Secondary metabolites of the cyanobacterium Microcoleus lyngbyaceus and the sea hare Stylocheilus longicauda: palatability and toxicity

The cyanobacterium Microcoleus lyngbyaceus (Kütz.) Crouan (Lyngbya majuscula Gomont) produces a variety of nitrogen-containing secondary metabolites. M. lyngbyaceus is eaten by the specialist sea hare Stylocheilus longicauda, which sequesters secondary metabolites from its diet and transforms some sequestered metabolites into related compounds. We examined the palatability and toxicity of the metabolites malyngamide A, malyngamide B, and a mixture of majusculamides A and B (from M. lyngbyaceus), and malyngamide B acetate (from S. longicauda, derived from malyngamide B), in order to explore a series of hypotheses about why M. lyngbyaceus produces secondary metabolites and why S. longicauda sequesters and transforms them. All three M. lyngbyaceus metabolites significantly reduced feeding by omnivorous pufferfish (Canthigaster solandri) and crabs (Leptodius spp.). Direct comparisons indicated that neither malyngamide A nor the majusculamide mixture differed significantly in palatability from malyngamide B. The S. longicauda metabolite malyngamide B acetate did not significantly deter feeding by either consumer. Direct comparisons indicated that pufferfish found malyngamide B acetate more palatable than malyngamide B, and that crabs showed a non-significant trend in this direction. Palatability of the metabolites did not correlate with toxicity. Although no significant differences were observed among the M. lyngbyaceus metabolites in their palatability, two toxicity assays consistently ranked malyngamide A more toxic than malyngamide B, which was more toxic than the majusculamide mixture. The majusculamide mixture did not significantly differ in toxicity from malyngamide B acetate, even though the two differed strongly in palatability.     

Dietary preferences of the opisthobranch mollusc Stylocheilus longicauda for secondary metabolites produced by the tropical cyanobacterium Lyngbya majuscula

Pure compounds isolated from the cyanobacterium Lyngbya majuscula Gomont were evaluated in an artificial diet for their influence on the feeding preferences of the sea hare Stylocheilus longicauda (Quoy and Gaimard, 1824), which lives in and feeds on this filamentous cyanobacterium (blue-green alga). Microcolin B, ypaoamide, malyngolide and other natural products acted as feeding deterrents at natural concentrations. At lower concentrations, sea hares were indifferent to ypaoamide and malyngolide in their diets. In contrast, barbamide stimulated sea hare feeding at the concentrations normally found in L. majuscula. Malyngamides and majusculamides, the most common natural products found in samples of L. majuscula from Guam, increased sea hare feeding at low concentrations and inhibited feeding at the higher concentrations that occurred in some collections of L. majuscula. Dietary selection of cyanobacteria by S. longicauda may be regulated by the concentration of specific chemical cues produced by L. majuscula. Received: 28 January 1997 / Accepted: 24 April 1998     

Niche separation in two species of hare. Metabolic costs of plant phenolics

Summary The distributions of mountain hare (Lepus timidus) and European hare (L. europaeus) overlap in central Sweden, but they occupy separate food niches in winter. In Scandinavia, the European hare is mainly a grazer while mountain hare is a predominant browser. Browse contain high amounts of secondary metabolites, such as phenols, compared to grass. This raises the question if the two hare species differ in their metabolic tolerance of plant phenols and that these differences influence their food choice.Phenolic excretion in urine increased significantly with phenolic intake in both species. Excretion of glucuronic acid conjugates, one of the major pathways of elimination of phenols in both hare species, is positively correlated to phenolic intake and excretion. However, the extent of excretion of phenolics by this route was different in the two species of hare. European hares excreted substantially more glucuronic acid per amount of phenolics than mountain hare. The phenols were metabolized to a larger extent in the mountain hare, indicating a higher detoxification capacity. From these results it is likely that European hare have a higher cost for the detoxification of plant phenols compared to mountain hare. This cost and negative effect on sodium balance when feeding on browse may prevent exploitation of forest habitats by European hares, whereas mountain hare are better able to do so.     

