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     

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     

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.     

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     

Distribution patterns of sea urchins and barrens in shallow Mediterranean rocky reefs impacted by the illegal fishery of the rock-boring mollusc<Emphasis Type="Italic"> Lithophaga lithophaga</Emphasis>

Shallow rocky habitats in SW Apulia (SE Italy, Mediterranean Sea) were surveyed in late spring 2002 to assess distribution patterns of sea urchins (Paracentrotus lividus and Arbacia lixula) and barren habitats (coralline barrens and bare substrates) in rocky reefs impacted by the destructive fishery of the rock-boring date-mussel Lithophaga lithophaga. Sea urchin density, test size-structure and biomass, and the percent cover of barrens were evaluated at four locations (5–6 km apart from each other), two heavily impacted by the date-mussel fishery and two controls. Sea urchin density and barren habitat cover were assessed at two and three sites (100–300 m apart), respectively, within each location. Sea urchin biomass was evaluated only at the scale of locations. Average density of P. lividus did not significantly change between impacted locations and controls, whereas A. lixula showed a greater density at the impacted locations. Distribution patterns of A. lixula, in addition, differed at the spatial scale of a few metres between impacted locations and controls, being generally more aggregated at the controls. The size-frequency distribution (test diameter) of P. lividus showed a mode at 3–4 cm at the impacted locations compared to a mode at 2–3 cm in the controls. The size-frequency of A. lixula was bimodal at the damaged locations (with modes at 1–2 and 4–5 cm, respectively) and unimodal (with the mode at 4–5 cm) at the controls. Average biomass of both sea urchins (P. lividus and A. lixula) was two- to fourfold greater at the impacted locations (~600 g wet wt m^–2) than at the controls (150–250 g wet wt m^–2). Barren habitats had a far greater average cover (mainly of macroalgae) at the impacted locations (from 79% to 96%) than at control locations (from 7% to 21%). These results show that the date-mussel fishery may have the potential to affect distribution patterns of sea urchins and to greatly enhance the percent cover of barren grounds in shallow Mediterranean rocky reefs.Communicated by R. Cattaneo-Vietti, Genova     

Differences in somatic and gonadic growth of sea urchins ( Stronglyocentrotus droebachiensis ) fed kelp ( Laminaria longicruris ) or the invasive alga Codium fragile ssp. tomentosoides are related to energy acquisition

The rocky subtidal community off the Atlantic coast of Nova Scotia has historically undergone a cyclical transition between Laminaria-dominated kelp beds and sea urchin-dominated barrens. Since the introduction of the invasive alga Codium fragile ssp. tomentosoides, a third community state has emerged: Codium-dominated algal beds. We conducted a 42-week feeding experiment in the laboratory, which mimicked the quantity and quality of food available to urchins (Strongylocentrotus droebachiensis) in each of these community states. Feeding rate, growth, reproduction, and survival of urchins fed either Laminaria longicruris or C. fragile ad libidum, or L. longicruris 2 days per month, were measured. Although the ad libidum feeding rate on C. fragile was higher than that on kelp, energy intake was lower. Urchins in the ad libidum kelp treatment were larger and had larger gonads than those in the C. fragile treatment. Urchins fed kelp infrequently exhibited little somatic and gonadic growth over the course of the experiment. Regression analysis revealed that urchin performance on these diets was strongly related to energy intake. Diet treatment had no effect on survival or gonad maturation. Although urchins can consume substantial amounts of C. fragile, it appears that they cannot, or do not, feed quickly enough to compensate for its lower nutritional value. Our results suggest that, although urchins feeding on C. fragile are capable of surviving, growing, and reproducing, the replacement of kelp by C. fragile in some areas might negatively affect urchin populations as they continue to repopulate the shallow subtidal zone.     

Particularités microstructurales du squelette de Paracentrotus lividus et Arbacia lixula: Rapports avec l'écologie et l'éthologie de ces échinoïdes

