Feeding aggregations of sea stars (<Emphasis Type="Italic">Asterias</Emphasis> spp. and <Emphasis Type="Italic">Henricia sanguinolenta</Emphasis>) associated with sea urchin (<Emphasis Type="Italic">Strongylocentrotus droebachiensis</Emphasis>) grazing fronts in Nova Scotia

Migrating feeding aggregations (or fronts) of sea urchins can dramatically alter subtidal seascapes by destructively grazing macrophytes. While direct effects of urchin fronts on macrophytes (particularly kelps) are well documented, indirect effects on associated fauna are largely unknown. Secondary aggregations of predators and scavengers form around fronts of Strongylocentrotus droebachiensis in Nova Scotia. We recorded mean densities of the sea stars Asterias spp. (mainly A. rubens) and Henricia sanguinolenta of up to 11.6 and 1.7 individuals 0.25 m⁻² along an urchin front over 1 year. For Asterias, mean density at the front was 7 and 15 times greater than in the kelp bed and adjacent barrens, respectively. There was strong concordance between locations of peak density of urchins and sea stars (Asterias r = 0.98; H. sanguinolenta r = 0.97) along transects across the kelp–barrens interface, indicating that sea star aggregations migrated along with the urchin front at rates of up to 2.5 m per month. Size–frequency distributions suggest that Asterias at the front were drawn from both the barrens (smaller individuals) and the kelp bed (larger individuals). These sea stars fed intensively on mussels on kelp holdfasts and in adjacent patches. Urchin grazing may precipitate aggregations of sea stars and other predators or scavengers by incidentally consuming or damaging mussels and other small invertebrates, and thereby releasing a strong odor cue. Consumption of protective holdfasts and turf algae by urchins could facilitate feeding by these consumers, which may obtain a substantial energy subsidy during destructive grazing events.     

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.     

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     

Chemical defense in the eggs and embryos of antarctic sea stars (Echinodermata)

Fifteen species of reproductively mature echinoderms (11 sea stars, 3 sea urchins, 1 sea cucumber) were collected from McMurdo Sound, Antarctica, during austral spring and summer of 1985 and 1986; eggs and embryos were obtained, and were tested for ichthyonoxicity using the common marine killifishFundulus grandis as a model predator. Chemical deterrents occurred in the large, yolky eggs of the pelagic lecithotrophic sea starPerknaster fuscus and the planktotrophic sea starPorania antarctica. Brooded embryos of the sea starsDiplasterias brucei andNotasterias armata were also noxious. Significant ichthyonoxicity was not detected in the remaining 7 species of sea stars, 3 sea urchins, and 1 sea cucumber. Chemical deterrents were generally effective at concentrations below a single egg or embryo per agar test-pellet. Although chemicals found in these eggs and embryos are noxious to an allopatric fish, they may not be noxious to sympatric fish.     

Experimental evaluation of aggregation behavior in the sea urchin Strongylocentrotus droebachiensis

Defensive aggregation by Strongylocentrotus droebachiensis has been invoked as the fulcrum for the transformation of macroalgal beds into coralline barrens in the northwest Atlantic. We critically examined some of the mechanisms contributing to aggregation behavior by experimentally manipulating sea urchins, purported predators and food during autumn 1983 and spring 1984 both in the laboratory and field. We utilized several approaches to examine a range of sea urchin responses to the presence of food, tethered predators, caged predators, crushed con-specifics in the field and predators in laboratory tanks. Some of the field cages had the property of allowing free passage to sea urchins while retaining lobsters; this allowed distinctions to be made between artifacts caused by cage walls or topographic barriers and unrestricted behavioral responses of sea urchins. The results falsified the hypothesis that aggregations of S. droebachiensis are elicited by predators. Except in the presence of algae, sea urchins always avoided decapod predators (but not sea stars) and fled from the vicinity of injured conspecifics. However, avoidance behavior was subordinate to feeding responses, demonstrated by mass migrations of sea urchins into cages (with lobsters) to feed on algae. We reaffirmed by observation and manipulation previous studies which showed that sea urchins aggregate only in the presence of food. Two types of sea urchin groupings were delineated: (1) surficial two dimensional associations, often caused by topographic or other features which inhibited dispersal and (2) cohesive three-dimensional aggregations induced by food.     

