Temporal and spatial variability in settlement of the sea urchin<Emphasis Type="Italic"> Paracentrotus lividus</Emphasis> in the NW Mediterranean

Settlement into the benthic habitat may be an important process in regulating sea urchin abundance, which potentially modifies the structure of benthic communities. Strong settlement events may increase sea urchin abundance beyond a certain threshold, leading to the formation of coralline barrens (overgrazed communities with a dominance of encrusting coralline algae). To understand the role of settlement in regulating sea urchin populations we first need to determine settlement variability. Temporal variation in settlement of the sea urchin Paracentrotus lividus was monitored at three sites in the Medes Islands, NW Mediterranean, during three settlement seasons (March 1998 through October 2000). Spatial variation in settlement was studied in 1999 at 50 sites along a gradient of exposures to waves and currents, inside and outside the archipelago, and separated by distances from tens to thousands of meters. Bathymetric distribution of settlement was also studied in 2000 at six sites at 5, 10, 15 and 20 m depths. Settlement of P. lividus occurred in a single annual peak within 3 weeks in May–June. Differences in settlement between years were more than two orders of magnitude. Spatial variability was found at all scales investigated, showing strong patchiness at the smallest spatial scales (tens of meters). Sea urchins settled preferentially at depths between 5 and 10 m. Substratum type, level of protection, and adult population densities were not significant in determining settlement. However, settlement was found to be related to the degree of exposure to waves and currents, indicating that physical processes are very important at the spatial scales investigated. This greatly variable settlement is a necessary, although not sufficient, condition to create gradients of adult P. lividus abundance. Further studies should be designed to investigate the interaction between settlement strength and post-settlement mortality.Communicated by O. Kinne, Oldendorf/Luhe     

Settlement and recruitment of sea urchins (Strongylocentrotus spp.) in a sea-urchin barren ground and a kelp bed: are populations regulated by settlement or post-settlement processes?

I sampled recruitment of very small sea urchins (Strongylocentrotus spp.) by using the anesthetic magnesium chloride to remove individuals from substrata collected in sea-urchin barren grounds (barrens) and kelp beds at Naples Reef near Santa Barbara, California, USA. Preliminary sampling found low numbers of newly settled individuals(<0.6 mm test diam) from April–July in 1984 and 1985, and in April, 1986. In early May, 1986, I found many newly settled seaurchins (0.3 to 0.6 mm, 5 to 17 d old), and I compared the densities of the cohort on several types of natural substrata in barrens and kelp-bed habitats. Newly settled individuals of both purple sea urchins (S. purpuratus) and red sea urchins (S. franciscanus) were present in similar, high densities (1 000 S. purpuratus m^-2) on foliose red algal turf, a dominant substratum ofthe kelp bed, and on crustose coralline algae, the dominant substratum of an adjacent barrens. Larvae of S. purpuratus reared and tested in the laboratory showed high rates of settlement on both red algal turf and on crustose coralline algae, but significantly lower rates on rock. Larvae also settled in response to a partiallypurified extract of coralline algae. The reduced settlement on natural rock surfaces relative to either algal treatment and the significant settlement in response to the extract of coralline algae indicate that larvae discriminate between natural substrata and probably respond to a settlement cue other than, or in addition to, a simple microbial (bacterial) film. The similar densities of young recruits of S. purpuratus on dominant substrata of barrens and kelp bed show that, at least in this case, differential settlement cannot explain the high densities of sea urchins in the barrens habitat. Movement between barrens and kelp bed is unlikely given the small sizes of the newly recruited sea urchins relative to the large distances often involved. Reduced post-settlement mortality of newly settled individuals in the barrens remains the most likely mechanism leading to the higher densities of sea urchins in barrens relative to kelp-bed habitats.     