Scanning-electron-microscope investigations on the test microstructure of two regular sea urchins, Paracentrotus lividus (Lamarck) and Arbacia lixula Linné have been especially devoted to the sutures between test plates, and the spines. Some features of both these components may be related to ecological and ethological differences between the two species. It is well documented that the structure of the sea urchin's test plates consists of a meshwork of calcareous trabecules embedded within the mesenchyme. The suture between the two lines of plates of each radial or intertradial zone exhibits a gap which plays some part in the process of the plates' growth. However, the gap may also constitute a complementary stress-breaker of mechanical forces (such as waves) which are exerted upon the test in the natural environment. In A. lixula this gap is so wide and the height: diameter ratio so low, that this species is particularly well fitted to bear physical stress and force on its apical region: since A. lixula almost exclusively inhabits vertical or subvertical rocky substrates at 2 to 15 m depth, where wave action is mainly exerted perpendicular to the substrate, these particular features of sutures and test shape may be a morphofunctional adaptation to this habitat. The sutures of P. lividus exhibit a narrower gap, making this species less able to bear strong apical pressure; consequently, P. lividus usually occurs on exposed horizontal or gently inclined substrates, but also inhabits sea-grass beds as deep as 15 to 20 m. The spines of P. lividus bear deep longitudinal grooves with lateral teeth, which seem especially fitted for collecting and transporting organic particles from the top of the spines to the apical region of the test where they are digested and assimilated by coelomocytes and epithelial cells. In the natural environment, the slightest water motion provides the spines with suspended particles; in extremely sheltered places or in aquaria however, the spines actively collect particles. Therefore, P. lividus populations can thrive in places where food resources other than suspended particles are scarce. The grooves on the spines of A. lixula are less marked, and thus unsuitable for collection of suspended particles. This species can therefore fulfil its energy requirements only by grazing and absorption of dissolved material. It appears that although belonging to the same biocoenosis, P. lividus and A. lixula do not occupy the same ecological niche. Their potential utilization in either urban or chemical pollution monitoring studies is discussed.     

Diet,behaviour and reproduction of the whelk Buccinum undatum in the northern Gulf of St. Lawrence,eastern Canada

A laboratory experiment was used to examine whether a decrease in the width of sutures is the basis for shrinking of the test in echinoids under conditions of low food availability and whether the body condition of sea urchins is likely to be retained when shrinking occurs. This experiment was performed with H. erythrogramma specimens collected in January 1987 from Pt. Lillias in Corio Bay, Australia. Suture widths in the test and gonad volume were significantly less (relative to body volume) in Heliocidaris erythrogramma (Val.) in the low food treatment than the high food treatment after 4 mo. These results imply that sea urchins which shrink may have a poorer body condition and be less able to reproduce than sea urchins that do not shrink. A comparison of the expected to the observed changes in suture width showed that a decrease in suture width is involved in the shrinking of the test in H. erythrogramma and may be of sufficient magnitude to fully explain the observed reduction in test size. This suggests that the amount a sea urchin may shrink under low food conditions is constrained by the degree to which the sutures can be reduced in width.     

Reproductive dynamics of the sea urchin Paracentrotus lividus on the Galicia coast (NW Spain): effects of habitat and population density

We studied the spatial variability in the size at first maturity and the reproductive cycle of Paracentrotus lividus in Galicia, contributing key information for the exploitation and management of this resource. The size at maturity varied between 20.4 (±1.2 SE) mm and 27.9 ± 1.2 mm and was smaller in areas of low population density where sea urchins do not form patches. Using a nonlinear model, we analysed the effect of depth, body size, sex and population density on the temporal pattern of the gonad index. The maximum and minimum indices were obtained at 4 m depth in the months before and after the spring spawning, respectively. The depth also affected the cycle phase, and the sea urchins at 4 m depth spawned 9.4 ± 3.0 days later than the sea urchins at 8 m depth and 20.5 ± 3.0 days later than those at 12 m depth. Moreover, the sea urchins living in patches showed a slight increase in gonad size as a consequence of the better-quality habitat. This shows that there is no intraspecific competition in this area despite the high population densities reached (18.5 kg m^?2).     

Annual reproductive cycles of the commercial sea urchinParacentrotus lividus from an exposed intertidal and a sheltered subtidal habitat on the west coast of Ireland