Susceptibility of non-indigenous ascidian species in British Columbia (Canada) to invertebrate predation

Non-indigenous ascidians are known to significantly alter the structure and composition of benthic communities and adversely affect shellfish aquaculture by fouling both the cultured species and the infrastructure. The ability of these species to persist in new locations and their current and potential distributions are dependent upon physiological tolerances to environmental factors and biotic resistance to competition and predation. Despite significant data on global invasion patterns, potential biotic resistance to non-indigenous ascidians is poorly understood. We identified potential predators of four non-indigenous ascidians (Styela clava, Botryllus schlosseri, Botrylloides violaceus, and Didemnum vexillum) in British Columbia (BC), Canada in order to: (1) assess the potential for biotic interference to limit the establishment and/or spread of these ascidian species in BC, and (2) identify candidate species to be used as ascidian biofouling control agents in shellfish aquaculture. Using a series of single- and multiple-choice laboratory experiments, potential benthic predators (including various species of molluscs, echinoderms, and arthropods) were offered non-indigenous ascidians as prey. The sea urchins Strongylocentrotus droebachiensis and Strongylocentrotus franciscanus, the sea stars Dermasterias imbricata and Evasterias troschelii, the nudibranch Hermissenda crassicornis, and the crabs Cancer productus and Carcinus maenas were found to consume one or more species of non-indigenous ascidians in single-choice experiments. However, when provided a choice, all predators chose their respective preferred food over ascidians. Thus, predation alone is unlikely to prevent large-scale establishment and spread of non-indigenous ascidians in BC, but it may have the potential to significantly reduce localized populations of ascidians. Green sea urchins, S. droebachiensis, were found to be efficient grazers of all four ascidian species, consuming 12.7 ± 5.14 cm² (mean ± SD) of adult B. violaceus over a 3-day period, 15 ± 3.7 juvenile colonies of B. violaceus over a 2-day period, and 63 ± 28.8 juvenile colonies of B. schlosseri over a 2-day period. Using sea urchins as biological control organisms may significantly reduce ascidian fouling in shellfish aquaculture.     

Spatiotemporal coupling/decoupling of planktonic larvae and benthic settlement in decapods in the Scottish east coast

Settlement patterns and the relationship between meroplanktonic larvae and settlement in decapods were studied on the Scottish east coast. Artificial settlement substrates (ASS), deployed at two locations (sandy vs. rocky sea substrates), were employed to collect megalopae and newly settled juveniles. Abundance of meroplanktonic larvae was used as an indicator of larval supply. The results showed a clear seasonality in settlement rates, and in some cases, significant differences between sites were detected. Nevertheless, the interference of the ASS with the surrounding habitat limits the study of spatial variability in settlement rates. Significant cross-correlation was found between the abundance of megalopae and juveniles in the collectors and planktonic larval abundance a month earlier. For individual species, this relationship was observed only in Pisidia longicornis. Complexities caused by the great variety of pre- and post-settlement processes, alongside effects of secondary dispersals of early juveniles may have obscured the relationship between meroplanktonic larvae and juveniles in other species.     

