A unique algal ridge system in the Exuma Cays, Bahamas

 An algal ridge system discovered along the Exuma Cays, Bahamas constructs bioherms to a thickness of at least 1.5 m and is associated with modern intertidal stromatolites. These algal ridges are unique because they grow in atypical environments characterized by relatively low wave energy, high rates of sedimentation and low rates of herbivory. They also are composed primarily of the branching crustose coralline alga, Neogoniolithon strictum, which heretofore was not known to form algal ridges. Lateral growth rates of crusts, vertical growth rates of branches and survivorship of transplanted N. strictum were greatest in the shallow fore reef zone of the algal ridge. The alga is also capable of surviving and growing when covered with sediment for at least 100 days. Under such conditions it transforms from a branched to an unbranched morphology. Parrotfish grazing, which is said to limit the abundance of branched corallines and algal ridges, was two orders of magnitude lower than in published accounts from other reef systems of the Caribbean and one order of magnitude less than that found on nearby coral reefs of the west Exuma Sound. Neogoniolithon strictum, a delicate and open-branched coralline, persisted for over a year without grazing damage when transplanted to a depth of 2.3 m. This algal ridge-building coralline becomes a well-indurated limestone following submarine lithification of sediment that infills the open branch framework. As a result, N. strictum ridges are comparable to the dense frameworks associated with most algal ridges. Observations of N. strictum -associated bioherms along Central America suggest this ridge system may exist elsewhere under conditions similar to those described for the Bahamas. Accepted: 23 March 1996     

Herbivory by crabs and the control of algal epibionts on Caribbean host corals

Summary A short-term experiment was conducted to examine the relationships among the branching coral Porites porites, algal epibionts, and a facultative crab associate Mithrax sculptus in Belize, Central America. Initial field observations suggested that coral colonies supporting resident crabs generally had lower algal cover than colonies without crabs. The hypothesis was tested that Mithrax significantly depresses host coral algal cover and thereby indirectly affects host survivorship and growth. Crab accessibility to an array of coral colonies, similarly covered with algal epibionts, was manipulated in three treatments. Results strongly support the hypothesis, with significant differences in algal cover (primarily Dictyota spp.) noted among treatments after only one month. Caged heads with crabs included and uncaged natural controls allowing crabs free access averaged less than 10% cover, whereas mean algal cover exceeded 75% where crabs were excluded. The uncaged treatment, in which crabs were allowed free access to Porites heads was not significantly different from the crab inclusion treatment. Collectively, these results demonstrate that under natural conditions, crabs can have pronounced effects on host corals by reducing fouling algal epibionts. Furthermore, these facultative coral associates may have more important, albeit localized effects on Caribbean corals than has been suggested previously.     

Diet composition influences the fitness of the herbivorous crab Grapsus albolineatus

The tropical rocky shore crab Grapsus albolineatus feeds primarily on filamentous algae but eats animal matter whenever it is available. During the summer the crab's diet switches to encrusting algae due to a die-off of filamentous algae. As a result of the switch the nutrients in the diet of the crab vary seasonally and may influence the fitness of the crab. Maintenance, growth, reproductive performance and nutrient storage of crabs were examined under four dietary regimes of increasing nutritional value ranging from low organic to high protein content. The nutritional quality of these diets significantly affected crab survival and moulting. Crabs fed on the nutritionally superior diet of algae and meat exhibited enhanced growth, higher levels of energy in the reproductive organs and stored more energy in the hepatopancreas than did individuals on the shore and crabs fed only on algal diets in the laboratory. Filamentous algae were a better food source than other algae, resulting in fewer deaths and superior levels of maintenance and growth. Growth and maintenance can occur on a pure algal diet, but reproductive performance and nutrient storage require some degree of added nutrients in the form of animal matter in the diet. Crabs fed coralline or foliose algae had higher mortality and fewer successful moults than crabs fed the other two diets. The fitness of G. albolineatus appears to be limited by the amount of extra nutrients obtained from animal matter. The opportunistic consumption of animal material in the form of carrion, or of animals associated with dietary algae, could be a key factor in the reproductive success of this crab.     

