvenile and adult G. locusta could consume both Ulva spp. and Ectocarpales, but that consumption of Ulva spp. was significantly higher. Cannibalism was common in individual treatments involving multiple size-classes of G. locusta, but only large, male gammarids consumed smaller juveniles in the presence of Ulva spp. as an alternative food source. However, no negative effects of cannibalism were found on total grazing impact. A model using size-specific grazing rates and growth rates of G. locusta and of Ulva spp. suggests that approximately 62 young juvenile, or 27 adult G. locusta are needed per gram DW of Ulva spp. to control the algal growth during ambient nutrient conditions, and approximately 2.6 times as many gammarids during enhanced nutrient conditions. On the Swedish west coast, densities and mean sizes of G. locusta in eelgrass beds are below these critical values, suggesting that the gammarids will not be able to control the growth of the filamentous macroalgae. However, in the field cage experiment, immigration of juveniles and reproduction of encaged adult G. locusta resulted in unexpectedly high densities of G. locusta (>4,000 individual m⁻²), and very low biomass of Ulva spp. in both ambient and nutrient-enriched treatments. Although the high numbers of juveniles in all cages precluded any significant treatment effects, this suggests that in the absent of predators, the population of G. locusta can grow significantly and control the biomass of Ulva spp. Furthermore, low grazing of Ectocarpales in the laboratory and high biomass of these filamentous brown algae in the field indicate a preference for the more palatable green algae Ulva spp. This study indicates that the high grazing capacity of G. locusta, in combination with high reproduction and growth rates, would allow the amphipod to play a key role in Z. marina ecosystems by controlling destructive blooms of filamentous green algae. However, high predation pressure appears to prevent large populations of G. locusta in eelgrass beds on the Swedish west coast today.     

Turbinaria ornata as an herbivory refuge for associate algae

Habitat associations are an integral part of coral reef community structure. Commonly, one organism lives in such close association within or near another that a spatial refuge occurs, whereby one of the organisms provides protection to the other. This is often the result of defenses of the host deterring an associate organism’s consumers. In Moorea, French Polynesia, the range and abundance of the brown macroalga, Turbinaria ornata, have increased drastically since 1980 such that dense aggregations of this macroalga are a dominant component of the backreef habitat. Turbinaria ornata is both mechanically and chemically defended from herbivores. Other species of macroalgae grow within aggregations of Turbinaria and may benefit from these defenses. This study investigates whether aggregations of Turbinaria create a refuge from herbivory for associate macroalgae. When Turbinaria aggregations were removed experimentally, there was a significant increase in the number of associate algal species. Moreover, an herbivory assay using the palatable local alga Acanthophora spicifera identified herbivory as the mechanism for lower diversity on bommies lacking Turbinaria aggregations. The local increase in algal richness due to the refuge from herbivory afforded by Turbinaria may be an important contribution to macroalgal and community dynamics on reefs in Moorea, French Polynesia.     

Habitat modification affects recruitment of abalone in central New Zealand

The habitat experienced during early life-history stages can determine the number and quality of individuals that recruit to adult populations. In a field experiment, biogenic habitat complexity was manipulated (presence or absence of foliose macroalgae) at two depths (2–3 m and 5–6 m) and the habitat-dependent effects on recruitment of the black foot abalone (Haliotis iris) were examined at three field sites along the south coast of Wellington, New Zealand (41°20′S, 174°47′E), between July and November 2005. Recruit density (<5 weeks post-settlement) was measured on cobbles covered with crustose coralline algae. Habitats of low complexity (barrens treatments) had consistently greater densities of recruits than habitats of high complexity (algae treatments). However, recruits in algae habitats were larger, and for deep habitats, there was greater survival in algae habitats compared with barrens habitats. While depth had no significant effect on early recruit (<2 weeks post-settlement) density, late recruit (<5 weeks post-settlement) density was greater in shallow habitats, and so it seems recruit survival was greater in shallow habitats. In this experiment, algal habitat complexity had strong effects on early recruit abundance, but habitat-dependent variations in recruit growth and survival may modify initial patterns of abundance and determine recruitment to adult abalone populations.     

