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     

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.     

Allocation of45calcium to body components of starved and fed purple sea urchins (Strongylocentrotus purpuratus)

Several lines of evidence in the literature indicate that environmental stress such as starvation may initiate reallocation of sea urchin endoskeletal tissue. For example, Aristotle's lantern enlarges under conditions of starvation, and sea urchins tagged with tetracycline and then fed develop a distinct growth line, while starved individuals develop a diffuse pattern. We designed anin vivo system to examine stress-related changes in calcification in the purple sea urchinStrongylocentrotus purpuratus. SmallS. purpuratus (ca. 2 cm test diam) were collected from the Mission Bay jetty or Imperial Beach (San Diego, California, USA) in 1987.⁴⁵Ca was incorporated from seawater into all body fractions including the organic tissue/coelomic fluid. In an initial experiment, sea urchins were fed or starved for 4 wk and then post-incubated in isotope. Overall, starved individuals deposited new calcite more slowly than did fed individuals; however, allocation was very different and calcification of teeth of starved sea urchins was nearly as great as in fed individuals. In a second experiment,S. purpuratus were first pre-labeled with isotope and then treated by feeding or starving. More of the labeled calcium was mobilized from the soft tissues and coelomic fluid into calcite in fed than in starved individuals. Growth of the teeth in starved sea urchins was significantly greater than in those fed. We conclude that starvation changes the metabolism of calcium in order to preferentially build teeth. However, we also found no evidence that calcium was resorbed from old skeletal calcite in order to build new skeleton.     

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     

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.     

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

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

Effects of salinity on respiration and nitrogen excretion in two species of echinoderms

The 30-d survival limit of Eupentacta quinquesemita and Strongylocentrotus droebachiensis is 12–13 S. The activity coefficient (1 000/righting time in seconds) of stepwise acclimated sea urchins declined from 16.3 at 30 S to 3.5 at 15 S. Oxygen consumption rates (QO₂) of both species held at 30 S and 13°C were highest in June and lowest in December. During the summer, when environmental salinity is most variable in southeastern Alaska, the QO₂ of both species held at 30, 20 and 15 S varied directly with salinity. Perivisceral fluid PO₂ varied directly with acclimation salinity in sea urchins, but not in sea cucumbers. Perivisceral fluid oxygen content of acclimated sea urchins was significantly lower at 15 and 20 S than at 30 S due to reduced PO₂ and extracellular fluid volume at the lower salinities. The QO₂ of both species varied directly with ambient salinity during a 30-10-30. semidiurnal pattern of fluctuating salinity. No change occurred in the average QO₂ of either species over a 15-30-15. semidiurnal pattern of fluctuating salinity. Sea urchin perivisceral fluid PO₂ declined as ambient salinity fluctuated away from the acclimation salinity in both cycles and increased as ambient salinity returned to the acclimation salinity. Total nitrogen excretion of stepwise acclimated sea cucumbers declined significantly from 30 to 15 S, but there was no salinity effect on total nitrogen excretion in sea urchins. Ammonia excretion varied directly with salinity in stepwise acclimated sea cucumbers (67–96% of total nitrogen excreted), but there was no salinity effect on ammonia excretion (89–95% of total nitrogen excreted) of sea urchins. Urea excretion did not vary with salinity in sea cucumbers (2–4% of total nitrogen excreted) or sea urchins (2–9% of total nitrogen excreted). Primary amines varied inversely with salinity in sea cucumbers (2–30% of total nitrogen excreted), but did not vary with salinity in sea urchins (2–4% of total nitrogen excreted). The oxygen: nitrogen ratio of both species indicated that carbohydrate and/or lipid form the primary catabolic substrate. The O:N ratio did not vary as a function of salinity. Both species are more tolerant to reduced salinity than previously reported, however, rates of oxygen consumption and/or nitrogen excretion are modified by salinity as well as season.     

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     

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.     