Geographic Variation in Camouflage Specialization by a Decorator Crab

In North Carolina, the decorator crab Libinia dubia camouflages almost exclusively with the chemically noxious alga Dictyota menstrualis. By placing this alga on its carapace, the crab behaviorally sequesters the defensive chemicals of the plant and gains protection from omnivorous consumers. However, Dictyota is absent north of North Carolina, whereas Libinia occurs as far north as New England. Crabs from three northern locations where Dictyota is absent (Rhode Island, Connecticut, and New Jersey) camouflaged to match their environment, rather than selectively accumulating any one species. When D. menstrualis was offered to crabs from northern sites, they did not distinguish between it and other seaweeds for camouflage, whereas crabs from Alabama and two locations in North Carolina used D. menstrualis almost exclusively. In addition, in winter and spring, when Dictyota was seasonally absent in North Carolina, Libinia selectively camouflaged with the sun sponge Hymeniacidon heliophila, which was chemically unpalatable to local fishes. Thus, southern crabs were consistent specialists on chemically defended species for camouflage, while northern crabs were more generalized. The geographic shift in crab behavior away from specialization coincides with a reported decrease in both total predation pressure and the frequency of omnivorous consumers. These shifts in the nature and intensity of predation pressure may favor different camouflage strategies (generalist vs. specialist), contributing to the observed geographic differences in camouflage behavior.     

Predictable spatial escapes from herbivory: how do these affect the evolution of herbivore resistance in tropical marine communities?

Summary Between-habitat differences in macrophyte consumption by herbivorous fishes were examined on three Caribbean and two Indian Ocean coral reefs. Transplanted sections of seagrasses were used as a bioassay to compare removal rates in reef-slope, reef-flat, sand-plain, and lagoon habitats. Herbivore susceptibility of fifty-two species of seaweeds from these habitats was also measured in the field. Seagrass consumption on shallow reef slopes was always significantly greater than on shallow reef flats, deep sand plains, or sandy lagoons. Reef-slope seaweeds were consistently resistant to herbivory while reef-flat seaweeds were consistently very susceptible to herbivory. This pattern supports the hypothesis that defenses against herbivores are costly in terms of fitness and are selected against in habitats with predictably low rates of herbivory.Sand-plain and lagoon seaweeds showed a mixed response when placed in habitats with high herbivore pressure; most fleshy red seaweeds were eaten rapidly, most fleshy green seaweeds were eaten at intermediate rates, and most calcified green seaweeds were avoided or eaten at very low rates. Differences in susceptibility between red and green seaweeds from sand-plain or lagoon habitats may result from differential competitive pressures experienced by these seaweed groups or from the differential probability of being encountered by herbivores. The susceptibility of a species to removal by herbivorous fishes was relatively consistent between reefs. Preferences of the sea urchin Diadema antillarum were also similar to those of the fish guilds.Unique secondary metabolites were characteristic of almost all of the most herbivore resistant seaweeds. However, some of the herbivore susceptible species also contain chemicals that have been proposed as defensive compounds. Genera such as Sargassum, Turbinaria, Thalassia, Halodule, and Thalassodendron, which produce polyphenolics or phenolic acids, were consumed at high to intermediate rates, suggesting that these compounds are not effective deterrents for some herbivorous fishes. Additionally, potential for the production of the compounds caulerpin, caulerpicin and caulerpenyne in various species of Caulerpa did not assure low susceptibility to herbivory.Heavily calcified seaweeds were very resistant to herbivory, but all of these species also produce toxic secondary metabolites which makes it difficult to distinguish between the effects of morphological and chemical defenses. Predictions of susceptibility to herbivory based on algal toughness and external morphology were of limited value in explaining differing resistances to herbivory.     