Measuring settlement intensity of echinoderms on coral reefs

A pilot study was conducted at Davies Reef on the central Great Barrier Reef between 20 November 1991 and 20 January 1992 to assess the suitability of settlement collectors for measuring larval settlement rates of echinoderms on coral reefs. The collectors were deployed for two months during summer when many echinoderm species are known to spawn. A total of 657 newly settled echinoderms were recovered from just 47 collectors, each having a volume of just 0.005 m³ but with a utilisable surface area of about 1.44 m², indicating that competent-to-settle larvae were very abundant during the sampling period. Echinoids (7.8 sampler^-1) and ophiuroids (5.6 sampler^-1) were the most abundant groups on the collectors. Asteroids (0.2 sampler^-1) and crinoids (0.3 sampler^-1 were less common. The asteroids were all identified to species and included the crown-of-thorns starfish Acanthaster planci, Choriaster granulatus and Culcita novaeguineae. The newly settled echinoids could not be distinguished from each other but included Echinometra mathaei and Mespilia globulus. The abundance of each of the five different classes of echinoderms on the samplers was correlated with their abundance on the natural substratum. Significant spatial variability was found in settlement rates of echinoids over 1000s of metres and ophiuroids over 100s of metres, but not over smaller spatial scales. It is concluded that the collectors can provide a useful tool for monitoring spatial and temporal variability in settlement rates of echinoderms on coral reefs and for testing hypotheses about patterns of larval dispersal.     

Bioerosion in coral reef communities in southwest Puerto Rico by the sea urchin<Emphasis Type="Italic"> Echinometra viridis</Emphasis>

Bioerosion is one of the most important structuring forces in coral reef communities. The bioerosion impact of several species of fish, sponges and sea urchins have been estimated in the Caribbean; however, there is no information for one important species, the red sea urchin Echinometra viridis. This species can be found in high densities in many localities. In this study, bioerosion rates for E. viridis were estimated in two patch reefs off La Parguera, southwest Puerto Rico, using the population size-class distribution, average densities, and the CaCO ₃ content in fecal pellets produced over 24 h. Average densities of urchins along four depth intervals were estimated using 40-m transect lines and 1-m ² quadrats. Average size and size-structure distribution were estimated by measuring the diameter of 180–220 urchins haphazardly collected at each of the four depth intervals. The ignition–loss method was used to estimate the daily rate of bioerosion. Fecal pellets produced by the urchins over a 24 h period were collected in buckets, rinsed in fresh water, dried for 24 h at 70°C, and then burned in a furnace at 550°C, first to eliminate organics, and then at 1000°C until constant weight to determine the amount of calcium carbonate (CaCO ₃) in the fecal pellets. HCl (10%) was then added to the remainder of the sample to test for presence of CaCO ₃. Average individual CaCO ₃ bioerosion rates were estimated at 0.181±0.104 g day ^-1. Average densities (0.77–62.0 ind. m ^-2), size (2.01–2.44 cm) and average bioerosion rates (0.114–4.14 kg m ^-2 year ^-1) were significantly higher in shallow areas (1–3 m) in both reefs. Bioerosion rates were low compared to those reported for parrotfish, endolithic sponges and the black sea urchin D. antillarum, but they were higher than those reported for other small-sized sea urchins in the Caribbean and the Indo-Pacific.     

Genetic structure of the subtidal red alga Delisea pulchra

We used random amplified polymorphic DNA (RAPDs) to examine small-scale spatial genetic structure in the red alga Delisea pulchra (Greville) Montagne at two locations near Sydney, Australia. We examined genetic structure among plants at four spatial scales ranging from 2 km apart down to <50 cm apart between locations, among sites within locations, among quadrats within sites, and among plants within quadrats. Haploid stages of D. pulchra were absent from the populations studied, suggesting that they are maintained through asexual reproduction of diploid plants. Consistent with this, we found that 19 RAPD phenotypes scored in this study had multiple individuals, indicating the presence of clones in these populations. However, there were no RAPD phenotypes common to two locations separated by only 2 km. Analysis of molecular variance revealed that strong genetic differences occurred between plants from these two locations, with 46.3% of the total genetic variation occurring at this scale, most probably reflecting limited gene flow. Within each location, <25% of the genetic variation was attributable to differences among sites or quadrats, indicating gene flow at those smaller scales. Most of the variation within each location occurred at the smallest spatial scale, among plants within 0.25 m² quadrats. Nonetheless, some pairwise genetic distances (φST) between sites or quadrats within locations were large, indicating some genetic divergence on smaller scales. Genetic distance was independent of spatial distance within both locations, suggesting that fine-scale differences within locations were most probably caused by variation in fine-scale patterns of water movement or fine-scale natural selection. We assessed the impact of one potential selective agent, grazing sea urchins, on the fine-scale genetic structure of D. pulchra. There was no evidence that grazing by sea urchins affected the genetic structure of D. pulchra. In combination with demographic data, our results indicated that local populations of D. pulchra within locations were relatively open and that fine-scale genetic structure was probably constrained by gene flow. At the larger scale however, strong genetic differentiation indicated little gene flow between locations and restricted dispersal of spores. Received: 22 April 1999 / Accepted: 29 November 1999     