Temporal variation in community interfaces: kelp-bed boundary dynamics adjacent to persistent urchin barrens

We examined, over 2 years, factors affecting the temporal stability of the lower limit of kelp beds (Alaria esculenta) at five subtidal sites in the Mingan Islands, northern Gulf of St. Lawrence. The position of the lower limit of the beds varied markedly among sites and over time and was largely controlled by the green sea urchin, Strongylocentrotus droebachiensis, which formed dense (up to 500 individuals m^–2) feeding fronts at the lower edge of the beds. These aggregations advanced over the kelp most rapidly during the summer (at rates as high as 2.5 m month^–1), and there appeared to be a threshold urchin biomass of ~5 kg m^–2 below which the fronts could not substantially reduce the limit of the beds. The fronts consisted mainly of large individuals, whereas smaller urchins predominated in the barrens zone below the kelp beds. At one site, we recorded large seasonal shifts in overall urchin densities, with large increases and decreases during the summer and winter, respectively. An urchin exclusion experiment indicated that algal recruitment in the barrens was two orders of magnitude greater in the absence than in the presence of urchins. The kelp Agarum cribrosum greatly restricted urchin movements, and the greater temporal stability of the kelp bed at one site appeared related to the gradual replacement of Alaria esculenta in the lower kelp bed by a large stand of Agarum cribrosum. We propose that perturbations by abiotic factors (e.g., ice scouring and water motion) trigger important but localized changes in urchin densities that, in turn, largely determine the limits of kelp bed distribution in this region of the Atlantic where urchin barrens are a persistent community state.Communicated by R.J. Thompson, St. Johns     

Settlement, recruitment and potential predators and competitors of juvenile echinoderms in the rocky subtidal zone

Recruitment patterns of marine invertebrates are affected both by settlement and early post-settlement events. This study examined the settlement and recruitment patterns of echinoderms at three sites in the rocky subtidal zone of Bocabec Cove, Bay of Fundy, Canada using artificial turf collectors and quadrats on the natural substrate. Potential predators were quantified at two of the sites along transects and in 1-m² quadrats. Both potential predators and competitors were quantified in 0.0625-m² quadrats. Settlement varied across sites (1.5–3 km apart) and two years of sampling (2004, 2005). The site of most potential settlement differed for the three groups of echinoderms: sea urchins (Strongylocentrotus droebachiensis), sea stars (Asterias spp.) and sea cucumber (Psolus fabricii). Settlement densities on the artificial turf collectors tended to be greater than the densities of settlers on the natural substrate. On the natural substrate, the only significant difference between densities of juveniles over time was that newly settled sea stars were found in July and were not found the following October. Large lobsters and carnivorous worms were potential predators with densities that varied between sites. Potential competitors that differed in abundance between sites were herbivorous gastropods and conspecifics for sea urchins; and carnivorous worms for sea stars. This study suggests that patterns of recruitment are either set up by patterns of settlement or by events during the first few weeks/months on the benthic substrate for these echinoderms.     

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.     

General ecology of the cassiduloid urchin Cassidulus caribbearum

Cassidulus caribbearum, a representative of the echinoid order Cassiduloida, occurs in localized populations of high densities (up to 100 m^-2) among the islands of the Puerto Rico bank in the eastern Caribbean Sea. These small (up to 35 mm) urchins are burrowers, principally in shallow-water areas with coarse sand bottom. Locomotion is achieved by ditaxic waves, passing from the front to the rear of large movable spines on the lateral portions of the ventral surface; this mechanism is unique among echinoids. Digging was most effective in sand grain sizes most closely approximating those of the environment (0 to 1). These urchins are somewhat selective deposit feeders, which ingest substrate particles primarily in the size range 0 to 1. Individuals ingest sand more or less continuously; complete passage of sand through the gut takes an average of 6 h in the laboratory. Aristotle's lantern is present in young juveniles (test length <1.0 mm), but is resorbed before the urchins reach a test length of 5.0 mm. The gut shows similarities in gross morphology to the spatangoid gut. Oxygen consumption is comparable to that of other tropical echinoids of similar wet weight. C. caribbearum broods its large (350 m) yolky eggs among the aboral spines for 10 to 12 days, at which time young urchins crawl off into the sand. Successive, overlapping broods are produced, so that frequently two broods of different developmental stages are present on a single urchin. Single urchins may brood continuously for up to 4 months or more. Throughout the year, at least 50% of all individuals over 18 mm were found brooding, with a maximum of 85% of all individuals brooding in mid-summer. The sex ratio of the population in all areas sampled was greater than 5 females to 1 male. No evidence of protandric hermaphroditism was found. Growth rate, measured from tagged indivuduals and from increase in median size in population samples, was between 0.5 and 1.0 mm month^-1. Mortality was due primarily to predation by the gastropod Cassis tuberosa, although occasional, episodic mortality was caused by physical factors such as heavy storm swells.     