Unveiling commonalities in understudied habitats of boulder-reefs: life-history traits of the widespread invertebrate and algal inhabitants

Compared to stable reef habitats, dynamic boulder-reefs (commonly called boulder-fields when intertidal) host many habitat specialist species. Most occur underneath boulders where they are largely hidden from view; only limited research has assessed their life-histories despite their widespread importance for biological diversity. But some abundant under-boulder species likely structuring this system are habitat generalists widely researched elsewhere. Here we review this research, focusing on three widespread under-boulder sessile taxa: spirorbids, serpulids (tubeworms) and nongeniculate coralline algae, and three mobile taxa: sea urchins, chitons and crabs. Spirorbids have extensive reproductive/colonization capabilities but are readily out-competed. We thus characterize spirorbids as mostly early-successional, while serpulids often have greater competitiveness. Nongeniculate corallines occur underneath boulders where light reaches, although they can withstand low levels of that and most other resources. Such traits characterize nongeniculate corallines as late-successional. Thus, succession underneath boulders may shift deterministically from early tubeworms to late nongeniculate corallines. Habitat generalist sea urchin and chiton species often have strong inter-specific interactions in exposed habitats. Future experiments may find that under-boulder aggregations of these taxa, and also crabs, are impacting algal and invertebrate assemblages. These experiments will be required if dynamic boulder-reefs are to be as thoroughly understood as other benthic systems.     

Patterns in shallow subtidal marine assemblages along the coast of New South Wales

Shallow subtidal areas of rocky reefs in central and southern New South Wales may best be described as a mosaic of habitats, the distributions of which are seemingly related to depth, wave exposure and a number of biological processes, particularly herbivory. The Fringe habitat is generally found only in the most shallow waters. Forests of the laminarian alga Ecklonia radiata are often found at intermediate depths. In deeper, or more sheltered water, sponges, ascidians and red algae are more abundant and the abundances of sea urchins and other invertebrate grazers decline. Overlying this broad-brush pattern are patches of crustose coralline algae (the Barrens habitat), the distributions of which are not clearly related to depth. Invertebrate herbivores, and sea urchins in particular, are abundant in the Barrens habitat. The Barrens habitat was most represented at the more southern locations. At the two most northern locations, reefs were shorter in length and dominated by ascidians (Pyura species).     

Variation on a theme of herbivory: Corallina–hermit crab relationship on a temperate‐subtropical rocky shore

Elucidation of species interactions involving multiple processes is an important task in ecology, as different processes may operate in non‐apparent, opposing directions. While herbivory is a major form of interspecific interaction affecting plants, calcareous algae are generally thought to be protected from herbivory due to CaCO₃ deposition. However, field observations and experiments in temperate‐subtropical intertidal rockpools showed that hermit crabs (Paguroidea, Decapoda) did not ingest Corallina but cut short its fronds. We tested the implications of this frond‐cutting behaviour with field experiments where hermit crab density and Corallina frond length were artificially manipulated. Hermit crab guts were fuller and contained more microscopic animals when Corallina fronds were shorter, suggesting that frond‐cutting affects accessibility to food resources that exist in the basal part of algal mat. Corallina showed compensatory growth when fronds were artificially reduced in length/density, simulating the cutting by hermit crabs. Frond density increased, probably due to increased exposure to light of the basal part of Corallina coupled with a reduction in apical dominance. The present study revealed a direct, non‐trophic relationship between hermit crabs and Corallina in which the former affected the latter for gaining access to food resources. Overall, the combination of positive and negative effects of hermit crab led to a positive, facilitative effect on Corallina. In terms of the effect on algae or plants, this relationship is functionally analogous to herbivory, without involving direct consumption. We discuss the problem of extending the concept of ecological engineering to herbivory and herbivory‐like phenomena and suggest the term ‘pseudo‐herbivory’ to describe relations in which animals have non‐trophic but herbivory‐like effects on plants/algae, as seen in the hermit crab–Corallina relationship.     

The Role of Encrusting Coralline Algae in the Diets of Selected Intertidal Herbivores

