Kelp and sea urchin settlement mediated by biotic interactions with benthic coralline algal species

Species interactions can influence key ecological processes that support community assembly and composition. For example, coralline algae encompass extensive diversity and may play a major role in regime shifts from kelp forests to urchin-dominated barrens through their role in inducing invertebrate larval metamorphosis and influencing kelp spore settlement. In a series of laboratory experiments, we tested the hypothesis that different coralline communities facilitate the maintenance of either ecosystem state by either promoting or inhibiting early recruitment of kelps or urchins. Coralline algae significantly increased red urchin metamorphosis compared with a control, while they had varying effects on kelp settlement. Urchin metamorphosis and density of juvenile canopy kelps did not differ significantly across coralline species abundant in both kelp forests and urchin barrens, suggesting that recruitment of urchin and canopy kelps does not depend on specific corallines. Non-calcified fleshy red algal crusts promoted the highest mean urchin metamorphosis percentage and showed some of the lowest canopy kelp settlement. In contrast, settlement of one subcanopy kelp species was reduced on crustose corallines, but elevated on articulated corallines, suggesting that articulated corallines, typically absent in urchin barrens, may need to recover before this subcanopy kelp could return. Coralline species differed in surface bacterial microbiome composition; however, urchin metamorphosis was not significantly different when microbiomes were removed with antibiotics. Our results clarify the role played by coralline algal species in kelp forest community assembly and could have important implications for kelp forest recovery.     

Seasonal changes in subarctic sea urchin populations from different habitats

This paper documents seasonal variation in certain sea urchin (Strongylocentrotus polyacanthus) characteristics in habitats of varying environmental conditions. At Shemya Island, Alaska, three habitat types [dense kelp beds, intermediate kelp beds, and algal barrens (low to no foliose algal cover)] were monitored seasonally from September 1995 to August 1996, for live and drift foliose algae. In general, drift algal abundance was greater in areas with more attached kelp, but this varied with season. Along with drift algae, sea urchin density, test size diameter, gonad and nutrition indices, and mobility were seasonally sampled within each habitat. Densities were highest in the algal barrens and lowest in the kelp beds. Seasonally, densities varied between summer/fall, and winter/spring, with lower numbers in the winter/spring. Test size was largest in the kelp habitats when compared to the intermediate or barren sites. Test size was seasonally consistent in the kelp habitats but not in the intermediate or barren sites. Here, test size did vary depending on season (larger urchins were found in winter). The gonad index showed much seasonal variation at the kelp and intermediate kelp sites, but was relatively more stable over time in the barren habitats. Between habitats, gonad and nutrition indices were larger in areas with kelp. Urchin movement varied seasonally between habitats, with more overall movement and variation in barren habitats. These results illustrate the importance of small-scale temporal and spatial variation. Monitoring for 1 year demonstrated that certain parameters varied more in areas of higher foliose algal cover (gonad indices), while other parameters varied more in low kelp areas (test size and movement). These results suggest that studies involving urchins should consider both time of year and overall algal community composition when conducting any type of experimental or monitoring work.     

The potential for seaweed resource development in subarctic Canada; Nunavik,Ungava Bay

Ungava Bay is ice covered 6–7 months of the year and evidence of ice scouring of seaweeds is extensive in the intertidal and shallow subtidal. Maximum tidal amplitudes of 16 m, among the highest in Canadian waters, compound this impact. Despite this level of annual perturbation, very extensive and dense beds of fucoids in the intertidal and laminarians in the subtidal are common on the western shores of Ungava Bay. Ground surveys of 24 intertidal stations combined with satellite images delineated 82,000 tons standing crop of Fucus vesiculosus and Fucus evanescens in Payne Bay, of which 36,000 tons were considered harvestable. Subtidally, kelp cover reached peak biomass at 5–10 m consisting of three primary species, Saccharina longicruris, Laminaria digitata, and Laminaria solidungula. In the area of Payne Bay, kelp beds of 100 ha were common, averaging 9–12 kg m⁻² wet weight. The productivity of brown algae at these latitudes has been assumed to be low relative to southern latitudes. Direct measurement of lineal growth indicates productivity is intermediate between arctic and temperate populations. The potential for medium level industrial harvest exists under conservative management strategies within the constraints of subarctic logistics.     