Growth response of an early successional assemblage of coralline algae and benthic diatoms to ocean acidification

The sustained absorption of anthropogenically released atmospheric CO₂ by the oceans is modifying seawater carbonate chemistry, a process termed ocean acidification (OA). By the year 2100, the worst case scenario is a decline in the average oceanic surface seawater pH by 0.3 units to 7.75. The changing seawater carbonate chemistry is predicted to negatively affect many marine species, particularly calcifying organisms such as coralline algae, while species such as diatoms and fleshy seaweed are predicted to be little affected or may even benefit from OA. It has been hypothesized in previous work that the direct negative effects imposed on coralline algae, and the direct positive effects on fleshy seaweeds and diatoms under a future high CO₂ ocean could result in a reduced ability of corallines to compete with diatoms and fleshy seaweed for space in the future. In a 6-week laboratory experiment, we examined the effect of pH 7.60 (pH predicted to occur due to ocean acidification just beyond the year 2100) compared to pH 8.05 (present day) on the lateral growth rates of an early successional, cold-temperate species assemblage dominated by crustose coralline algae and benthic diatoms. Crustose coralline algae and benthic diatoms maintained positive growth rates in both pH treatments. The growth rates of coralline algae were three times lower at pH 7.60, and a non-significant decline in diatom growth meant that proportions of the two functional groups remained similar over the course of the experiment. Our results do not support our hypothesis that benthic diatoms will outcompete crustose coralline algae under future pH conditions. However, while crustose coralline algae were able to maintain their presence in this benthic rocky reef species assemblage, the reduced growth rates suggest that they will be less capable of recolonizing after disturbance events, which could result in reduced coralline cover under OA conditions.     

Foods and predators of the green sea urchin Strongylocentrotus droebachiensis in Newfoundland waters

Gut analyses of the green sea urchin Strongylocentrotus droebachiensis (O. F. Müller) demonstrated that perennial phaeophytes, mostly fucoids and Alaria esculenta, were predominant in the diet. Ephemeral species, coralline algae and animals, were consumed in smaller amounts when available. Grazing by the urchins is evidently responsible for the dearth of non-coralline sublittoral algae in Newfoundland waters. Lobsters, rock crabs, purple sea stars, other urchins, and a variety of fishes and birds feed on S. droebachiensis, but predation is apparently not effective in limiting the abundance of the urchin.Studies in Biology from the Memorial University of Newfoundland No. 234.Contribution from the Marine Sciences Research Laboratory No. 66.     

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.     

Bacteria on the surface of crustose coralline algae induce metamorphosis of the crown-of-thorns starfish Acanthaster planci

The crustose coralline alga Lithothamnium pseudosorum induces high rates of settlement and metamorphosis of larvae of the coral-eating crown-of-thorns starfish (Acanthaster planci). In cases where crustose coralline algae (CCA) induce metamorphosis of marine invertebrate larvae it is normally assumed that the inductive molecules are produced by the alga, but an alternative is that they originate from bacteria on the plant surface. Bioassays using shards of L. pseudosorum treated with several antibiotics, whereby some shards were reinfected with bacteria from the alga, showed that if bacteria populations are depleted then settlement and metamorphosis of larvae of A. planci are inhibited. This demonstrates that bacteria are necessary for induction and suggests that morphogenic substances are produced by bacteria on the surface of the alga and not directly by the alga itself. However, surface bacteria are not inductive if they are isolated from soluble algal compounds, suggesting either that they require a substrate from the alga to produce the inductive agents or, alternatively but less likely, that compounds from both the alga and bacteria are required. There is no evidence that inductive compounds derive from the alga, since algal cell debris and soluble extracts prepared from the alga do not induce metamorphosis of A. planci. This is the first time that induction of metamorphosis in a marine invertebrate by CCA has been shown to be mediated by bacteria associated with the alga.     