Kalk Bay, South Africa, has a typical south coast zonation pattern with a band of seaweed dominating the mid-eulittoral and between two molluscan-herbivore dominated upper and lower eulittoral zones. Encrusting coralline algae were very obvious features of these zones. The most abundant herbivores in the upper eulittoral were the limpet, Cymbula oculus (10.4 ± 1.6 individuals m⁻²; 201.65 ± 32.68 g.m⁻²) and the false limpet, Siphonaria capensis (97.07± 19.92 individuals m⁻²; 77.93 16.02 g.m⁻²). The territorial gardening limpet, Scutellastra cochlear, dominated the lower eulittoral zone, achieving very high densities (545.27 ± 84.35 m⁻²) and biomass (4630.17 ± 556.13 g.m⁻²), and excluded all other herbivores and most seaweeds, except for its garden alga and the encrusting coralline alga, Spongites yendoi (35.93 ± 2.26% cover). In the upper eulittoral zone, encrusting coralline algae were only present in the guts of the chiton Acanthochiton garnoti (30.5 ± 1.33%) and the limpet C. oculus (2.9 ± 0.34%). The lower eulittoral zone limpet, Scutellastra cochlear also had a large percentage of encrusting coralline algae in its gut with limpets lacking gardens having higher (45.1 ± 1.68%) proportions of coralline algae in their guts than those with gardens (25.6 ± 0.8%). Encrusting coralline algae had high organic contents, similar to those of other encrusting and turf-forming algae, but higher organic contents than foliose algae. Radula structure, grazing frequencies as a percentage of the area grazed (upper eulittoral 73.25 ± 3.60% d⁻¹; lower eulittoral 46.0 ± 3.29% d⁻¹), and algal organic content provided evidence to support the dietary habits of the above herbivores. The data show that many intertidal molluscs are actively consuming encrusting coralline algae and that these seaweeds should be seen as an important food source.     

Predation risk predicts use of a novel habitat

Predator–prey interactions are often highly co‐evolved, with selection over time for prey with morphological and behavioral traits that minimize predation risk. Consequently, in many environments prey choose among potential habitats according to their refuge value. It is unclear, however, when presented with new habitats, if prey are able to evaluate the predation risk of these relative to familiar habitats and utilize these in accordance with their value. We tested whether, along the east coast of the USA, native mud crabs Panopeus herbstii utilize the non‐native alga Gracilaria vermiculophylla according to its relative refuge value. Experiments examining predation by blue crabs Callinectes sapidus on mud crabs revealed that the non‐native alga had an intermediate refuge value relative to native oysters, which were the most protective, and unvegetated sediment, which was the least. In subsequent choice experiments, mud crabs selected oysters over alga over unvegetated sediment, in accordance with habitat refuge values. Further, in field experiments, the use of Gracilaria by mud crabs was inversely related to the proximity of the alga to the preferred habitat type, oysters, and was reduced by the presence of a blue crab predator. Consequently, mud crabs are utilizing the non‐native alga Gracilaria in accordance with its intermediate refuge value. The relative refuge value of non‐native vs native habitat‐forming species may provide a baseline expectation against which to measure the speed of learning and opportunism in the response of native prey to novel protective habitats.     

Seaweeds preferred by herbivorous fishes

Marine macrobenthic algae (or seaweeds), epiphytic microalgae, and other aquatic plants constitute the main food items of marine herbivorous fishes. About 5% of all fish species are herbivorous; only 30% of these are marine, most of them living in coral reefs. An analysis was performed on all the seaweeds that formed part of the natural diet of these fishes, based on information contained in FishBase (). The results showed that many coral-reef-associated marine herbivorous fishes, such as the families Blennidae, Kyphosidae and Siganidae, fed selectively on filamentous and turf fleshy seaweeds, which they prefer over calcareous coralline and encrusting species. In particular, Chlorophyceae of the genera Cladophora, Enteromorpha and Ulva were preferred by Scartichthys viridis (Blennidae), Girella spp. (Kyphosidae), Sarpa salpa (Sparidae), and Phaeophyceae in the genera Sargassum and Dictyota were preferred by Kyphosus spp. (Kyphosidae) and Siganus spp. (Siganidae). A web-based tool was developed to provide information on plants (algae, seagrasses, terrestrial plants and fruits) preferred as food by herbivorous fishes (). The tool is intended to assist aquaculturists, conservationists and ecosystem-based fisheries managers.     