Comparing macroalgal food and habitat choice in sympatric, tube-building amphipods, Ampithoe lacertosa and Peramphithoe humeralis

For small tube-building amphipods that live on the algae they consume, food and habitat are tightly linked. This study compared two closely related amphipods to determine whether the species’ algal preferences are based on the food value of the algae or on some other aspect of their algal habitat. Ampithoe lacertosa and Peramphithoe humeralis are both abundant on Shannon Point beach (Anacortes, Washington, USA; 48°30.542′ N, 122°41.070′ W) but specialize on different algae. In observations and laboratory experiments conducted July–September 1997, 2007, and 2008, the two species exhibited markedly different choices of food and habitat when offered six common macroalgae. Ampithoe lacertosa ate all algae offered, but preferentially built tubes on the green alga Ulva lactuca. Survival was relatively low among juveniles maintained on single species diets, except when they were fed Mazzaella splendens. Conversely, P. humeralis consumed primarily the brown kelp Saccharina latissima, Alaria marginata, and Desmarestia ligulata and preferred those species for tube building. Juvenile P. humeralis could not survive on a diet of U. lactuca or M. splendens. While A. lacertosa builds simple, temporary tubes and relocates frequently, P. humeralis is a highly thigmotactic species that builds long-term, complex tubes on the alga it prefers to eat. Feeding and habitat preferences of the two species were not clearly linked to nitrogen content of the algae, C:N ratio, or toughness of the algal tissue. Instead, preferences of the species may be related to their mobility and the permanence of the tubes they build. Ampithoe lacertosa and P. humeralis also use different feeding strategies; the former appears to mix algae to produce a high-quality diet, while the latter is more selective and has a capacity for compensatory feeding. The species are abundant on the same protected rocky shores, but specialize on different algae for habitat and food. Results suggest that the nutritional requirements of these sympatric mesograzers differ considerably and even closely related species can exhibit divergent behavioral strategies.     

Early life histories in the bryozoan<Emphasis Type="Italic"> Schizobrachiella sanguinea</Emphasis>: a case study

Mediterranean populations of Schizobrachiella sanguinea seem to span a wide range of contrasting benthic habitats. We have studied adult, larval, and recruit populations of S. sanguinea from sublittoral (approximately 10 m depth) communities at Blanes (Spain, NW Mediterranean). At the study site, the species colonises semi-obscure caves and, to a lesser extent, communities of sciaphilic algae. Our first goal was to show the periods of brooding, larval release, and recruitment. Plankton was collected about twice a week between July 1998 and June 2000 over an artificial reef. A total of 303 larvae were collected in 23 of the 102 days of sampling. At the study site the species shows a single, annual larval release period, which occurs from late March to June–July every year. Recruits were found from late May to late August 1999 and in late May 2000. A recruitment peak was observed in July 1999 and again in late May 2000. The presence of brooding adults corresponded to the periods of larval supply and recruitment. Our second goal was to investigate the causes of the variable distribution of the species along several communities in the north-west Mediterranean Sea. We describe adult distribution and the dynamics of larval supply, recruitment of early (15 days from settlement) and late (4 months from settlement) recruits. Patterns of larval presence and early recruit distribution along the communities on the reef did not reflect the adult distributions, whereas this was closely matched by the distribution of late recruits. Post-recruitment mortality strongly affected recruits of the PA (photophilic algae) and SA (sciaphilic algae) communities, whereas it was very low in the communities of SOC (semi-obscure caves). Mortality of recruits was density independent and colony survivorship in the SA communities, where recruitment was the highest, appeared strongly affected by competition with brown algae. Like the majority of cheilostomate bryozoans, S. sanguinea releases a coronate larva capable of active swimming and positive reaction to light. Such swimming abilities may allow larvae to actively select the substrate on which to settle and to avoid unsuitable substrates. It seems that larval behaviour does not explain the preference of S. sanguinea for sciaphilic habitats. High post-recruitment mortality due to various factors, especially competition with fast-growing algae, seems particularly relevant in determining the adult distributions observed.Communicated by S.A. Poulet, Roscoff     