Resource allocation and growth rates in the sea urchin Evechinus chloroticus (Echinoidea: Echinometridae)

The morphometry of the sea urchin Evechinus chloroticus from habitats of contrasting algal abundance but of similar urchin density was examined at two localities in southern New Zealand during 1993. Populations from habitats of high algal abundance (Dusky Sound) had similar relationships of demipyramid (jaws) to test diameter, but individuals had significantly smaller jaws relative to their test diameter than those from a locality where algal abundance was low (Arapawa Island). The body wall mass (in relation to total wet weight) was similar for populations from both localities but, for Dusky Sound populations, individuals from exposed sites had greater relative body mass than those from sheltered sites. The ratios of gonad weight:total weight were similar between localities. However, E. chloroticus from Arapawa Island reached reproductive maturity at a smaller size than those from Dusky Sound. Growth rates of E. chloroticus varied among localities in Dusky Sound. Growth was modelled by the Tanaka function, which allows for rapid early growth until reproductive maturity is reached and declining growth rates thereafter. The results show that sea urchins respond to low resource availability by increasing the size of the food-gathering apparatus, maturing at a smaller size, and growing to a smaller size than individuals from food-rich habitats. Received: 3 December 1996 / Accepted: 29 January 1997     

Selective accumulation of mycosporine-like amino acids in ovaries of the green sea urchin Strongylocentrotus droebachiensis is not affected by ultraviolet radiation

Field sampling and laboratory experiments examined whether ultraviolet radiation (UVR) affects the reproduction or the accumulation of mycosporine-like amino acids (MAAs) and ascorbic acid in ovaries of the green sea urchin Strongylocentrotus droebachiensis (Müller). Ovaries of sea urchins sampled across a depth gradient (0.5–10 m) in March 1998 did not differ in their gonadal index, or in concentrations of MAAs, or ascorbic acid. Concentrations of MAAs and ascorbic acid in ovaries were higher in sea urchins collected from a kelp bed compared with those collected from a community of crustose coralline algae. The concentrations of MAAs in ovaries varied seasonally, peaking in March, when sea urchins had high gonadal indices just before spawning. Ovaries of sea urchins maintained on controlled diets from October 1997 to April 1998 accumulated significantly higher concentrations of MAAs when fed a diet rich in MAAs than did ovaries of sea urchins fed an alga lacking MAAs, but the gonadal indices did not differ between diets. Sea urchins accumulated principally one MAA, shinorine, but not others that were available in high concentrations in their diet. Neither the gonadal index nor the ovarian concentrations of MAAs were affected by daily exposure of adult urchins to UVR for 6 months. Concentrations of ascorbic acid in ovaries did not differ among diets or UV-treatments. The percentages of nutritive phagocytes and gametic cells were not affected by diet or UVR, and did not co-vary with concentrations of MAAs or ascorbic acid in ovaries. These data support previous demonstrations that female sea urchins accumulate MAAs from their diet of macroalgae, but further show that the accumulation is selective for specific MAAs, particularly shinorine, and that adult S. droebachiensis do not accumulate MAAs in their ovaries or eggs in response to UV-exposure. These are also the first experimental studies to address whether MAAs are affected by or regulate gametogenesis, and indicate that they do not. Received: 5 May 2000 / Accepted: 29 September 2000     

Population dynamics of the sea urchin Strongylocentrotus purpuratus in a Central California kelp forest: recruitment,mortality, growth,and diet

The population dynamics of Strongylocentrotus purpuratus inhabiting dense mats of geniculate coralline algae in a shallow central California Macrocystis pyrifera forest was examined. Sea otters had occupied the area for over two decades. Sea urchin density and size distribution were sampled during 1984–1986 to determine recruitment and mortality patterns. Growth rates were obtained from tetracycline-labeled individuals and changes in sizefrequency modes. Periodic collections were made to examine gut contents and the relationship of diet to food availability. The interaction of recruitment and mortality resulted in a dynamic population structure. Mean densities ranged from 6.5 to 12.7 urchins 0.25 m^-2. The population consisted primarily of urchins up to 40 mm in test diameter in a size distribution which changed from unimodal to bimodal and back over the 2 yr study. Mortality was temporally variable and related to test diameter. Growth rates were somewhat lower than most previous reports for the species and suggested that most of the population was made up of 1 and 2 yr-old individuals. Diet consisted largely of geniculate coralline algae, with fleshy brown algae becoming important when available as drift. This study showed high densities of small urchins can exist in a California kelp forest inhabited by sea otters, but regular recruitment may be necessary to maintain such populations.     