Seaweed secondary metabolites as antifoulants: effects of Dictyota spp. diterpenes on survivorship, settlement, and development of marine invertebrate larvae

Summary. A recent investigation showed that the brown seaweed Dictyota menstrualis was unfouled relative to co-occurring seaweeds, and that larvae of fouling invertebrates avoided settling on D. menstrualis due to chemicals on its surface. The secondary metabolites dictyol E and pachydictyol A are among the compounds found on this alga's surface. In the present study, we tested the effects of specific diterpenes from Dictyota on the survivorship, growth, and development of invertebrate larvae and developing juveniles that could foul seaweeds. Exposure to dictyol E, dictyol B acetate, pachydictyol A, and dictyodial from Dictyota menstrualis and D. ciliolata caused significant larval mortality, abnormal development, and reduce growth rates for three species of co-occurring invertebrates when their larvae were forced into contact with these metabolites. Larvae were damaged at metabolite concentrations as low as 5% of maximum possible surface concentrations of these compounds for the populations of Dictyota we studied. The negative effects of these secondary metabolites on potential foulers, in conjunction with data demonstrating larval avoidance of dictyol-covered surfaces, suggest that these compounds could function as chemical defenses against fouling, and could select for larvae that avoid hosts producing these metabolites. Received 25 May 1998; accepted 22 June 1998.     

Effect of macrophyte secondary metabolites on feeding preferences of the herbivorous parrotfish Sparisoma radians

Experiments were conducted to test the role of secondary metabolites in determining the natural feeding preference hierarchy of the bucktooth parrotfish Sparisoma radians. The two least preferred food genera of S. radians, Halimeda and Penicillus, both contain 1,4-diacetoxy-1,3-butadiene terpenes, while the most preferred species, Thalassia testudinum, does not. Experiments with agar cylinders containing macrophyte homogenates showed that macrophyte biteability was not a factor. Instead preference could be altered by the application of the diacetoxybutadiene containing terpenes 4,9-diacetoxyudoteal and caulerpenyne or fractions or extracts containing them at naturally occurring concentrations. Concentration of the active terpenes affected the intensity of the fish's preference for the control in pairwise comparisons. Extracts and fractions which did not contain 4,9-diacetoxyudoteal or caulerpenyne did not affect fish feeding preferences at naturally occurring concentrations. Experiments in which S. radians were given no plant choice showed that coating T. testudinum with H. incrassata organic crude extract reduced the number of bites consumed and the biomass consumed to a level equivalent to that obtained for H. incrassata plants.     

Prey defense,predator preference,and nonrandom diet: The interactions between Pycnopodia helianthoides and two species of sea urchins

Interactions between the predatory sea star Pycnopodia helianthoides (Brandt, 1835) and two of its natural prey, the sea urchins Strongylocentrotus purpuratus (Stimpson, 1857) and S. franciscanus (Agassiz, 1863), are examined with regard to predator preference, predator diet, and prey defenses. The sea star is able to detect both species of sea urchin upstream in a Y-trough, but does not consistently choose one over the other (i.e., no preference). However, when the sea star is presented with equal numbers of similar-sized specimens of the two species of sea urchin, its diet is markedly nonrandom, since S. purpuratus is eaten almost 98% of the time. The defensive responses of the two species of sea urchin differ in form and effectiveness. S. franciscanus employs its long spines as defensive weapons, pinching the rays of an attacking sea star. This defensive response is more effective than the pedicellarial response used by S. purpuratus. The nonrandom diet of the predator seems to result primarily from prey defensive responses that differ in effectiveness, rather than from an intrinsic, behavioral preference of the predator at an earlier stage in the predator/prey interaction.     