Inhibition of larval settlement to a soft bottom benthic community by drifting algal mats: An experimental test

The hypothesis that drifting red algal mats inhibit settlement of planktonic larvae was tested in a field experiment in 1986. Substratum free of algae (caged boxes) was compared with substratum covered with algae (natural substratum and open boxes). Whereas settling densities of 1500 to 5500 ind. m^-2 of the bivalves Macoma balthica, Cardium glaucum and Mya arenaria were observed in sediment without algal cover, no recruits of the same species were found beneath the algae during the period of peak settlement (June to July). The same difference was observed for the polychaete Nereis diversicolor, although in this case occasional individuals were found beneath the algae. The results demonstrate that algae mats may be efficient inhibitors of larval settlement to benthic soft-substratum communities. It is concluded that algae act as a larval filter.     

Tissue type matters: selective herbivory on different life history stages of an isomorphic alga

Selective grazing by herbivores can have large effects on the population dynamics and community structure of primary producers. However, the ecological impacts of within-species herbivore preference for tissues of different phases (e.g., ploidy levels) or reproductive status remain relatively poorly known, especially among algae and other species with free-living haploid (gametophyte) and diploid (sporophyte) phases. We tested for herbivore selectivity among tissue types of the isomorphic (identical haploid and diploid free-living stages) red alga Mazzaella flaccida. Laboratory feeding assays demonstrated that the snail Tegula funebralis exhibited more than a threefold preference for gametophyte reproductive tissue over other tissue types, due to morphological differences. In contrast, the urchin Strongylocentrotus purpuratus did not distinguish as clearly between gametophytes and sporophytes; but it did prefer sporophyte reproductive to nonreproductive tissue, due to differences in water-soluble chemicals. Field surveys of grazer damage on M. flaccida blades were consistent with these laboratory preferences, with more damage found on gametophytes than sporophytes and reproductive than nonreproductive tissues. Differential fecundity can contribute to a skew in relative frequencies of phases in the field, and our results suggest that differential grazing by snails may contribute to this pattern and thus play a role in algal population biology.     

Modelling the effects of green macroalgae blooms on the population dynamics of Cyathura carinata (Crustacea: Isopoda) in an eutrophied estuary

A population dynamics model was developed to simulate the effects of benthic macroalgae blooms (mostly Enteromorpha spp.) on the productivity of Cyathura carinata (Crustacea: Isopoda), a possible keystone species in the benthic communities of the Mondego estuary. The model describes C. carinata population dynamics, as well as the relationships between Enteromorpha biomass, Enteromorpha decaying rates, organic matter content in the sediments and detritus consumption by C. carinata, a detritic feeder. Model results support the idea that seasonal blooms of Enteromorpha determine a significant increase of organic matter content in the sediments, due to macroalgae decay, which initially contributes to enhance C. carinata consumption and growth rates, determining a significant increase in the biomass. Nevertheless, later, following the algae bloom, C. carinata biomass decreases, and reaches its lowest value, close to 0, when the algae crash. This effect is probably related with strong anoxic conditions, especially during night, due to high algal decomposition rates. In accordance with the model, migration of new individuals from adjacent areas must occur in order to recolonise the area affected by the algae bloom. Therefore, it seems reasonable to conclude that macroalgae blooms that are limited in space may favour C. carinata populations, but extensive blooms affecting the whole area of distribution of this species will determine its disappearance.     