Increased macroalgal abundance following mass mortalities of sea urchins (Strongylocentrotus droebachiensis) along the Atlantic coast of Nova Scotia

Summary Recurrent outbreaks of disease between 1980 and 1983 caused catastrophic mortality of sea urchins (>260,000 t fresh weight) along 280 km (straight line distance) of the Atlantic coast of Nova Scotia. The complete elimination of sea urchins and concomitant development of fleshy macroalgal communities have occurred along different parts of this coast in different years. Macroalgal communities in areas where sea urchins died off 1, 3 and 4 years previously are compared to existing sea urchin-dominated barren grounds and to a mature kelp bed without sea urchins. Changes in macroalgal cover and species composition, and increases in biomass, density and size of kelp (Laminaria) species, characterize the succession from barren grounds to 3- and 4-year-old kelp beds. The greatest change occurred between one and three years following sea urchin mass mortality. Within 3 years, kelp beds attained a level of biomass (7.6 kg m^-2) comparable to that of mature beds. Recovery of sea urchin populations via recruitment of planktonic larvae has been slow and spatially variable. Large-scale reciprocal fluctuations in kelp and sea urchin biomass may characterize the trajectory of a dynamic system which cycles between two alternate community states: kelp beds and sea urchin-dominated barren grounds. Periodic decimation of sea urchin populations by disease may be an important mechanism underlying this cyclicity.     

Morphological,ultrastructural, and genetic characterization of coalescence in the intertidal and shallow subtidal kelps Lessonia spicata and L. berteroana (Laminariales,Heterokontophyta)

Coalescing macroalgae may fuse with conspecifics, forming genetically heterogeneous entities known as chimera. This process has been shown in taxa from roughly half the red algal orders and in the Codium species, a green alga. Field observations indicate that common and dominant kelps along central Chile exhibit a fused holdfast. We evaluated whether such fusions are true coalescence processes in Lessonia spicata and Lessonia berteroana. To this end, we characterized the ultrastructural event involved in holdfast fusion in the laboratory. Additionally, coalescence in natural populations was quantified by measuring the frequency of individuals with genetically heterogenic stipes within the same holdfast. Results indicate that coalescence appears as a frequent process in laboratory, mostly restricted to intraspecific fusions. During fusion, the meristodermatic cells located in the contact area modify their morphology and reduce the number of plastids, mitochondria, and cell inclusions. The cell wall becomes much thinner and develops plasmodesmata, enhancing communication with equivalent cells of the other coalescencing individual. Stipe genotyping indicates that there is a widespread occurrence of chimerism in both species and genetic heterogeneity is increasing directly with the increasing number of stipes. The combination of results suggests that kelp frequently coalesce in the field, and the histological response observed approaches that of red algae. Since kelps are part of the dominant vegetation in low intertidal and shallow subtidal beds, the adaptive values of coalescence in these species should be evaluated. It is concluded that coalescence and chimerism are evolutionary convergent processes, occurring in all three major groups of seaweeds.     

Climate‐driven disparities among ecological interactions threaten kelp forest persistence

The combination of ocean warming and acidification brings an uncertain future to kelp forests that occupy the warmest parts of their range. These forests are not only subject to the direct negative effects of ocean climate change, but also to a combination of unknown indirect effects associated with changing ecological landscapes. Here, we used mesocosm experiments to test the direct effects of ocean warming and acidification on kelp biomass and photosynthetic health, as well as climate‐driven disparities in indirect effects involving key consumers (urchins and rock lobsters) and competitors (algal turf). Elevated water temperature directly reduced kelp biomass, while their turf‐forming competitors expanded in response to ocean acidification and declining kelp canopy. Elevated temperatures also increased growth of urchins and, concurrently, the rate at which they thinned kelp canopy. Rock lobsters, which are renowned for keeping urchin populations in check, indirectly intensified negative pressures on kelp by reducing their consumption of urchins in response to elevated temperature. Overall, these results suggest that kelp forests situated towards the low‐latitude margins of their distribution will need to adapt to ocean warming in order to persist in the future. What is less certain is how such adaptation in kelps can occur in the face of intensifying consumptive (via ocean warming) and competitive (via ocean acidification) pressures that affect key ecological interactions associated with their persistence. If such indirect effects counter adaptation to changing climate, they may erode the stability of kelp forests and increase the probability of regime shifts from complex habitat‐forming species to more simple habitats dominated by algal turfs.     

