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.     

Removal of glycine from solution by the sea urchin Strongylocentrotus purpuratus

Radioautography and liquid-scintillation counting of juvenile sea urchins — Strongylocentrotus purpuratus (Stimpson) — exposed to ³H- and ¹⁴C-labeled glycine showed that the ³H is rapidly translocated throughout the animal and lost, while the ¹⁴C is slowly translocated into the interior of the urchin and incorporated into insoluble material.     

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.     

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

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

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.     

Long-term and trans-life-cycle effects of exposure to ocean acidification in the green sea urchin Strongylocentrotus droebachiensis

Anthropogenic CO₂ emissions are acidifying the world’s oceans. A growing body of evidence demonstrates that ocean acidification can impact survival, growth, development and physiology of marine invertebrates. Here, we tested the impact of long-term (up to 16 months) and trans-life-cycle (adult, embryo/larvae and juvenile) exposure to elevated pCO₂ (1,200 μatm, compared to control 400 μatm) on the green sea urchin Strongylocentrotus droebachiensis. Female fecundity was decreased 4.5-fold when acclimated to elevated pCO₂ for 4 months during reproductive conditioning, while no difference was observed in females acclimated for 16 months. Moreover, adult pre-exposure for 4 months to elevated pCO₂ had a direct negative impact on subsequent larval settlement success. Five to nine times fewer offspring reached the juvenile stage in cultures using gametes collected from adults previously acclimated to high pCO₂ for 4 months. However, no difference in larval survival was observed when adults were pre-exposed for 16 months to elevated pCO₂. pCO₂ had no direct negative impact on juvenile survival except when both larvae and juveniles were raised in elevated pCO₂. These negative effects on settlement success and juvenile survival can be attributed to carry-over effects from adults to larvae and from larvae to juveniles. Our results support the contention that adult sea urchins can acclimate to moderately elevated pCO₂ in a matter of a few months and that carry-over effects can exacerbate the negative impact of ocean acidification on larvae and juveniles.     

Dietary accumulation of UV-absorbing mycosporine-like amino acids (MAAs) by the green sea urchin (Strongylocentrotus droebachiensis)

The bioaccumulation of ultraviolet radiation-absorbing mycosporine-like amino acids (MAAs) by Strongylocentrotus droebachiensis (Müller) was determined in specimens maintained on MAA-rich and MAA-deficient diets. Individuals were fed either the red alga Mastocarpus stellatus (Stackhouse), which has a high concentration of the MAA shinorine (6.98 nmolmg^–1 dry wt), and trace amounts of the MAAs porphyra-334, asterina-330 and palythine, or the brown alga Laminaria saccharina (Lamouroux), which contains no detectable MAAs. Reproductively spent urchins were fed ad libitum during a 9 mo period in 1992 to 1993 until they were once again gravid. To monitor accumulation, samples of urchin tissues and ingested food were taken throughout the 9 mo period from males and females maintained on each diet, and the concentrations of MAAs determined using high performance liquid chromatography. Urchins maintained on a diet of M. stellatus showed an ovarian shinorine concentration (8.33 nmol mg^–1 dry wt) 25 times higher than those fed L. saccharina. There was no change in MAA content of testes or somatic tissues in either feeding group. More than 99% of the MAAs present in an experimental meal of M. stellatus were removed during passage through the gut. Previous studies have inferred dietary accumulation of MAAs by comparing MAAs present in animal tissues with the consumer's potential diet in the field. This is the first study to have monitored MAA accumulation in animal tissues in conjunction with a controlled diet having a known MAA composition. Concentrating MAAs in the ovaries may provide eggs released into the water column with protection from damaging solar ultraviolet radiation.     