Severe storm disturbances and reversal of community structure in a southern California kelp forest

Regular observations made over a period of 5 yr in four permanent transects provided data on plant, sea urchin, and fish densities which indicate that two unusually severe winter storms in 1980 (Storm I) and 1983 (Storm II) had different effects on a southern California kelp-forest community. Storm I removed all canopies of the giant kelp Macrocystis pyrifera, but spared most understory kelps, mainly Pterygophora californica. Hence, the previously large accumulation of detached drift kelp, mostly M. pyrifera, disappeared. Denied their preferred diet of drift kelp, the sea urchins Strongylocentrotus franciscanus and S. purpuratus then emerged from shelters to find alternative food. Without effective predators, they consumed most living plants, including the surviving understory kelps. This weakened the important detritus-based food chain, as indicated indirectly by declining abundances of algal turf and fish (Embiotocidae) that eat small animals living in turf. In 1983, Storm II reversed the process by eliminating exposed urchins, while clearing rock surfaces for widespread kelp settlement and growth. By summer 1984, the kelp grew to maturity to form extensive canopies despite elevated water temperatures during summer and fall of 1983. Thus, severe storms may have vastly different effects on community structure, depending on the state of the community before the disturbance.     

Abalones and sea urchins in an area inhabited by sea otters

Abalones (Haliotis spp.) and sea urchins (Strongylocentrotus spp.) are part of the subtidal fauna in the kelp bed off Hopkins Marine Station, Pacific Grove, California, USA, a protected marine reserve. Although these animals have been preyed upon by sea otters for over 10 years, their densities are gubstantial (Haliotis spp.: 0.21/m²; Strongylocentrotus spp.: 0.22/m²), and two species, H. rufescens (Swainson) and S. purpuratus (Stimpson), have wide size distributions indicating broad age ranges. Most of the adult abalones and sea urchins occur clustered deep in crevices, either as a direct result of sea-otter predation or because ample drift of algal food reduces foraging activities. Abalones tend to occur in larger crevices than sea urchins, and competition for suitable crevice space may exist among these large, invertebrate, algal-drift feeders.     

Fish predators and scavengers of the sea urchin Paracentrotus lividus in protected areas of the north-west Mediterranean Sea

Direct observations of predation on 436 individuals of the sea urchin Paracentrotus lividus (Lamarck) were carried out in infralittoral rocky bottoms (between 5 and 20 m deep) in three Mediterranean marine reserves. The predator guild was composed of six fish species, the sparids Diplodus sargus and D. vulgaris being the main predators, and the labrid Coris julis a major predator of juvenile sea urchins. Four species attempted but failed to open sea urchins. The scavenger guild was most rich in species, with 17 species observed. Predation was size-dependent; the size of predators increased with increasing size of the sea urchins. The presence of two feeding guilds is suggested, one composed of sparids (Diplodus spp.), able to kill juvenile and adult sea urchins, and the other composed of labrids (mainly C. julis), which feed on juvenile sea urchins. To avoid the extension of overgrazed, barren areas created by P. lividus populations, fisheries' regulations should focus on major sea-urchin predators, chiefly D. sargus, D. vulgaris and C. julis. Received: 23 April 1997 / Accepted: 30 May 1997     