Larval bloom of the oviparous sponge Cliona viridis: coupling of larval abundance and adult distribution

 We performed an intensive year-round sampling with the aim of studying the abundance of sponge larvae in four Mediterranean benthic communities: photophilic algae, sciaphilous algae, semi-obscure (i.e. low light-intensity) caves and sandy bottoms. We record here for the first time, a larval bloom of Cliona viridis (Schmidt 1862), the most common excavating sponge in the Mediterranean, which took place simultaneously in several rocky communities of the Blanes sub-littoral (NE Spain), and discuss the role of restricted larval dispersal in the distribution of adult sponges. In the communities studied, C. viridis larvae bloomed synchronously once, in June. Spawning and consequent embryo development presumably occurred in May, when water temperature was 16 °C. The free larva is a small, evenly ciliated, weakly swimming parenchymella with low dispersal capabilities. The number of larvae m⁻³ and sponge abundance (as percent cover and biomass) were significantly higher in the community of sciaphilous algae than in the other communities studied. Because of limited larval dispersal, larval and adult abundance in the communities were positively correlated. Larvae developed into juvenile sponges 10 to 15 d after settlement. Settlers displayed distinctive features: a peripheral cuticle, vacuolar etching-like cells at the sponge base, absence of oscular chimneys, and the presence of zooxanthellae, which were presumably transmitted during oocyte maturation. Received: 24 January 2000 / Accepted: 4 July 2000     

Macroalgal morphology mediates particle capture by the corallimorpharian <Emphasis Type="Italic">Corynactis californica</Emphasis>

The shallow kelp forest at Santa Catalina Island, California (33.45 N, −118.49 W) is distinguished by several canopy guilds ranging from a floating canopy (Macrocystis pyrifera), to a stipitate, erect understory canopy (Eisenia arborea), to a short prostrate canopy just above the substratum (Dictyopteris, Gelidium, Laminaria, Plocamium spp.), followed by algal turfs and encrusting coralline algae. The prostrate macroalgae found beneath E. arborea canopies are primarily branching red algae, while those in open habitats are foliose brown algae. Densities of Corynactis californica, are significantly greater under E. arborea canopies than outside (approximately 1,200 versus 300 polyps m⁻² respectively). Morphological differences in macroalgae between these habitats may affect the rate of C. californica particle capture and serve as a mechanism for determining polyp distribution and abundance. Laboratory experiments in a unidirectional flume under low (9.5 cm s⁻¹) and high (21 cm s⁻¹) flow speeds examined the effect of two morphologically distinct macroalgae on the capture rate of Artemia sp. cysts by C. californica polyps. These experiments (January–March 2006) tested the hypothesis that a foliose macroalga, D. undulata, would inhibit particle capture more than a branching alga, G. robustum. G. robustum, found predominantly under the E. arborea canopy did not affect particle capture. However, D. undulata, found predominantly outside of the canopy, inhibited particle capture rates by 40% by redirecting particles around C. californica polyps and causing contraction of the feeding tentacles. These results suggest that the morphology of flexible marine organisms may affect the distribution and abundance of adjacent passive suspension feeders.     

Community development on subtidal temperate reefs: the influences of wave energy and the stochastic recruitment of a dominant kelp

Patterns of community development on subtidal rocky reefs in Marmion Lagoon, southwest Australia, were investigated with a settlement panel experiment. We tested the hypothesis that community development would differ between outer and inner reefs lines, because exposure to swell and wave energy was significantly greater on outer reefs. Following a 14-month deployment, we recorded pronounced variability between panels and sites, but did not detect any effect of wave exposure on the structure of panel assemblages. Subsequent data exploration suggested the importance of the presence of kelp recruits (Ecklonia radiata) in structuring the overall assemblage. Panel assemblages with kelp recruits were significantly different in structure to those without, principally because of greater space coverage of encrusting coralline algae and less coverage of red turfing algae, spirorbids, and bryozoans. Mature E. radiata act as ecosystem engineers in subtidal rocky reefs in southwest Australia. Our results suggested the importance of young, recruiting kelps in determining patterns of early community development on newly available hard substrata.     