Mycosporine-like amino acids: possible UV protection in eggs of the sea hare Aplysia dactylomela

We investigated mycosporine amino acid (MAA) involvement as protective sunscreens in spawn of the sea hare Aplysia dactylomela to determine if adult diet and ultraviolet (UV) exposure affected the UV sensitivity of developing embryos. Adults were fed a red alga rich in MAAs (Acanthophora spicifera) or a green alga poor in MAAs (Ulva lactuca). Adults on each diet were exposed for 2 wk to ambient solar irradiance with two types of acrylic filters; one allowed exposure to wavelengths >275 nm (designated UV) and one to wavelengths only >410 nm (designated NOUV). Spawn from each adult group was likewise treated with UV or NOUV and monitored during development for differences in mortality and metabolic rate (measured as oxygen consumption: V˙ _O2). Also recorded were number of eggs or embryos per capsule, times to hatching, hatching success, size at hatching, and V˙ _O2 of adults. Spawn from adults eating red algae was almost twice as rich in MAAs as spawn from adults eating green algae, suggesting that MAA content is diet-related. Although overall quantities of MAAs in the spawn reflected MAA contents of the adult diet, specific MAAs were differentially sequestered in the spawn. Thus, porphyra-334, found in high concentration in Aplysia dactylomela's preferred red algal food, was present in only low concentration in the spawn. Conversely, mycosporine-glycine, in low concentration in red algal food, was the most abundant MAA in the spawn. UV treatment of adults had no effect on quantities of MAAs in the spawn. Adults exposed to UV had significantly higher V˙ _O2s and spawned twice as often. The UV-treated adults produced spawn with significantly higher V˙ _O2s and their embryos developed to hatching sooner. The only significant effect of UV exposure of the spawn was to reduce the percentage of veligers hatching from 71 to 50%. There was no significant effect on hatching time or size of the veligers at hatching, nor on number of eggs per capsule. Received: 13 May 1997 / Accepted: 27 June 1997     

Lack of sequestration of host plant glucosinolates in <Emphasis Type="Italic">Pieris rapae</Emphasis> and <Emphasis Type="Italic">P. grarricae</Emphasis>

Summary. Sequestration of plant toxins in herbivores is often correlated with aposematic coloration and gregarious behaviour. Larvae of Pieris brassicae show these conspicuous morphological and behavioural characteristics and were thus suggested to sequester glucosinolates that are characteristic secondary metabolites of their host plants. P. rapaeare camouflaged and solitary, and are thus not expected to sequester. To test this hypothesis and to check the repeatabi-lity of a study that did report the presence of the glucosinolate sinigrin in P. brassicae, larvae were reared on three species of Brassicaceae (Sinapis alba, Brassica nigra and Barbarea stricta), and different leaf and insect samples were taken for glucosinolate analysis. The major host plant glucosinolates could only be found in traces or not at all in larval haemolymph, bled or starved larvae, faeces or pupae of both species or P. brassicae regurgitant. Haemolymph of both Pieris spp. was not rejected by the ant Myrmica rubra in dual-choice assays; the regurgitant of P. brassicae was rejected. This suggests the presence of compounds other than glucosinolates that might be sequestered in or produced by P. brassicae only. In faeces of both Pieris spp. a compound which yielded 4-hydroxybenzylcyanide (HBC) upon incubation with sulfatase was detected in high concentrations when larvae had been reared on S. alba. This compound may be derived from hydrolysis of sinalbin, the main glucosinolate of that plant. The unidentified HBC progenitor was apparently not sequestered in the two Pieris spp., and was not detected in faeces of larvae reared on B. nigra or B. stricta. Received 18 July 2002; accepted 11 September 2002.     

Sources of variation in herbivore preference: among-individual and past diet effects on amphipod host choice

