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.     

Changes in sea urchin populations after the destruction of kelp beds

In St. Margaret's Bay, Nova Scotia, Canada, there are large areas in which sea urchins (Strongylocentrotus droebachiensis) have eliminated beds of kelp (Laminaria spp.). Sites were identified where destruction of kelp beds had taken place 1, 2, 3, 3.5 and 4 years ago. With increase of time since kelp disappearance, the sea urchins showed decreased growth rate, reduced gonad size, but an increase in numbers resulting from high recruitment rates in the first two years after kelp bed destruction. These sea urchin populations, by their browsing, effectively prevent the regeneration of kelp. There is, as yet, no evidence of the sea urchin populations being starved out to allow the kelp to return.     

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.     

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     

The association of N2-fixing bacteria with sea urchins

Dinitrogen fixation associated with bacteria in the gastrointestinal tract of sea urchins appears to be a widespread phenomenon: sea urchins from the tropics (Diadema antillarum, Echinometra lacunter, Tripneustes ventricosus), the temperature zone (Strongylocentrotus droebachiensis) and the arctic (S. droebachiensis) exhibited nitrogenase activity (C₂H₂ reduction). Pronounced seasonal variation was found in nitrogenase activity of temperate sea urchins feeding on kelp (Laminaria spp.) and eelgrass (Zostera marina). The mean monthly nitrogenase activity was inversely correlated with the nitrogen content of the sea urchin's food, which varied up to fivefold over the course of a year. The highest rate of nitrogenase activity recorded for a temperate sea urchin during the 14 month sampling period was 11.6g N fixed g wet wt^-1 d^-1, with a yearly mean activity of 1.36 g N fixed g wet wt^-1 d^-1. Studies with ¹⁵N confirmed the C₂H₂ reduction results and showed incorporation of microbially-fixed nitrogen into S. droebachiensis demonstrating that N₂ fixation can be a source of N for the sea urchin. Laboratory experiments indicated that part of the sea urchin's (S. droebachiensis) normal gastrointestinal microflora is responsible for the observed nitrogenase activity.     

Changing lobster abundance and the destruction of kelp beds by sea urchins

In a study area in Nova Scotia, Canada, abundance of the lobster Homarus americanus decreased by nearly 50% in 14 years. The lobster is a major predator of sea urchins, and during the past 6 years the sea urchin Strongylocentrotus droebachiensis has destroyed 70% of the beds of Laminaria spp. in the area. Implications for management are discussed.     

Enlarged lantern size in similar-sized,sympatric, sibling species of Strongylocentrotid sea urchins: from phenotypic accommodation to functional adaptation for durophagy

Strongylocentrotus droebachiensis is a well known omnivorous sea urchin with an unrivalled capacity to destroy North Atlantic kelp forests. S. pallidus is a lesser known, morphologically similar, and closely related species with no record of destructive grazing, despite its larger lantern size. I quantify the lantern size of both species using bivariate allometric analysis, and test the hypothesis that enlarged lantern size facilitates durophagy, the consumption of hard prey, by measuring the feeding capacity of urchins with different lantern sizes when offered a hard-shelled prey, the blue mussel, Mytilus edulis. The results suggest that S. droebachiensis has a limited capacity for durophagous feeding irrespective of lantern size, whereas in S. pallidus the ability to exploit hard shelled prey is positively related to lantern size. This is apparently the first evidence of a relationship between trophic morphology and diet in regular sea urchins. The hypothesis of systematic latitudinal variation in the lantern size of S. pallidus is reappraised and rejected. S. droebachiensis had larger gonads than S. pallidus in field samples, confirming that its small lantern is not impeding nutrient acquisition in shallow habitats.     

Relation of food preference to fitness for the green sea urchin,Strongylocentrotus droebachiensis

We maintained 30 to 35 mm sea urchins (Strongylocentrotus droebachiensis) on 17 different macroalgal diets in the field over a 130-d period and correlated the resulting changes in somatic and gonadal tissues with measures of food preferences. The varions algae fell into there distinct categories according to their ability to support growth. The most striking effect of diet was on gonadal mass, there being a 14-fold increase for the best algal diet, compared to initial gonadal mass, and nearly a threefold decrease for the worst diet. The relation of food preference to somatic growth was nonlinear. A sharp increase in the growth of test diameter, total mass and test mass occurred between the nonpreferred and intermediately preferred algae, but growth rates were similar for intermediate and preferred algae. Gonadal growth tended to increase exponentially with an important measure of food preference, feeding rate (g ingested d^–1). These observations suggested that urchins only channel food resources into somatic growth up to a given point, additional reserves being used for producing gametes. The positive relationship of food preference to growth, and especially the strong correlation with gonadal production, indicates that the highly selective feeding of the urchin contributes to ils fitness.     