A new apparatus for force measurement in marine bioadhesion

In an experimental study on the effect of parrotfish (probably Scarus taeniurus) grazing on the structure of benthic reef communities, fishes in densities of 0.6 to 1.5 parrotfish per m² or 9 to 17 g wet weight of fish per m² of feeding surface were found to have an optimum effect, resulting in the greatest benthic species richness and biomass on 2-dimensional surfaces. The presence of refuges (3-dimensional habitats), however, has a greater impact on bemthic community structure (number of species and biomass) than does just the density of parrotfish in such an experimental system. Coral recruitment is enhanced by the presence of refuges and, like coralline algae, is more successful under increased grazing pressure. These optimum densities of parrotfishes relate well to observed field densities where, in a collection from a Hawaiian patch reef, there were 1.1 fish or 10.8 g wet weight of parrotfish per square meter of collection area. The success of coralline algae and corals under high grazing pressure may have important consequences for the stability and structure of modern coral reefs.     

The larval settling response of the lined chiton Tonicella lineata

The lined chiton Tonicella lineata (Wood, 1815) is found on enerusting coralline algae in the lower rocky intertidal zone of Oregon and San Juan Island, Washington, USA. An encrusting coralline alga is the major food item of this chiton. Experiments were performed to test the settling response of T. lineata larvae to various algae and other substrata. In these experiments, the larvae would settle only on pieces of encrusting coralline algae and piecesof ceramic roofing tile soaked in a coralline algal extract. The settling stimulus is probably chemical in nature, and is inactivated by boiling. In laboratory cultures, normal development stops at the trochophore stage (110 to 160 h, depending on the temperature). Metamorphosis and further development will take place only after stttlement on encrusting coralline algae.     

Trophic cascades result in large-scale coralline algae loss through differential grazer effects

Removal of predators can have strong indirect effects on primary producers through trophic cascades. Crustose coralline algae (CCA) are major primary producers worldwide that may be influenced by predator removal through changes in grazer composition and biomass. CCA have been most widely studied in Caribbean and temperate reefs, where cover increases with increasing grazer biomass due to removal of competitive fleshy algae. However, each of these systems has one dominant grazer type, herbivorous fishes or sea urchins, which may not be functionally equivalent. Where fishes and sea urchins co-occur, fishing can result in a phase shift in the grazing community with subsequent effects on CCA and other substrata. Kenyan reefs have herbivorous fishes and sea urchins, providing an opportunity to determine the relative impacts of each grazer type and evaluate potential human-induced trophic cascades. We hypothesized that fish benefit CCA, abundant sea urchins erode CCA, and that fishing indirectly reduces CCA cover by removing sea urchin predators. We used closures and fished reefs as a large-scale, long-term natural experiment to assess how fishing and resultant changes in communities affect CCA abundance. We used a short-term caging experiment to directly test the effects of grazing on CCA. CCA cover declined with increasing fish and sea urchin abundance, but the negative impact of sea urchin grazing was much stronger than that of fishes. Abundant sea urchins reduced the CCA growth rate to almost zero and prevented CCA accumulation. A warming event (El Ni?o Southern Oscillation, ENSO) occurred during the 18-year study and had a strong but short-term positive effect on CCA cover. However, the effect of the ENSO on CCA was lower in magnitude than the effect of sea urchin grazing. We compare our results with worldwide literature on bioerosion by fishes and sea urchins. Grazer influence depends on whether benefits of fleshy algae removal outweigh costs of grazer-induced bioerosion. However, the cost-benefit ratio for CCA appears to change with grazer type, grazer abundance, and environment. In Kenya, predator removal leads to a trophic cascade that is expected to reduce net calcification of reefs and therefore reduce reef stability, growth, and resilience.     