Comparative study of vanadium biokinetics in three species of echinoderms

Vanadium-48 (as vanadate) was used to study the uptake, tissue distribution, depuration and food-chain transfer of vanadium through 3 species of echinoderms: the seastar Marthasterias glacialis L., the sea urchin Paracentrotus lividus Lmk. and the holothurian Holothuria forskali D.Ch.; all were collected from the littoral zone near Monaco. Uptake by all species was relativelyslow; after 3 wk exposure, isotopic equilibrium had not been reached and whole-body concentration factors ranged from 5 and 7 in the holothurian and sea urchin, respectively, to 18 in the seastar. Sixty-three to 77% of the incorporated radiotracer was associated with the body wall or test, suggesting surface sorption as the principal mechanism governing uptake from water. Stable vanadium measurements confirmed the preponderance of this element in the external hard parts of the echinoderms; however, concentration factors based on stable vanadium levels were significantly higher than those measured experimentally. Subsequent vanadium depuration rates were also species-dependent, with biological half-times for loss ranging from approximately 50 d in the sea urchin and holothurian to 123 d in the seastar. Food-chain transfer experiments indicated that seastars can assimilate and retain a large fraction of the vanadium ingested with food whereas sea urchins appear to lack this capability. The relative importance of the water and food input pathway in achieving vanadium levels in echinoderms is discussed in light of results of ⁴⁸V distribution in experimental individuals and stable vanadium distribution in samples from the natural environment.     

Larval recruitment in response to manipulated field flows

Settlement responses to boundary-layer flow of several invertebrate taxa, including the hydroid Tubularia crocea, the bryozoans Bugula turrita and Schizoporella unicornis, and the tube-building polychaete Hydroides dianthus were studied in manipulated field flows in Great Harbor, Massachusetts, USA. During three experiments in 1989 and two in 1990, densities of newly-recruited larvae were measured on flat plates, whose flow regimes had been manipulated by altering the leading-edge configurations. Settlement responses to flow were strongly species-specific, with T. crocea preferring regions of both high turbulence and strong shear stress, and S. unicornis settling a most exclusively in regions of high shear stress. B. turrita settled most prominently in regions of reduced shear stress, exhibiting settlement patterns that closely approximated predictions from a model of passive particle contact. H. dianthus showed a moderate avoidance of regions with high shear stress. These results indicate that boundary-layer flows affect settlement of several common encrusting species, a probable consequence of larval behaviors such as substrate rejection or exploration in response to flow. These responses are likely to generate patchiness during initial colonization of natural habitats, and certainly affect colonization of settlement panels commonly used in marine ecological studies.     

Morpho-functional defences of Mediterranean sea urchins,<Emphasis Type="Italic"> Paracentrotus lividus</Emphasis> and<Emphasis Type="Italic"> Arbacia lixula</Emphasis>, against fish predators

Morpho-functional features potentially involved in defence mechanisms against fish predators (i.e. attachment tenacity, spine length, and test robustness and thickness) have been assessed in two Mediterranean sea urchins, Paracentrotus lividus and Arbacia lixula. All four morpho-functional features were significantly and positively related to individual size for both species of sea urchins. Test robustness (i.e. static load needed to break sea urchin tests) was significantly greater for A. lixula (from 3,450 to 15,000 g depending on size) than for P. lividus (1,180–11,180 g). Attachment tenacity (i.e. force needed to dislodge sea urchins from the rocky substrate) was greater in A. lixula (280–3,300 g) than in P. lividus (110–1,450 g), and the difference tended to decrease in relation to smaller sea urchin size. Spine length was greater in A. lixula (1.5–2.9 cm) than in P. lividus (0.5–2.3 cm), but the difference decreased for larger sea urchin size. Test thickness was slightly greater (but not significantly) in A. lixula (0.35–1.10 mm) than in P. lividus (0.12–0.90 mm). These results provide evidence that morpho-functional features of sea urchins could be involved in affecting predation rates by fishes upon P. lividus and A. lixula, with potential implications for the population structure and distribution patterns of the two sea urchins in shallow rocky reefs.Communicated by R. Cattaneo-Vietti, Genova     

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.     