Allozyme and mitochondrial DNA evidence of population subdivision in the purple sea urchin Strongylocentrotus purpuratus

Despite high potential for dispersal, the purple sea urchin Strongylocentrotus purpuratus was found to have significant genetic subdivision among locations. Ten geographic locations along the coast of California and Baja California were sampled between 1994 and 1995. Samples from some locations included both adult and recruit urchins. Allozyme analyses revealed a genetic mosaic, where differentiation over short geographic distances could exceed differentiation over much larger distances. Significant allozyme differentiation was found among subpopulations of adults (standardized variance, F _ST =0.033), among subpopulations of recruits (F _ST =0.037), and between adults and recruits from the same location. DNA-sequence data for the mitochondrial cytochrome oxidase I gene also showed significant heterogeneity among locations, with a mild break in haplotype frequencies observed 300 km south of Point Conception. California. Repeated sampling over time is necessary to determine whether these patterns of differentiation are stable and to begin to understand what forces produce them.     

Mycosporine-like amino acids prevent UVB-induced abnormalities during early development of the green sea urchin Strongylocentrotus droebachiensis

Experiments were performed to determine how ultraviolet radiation (UVR) in the environmentally relevant range affects development of the sea urchin Strongylocentrotus droebachiensis (Müller) and whether mycosporine-like amino acids (MAAs), present in the early life stages, reduce UV-induced damage. Eggs, embryos, and larvae contained five MAAs having absorption maxima ranging from 320 to 334 nm. Eggs contained principally shinorine and porphyra-334, which absorb maximally at 334 nm and half-maximally at 312 and 348 nm, spanning much of the environmental range of biologically effective UVR. Concentrations of MAAs remained constant in unirradiated embryos through the gastrula stage, but decreased significantly in two-armed pluteus larvae. Daily exposure to combined photosynthetically active radiation (PAR, 400–700 nm) and UVR did not affect the concentration of MAAs in these embryos up to the two-armed pluteus stage. Prism larvae of sea urchins and the sand dollar Echinarachnius parma (Lamarck) did not accumulate shinorine from the surrounding seawater. Daily exposure of embryos to UVA (320–400 nm) and UVB (295–320 nm) radiation in the presence of PAR induced delays and abnormalities during development, and removing UVB eliminated this effect. Abnormalities in embryos included thickening of the blastoderm wall, filling of the blastocoel by abnormal cells, exogastrulation, and formation of abnormal spicules. The percentage of embryos that developed normally was lower in batches of embryos exposed to PAR + UVA + UVB, except in embryos from urchins maintained on MAA-rich diets. In all cases, the percentage of PAR + UVA + UVB-exposed embryos that developed normally was positively related to the concentration of MAAs in eggs from which the embryos developed. Thus, the MAAs found in S. droebachiensis embryos protect them against UVB-induced abnormalities during their development to at least the four-armed pluteus larval stage. Received: 8 May 2000 / Accepted: 29 September 2000     

Fungal disease resistance of Caribbean sea fan corals (Gorgonia spp.)

An on-going, Caribbean-wide epizootic affecting sea fan corals (Gorgonia spp.) is caused by the fungus Aspergillus sydowii (Thom et Church). We examined the role of crude extracts in resistance of two species of sea fans, Gorgonia ventalina (L.) and G. flabellum (L.), against A. sydowii and a bacterial pathogen of fish, Listonella anguillarum (MacDonell et Colwell). Sea fans were collected in January 1997 from San Salvador, Bahamas, and in June 1997 and January 1998 from Alligator Reef, Florida Keys, USA. Crude extracts from both species were tested to determine concentrations inhibiting germination of A. sydowii spores. Crude extracts from both species inhibited spore germination at concentrations as low as 1.5 mg ml⁻¹; most samples were active at 5 to 10 mg ml⁻¹. These concentrations are within the range estimated in living tissue and were higher in healthy colonies suggesting their role in mediating disease susceptibility. We also detected within-colony gradients in antifungal activity, which varied with the disease state of the colony. In healthy sea fans, resistance was highest at colony edges and lowest in medial and central regions of the colony. Among sea fans with lesions in the colony center, resistance in tissue from proximal and medial regions was as high as tissue from the colony edge (i.e. distal region). The increase in antifungal activity suggests an inducible response by the coral host to the fungal pathogen. This response is most evident among sea fans with lesions in the colony center and not among colonies with lesions at the edge. Antibacterial activity of crude extracts against L. anguillarum was highest at the colony edge but did not vary with disease state or tissue location. Received: 10 March 1999 / Accepted: 10 November 1999     