Coral cavity sponges depend on reef-derived food resources: stable isotope and fatty acid constraints

The diet of cavity sponges on the narrow fringing reefs of Curaçao, Caribbean was studied. The origin and resources of the bulk food of these sponges, i.e., dissolved organic matter (DOM), were identified using stable carbon and nitrogen isotopes and fatty acid biomarkers. We found that phytoplankton and its derived DOM from the adjacent open sea and from reef overlying water is not the main source of food for most of the sponges examined nor is bacterioplankton. Interestingly, dual stable isotope signatures (δ¹³C_org, δ¹⁵N_org) and fatty acid biomarkers appoint coral mucus and organic matter derived from crustose coralline algae (CCA) as probable food sources for encrusting sponges. Mucus-derived DOM may contribute up to 66% to the diet of examined sponges based on results of dual isotope mixing model analysis. The contribution of CCA (as purported representative for benthic algae) was smaller with values up to 31%. Together, mucus- and CCA-derived substrates contributed for 48–73% to the diet of sponges. The presence of the exogenous fatty acid 20:4ω6 in sponges, which is abundant in coral mucus of Madracis mirabilis and in CCA, highlights these reef-derived resources as sources of nutrition for DOM feeding cavity sponges. The relatively high concentrations of exogenous 20:4ω6 in all sponges examined supports our hypothesis that the bulk of the food of the cavity sponge community is reef-derived. Our results imply that cavity sponges play an important role in conserving food and energy produced within the reef.     

The monopolization of understorey habitat by subtidal encrusting coralline algae: a test of the combined effects of canopy-mediated light and sedimentation

Encrusting-coralline-algae habitat monopolized the substrata (>74% cover) under canopies of Ecklonia radiata at four sites (~1 km apart) within each of four localities (>100 km apart) spanning >1000 km of continuous South Australian coastline. This monopolization was analyzed experimentally to test whether canopy-mediated shade and sedimentation account for this canopy–understorey association. I tested the hypothesis that initially different habitat types (turf-forming-algae habitat versus habitat dominated by encrusting coralline algae) will converge to become like those under E. radiata if subjected to lower light and accumulation of sediment in the absence of E. radiata. Convergence in the absence of canopies, but in physically similar environments as under canopies, provides strong evidence that understorey habitats are primarily dependent on the manipulated environments. The experiment was provided with sufficient time (338 days) to adequately test for convergence as evidenced by indistinguishable percentage covers of encrusting coralline algae between canopies and treatments of shade on previously unoccupied rock and between habitat types transplanted to canopies. Convergence of habitat types, however, did not occur under treatments of shade and sedimentation. The effect of lowering light accounted for ~52% of the effect of canopies on percentage cover of encrusting-coralline-algae habitat. The effect of reducing sediment accumulation under shade was negligible. The magnitude of unaccounted effects (~48%) highlights a need to assess alternative factors that act to exclude taxa from or include taxa in the understorey assemblage. It is clear that canopies place strong constraints on the presence and abundance of many taxa, but not encrusting-algae habitats which beneficially coexist as understorey. A more complete identification of such positive and negative effects is needed to improve our understanding of the conditions that produce readily recognizable canopy–understorey associations that are repeated with great fidelity on a regional scale.     