Reproduction of the commercial sea urchinParacentrotus lividus (Lamarck) from contrasting habitats on the west coast of Ireland was examined from May 1986 through August 1988. Urchins were collected intertidally from an exposed rocky shore and subtidally from a protected bay. Monthly measurements of the gonad index and histological examination of the gonads demonstrated thatP. lividus has an annual reproductive cycle. Although the two populations exhibited similar reproductive patterns over three breeding seasons, there were inter-annual and inter-population differences in the amplitude of gonadal growth and in the time when spawning started. The subtidal urchins had significantly larger gonads and exhibited a longer period of reproductive maturity compared with the intertidal urchins. This difference was also evident in the histology of the ovaries at the beginning of breeding, when most of the subtidal females contained mature ovaries, whereas most of the intertidal females contained premature ovaries. An inter-annual difference in the onset of spawning was observed, with the start of gamete release differing by as much as four weeks between years. It appears that inter-annual differences in sea temperature influence the temporal variation in spawning byP. lividus and that increasing temperature may serve as an exogenous cue for gamete release. The inter-annual variability in the onset of spawning suggests that photoperiod does not cue gamete release. Gonadal growth, on the other hand, occurs during the coldest months of the year and during the period of shortest days, suggesting that temperature and photoperiod may both influence gonadal growth during the winter. Oogenesis and spermatogenesis ofP. lividus were examined histologically. During the post-spawning recovery and growth stages, the gonads gained weight through growth of the nutritive phagocytes and the accumulation of periodic acid Schiff (PAS) + droplets by these cells. The PAS + material appears to play a nutritive role in gametogenesis. For the females, the frequencies of six ovarian maturity stages was assessed at approximately monthly intervals. Small oocytes were present throughout the year and clusters of early oocytes were most apparent during the spent and recovery stages. With the onset of vitellogenesis and subsequent accumulation of ova, the nutritive phagocytes and their PAS + droplets became depleted. The ovarian condition at the onset of breeding was variable, due to differences in the number of vitellogenic oocytes, differences in the number of ova in storage, and differences in the amount of PAS + material. In general, the nutritive phagocyte tissue is reduced by the onset of spawning and is exhausted by the termination of breeding. A similar series of events occurs during spermatogenesis. The relevance of this study for the use ofP. lividus as a brood-stock organism for mariculture is discussed.     

Patterns of abundance, population size structure and microhabitat usage of Paracentrotus lividus (Echinodermata: Echinoidea) in SW Portugal and NW Italy

In coastal habitats, wave exposure influences several aspects of the life history of marine organisms. Here, we assess how hydrodynamic conditions can generate variation in density, size structure and microhabitat usage of Paracentrotus lividus and whether these effects are consistent between regions that are markedly different for oceanic climate, such as the coasts of SW Portugal and NW Italy. The abundance of P. lividus was ~4 times higher in SW Portugal than in NW Italy, but within each region, there was no effect of wave exposure. In SW Portugal, higher urchin abundances were found at shallower depths, while no effect of depth on urchin abundance emerged in NW Italy. Most of the variation in urchin abundance occurred at small spatial scales (metres), and our results suggest that habitat complexity, that is, the presence of cracks and crevices, is an important determinant of patterns of distribution of this species. The population in NW Italy was characterized by a unimodal size distribution, with a higher proportion of medium-sized individuals. In contrast, in SW Portugal, smaller individuals represented a large proportion of the populations. Size structure varied between exposed and sheltered habitats in SW Portugal, suggesting that the proportion of individuals from different size cohorts may vary along wave-exposure gradients as a result of direct or indirect effects of hydrodynamic forces. In SW Portugal, most urchins occurred in burrows, while in NW Italy, urchins were mainly observed in crevices. These results suggest that creating/occupying burrows might be an adaptive behaviour that allows sea urchins to better withstand stressful hydrodynamic conditions and, therefore, are more common on exposed Atlantic coasts. Overall, our study suggests that the effects of hydrodynamic forces on sea urchin populations are context dependent and vary according to background oceanic climate.     

消油剂与120#燃料油对马粪海胆(Hemicentrotus pul-cherrimus)抗氧化酶活性的影响

在溢油事故应急处置中,消油剂的使用备受争议。为探究消油剂和溢油对海洋底栖模式生物海胆(Hemicentrotus pulcherrimus)复合毒性效应,通过WAFs(Water-accommodated fractions)和CEWAFs(Chemically enhanced water-accommodated fractions)的96 h暴露实验,测定海胆肠和性腺中过氧化氢酶(CAT)、超氧化物歧化酶(SOD)、谷胱甘肽硫转移酶(GST)、谷胱甘肽过氧化物酶(GPx)活性的变化。实验结果表明:随油水配比浓度的增加,4种酶活性呈现先升高后降低趋势,且酶活性峰值均极显著高于海水对照组水平(P0.01)。肠中4种酶活性最大诱导倍数均高于性腺。相同暴露浓度下,CEWAFs组4种酶活性诱导程度均高于WAFs组。消油剂对照组和海水对照组间4种酶活性则无显著性差异(P0.05)。     

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.     