Understanding which factors affect the feeding preferences of herbivores is essential for predicting the effects of herbivores on plant assemblages and the evolution of plant–herbivore interactions. Most studies of marine herbivory have focussed on the plant traits that determine preferences (especially secondary metabolites), while few studies have considered how preferences may vary among individual herbivores due to genetic or environmental sources of variation. Such intraspecific variation is essential for evolutionary change in preference behaviour and may alter the outcome of plant–herbivore interactions. In an abundant marine herbivore, we determined the relative importance of among-individual and environmental effects on preferences for three host algae of varying quality. Repeated preference assays were conducted with the amphipod Peramphithoe parmerong and three of its brown algal hosts: Sargassum linearifolium, S. vestitum and Padina crassa. We found no evidence that preference varied among individuals, thus constraining the ability of natural selection to promote increased specialisation on high-quality S. linearifolium. Most of the variation in preference occurred within individuals, with amphipod preferences strongly influenced by past diet. The increased tendency for amphipods to select alternate hosts to that on which they had been recently feeding indicates that amphipods are actively seeking mixed diets. Such a feeding strategy provides an explanation for the persistence of this herbivore on hosts in the field that support poor growth and survival if consumed alone. The effects of past diet indicate that herbivore preferences are a function of herbivore history in addition to plant traits and are likely to vary with the availability of algae in space and time.     

Growth,reproduction and mortality of the sea hare Dolabella auricularia (Gastropoda: Aplysiidae) in the Central Visayas,Philippines

The growth parameters W and K of the von Bertalanffy growth equation were estimated from size-frequency data of the sea hare Dolabella auricularia using an objective, computer-based method. The results obtained from two locations in the central Philippines were comparable, although based on only a few individuals; the means for both sites were W=493 g and K=0.9 (yearly basis). A value of Z=3.66, corresponding to an annual survival rate of 2.6%, was estimated for the juveniles and adults from a length-converted catch curve. Spawning and recruitment were found to occur throughout the year with peaks in May to July and September to October.ICLARM Contribution No. 170     

Atypical plant–herbivore association of algal food and a kleptoplastic sea slug (Elysia clarki) revealed by DNA barcoding and field surveys

The identity of food sources and feeding preferences of specialist herbivores have been commonly inferred from spatial associations between consumer and food items. However, such basic information for well-known marine herbivores, sacoglossans (sea slugs), and their algal diets remains disappointingly lacking, especially from field studies. The sacoglossan, Elysia clarki (Pierce et al. in Molluscan Res 26:23–38, 2006), is kleptoplastic and sequesters chloroplasts from algal food to photosynthesize, so DNA identification of sequestered chloroplasts was employed to verify the algal species fed upon by the slug across its geographic range. The molecular information on the algae consumed by E. clarki was combined with field surveys of slugs and algae in slug habitats in the Florida Keys in July and August of 2008 in order to evaluate whether the diet of this herbivore could be predicted based on its spatial association with algae in the field. A considerable mismatch between food availability and kleptoplast identity was recorded. E. clarki commonly occupied areas devoid of potential food and often contained symbiotic plastids from algal species different from those most frequently found in the surveyed habitats. In three of the four study sites, algal species present were poor predictors of slug diet. These findings suggest that the photosynthetic capability of E. clarki may release the slug from the constraint of requiring proximity to its food sources and may allow for the potential lack of spatial coupling between this herbivore and its algal food. This combination of field surveys and DNA barcoding provided critical and previously unavailable information on herbivore feeding in this marine system.     

Food availability and diet composition of three coexisting Mediterranean limpets (Patella spp.)

The gut contents of three intertidal patellid limpets were analysed by collecting foraging specimens on a breakwater on the Tyrrhenian coast (central Italy) between May 1988 and October 1989. The three species coexist there showing a different, but partially overlapping zonation: Patella aspera dominates the infralittoral fringe; the majority of P. caerulea inhabits the lower midlittoral, while P. rustica is most abundant in the upper midlittoral. The algae present on slivers of substratum over which each limpet collected was moving were identified. Moreover, floristic surveys were made along the shore in order to characterize the algal cover of the different zonal belts. The floristic study revealed that the basic elements of algal communities typical of western Mediterranean rocky shores are present in the study area. The algae found on the slivers under the foraging limpets were generally representative of the algal community typical of the same zone. There was a marked difference between the diets of P. rustica and P. aspera due to the fact that the first species forages on a few low lying epilithic and endolithic Cyanophyceae, while P. aspera feeds on a large number of species belonging to all the main algal classes and life forms considered, including frondose epilithics and epiphytics. The diet of P. caerulea resembles that of P. aspera in algal heterogeneity, but is dominated by Cyanophyceae as in P. rustica. A detailed analysis of the differences between gut contents of each limpet species and the relative slivers showed an obvious general correspondence, but revealed also that the diets of the three species do not completely reflect the availability of algae. These findings suggest that the basic diet segregation mechanism between the three populations is their zonal separation. However, the difference in gut contents of heterospecific limpets foraging in the same zone suggests the existence of supplementary morphy-functional or behavioural mechanisms for diet segregation between the three species.     