Tidal height,rather than habitat selection for conspecifics,controls settlement in mussels

Settlement is a major determinant of intertidal populations. However, the energy costs of lost larvae are very high. Accordingly, arrival and attachment on suitable substrata are essential requirements for species’ survival. On the intertidal, the presence of cues left by adult or juvenile conspecifics could be vital for the successful establishment of larvae arriving on the shore. Two mussel species, the indigenous Perna perna and the invasive Mytilus galloprovincialis, co-occur on the lower eulittoral zone on the south coast of South Africa. P. perna dominates the low and M. galloprovincialis the high mussel zones, with co-existence in the mid mussel zone. This study tested the hypothesis of settlement selectivity for conspecifics in these two mussel species, to understand whether the final adult distribution of mussels on the shores is determined by active behavioural and chemical mechanisms. Preferential selection by larvae for conspecifics was tested in the field during the peak settlement period in 2004 in natural mussel beds across zones and through manipulative experiments in the mid-zone where the species co-exist. On natural beds, settlement was determined by counts of settlers attached over 48 h onto artificial collectors. Collectors were placed on beds of P. perna and M. galloprovincialis present at both high- and low-adult densities, as well as in mixed beds. On such natural beds, settlers of both species consistently favored low-zone P. perna beds. Settlement patterns over 24 h onto experimentally created mussel patches consisting of P. perna, M. galloprovincialis or the two species combined beds, set in the mixed zone, did not conform with the results of the natural beds study: settlers of both species settled with no discrimination among different patches. The results indicate that mussels, which are sedentary, lack attraction to conspecifics at settlement. This highlights the importance of tidal height in setting settlement rates, and of post-settlement events in shaping populations of these broadcast spawners.     

The role of behavioral responses to predators in modifying urchins' (Strongylocentrotus droebachiensis) destructive grazing and seasonal foraging patterns

We documented spatial and temporal patterns of urchins (Strongylocentrotus droebachiensis) and periwinkles (Littorina littorea) in three habitats: a persistent Laminaria longicruris and L. digitata bed; an urchin dominated barrens, and the edge of the kelp bed that formed a boundary between the two. Urchins were rare in the kelp and, when present, always large and well hidden, a pattern we interpret as a response to crab and lobster predation. Urchins were abundant in the barrens, and, in the summer when predaceous fish were active during the day, foraged only at nigh. We observed the formation of a dense urchin feeding front along the kelp bed edge, and these urchins remained exposed and feeding even during the summer. Laboratory experiments demonstrated that aggregations are an effective defense against some predators, and that the presence of crabs increases the tendency of large urchins to aggregate. We hypothesize that healthy Laminaria spp. beds persist because kelp bed associated predators keep urchins at low densities and in hiding. A reduction in predation pressure permits urchin densities to increase to the point where they form aggregations, which provide better defense than hiding. These aggregations then graze destructively on Laminaria spp., forming barrens. These barrens seem to be a new, stable configuration of the system.     

Crushing predation of the spiny star Marthasterias glacialis upon the sea urchin Paracentrotus lividus

Literature data report that only fish predators are able to crush sea urchin tests in Mediterranean rocky reefs. This experimental study showed that the spiny star Marthasterias glacialis is able to break Paracentrotus lividus tests and that the breaking event is more likely to occur for small-sized sea urchins than for big ones. Our results show that the role of M. glacialis in regulating P. lividus population density can be important in specific locations. They may have important implications, moreover, for the use of tethering techniques aimed at identifying predator types of sea urchins.