Human impact on <Emphasis Type="Italic">Paracentrotus lividus</Emphasis>: the result of harvest restrictions and accessibility of locations

The sea urchin Paracentrotus lividus is common in the Mediterranean Sea in shallow subtidal rocky habitats, and it is intensely harvested for commercial and recreational purposes. This study is aimed at investigating whether the effects of harvest restrictions of P. lividus in rocky reef habitats interact with the accessibility of locations in structuring sea urchin population (total and commercial-sized individuals). These results are important for generating hypotheses about the influence of human harvesting on P. lividus and for addressing suitable measures of conservation. Paracentrotus lividus was sampled after the end of the sea urchin harvesting period (May–July 2007) within the Gulf of Alghero (North West Sardinia), where the Capo Caccia–Isola Piana Marine Protected Area (MPA) was established since 2002. Paracentrotus lividus was sampled at sixteen locations and attributed in groups of four to 4 combinations of harvest restrictions (Restricted Harvest, RH, vs. Unlimited Harvest, UH) and accessibility (Boat vs. Car), which correspond to a gradient of potential human activity on P. lividus in the ranked order of very low (RHBoatfar), low (RHBoatclose), moderate (RHCar) and high (UHCar). At each location, two depth ranges of 3–7 and 8–12 m were considered. At each of these depths, two areas of about 100-m² size were chosen. The density of P. lividus was assessed in ten quadrats of 1 × 1 m, and the size of 100 individuals (test diameter) was considered. Human activity has been found to significantly affect population structure of P. lividus influencing the proportion of individuals larger than 50 mm. Although harvest was restricted by MPA regulations, a significantly lower abundance of large individuals was found at sites accessible by car. This result highlights that there is an effect of harvest restrictions in relation to accessibility and emphasizes the need to carefully address the enforcement of the MPA toward easily accessible sites. Thus, surveillance and investment in enforcement of marine reserves seem crucial points that may provide the greatest return on maintaining the ecological benefits to the fishery activities.     

Leading role of the sea urchin <Emphasis Type="Italic">Arbacia lixula</Emphasis> in maintaining the barren state in southwestern Mediterranean

Sympatric sea urchin species are usually considered to belong to the same grazer guild. Nevertheless, their role in community dynamics may vary due to species-specific morphological traits, feeding preferences and foraging behavior. In the Mediterranean Sea, the two species Paracentrotus lividus and Arbacia lixula co-occur in barren areas. Whereas P. lividus is usually considered responsible for creating a barren ground, the roles of the two sea urchin species in its maintenance are currently unclear. The relative and combined effects of P. lividus and A. lixula on maintaining the benthic community in the barren state were tested experimentally, using orthogonal exclusion of the two species. Results show that exclusion of A. lixula, regardless of the presence of P. lividus, led to a significant decrease in the surface of bare rock and a significant change of the algal assemblages, thus demonstrating the major role of this species in maintaining the barren state.     

Juvenile–adult associations in sea urchins ( Strongylocentrotus franciscanus and S. droebachiensis ): protection from predation and hydrodynamics in S. franciscanus

Juvenile red sea urchins, Strongylocentrotus franciscanus, aggregate under adult conspecifics, whereas sympatric juvenile green sea urchins, Strongylocentrotus droebachiensis, are typically more solitary and dispersed. Neither the potential advantage of juvenile sheltering nor the differences in post-settlement behavior between the two species has been demonstrated experimentally, but may be related to protection from predators and/or hydrodynamics. In predation experiments, juvenile vulnerability differed in the two species as the seastar Pycnopodia helianthoides consistently chose juvenile S. franciscanus over S. droebachiensis (100% vs. 0%). When associated with adults, juvenile mortality decreased dramatically in S. franciscanus (90% alone vs. 5% with adults), but very little in S. droebachiensis (85% vs. 75%). Not surprisingly, juvenile behavioral responses in the two species reflect this difference in vulnerability. Juvenile S. franciscanus sheltered under adults when predation risk was high, but not when risk was low (44% vs. 13%), whereas sheltering in S. droebachiensis was infrequent and not related to predation risk (7% for high risk versus 5% for low risk). From a hydrodynamic perspective, the presence of an adult led to the creation of a hydrodynamic refuge for juveniles, where average water velocities were reduced by > 60% around the adult urchin. Again, striking differences in sheltering rate were apparent in S. franciscanus (52% vs. 13% for high flow and low flow, respectively), but not S. droebachiensis (5% for high flow versus 4% for low flow). Sheltering behavior was also species-specific as juveniles did not shelter at high rates under adults of the opposite species (≤ 16%). A field survey confirmed these finding in that juvenile S. franciscanus abundance was associated with both adults and water motion (R ² = 0.80, P = 0.008, best-subsets regression). These results suggest that sheltering confers juvenile S. franciscanus with a degree of protection from predators and water motion, and that species-specific differences in this post-settlement behavior may be related to the differences in the protection afforded by adults.     