Morphology, molecular phylogeny and photosynthetic activity of the sacoglossan mollusc, Elysia nigrocapitata, from Korea

The morphology, external coloration as well as the life span of a kleptoplastic mollusc, Elysia nigrocapitata, was affected by its algal diet. Among algal diets, Chaetomorpha moniligera was the best for growth but not for animal longevity. TEM studies showed several distinctive layers composed of different cell types in sectioned parapodia. The chloroplast-containing digestive cells were located beneath the layer of vacuolated cells. The digestive cells contained 10–15 chloroplasts, in varying states of intactness, and several nuclei. Chloroplasts were not enclosed by any membranous structure in the host cytosol. Chlorophyll a fluorometry showed that the photosynthetic activity of kleptoplasts in E. nigrocapitata could be maintained for a long time only when animals were kept in the dark. The photosynthetic activity of kleptoplasts lasted 3–4 days when the animals were exposed to continuous illumination of 200 μmol photons m^?2 s^?1. These results suggested that the contribution of kleptoplasts to the survival of the animals might be minimal if the chloroplasts are not sequestered continuously. Cox I, 16S rDNA, and 28S rDNA sequence data have been obtained in order to phylogenetically place the new species of Elysia found in Korea.     

Effects of environmental stressors on blood-glucose levels in sea hares,Aplysia dactylomela

Effects of natural stressors such as tidepool strandings, air exposure, and low salinity on blood-glucose levels were investigated in the sea hare Aplysia dactylomela from shallow areas of Discovery Bay, Jamaica. All treatments produced large and significant elevations in blood-glucose titre, 1.5-to 2.3-fold above baseline levels of 25 to 35 g glucose ml^-1. Response times were variable, with significant elevations being manifested within 30 to 120 min from initiation of the stressor. Recovery was swift, within an hour or two of restoration to pretreatment conditions, and often involved an undershoot to levels below control values. In two experiments involving tidepool strandings and associated high body-temperatures, excessively low blood-glucose titres were followed by death of all test individuals. When sea hares were exposed to 75 and 50% seawater (100%=33S) for 1 h, maximum elevation in blood-glucose concentrations occurred 1 to 2 h from onset of exposure, coincidental with maximum dilution of the body fluids of test individuals. The responsiveness of blood-glucose titres to relatively small temperature or salinity changes, or to short-duration air exposures, suggests that monitoring this physiological parameter may be a useful and sensitive means of diagnosing a wide variety of stressors in marine gastropods.     

Antimicrobial activity of extracts of Caribbean gorgonian corals

Extracts of 39 species of Caribbean gorgonians were tested for antimicrobial activity against 15 strains of marine bacteria. The bacteria consisted of three opportunistic pathogens, Vibrio parahaemolyticus, Leucothrix mucor, and Aerococcus viridans, and 12 strains isolated from either healthy or decayed gorgonians. Overall, only 15% (79 out of 544) of the tests resulted in antibacterial activity with 33% (13 out of 39) of the gorgonians inhibiting only one bacterial strain and 23% (9 out of 39) showing no activity. The extracts of four Pseudopterogorgia species showed relatively high levels of activity, inhibiting 43 to 86% of the bacterial strains. The potency of the active Pseudopterogorgia species was variable, however, and three additional Pseudopterogorgia species were inactive against all bacterial strains. With the exception of one sensitive strain, Vibrio species were resistant to gorgonian metabolites. Our results indicate that organic extracts of most Caribbean gorgonians do not possess potent, broad-spectrum antibacterial activity inhibitory to the growth of opportunistic marine pathogens and bacteria associated with healthy and decayed gorgonian surfaces. These findings suggest that the inhibition of bacterial growth is not the primary ecological function of gorgonian secondary metabolites and that bacteria may not be important selective agents in the evolution of gorgonian secondary chemistry.     