Seaweed-herbivore-predator interactions: host-plant specialization reduces predation on small herbivores

Summary Because feeding specialization among marine herbivores is rare, marine communities provide a simplified system for identifying factors selecting for specialization. On Australia's Great Barrier Reef, we investigated interactions among the chemically-defended seaweed Chlorodesmis fastigiata, herbivores specialized on this alga, and potential predators of these herbivores. Chlorodesmis is a low preference food for reef fishes but appears to be the only food of the crab Caphyra rotundifrons and the ascoglossan gastropods Elysia sp. and Cyerce nigricans. The crab is found only in patches of Chlorodesmis, feeds solely on the alga, and selectively shelters in it in laboratory choice experiments. Crab grazing on the red seaweed Acanthophora spicifera was stimulated when this alga was coated with increasing concentrations of the cytotoxic diterpenoid chlorodesmin, the major secondary metabolite of Chlorodesmis. Crabs did not sequester Chlorodesmis metabolites but avoided predators by sheltering in the unpalatable alga. All crabs tethered on the reef without access to Chlorodesmis patches were rapidly eaten; those with access to Chlorodesmis patches were much less susceptible to predation. The cryptic ascoglossan Elysia sp. was found exclusively in patches of Chlorodesmis and sequestered metabolites from the alga. Living Elysia were unpalatable to the common wrasse Thalassoma lunare in laboratory assays, but the crude organic extract of Elysia did not significantly deter feeding by Thalassoma. Elysia sequestered chlorodesmin, which deterred feeding by reef fishes in field assays but was ineffective against Thalassoma in laboratory assays at 5% food dry mass. Unlike Elysia, the aposematically colored ascoglossan Cyerce nigricans sequestered Chlorodesmis metabolites in relatively small amounts, but produced larger amounts of unrelated polypropionate compounds. Cyerce were never attacked by fishes and the crude organic extract of this slug strongly deterred feeding by wrasses in laboratory assays. The dorid nudibranch Gymnodoris sp. was found only in Chlorodesmis patches and appeared to be a specialized predator on Elysia; it would not prey on Cyerce. Data from this and other recent investigations demonstrate that some small marine herbivores feed selectively or exclusively on seaweeds that are chemically defended from fishes. This association reduces predation on the herbivores and suggests that escape from and deterrence of predation may be a dominant factor selecting for specialization among these herbivores.     

Refuge function of marine algae complicates selection in an intertidal snail

Species with restricted gene flow often show trait-shifts from one type of environment to another. In those rock-dwelling marine gastropods that lack larval dispersal, size generally decreases in wave-exposed habitats reducing risk of dislodgement, while increases in less exposed habitats to resist crab-crushing. In Littorina fabalis, however, snails of moderately exposed shores are generally much larger (11–14 mm) than snails of sheltered shores (5–8 mm). Observations from the White Sea (where crabs are not present) indicate that in the absence of crabs snails are small (6–7 mm) in both habitats. We assumed that the optimal size for L. fabalis in the absence of crabs is less than 8 mm, and thus that increased size in moderately exposed habitats in areas with crabs might be a response to crab predation. In a crab-rich area (Sweden) we showed that crab predation is an important mortality factor for this snail species in both sheltered and moderately exposed habitats. In sheltered habitats, snails were relatively more protected from crab-predation when dwelling on their habitual substrate, fucoid algae, than if experimentally tethered to rocks below the algae. This showed that algae function as snail refuges. Snail dislodgement increased, however, with wave exposure but tethering snails in moderately exposed habitats showed that large snails survived equally well on rocks under the algae as in the canopy of the algae. Thus in sheltered habitats a small snail size is favored, probably due to life-history reasons, while increased risk of being dislodged from the algae refuges promotes a large size in moderately exposed habitats. This study shows an example of selection of a trait depends on complex interactions of different factors (life-history optimization, crab predation, wave induced dislodgement and algal refuges).     

Experimental effects of kelp canopies on subtidal coralline algae

Plants are often grouped as canopy species or understorey species because it is thought that that these sets of taxa interact in predictable ways. Mensurative experiments in southern Australia demonstrated that the percentage cover of encrusting coralline algae was greater, and articulated (branching) coralline algae less, on boulders under a canopy of dense kelp (>7 plants per m²), Ecklonia radiata, than on boulders without kelp. Experimental clearances of kelp and reciprocal transplants of boulders between patches of E. radiata and patches without kelp showed that canopies maintained and facilitated the growth of encrusting coralline algae and reduced the cover of articulated coralline algae. Potential artefacts associated with clearing kelp and transplanting boulders were not detected when tested with a series of translocation controls. These results reject the model that the co‐occurrence of E. radiata and encrusting corallines is just an assemblage of plants caused by spatial and temporal coincidence. Instead, they support the model that kelp facilitates the growth and survival of understorey algae.     