Grazing by sea urchins at the margins of barren patches on Mediterranean rocky reefs

The role played by the urchins, Paracentrotus lividus and Arbacia lixula, in the formation and persistence of barren areas dominated by encrusting coralline macroalgae is yet to be fully elucidated. This study, carried out in the NW Mediterranean (43° 30′N, 10° 20′E) between February 2005 and April 2006, investigated how the loss or density decrease in one or both urchin species influences the recovery of erect macroalgal stands (dominated by filamentous forms) at the margins of barren areas. At a depth of 4–6 m, three barren patches were assigned to each of the following treatments: (1) control (natural densities of A. lixula and P. lividus); (2) 50 % of the natural density of A. lixula and natural density of P. lividus; (3) total removal of A. lixula and natural density of P. lividus; (4) 50 % of the natural density of P. lividus and natural density of A. lixula; (5) total removal of P. lividus and natural density of A. lixula; (6) 50 % of the natural densities of both A. lixula and P. lividus; (7) total removal of both A. lixula and P. lividus. The effects of the herbivore treatments were evaluated either in the presence or the absence of encrusting corallines. The partial or total removal of A. lixula, P. lividus or both favored the proliferation of filamentous macroalgae at the margins of barren patches. The presence of encrusting corallines reduced the development of these macroalgae. The results of this study suggest that a moderate decrease in the density of just one of the two species can decrease the ability of the herbivore assemblage to control the proliferation of filamentous macroalgae at the margins of barren patches. The extent of barren areas appears, therefore, to be regulated by the outcome of density-dependent interactions between the two species of sea urchins.     

Effect of sunlight intensity and albinism on the covering response of the Caribbean sea urchin<Emphasis Type="Italic"> Tripneustes ventricosus</Emphasis>

At Discovery Bay, Jamaica, Tripneustes ventricosus lives in beds of the turtle grass Thalassia testudinum. Especially during daylight hours, it covers its aboral surface with fragments of this plant and other objects. Normally pigmented, wild-type sea urchins covered themselves significantly less with Thalassia when sunlight was experimentally decreased to 66% or 32% ambient intensity. Consistent with this result, naturally occurring sea urchins exhibited significantly less covering at a deep (3.5 m) site than at a shallow (1 m) site, where light intensities at the bottom were 619 and 946 mol s^–1 m^–2, respectively. The graded covering response to light intensity suggests that covering is a defense against damaging solar radiation. Albino sea urchins covered themselves significantly more with Thalassia than wild-type sea urchins in both full and 66% sunlight. In addition, at the shallow site where they accounted for about 4% of the population, they showed significantly greater covering than wild-type urchins. The greater covering response of albino sea urchins suggests a greater susceptibility to solar radiation.Communicated by P. W. Sammarco, Chauvin     

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.     

The effects of predator abundance and habitat structural complexity on survival of juvenile sea urchins

We studied the effect of the abundance of predatory fishes and structural complexity of algal assemblages on the survival of juveniles of the sea urchin Paracentrotus lividus on Mediterranean infralittoral rocky bottoms. Post-settlement juveniles (2–10 mm) were placed on four distinct natural substrates with increasing structural complexity (coralline barren, algal turf, erect fleshy algal assemblages and small crevices) inside and outside the Medes Islands Marine Reserve. Predation on these sea urchins increased at greater abundance of predatory fishes, and decreased with greater structural complexity. The refuge provided by structural complexity, however, decreased with increasing size of sea urchin recruits. Predation on the smallest post-settlers was carried out almost exclusively by small fishes (<20 cm), mainly the labrid Coris julis, while the dominant predator of larger juveniles was the sparid Diplodus sargus. Our results demonstrate the cascading effects caused by the prohibition of fishing in marine reserves, and highlight the potential role of small predatory fishes in the control of sea urchin populations.Communicated by O. Kinne, Oldendorf/Luhe