Heat waves,baby booms,and the destruction of kelp beds by sea urchins

Large populations of sea urchins, Strongylocentrotus droebachiensis (Müller), destroyed kelp beds along the Atlantic coast of Nova Scotia in the 1960's and 1970's. The origin of these large sea urchin populations is not understood. We have investigated the potential influence of variable growth and development of the planktonic larvae of sea urchins (in response to temperature and food abundance) on recruitment of benthic juveniles. The adult sea urchins were collected at Sandy Cove, Digby County, Nova Scotia, Canada, in December 1986. Temperature strongly affected larval size and the growth of the echinus rudiment within the range 3° to 9°C, and larvae grew most rapidly at 14°C. Food abundance had a smaller effect on larval growth, and these effects were apparent only at high temperature. Larvae fed the same concentration of two different algal food species grew and developed similarly. Correspondence between spring temperature variation and qualitative variation in sea urchin recruitment, as well as strong temperature effects on larval growth in culture, and the occurrence of a large, positive temperature anomaly in June 1960, all suggest that temperature effects on larval growth and development may have led to intense sea urchin recruitment in 1960 and the appearance of large adult populations 4 to 6 yr later. This result invites further research.     

Comparative feeding preference of Strongylocentrotus droebachiensis (Echinoidea) for the invasive seaweed Codium fragile ssp. tomentosoides (Chlorophyceae) and four other seaweeds

Laboratory experiments conducted during 1987 on Appledore Island, Maine, USA, tested whether feeding preference or the absence of an attractant was the cause for the occurrence of beds of Codium fragile ssp. tomentosoides (herein referred to as Codium fragile) within rocky barrens grazed clear of kelp by the sea urchin Strongylocentrotus droebachiensis. Consumption of C. fragile in single-diet experiments (1 seaweed/sea urchin) was highly variable and was not significantly different from that for several other seaweeds (Agarum cribrosum, Ascophyllum nodosum, Chondrus crispus, and Laminana saccharina) important in the field diet of the green sea urchin. In multiple-diet experiments (5 seaweeds/sea urchin) significantly less Codium fragile was eaten than Chondrus crispus, but significantly more Codium fragile was eaten than A. cribrosum. Chemosensory experiments suggest that C. fragile does not attract the sea urchin. Sea urchins are unable to detect C. fragile but will eat it when they come in contact with it.     

Anaerobic and aerobic energy metabolism in ovaries of the sea urchin Strongylocentrotus droebachiensis

The gonads of sea urchins undergo large changes in mass during their gametogenic cycle. In addition, they have relatively low aerobic capacities and are poorly perfused by the circulatory system and thus are continually hypoxic or anoxic. The present study of Strongylocentrotus droebachiensis investigates seasonal changes in the relationships among mass of the ovaries, pH and P_O2 of the perivisceral coelomic fluid which bathes the ovaries, and partitioning of ovary energy metabolism into its anaerobic and aerobic components. S. droebachiensis were collected at Blue Hill Falls, Maine, USA, from August 1982 to March 1984. We found that from 76 to 92% of the heat dissipated by isolated ovaries of the sea urchin S. droebachiensis derives from anaerobic energy metabolism at partial pressures of oxygen prevailing in vivo. Ovaries from S. droebachiensis have the capacity to produce large amounts of lactate under imposed anoxia, but lactate accounts for only 37% of the total anoxic heat dissipation, which suggests that other end products of anaerobiosis are present. Seasonal changes in pH and P_O2 of the perivisceral coelomic fluid can be explained by a complex relationship among changes in temperature, reproductive condition, and anaerobic and aerobic metabolism in the ovaries, gut and body wall. Seasonal changes in the buffering capacity of the perivisceral coelomic fluid must be determined before the effects of respiratory and metabolic acid production on the acid-base status of the coelomic fluid can be fully understood.     

Incidence,occurrence and distribution of the nematode Echinomermella matsi in its echinoid host,Strongylocentrotus droebachiensis,in northern Norway