Structure of and relationships within and between the littoral,rock-substrate fish communities off four islands in the Canarian Archipelago

A total of 577 visual surveys (each of 5 min in duration and 100 m² in area) were conducted throughout 1990 and 1991 at 32 locations off four Canary islands (i.e., Alegranza, Fuerteventura, Gran Canaria and Tenerife) with the objects of describing the coastal fish community, comparing the differences in the fish fauna within and between these islands, and determining the biotic and abiotic factors related to the structure of the fish communities. A total of 76 species were recorded; the most common were Abudefduf luridus, Canthigaster rostrata, Chromis limbatus, Sparisoma cretense and Thalassoma pavo (94.28, 86.48, 52.34, 73.31, and 94.10% frequency of occurrence, respectively). The abundance and average size of the commercially important species was greater in those locations where there was less fishing pressure. The stepwise linear regression models were capable of explaining only a low amount of variation in the dependent variables (i.e., number of species, number of individuals, average size and species diversity) of the fish community. The independent variables recorded were date, time of day, depth, slope of the substrate, substrate type, percentage of sand, percentage of algae, algal height, number of sea urchins (Diadema antillarum) and the individual islands. An ANOVA, using the islands only as independent variables, indicated that each island contributed significantly to the variation in the four dependent variables and there were significant differences among the islands. Detrended correspondence analysis (DCA) and a two-way indicator species analysis (TWINSPAN) determined associations between species and environmental attributes of the survey locations. The patterns in the TWINSPAN analysis indicated that localities had faunal resemblances based on the island off which where they were located.     

Selective consumption and facilitation by mesograzers in adult and colonizing macroalgal assemblages

Grazing pressure on macroalgae in littoral communities may vary with algal species, the age of an algal individual and grazer identity. Previous studies on alga–grazer interactions have shown that grazer preference for an algal species may release another one from interspecific competition. We measured the impacts of four common grazer taxa and the natural grazer guild on macroalgal communities at both their colonization and adult stages, and compared the impacts to grazer exclosures. The grazer effects were stronger on colonizing than on adult macroalgae; grazers did not reduce the total density of adult algae. Grazers both feed on propagules and indirectly facilitate other algae, depending on the grazer or algal species. Hydrobia species increased the settlement of spores of the red alga Ceramium tenuicorne. Similarly, the gastropod Theodoxus fluviatilis tended to facilitate one crustose algal species, but decreased the propagule density of annual filamentous algae, suggesting a preference for one species to the advantage of another. Effects of crustacean mesograzers on the studied macroalgae were weak. These results indicate that northern Baltic macroalgae are limited to grazing mainly during their colonization stage.     

Influence of a dominant consumer species reverses at increased diversity

Theory and experiments indicate that changes in consumer diversity affect benthic community structure and ecosystem functioning. Although the effects of consumer diversity have been tested in the laboratory and the field, little is known about effects of consumer diversity in the subtidal zone, one of the largest marine habitats. We investigated the grazing effects of sea urchins on algal abundance and benthic community structure in a natural subtidal habitat of the Galápagos Islands. Three species of urchins (Eucidaris, Lytechinus, and Tripneustes) were manipulated in inclusion cages following a replacement design with three levels of species richness (one, two, and three species) with all possible two-species urchin combinations. Identity was the main factor accounting for changes in the percentage of substrate grazed and benthic community structure. Two out of the three two-species assemblages grazed more than expected, suggesting a richness effect, but analyses revealed that this increased grazing was due to a sampling effect of the largest and commercially valued urchin species, Tripneustes. Benthic community structure in treatments with Eucidaris, Lytechinus, and Tripneustes alone was significantly different at the end of the experiment, suggesting that resource use differentiation occurred. Communities in Tripneustes enclosures were characterized by abundant crustose coralline algae and grazed substrate, while those without it contained abundant green foliose algae (Ulva sp.). An unexpected emergent property of the system was that the most species-rich urchin assemblage underyielded, grazing less than any other assemblage with Tripneustes, effectively reversing its dominant influence observed in the two-species treatments. While further experiments are needed to discern the mechanisms of underyielding, it may be related to changing interspecific interactions as richness increases from two to three species or to density-dependent Tripneustes grazing. This study highlights the general importance of evaluating consumer richness effects across the entire range of species richness considered, as the performance of the most species-rich consumer assemblage could not be predicted by manipulations of intermediate levels of consumer species richness.