Spatial variation in caddisfly grazing regimes within a northern California watershed

Ecologists seek better understanding of why species interactions change across space and time in natural communities. In streams, species effects on resources and community structure may change as physical characteristics of the stream environment change along drainage networks. We examined spatial and seasonal effects of armored grazers using a small-scale exclusion experiment that was replicated in streams of different drainage areas. Effects of grazing varied with stream size and were related to variation in grazer abundance and phenology. We identified three distinct grazing regimes and a stream size (drainage area [DA]) threshold corresponding to a shift from one to two functional trophic levels. In streams with DA < 1 km2, armored grazers did not reduce biomass of algal biofilms. In slightly larger streams (2-3 km2 DA), the armored grazer guild was dominated by bivoltine Glossosoma. These caddisflies persisted and limited algal biofilms throughout the summer in one of these streams. In the largest tributaries (DA > 10 km2), the grazer guild was dominated by univoltine caddisflies, and grazing limited algal biofilms in early summer, but not late summer, after caddisflies pupated. Drainage area is a useful predictor of spatial transitions in food web interactions within and among watersheds. Quantifying the drainage area threshold at which interactions change in catchments with differing geology, vegetation, hydrology, climate, land use, or species pools should help build the understanding we need to forecast ecological responses to environmental change.     

Habitat structure and complexity as determinants of biodiversity in blue mussel beds on sublittoral rocky shores

Habitat-forming, ecosystem engineer species are common in most marine systems. Still, much uncertainty exists about how individual and population-level traits of these species contribute to ecosystem processes and how engineering species jointly affect biodiversity. In this manipulative field experiment, we examined how biodiversity in marginal blue mussel beds is affected by blue mussel (1) body size, density and patch context and (2) presence of fucoid and algal structures. In the study area, bladder-wrack (Fucus vesiculosus), filamentous algae and blue mussels (Mytilus edulis) coexist at shallow depths in a variety of patch configurations and offer complex habitats with a high variability of resources. We hypothesized that complexity in terms of mussel bed structure and algal presence determines species composition and abundance. Results from the experiment were compared with macrofaunal communities found in natural populations of both engineering species. Results show that the physical structure and blue mussel patch context are important determinants for species composition and abundance. Results further show that the presence of algal structures positively affects diversity in blue mussel habitats due to increased surface availability and complexity that these algae offer. This study shows that blue mussel beds at the very margin of their distribution have an indisputable function for promoting and maintaining biodiversity and suggest that facilitative effects of habitat-modifying species are important on Baltic Sea rocky shores with fundamental importance to community structure.     

Biomass, diversity and production of rocky shore macroalgae at two nutrient enrichment and wave action levels

The littoral zone of temperate rocky shores is normally dominated by perennial macroalgae (e.g. Fucus, Ascophyllum, Laminaria), but nutrient enrichment and/or permanently decreased wave action may lead to structural community changes from dominance of perennials to increased amounts of annual opportunistic species (mainly green algae). Macroalgal biomass, diversity and production as well as relationships between the two latter were studied using Solbergstrand’s rocky shore mesocosms in SE Norway in connection with a long-term experimental manipulation of nutrient addition and wave action (high and low levels of both factors applied in a crossed way to eight outdoor basins). After more than 2 years of experimental treatment, the total standing stock of macroalgae was larger in low nutrient than in high nutrient treatments as well as in high wave compared to low wave treatments (in autumn only). For macroalgal functional groups, bushy and filamentous brown and filamentous red algae were generally favoured by low nutrient concentrations, while annual filamentous and sheet-like green algae were stimulated by the nutrient enrichment. There was only one significant interaction between nutrient enrichment and wave action (for brown filamentous algae in autumn) and also only one significant main effect of the wave treatment (for bushy brown algae in autumn). Surprisingly, the high nutrient treatments supported a higher diversity of macroalgae, whereas the low nutrient treatments generally showed higher production rates. Moreover, significantly negative correlations were found between macroalgal diversity and primary productivity in both summer and autumn. This study shows that it is the biological components of the communities subjected to external forcing (nutrient addition or decreased wave action) that regulate production and this contradicts the common misperception that resource production in natural systems simply can be fast-forwarded by fertilization. The negative relationships between diversity and productivity, although a consequence of unexpected results for diversity and production, are also novel and hint towards species identities having more important functional consequences than general species dominance patterns and the amount of species per se. These results also emphasise the context dependency of findings within the field of biodiversity and ecosystem functioning.