COMPOSITION,DISTRIBUTION, AND ABUNDANCE OF DEEP‐WATER (>30 m) MACROALGAE IN CENTRAL CALIFORNIA1

The deep‐water macroalgal assemblage was described at 14 sites off the central California coast during 1999 and 2000 from SCUBA and remotely operated vehicle sampling. The stipitate kelp Pleurophycus gardneri Setchell & Gardner, previously thought to be rare in the region, was abundant from 30 to 45 m, forming kelp beds below the well‐known giant kelp forests. Macroalgae typically formed three broadly overlapping zones usually characterized by one or a few visually dominant taxa: 1) the upper “Pleurophycus zone” (30–45 m) of stipitate kelps and Desmarestia spp. with a high percent cover of corallines, low cover of uncalcified red algae, and rare green algae; 2) a middle “Maripelta zone” (40–55 m) with other uncalcified red algae and infrequent corallines and green algae; and 3) a zone (55–75 m) of infrequent patches of nongeniculate coralline algae. The green alga Palmophyllum umbracola Nelson & Ryan, not previously reported from the Northeast Pacific, was found over the entire geographical range sampled from 35 to 54 m. Year‐round profiles of water column irradiance revealed unexpectedly clear water with an average K₀ of 0.106·m ^{? 1} Received 18 January 2002. Accepted 16 December 2002. . The low percent surface irradiance found at the average lower macroalgal depth limits in this study (0.56% for brown algae, 0.12% for uncalcified red algae, and 0.01% for nongeniculate coralline algae) and lack of large grazers suggest that light controls the lower distributional limits. The ubiquitous distribution, perennial nature, and similar lower depth limits of deep‐water macroalgal assemblages at all sites suggest that these assemblages are a common persistent part of the benthic biota in this region.     

Coralline algal maerl frameworks-Islands within the phaeophytic kelp belt

Summary In the subtropical belt highly productive ecosystems are formed by coral reefs in oligotrophic seas. Towards more eutrophic conditions, coral reefs diminish and are subsequently replaced by highly productive kelp forests. In high latitudes framework constructing carbonate production is enhanced by the growth of branching coralline algae which predominantly generate maerl-type deposits. On a global view, these coralline algal ecosystems show an island-like distribution pattern within the phaeophytic kelp belt. Compared to kelp ecosystems, coralline-algaldominated ecosystems have low rates of productivity. Therefore, it is reasonable to seek the pronounced competitive value of the extremely slow-growing corallines. Due to their low annual growth increment, the coralline algae studied are very endangered by abiotic physical disturbances and by overgrowth of rapidly growing filamentous algae or sessile invertebrates. To overcome fouling pressure and storm-triggered physical disturbances, coralline algae thrive well in wave-sheltered headlands or skerry areas and generate characteristic ‘denuded areas’ by intense herbivory. This general distributional pattern is also true for high-boreal to subarctic coralline algal bioherms in northern Norway. Such a complex biological feedback maintains a high potential of self-regulation or self-organization in the algal reef bioherms. The different proponents involved in feedback processes include bacterial colonization, diatom microfouling and selective induction of larval metamorphosis. The negative impact of diatom microfouling and the important role of herbivores are relevant activities in the feedback system on a microscopic scale. Macroscopically, intense herbivory on coralline algae create denuded conditions, which are a widespread phenomenon in coralline algal ecosystems.     

Effects of Macroalgal Chemical Extracts on Spore Behavior of the Antarctic Epiphyte Elachista antarctica Phaeophyceae

Most macroalgal species along the Western Antarctic Peninsula (WAP) are defended against predation, many using chemical defenses. These subtidal communities are also mostly devoid of free living filamentous algae. However, one endo/epiphyte, Elachista antarctica, is found growing exclusively out of the palatable rhodophyte Palmaria decipiens. To understand this unusual and exclusive epiphytization, we tested whether macroalgal secondary metabolites such as those responsible for deterring sympatric grazers, affect the behaviors of the epiphyte's spores. Settlement, germination, and swimming behaviors of the epiphyte's motile spores were quantified in the presence of fractionated lipophilic and hydrophilic extracts of host P. decipiens and other rhodophytes from the shallow subtidal. Host P. decipiens was the only alga tested that did not inhibit spore settlement or germination. We also examined whether extracts from these chemically rich algae affect spore swimming behaviors and found spores to be chemotactically attracted to seawater soluble extract fractions of host P. decipiens. These results indicate that chemosensory behaviors of the epiphyte's spores to metabolites associated with these chemically defended macrophytes can explain this exclusive epiphyte–host interaction.     