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     

Effects of food ration and feeding regime on growth and reproduction of the sea urchin Strongylocentrotus droebachiensis

To determine the effects food ration and feeding regime on growth and reproduction of Strongylocentrotus droebachiensis (Müller), sea urchins in laboratory aquaria were fed kelp (Laminaria longicruris) supplied at either a high (H, ad libidum daily) or a low (L, ad libidum 1 d wk⁻¹) ration in two successive 12-wk intervals during the reproductive period. After 24 wk, urchins fed the high ration continuously (HH) or for the last 12 wk only (LH) had a significantly greater mean gonad index [(gonad weight/total body weight) × 100] and body weight than urchins fed the low ration continuously (LL) or for the last 12 wk only (HL). Urchins in the HL treatment had a significantly greater gonad index than those in the LL treatment; there was no significant difference in gonad index between the LH and HH treatments. Females had a greater gonad index than males in the low ration (LL and HL) treatments at the end of the experiment; there was no significant difference between sexes in the high ration (LH, HH) treatments. Gametogenesis proceeded to maturation in all treatments and some individuals spawned at the end of the experiment. Females in the high ration (HH and LH) treatments had a greater proportion of nutritive phagocytes in their ovaries than females in the low ration treatments, but there was no effect of feeding treatment on oocyte or ovum size. Feeding treatment had no effect on the relative abundance of nutritive phagocytes in the testes, although the proportion of spermatocytes was higher (and that of spermatozoa lower) in the high ration than in the low ration treatments. Urchins in the high ration treatments had a lower mean jaw height index [(jaw height/test diameter) × 100] and greater mean test diameter than those in the low ration treatments at the end of the experiment, although these differences were not statistically significant. Feeding rate on kelp at the end of the experiment was significantly greater for urchins in the low ration than in the high ration treatments. Our experimental results show that even relatively low rations of kelp support somatic and gonadal growth in S. droebachiensis. Increasing the supply of kelp, particularly during the period of active gametogenesis, results in maximal rates of growth and reproduction. These results suggest that populations of S. droebachiensis in barrens may derive a substantial proportion of their nutrition from drift kelp, which may contribute to their persistence in these habitats. They also explain the large body size, high reproductive effort and fecundity of urchins grazing on kelp beds. These findings have important implications for understanding the dynamics of natural populations of S. droebachiensis and for development of effective aquacultural practices. Received: 17 February 1997 / Accepted: 5 March 1997     

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.     

Isolation and characterisation of zinc-binding proteins distinct from metallothionein from the eggs of the sea urchin Strongylocentrotus intermedius

Two low-molecular-weight zinc-binding proteins were purified from eggs of the sea urchin Strongylocentrotus intermedius (Echinodermata: Echinoidea) by gel-permeation and anion-exchange chromatography followed by high performance liquid chromatography. The characteristics of these proteins are distinct from those of metallothionein. Amino acid compositions show a low or intermediate content of cysteine and high amounts of acidic amino acids, glycine and histidine; they also contain methionine and aromatic residues. On the basis of these characteristics, the S. intermedius zinc-binding proteins appear similar to other metalloproteins isolated from both vertebrates and invertebrates. No typical metallothionein was detected in the extracts from S. intermedius eggs.     

Response of embryos of the sea urchin Strongylocentrotus purpuratus to aqueous petroleum waste includes the expression of a high molecular weight glycoprotein