Movement and feeding activity of red sea urchins (Strongylocentrotus franciscanus) adjacent to a kelp forest

Movement and feeding were studied in a population of red sea urchins, Strongylocentrotus franciscanus (Agassiz, 1863), found within and immediately seaward of a kelp forest offshore from Santa Cruz, California, USA. Mean sea urchin movements varied from 7.5 cm/day inside the kelp forest to over 50 cm/day at 15 and 100 m outside the kelp forest. The percentage of sea urchins feeding decreased from 66% inside the kelp forest to 16 and 15% at 15 and 100 m outside the kelp forest. These data indicate that movement by these sea urchins is a response to a low food supply.     

Induction of metamorphosis of two species of sea urchin from Sea of Japan

The induction of metamorphosis in mature larvae by selected chemical compounds and natural substrata was investigated in two species of sea urchins from the Sea of Japan, Strongylocentrotus intermedius and Scaphechinus mirabilis. Glutamine in crystalline form was added directly to water containing mature larvae, and this compound, at a final concentration of 10 g ml^-1, was an inducer of metamorphosis in S. intermedius (100% activity) and S. mirabilis (50% activity). Gutamine, or its natural mimetic molecules, may be an active component of the exogenous cue that induces metamorphosis of S. intermedius larvae. This exogenous cue was produced by the epiphytic calcareous algae, Melobesia spp. that colonized the older sea grass Zostera marina. Glutamic acid was also used as an inducer of metamorphosis for S. intermedius and S. mirabilis larvae (50 to 60% activity), but it was toxic to the larvae.     

Precision and accuracy of stable isotope signals in otoliths of pen-reared cod (<Emphasis Type="Italic">Gadus morhua</Emphasis>) when sampled with a high-resolution micromill

A high-resolution sampling technique for stable-isotope measurements in otoliths is now available, but little is known about the precision and accuracy of the results when using this technique. Otolith samples of pen-reared cod (Gadus morhua L.) with well-defined temperature history were obtained using this high-resolution sampling technique. Samples from different-aged cod otoliths were collected in two yearly increments corresponding to the calendar years 1994 and 1995, and the stable-oxygen (¹⁸O) and -carbon (¹³C) isotope compositions of the samples were measured. Otolith ¹⁸O values had a clear seasonal pattern in response to the seasonal water temperature. Otolith material deposited in the same calendar years of different-aged fish showed similar ¹⁸O values. High precision of the temperature estimate using otolith ¹⁸O values was obtained when comparing results from different otoliths. The accuracy of the temperature estimates, relative to measured mean monthly water temperature, was also high, but reduced sampling resolution in the otoliths significantly reduced the accuracy due to attenuation of the ¹⁸O signals. We found that the otolith ¹³C values had a cyclic pattern, roughly in phase with the ¹⁸O values. This was most likely caused by temperature-increased metabolism. Indications of age-specific otolith ¹³C values are also presented.Communicated by M. Kühl, Helsingør     

Effects of temperature and salinity acclimation of adults on larval survival,physiology, and early development of Lytechinus variegatus (Echinodermata: Echinoidea)

Larval survival and developmental rates of Lytechinus variegatus (Lamarck) were determined as a function of temperature and salinity in two experiments by: (1) directly transferring fertilized eggs to 35, 30, 27.5, 25, 20, 15, and 10S seawater at 18 and 23°C, and (2) acclimation of adult sea urchins to the conditions described above for 1 to 4 wk prior to spawning. Developmental rates and percent survival of larvae prior to metamorphosis decreased at salinities below 35 (Q₁₀ values for metamorphosis=0.380 to 0.384). Temperature and salinity significantly (P<0.05) affected metabolic rates of L. variegatus plutei. These results show that L. variegatus larvae are stenohaline when compared to larvae of other echinoderm species. LC₅₀ values (S), developmental rates, and survival to metamorphosis indicate that acclimation of adult sea urchins to lower salinity prior to spawing and fertilization does not enhance development or survival of embryos exposed to low salinity.     