Resource utilization for decorating in three intertidal majid crabs (Brachyura: Majidae)

 For three spider crabs (Tiarinia cornigera, Micippaplatipes and Pugettia quadridens quadridens), patterns of algal utilization for decorating were compared with the dynamics of algae on an intertidal rocky shore reef where the crabs co-occurred. T. cornigera and P. quadridens quadridens were most abundant from autumn to spring when the dominant algae (Boodleacoacta, Sargassum hemiphyllum, S. thunbergii and Corallina pilulifera) occurred in high coverage, while M. platipes was most abundant from spring to autumn. Monthly change of algae used for decorating was not correlated with algae growing in the crab habitat for T. cornigera, but for M. platipes, it was positively correlated for two algal species, and for P. quadridens quadridens, negatively correlated for one algal species. Each species of the spider crabs used some algal species preferentially for decoration. Decorating preference experiments conducted in the laboratory showed that M. platipes and P. quadridens quadridens exhibited similar preference to their algal utilization in the field, whereas for T. cornigera, algal preference in the experiment differed from utilization in the field. Comparisons between materials used for decoration and gut contents revealed that T. cornigera and M. platipes used algal species differently for decorating and feeding, while P. quadridens quadridens used the same algal species for both decorating and feeding. Different tactics for camouflage are discussed in terms of algal utilizations by the three majid species. Received: 25 September 1999 / Accepted: 22 June 2000     

Toxicity of atrazine and chlorsulfuron to algae

Sensitivity of three algal species, Chlorella vulgaris, Scenedesmus acutus and Pseudanabaena galeata to herbicides atrazine and chlorsulfuron was studied using single species toxicity tests. Organisms were exposed to different concentrations of these herbicides and the algal growth was measured by turbidity at 750 nm. Atrazine appeared to be the most inhibitory to algae growth. 96 hr E_bC₅₀ of atrazine was: 1.3, 0.014, 0.014 mg/1 for C. saccharophila, S. acutus and P. galeata, respectively and 96 hr E_bC₅₀ of chlorsulfuron was 74.5 mg/1 for C.saccharophila, 0.19 mg/1 for S. acutus and 21.1 mg/1 for P. galeata     

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.     

Herbivorous amphipods inhabit protective microhabitats within thalli of giant kelp <Emphasis Type="Italic">Macrocystis pyrifera</Emphasis>

Many small marine herbivores utilize specific algal hosts, but the ultimate factors that shape host selection are not well understood. For example, the use of particular microhabitats within algal hosts and the functional role of these microhabitats have received little attention, especially in large algae such as kelps. We studied microhabitat use of the herbivorous amphipod Peramphithoe femorata that inhabits nest-like domiciles on the blades of giant kelp Macrocystis pyrifera. The vertical position of nest-bearing blades along the stipe of the algal thallus and the position of the nests within the lateral blades of M. pyrifera were surveyed in two kelp forests in northern-central Chile. Additionally, we conducted laboratory and field experiments to unravel the mechanisms driving the observed distributions. Peramphithoe femorata nests were predominantly built on the distal blade tips in apical sections of the stipes. Within-blade and within-stipe feeding preferences of P. femorata did not explain the amphipod distribution. Amphipods did not consistently select distal over proximal blade sections in habitat choice experiments. Mortality of tethered amphipods without nests was higher at the seafloor than at the sea surface in the field. Nests mitigated mortality of tethered amphipods, especially at the seafloor. Thus, protective microhabitats within thalli of large kelp species can substantially enhance survival of small marine herbivores. Our results suggest that differential survival from predation might be more important than food preferences in determining the microhabitat distribution of these herbivores.     

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.