The incidence, occurrence and distribution of the nematode Echinomermella matsi (Jones and Hagen) in its host, the echinoid Strongylocentrotus droebachiensis Müller, were examined in Norwegian waters from Nordmre (63° N) to the Russian border (70° N) in a range of habitats (e.g. kelp cover and community type) from 1982 to 1992. The results are used to examine the likely influence of the parasite on fitness (individual growth and gonad production) and its potential in regulating the population size of its echinoid host. The nematode infections are restricted to areas between Vikna (Vega) and middle Troms, with the highest prevalence (40 to 88% infected) in the Bodø area, the focus of the distribution. The prevalence rates are similar on kelp beds and barren grounds. No difference in growth rates between infected and noninfected sea urchins was found. Nematode-infected sea urchins in kelp bed populations had gonads weighing about one-sixth of those of equivalent uninfected individuals. This indicates that nematodes are really parasites causing reduced reproductive capacity. Areas with high prevalence (>40% infected) of nematodes had low sea urchin densities. High nematode intensities were found in the areas with high prevalence. The mean abundance in high prevalence areas was highest (eight nematodes) in the 4-to 7-yr-old age group and decreased to zero among the oldest age groups. The ratio of variance/mean abundance was also highest among 5-to 8-yr-old sea urchins. At highly and slightly infected locations, the relationships between the log of the variance and the log of the mean abundance of parasites were 1.35 and 1.69, respectively. Measurements of the frequency distribution of nematodes in sea urchins and the corresponding estimates of the negative binomial distribution showed significantly higher counts in estimated than observed subpopulations exceeding 20 nematodes per sea urchin. These results may explain nematode-induced sea urchin mortality. The low density of the host in the Bodø area, which had high prevalence, may be a result of increased host mortality caused by nematodes, by lower recruitment because of reduced gonad production, or by other factors. Low prevalence among older sea urchins may be caused by nematodeinduced host mortality or by cycles in infection. However, studies on the life cycle of the nematode and long term investigations of host density and nematode intensity/prevalence will be needed before any true conclusions can can be drawn.     

Seasonal variation in movement,aggregation and destructive grazing of the green sea urchin (<Emphasis Type="Italic">Strongylocentrotus droebachiensis</Emphasis>) in relation to wave action and sea temperature

Hydrodynamic forces are an important determinant of subtidal community structure, particularly when they limit the distribution and foraging ability of mobile consumers. We examined the effect of wave action on the rate of movement and destructive grazing of a kelp bed by the green sea urchin (Strongylocentrotus droebachiensis) under field conditions. We measured density and rate of advance at fixed intervals along ∼100 m of a grazing front over 1 year, and quantified individual movement rates in the barrens 5–10 m behind the urchin front using a time-lapse videography. Seasonal variation in the mean rate of advance of the front (range: 0–4 m month⁻¹) was explained by changes in urchin density at the front (120–360 individuals m⁻²), which in turn varied inversely with significant wave height (0.5–2 m). Water temperature (0.8–17.6°C) had no effect on the rate of advance or on urchin density (aggregation) at the front, except when temperature exceeded 17°C. Movement of individual urchins also was affected by wave action: we observed a significant decrease in speed and displacement of urchins with increasing significant wave height. Wave action had no effect on the proportion of urchins moving or the degree of linearity of their movements. We propose that the decrease in urchin density at the front associated with increased wave action, results from de-aggregation, which reduces the risk of dislodgement, combined with a reduction in urchin movement in barrens, which supplies new urchins to the front.     

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.     

Effects of sea urchin disease on coastal marine ecosystems

Outbreaks of disease in herbivorous sea urchins have led to ecosystem phase shifts from urchin barrens to kelp beds (forests) on temperate rocky reefs, and from coral to macroalgal-dominated reefs in the tropics. We analyzed temporal patterns in epizootics that cause mass mortality of sea urchins, and consequent phase shifts, based on published records over a 42-year period (1970–2012). We found no evidence for a general increase in disease outbreaks among seven species of ecologically important and intensively studied sea urchins. Periodic waves of recurrent amoebic disease of Strongylocentrotus droebachiensis in Nova Scotia coincide with periods when the system was in a barrens state and appear to have increased in frequency. In contrast, following a major epizootic that decimated Diadema antillarum throughout the Caribbean in 1983, subsequent outbreaks of disease were highly localized and none have been reported since 1991. Epizootics of Strongylocentrotus in the NW Atlantic and NE Pacific, and Paracentrotus and Diadema in the eastern Atlantic, have been linked to climate change and overfishing of sea urchin predators. The spatial extent of recurrent disease outbreaks in these species, and the frequency of phase shifts associated with these epizootics, has decreased over time due to the expansion of the macroalgal state and its stabilization through positive feedback mechanisms. Longitudinal studies to monitor disease outbreaks in sea urchin populations and improved techniques to identify causative agents are needed to assess changes in the frequency and extent of epizootics, which can profoundly affect the structure and functioning of coastal marine ecosystems.     