Dispersion and phagokinesis in the echinoid Evechinus chloroticus (Val.)

Movement in the echinometrid sea urchin Evechinus chloroticus (Val.) was investigated in relation to the availability of a preferred food species, the laminarian alga Ecklonia radiata (C. Ag.). Patterns of microhabitat occupancy, dispersion, and movement were described in two habitats: Coralline Flats, dominated by encrusting coralline algae; and Sediment Flats, occupied by turfing corallines, Ecklonia and the mussel Modiolus areolatus (Gould). In both habitats, sea urchins were positively associated with encrusting coralline algae, despite the fact that encrusting corallines occupied a minor proportion of the substratum in the Sediment Flat habitat. Sea urchins were significantly clumped in both habitat types, and more so in sites within the Sediment Flats habitat. Movement was not directional in either habitat, but the magnitude of movement was almost twice as high in sites from the Coralline Flats habitat, where kelp was absent, as in the Sediment Flats sites, where kelp was present. Behaviour in response to the presence of Ecklonia in both habitats was investigated experimentally. In both habitats, urchins exposed to a “drift” kelp plant at a distance of 1–2 m exhibited net movement approximately twice as great as that shown by control urchins, not exposed to a kelp plant. Despite the relatively small distances the urchins were from the kelp, there was no evidence of directionality in movement. We conclude from the present study that further understanding of the feeding behaviour of urchins requires that a distinction be made between attached and drift algae, and between directionality and magnitude of movement, and that the interaction of these and other factors are best investigated under field conditions.     

Natural inducers for coral larval metamorphosis

 Coral gametes from Acropora millepora (Ehrenberg, 1834) and from multi-species spawning slicks provided larvae for use in metamorphosis assays with a selection of naturally occurring inducer chemicals. Four species of crustose coralline algae, one non-coralline crustose alga and two branching coralline algae induced larval metamorphosis. However, one additional species of branching coralline algae did not produce a larval response. Metamorphosis was also observed when larvae were exposed to skeleton from the massive coral Goniastrea retiformis (Lamarck, 1816) and to calcified reef rubble, demonstrating metamorphosis is possible in the absence of encrusting algae. Chemical extracts from these algae and the coral skeleton, obtained using either decalcification or simple methanol extraction procedures, also contained active inducers. These results extend the number of crustose algal species known to induce coral metamorphosis, suggest that some inducers may not necessarily be strongly associated with the calcified algal cell walls, and indicate that inducer sources in reef habitats may be more diverse than previously reported. Accepted: 21 May 1999     

Recruitment of the intertidal kelp Lessonia nigrescens Bory in northern Chile: successional constraints and opportunities

The strong 1982/83 El Niño event caused local extinction in populations of the low intertidal kelp Lessonia nigrescens Bory on northern Chilean coasts. The kelp has partially recolonized, but its recovery has apparently been low. This study documents the effect of biological factors that potentially decrease the velocity of kelp recolonization in northern Chile. A removal experiment showed that encrusting coralline algae, which dominate the lower intertidal, significantly reduce the recruitment of L. nigrescens in the presence of herbivores. Epithallium shedding by encrusting corallines is the most probable cause for this inhibition process. On the other hand, grazing on encrusting corallines by the chiton Enoplochiton niger (Barnes), a large-sized herbivore (10–20 cm length), could also affect kelp recruitment. The intestinal content of E. niger, where encrusting corallines are the main item (84.2%), revealed the presence of L. nigrescens. Among the natural substrata on which kelp recruits, measured at five different localities, the coarsely-branched alga Corallina officinalis (Decaisne) Kützing reached the highest frequency, despite its extremely low cover (< 1%) in the field. This suggests that settlement on turfs of C. officinalis allows the kelp to escape from herbivory, thus facilitating its recruitment. The roles of timing of kelp recruitment and seasonal grazing are also discussed.