The present study investigated the sublethal effects of the largest aqueous waste associated with offshore oil development, produced water (PW), on sea urchin embryo development. PW inhibited normal gastrulation in purple sea urchin (Strongylocentrotus purpuratus) embryos exposed at the hatching stage. The exposed embryos did not form complete archenterons, and secondary mesenchyme cells exhibited an unusual behavior with respect to the wall of the blastocoelic cavity. In addition, an abundance of extracellular matrix was observed in the blastocoelic cavities in the embryos exposed to higher concentrations of PW. An inhibition of the development of embryos through the pluteus stage was observed as a result of PW exposure. This was manifested by inhibition of normal spicule formation with concomitant abnormal pluteus morphology. To determine if a biochemical stress response occurs as a result of PW exposure, embryos at the early gastrula stage were exposed to PW for 2 h, followed by an additional 1 h in 3H-leucine and PW. These embryos expressed a 253 kDa protein as observed by one-dimensional SDS (sodium dodecyl sulfate) polyacrylamide gel electrophoresis and autoradiography. This protein could occasionally be observed in gels stained for protein, and was glycosylated as demonstrated by staining with periodic acid Schiff (PAS). The 70 to 73 kDa heat-shock proteins previously described in sea urchin embryos (and other organisms) as a result of stress were not synthesized as a result of PW exposure. Heat shock did not induce synthesis of the 253 kDa glycoprotein. Combined PW and heat-shock exposure elicited both the heat-shock response (73 kDa protein) as well as expression of the 253 kDa glycoprotein. These data demonstrate the use of expression of endogenous macromolecular markers to distinguish exposures of embryos to different perturbations. Arsenic (as sodium arsenite), a metal constituent of PW, also induced the 253 kDa glycoprotein in early gastrula-stage embryos. We suggest that this glycoprotein may be a marker for PW (and a metal constituent) exposure in sea urchin embryos, and may be related to morphological abnormalities.     

Sex and microhabitat influence the uptake and allocation of mycosporine-like amino acids to tissues in the purple sea urchin, Strongylocentrotus purpuratus

UVR-absorbing mycosporine-like amino acids (MAAs) were detected in tissues of Strongylocentrotus purpuratus and in ten species of Rhodophyte macroalgae (eight previously untested) collected from intertidal microhabitats in November and January 2006?C2007 in Central California (35°09??N, 120°45??W). In sea urchins, MAA concentrations were higher in ovaries than testes, while epidermal concentrations were similar between sexes. Ovaries and epidermal tissues had similar MAA signatures and broadband UVA/UVB absorbance, while testes had a narrower absorption ranges shifted toward higher energy wavelengths. Sea urchins occupying pits in the substrate exhibited lower MAA concentrations than those outside pits, suggesting adult microhabitat may impact UV protection. Light levels did not influence gonadal MAA concentrations, but correlated with elevated epidermal MAA concentrations for males in the sunniest microhabitat. This study suggests sex and habitat strongly influence MAA concentrations among individual S. purpuratus and that allocation of MAA sunscreens to tissues in response to UVR is sex-dependent.     

Effects of temperature and food ration on gonad growth and oogenesis of the green sea urchin, Strongylocentrotus droebachiensis

To assess the effects of both temperature and food ration on gonad growth and oogenesis of the green sea urchin, Strongylocentrotus droebachiensis (O.F. Müller), individuals collected December 1996 (winter experiment) and June 1997 (summer experiment) were maintained for 3 months in one of four experimental treatments: (1) 3 °C and fed ad libitum (high ration), (2) 3 °C and fed one-seventh of the maximum ration (low ration), and (3) 12 °C and fed the high ration; (4) 12 °C and fed the low ration. All individuals were fed an artificial diet and exposed to only 1 h of light every day. At the end of both experiments, mean gonad indices of sea urchins fed the high ration had increased significantly (11–24% and 6–19% in the winter and summer experiments, respectively), while the gonad indices of individuals fed the low ration did not change. At the high ration (both experiments), the increase in gonad index of sea urchins occurred primarily as the result of a significant increase in the mass of nutritive phagocytes, as revealed by histological analyses. Primary oocytes were significantly larger in individuals held at 3 °C than at 12 °C throughout the winter experiment, regardless of food ration; during the summer experiment, primary oocytes were significantly larger in individuals receiving the high ration, regardless of the temperature at which they were held. These results suggest that: (1) food availability is the most important factor regulating energy storage and the relative size of gonads throughout the year, (2) temperature affects the rate of growth and maturation of primary oocytes during the later stages of oogenesis, and (3) once gametogenesis has been initiated, mature ova will be produced, even under conditions of low food availability. Conditions of high food availability in summer and low temperature in winter would thus favor reproductive output in sea urchin populations. Received: 1 March 2000 / Accepted: 4 October 2000     

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.