Effects of food type and ration on reproductive maturation and growth of the sea urchin Strongylocentrotus droebachiensis

We investigated the effects of food quality and quantity on reproductive maturation and growth of juveniles of Strongylocentrotus droebachiensis (Müller) in a 22 month laboratory experiment in which we fed sea urchins four diets: (1) kelp (Laminaria spp.) for 6 d wk⁻¹ and mussel (Mytilus spp.) flesh for 1 d wk⁻¹ (KM); (2) kelp for 7 d wk⁻¹ (high ration, KH); (3) kelp for 1 d wk⁻¹ (low ration, KL) and (4) no food other than encrusting coralline algae (NF). At their first and second opportunity for reproduction, all sea urchins in the KM and KH treatments, and most in the KL treatment were reproductively mature, whereas all sea urchins in the NF treatment remained immature. Gonad index differed significantly among all fed treatments at first and second reproduction, and was highest in the KM and lowest in the KL treatment. Gonad index was similar in both sexes at first reproduction, but it was higher in females than in males at second reproduction. Diet had little or no effect on the relative abundance of spermatocytes, spermatozoa, or nutritive phagocytes in testes at first and second reproduction. In ovaries, nutritive phagocytes were significantly more abundant in females in the KM and KH treatments than in the KL treatment at first reproduction, and significantly more abundant in unfed (NF) than fed (KM, KH, KL) females at second reproduction. Mean oocyte size was similar in all fed females at first reproduction, but significantly larger in fed than unfed females at second reproduction. Mean ovum size was similar in all fed females in both reproductive periods. Increase in test diameter was greatest in the KM treatment and smallest in the KL treatment; sea urchins in the NF treatment decreased slightly in size. Survival was 95 to 100% in all fed sea urchins but significantly lower in unfed ones. The feeding rate on kelp was significantly greater in the KL than the KM and KH treatments. In the KM treatment, the feeding rate on kelp increased significantly over a 6 d period after mussel flesh was provided. Our results demonstrate that a diet of high food quality and quantity accelerates reproductive maturation and growth rate, and enhances gonad production and survival in juvenile and young adult S. droebachiensis. These findings contribute to our understanding of the reproductive ecology of S. droebachiensis in habitats with differing food supplies (e.g., kelp beds and barrens). Our results also can be used to improve aquacultural practices for sea urchins. Received: 3 June 1998 / Accepted: 2 February 1999     

Ecological energetics of the seaweed zone in a marine bay on the Atlantic coast of Canada. III. Energy transformations by sea urchins

An energy budget was constructed for a population of the sea urchin Strongylocentrotus droebachiensis in the nearshore area of St. Margaret's Bay, Nova Scotia, Canada. Of the 6 age classes identified, ages 1+and 2+accounted for about 1/2 the population energy flow. Population production efficiencies were: production/assimilation=0.28, production/consumption=0.04 to 0.13, and production/biomass=0.80. Although S. droebachiensis was the dominant herbivore in the seaweed bed, it utilized only 1 to 7% of seaweed production. As with other populations of sea urchins, however, it had a proportionately greater influence on seaweed biomass, and also presumably production, by clearing seaweed from large areas of substrate and maintaining it clear. Loss of dissolved organic matter, the only term in the energy budget not measured, was estimated by substracting the other terms in the energy budget from consumption. In laboratory individuals, this ranged from 40 to 80% of absorption (consumption-faeces). A critical review of energy budgets for 6 other species of marine benthic grazers also revealed large amounts of energy unaccounted for that might be attributed to loss of dissolved organic matter.Contribution to the International Biological Program CCIBP 192Bedford Institute Contribution.     

Effects of grazing by two species of sea urchins (Strongylocentrotus franciscanus and Lytechinus anamesus) on recruitment and survival of two species of kelp (Macrocystis pyrifera and Pterygophora californica)

We studied the effects of grazing by two species of sea urchins on two species of kelp (Macrocystis pyrifera and Pterygophora californica) in the San Onofre kelp bed in southern California from 1978 through 1981. Both red sea urchins, Strongylocentrotus franciscanus, and white sea urchins, Lytechinus anamesus, were abundant and lived in aggregations. The purple sea urchin (S. purpuratus) was rare at the study site and was not studied. The aggregations of red urchins were either relatively small and stationary (for over 3 yr) or relatively large and motile (advancing at about 2 m mo^–1). Both stationary and moving aggregations were observed at the same time, and within 100 m of one another. Stationary aggregations of red urchins probably subsisted mainly on drift kelp and had no effect on kelp recruitment or on adult kelp abundance. In contrast, red sea urchins in large, motile aggregations or fronts ate almost all the macroalgae in their path. The condition of their gonalds indicated that red urchins in fronts were starved relative to red urchins in the small, stationary aggregations. Large, motile aggregations developed after 2 yr of declining kelp abundance (probably due largely to storms). We propose that a scarcity of drift algae for food results in a change in the behavior pattern of the red urchins and thus leads to the formation of large, motile aggregations. The aggregations of white urchins, which occurred along the offshore margin of the kelp bed, were large, but relatively stationary. The white urchins rarely ate adult kelps, but grazed extensively on early developmental stages of kelps and evidently prevented seaward expansion of the bed. The spatial distribution of both types of red urchin aggregations appeared to be unrelated to predation pressure from fishes or lobsters.Please address all requests for reprints to the senior author at his present address.     

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.     

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.