Growth and mortality of subtidal red sea urchins (Strongylocentrotus franciscanus) at San Nicolas Island,California, USA: problems with models

Red sea urchins, Strongylocentrotus franciscanus, were tagged with tetracycline in 1990 at subtidal sites off San Nicolas Island, California, USA. After one year in the field, the sea urchins were collected and growth increments were measured based on tetracycline marks, which indicated initial slow growth, a maximum rate, and finally a prolonged period of very slow growth. Small red sea urchins (4 cm diam) were estimated to be 3 to 4 yr old, which is much older than has previously been reported. It is estimated that about 12 yr would be required to attain 10 cm diam. Survival has previously been modeled assuming a constant rate. If the population of red sea urchins is assumed to be stable and stationary, annual survival rate was between 71 and 77% yr^-1. Census data for the two years of study have permitted annual survival to be estimated without assuming stable and stationary population structure. Under these conditions, annual survival rate was between 79 and 86% yr^-1. Analysis of transitions in the size distributions from 1990 to 1991 suggested that annual survival may have been sizespecific: 91% yr^-1 for individuals 1.1 to 4.0 cm diam, 82% yr^-1 for individuals 4.1 to 7.0 cm diam, and 63% yr^-1 for those of 7.1 to 10.0 cm diam. An alternative explanation to size-specific survival in our study is sizespecific immigration.     

Population dynamics and reproductive biology of the commensal isopod Colidotea rostrata (Crustacea: Isopoda: Idoteidae)

Colidotea rostrata (Benedict, 1898) is the only known commensal idoteid isopod, living on and mimicking the color of two northeastern Pacific sea urchins of the genus Strongylocentrotus. The population dynamics and reproductive biology of C. rostrata on its host urchins were studied at a low rocky intertidal area in southern California (33°40N; 118°30W) from December 1984 to December 1986. Isopod populations remained relatively stable throughout the 2 yr study, with isopods inhabiting an average of 56.1% of the urchins at 6.1±0.6 (x±2 SE) isopods per urchin. Female isopods reach sexual maturity at a length of 6.8 mm. Fecundity in C. rostrata averaged 11.8±0.9 (x±2 SE) embryos per brood, and increased with female body size. Breeding occurs all year long in C. rostrata, with a main reproductive period between the warmer spring and summer months. Newly released mancas and juvenile isopods were present during all months of the study. C. rostrata differs from the free-living Idoteidae in its smaller maximum size, reduced fecundity, 1:1 sex ratio, and low juvenile mortality. These features may represent adaptations to a commensal life style that reflect a reduced mortality pressure on these isopods.     

Diet and trophic position of Atlantic bluefin tuna (<Emphasis Type="Italic">Thunnus thynnus</Emphasis>) inferred from stable carbon and nitrogen isotope analysis

Stable ¹³C and ¹⁵N isotope analyses of scale, bone, and muscle tissues were used to investigate diet and trophic position of North Atlantic bluefin tuna (Thunnus thynnus Linnaeus) during residency in the northwest Atlantic Ocean off the northeast coast of the United States. Adult bluefin tuna scales collected from fish between June and October 2001 were significantly enriched in ¹³C compared to both muscle and bone across all months, while muscle was significantly enriched in ¹⁵N compared to either bone or scale throughout the same period. In muscle tissue, there was evidence of a shift over the summer from prey with ¹³C values (–17 to –18) that were characteristic of silver hake (Merluccius bilinearis) to species with ¹³C values of –20 to –21 that were similar to Atlantic herring (Clupea harengus) and sandlance (Ammodytes americanus). Depletion of ¹⁵N values in adult scales and bone compared to muscle tissue may be explained by bone and scale samples representing juvenile or life-long feeding habits, isotopic routing, or isotopic differences in amino acid composition of the three tissue types. Adult bluefin tuna were estimated to be feeding at a trophic position similar to pelagic sharks in the northwest Atlantic Ocean, while the trophic positions of yellowfin tuna (Thunnus albacares), albacore tuna (Thunnus alalunga), and juvenile bluefin tuna were indicative of a diet of up to a full trophic position below adult bluefin tuna. The close relationship between the juvenile bluefin ¹⁵N values and those of suspension feeders suggests that nektonic crustaceans or zooplankton may contribute significantly to the diet of bluefin tuna, a food source previously overlooked for this species in the northwest Atlantic Ocean.Communicated by J.P. Grassle, New Brunswick