Detached kelps from distant sources are a food subsidy for sea urchins

Trophic subsidies link habitats and can determine community structure in the subsidised habitats. Knowledge of the spatial extents of trophic interactions is important for understanding food webs, and for making spatial management practices more efficient. We demonstrate trophic linkages between detached (drift) fragments of the kelp Ecklonia radiata and the purple sea urchin Heliocidaris erythrogramma among discrete rocky reefs separated by kilometres. Sea urchins were abundant at one inshore reef, where the biomass of drift was usually high. There, sea urchins trapped detached kelp at high rates, although local kelp abundance was low. Most detached kelp present on the reef was retained by sea urchins. Detached seagrass, which was abundant on the reef, was not retained by sea urchins in large quantities. Experiments with tethered pieces of kelp showed that sea urchins only consumed detached fragments, and did not consume attached kelps. Comparisons of the morphology of detached fragments of kelp collected from the inshore reef to attached kelps from reefs further offshore showed that a large proportion (30-95%, varying among dates) of the fragments originated at distant reefs (>/=2 km away). At the inshore reef, the sea urchin H. erythrogramma is subsidised by detached kelps, and detached kelp fragments have been transported across landscapes. Cross-habitat resource subsidies therefore link discrete reef habitats separated by kilometres of non-reef habitat.     

156 Effects of Light and Nutrients on Periphyton Community Composition in a Hawaiian Stream

The commercial harvest of floating kelps has promise as a business opportunity that could be especially valuable for small communities in southeast Alaska. The giant kelp, Macrocystis , is harvested mainly for the commercial herring roe-on-kelp harvest in Alaska. In addition, in southeast Alaska the two other species of floating kelps, Nereocystis luetkeana and Alaria fistulosa , have been commercially harvested since 1992 for use as agrochemicals by the Alaska KelpCo. The continued existence of this company and others that may wish to harvest seaweeds on a commercial basis is threatened by the lack of a kelp harvest management system in Alaska. The objective of this research is to develop a reliable and cost effective method for estimating the aerial extent and the standing crop biomass of the floating kelps in SE Alaska. We are employing an aerial multispectral imaging system that can be flown at varying altitudes to achieve spatial resolutions ranging from 0.5 to 2 meters. We are investigating whether the three species have unique spectral or textural characteristics in the multispectral imagery that can be used to differentiate the populations. The end products for this project will be kelp resource maps that delineate the spatial extent and biomass of kelp populations within the beds and a methodology for assessing kelp beds with mixed species composition. These products will find use by resource managers such as the Alaska Department of Fish and Game and the Department of Natural Resources.     

155 Mapping Floating Kelp Beds in Alaska With Remote Sensing

The commercial harvest of floating kelps has promise as a business opportunity that could be especially valuable for small communities in southeast Alaska. The giant kelp, Macrocystis, is harvested mainly for the commercial herring roe‐on‐kelp harvest in Alaska. In addition, in southeast Alaska the two other species of floating kelps, Nereocystis luetkeana and Alaria fistulosa, have been commercially harvested since 1992 for use as agrochemicals by the Alaska KelpCo. The continued existence of this company and others that may wish to harvest seaweeds on a commercial basis is threatened by the lack of a kelp harvest management system in Alaska. The objective of this research is to develop a reliable and cost effective method for estimating the aerial extent and the standing crop biomass of the floating kelps in SE Alaska. We are employing an aerial multispectral imaging system that can be flown at varying altitudes to achieve spatial resolutions ranging from 0.5 to 2 meters. We are investigating whether the three species have unique spectral or textural characteristics in the multispectral imagery that can be used to differentiate the populations. The end products for this project will be kelp resource maps that delineate the spatial extent and biomass of kelp populations within the beds and a methodology for assessing kelp beds with mixed species composition. These products will find use by resource managers such as the Alaska Department of Fish and Game and the Department of Natural Resources.     

Overgrazing of Kelp Beds Along the Coast of Norway

The aim of this study was to better understand the down-grazing of kelp beds by sea urchins (Strongylocentrotus droebachiensis) along the coast of Norway. Barren grounds were first observed in sheltered areas along the coast of the counties of Trø ndelag, Nordland and Troms in 1974. In the 1980s, the barren grounds spread to areas more heavily exposed to waves. In the 1990s, the kelp beds were re-established in some localities in southern Trø ndelag, initially in wave-exposed areas. In the northernmost parts of Norway, i.e. the counties of Troms and Finnmark, the barren ground areas may still be increasing. Crabs (Cancer pagurus) and common eiders are the most common predators on urchins. Predation on sea urchins in kelp beds is probably not among the factors that limit the sea urchin populations. Along the coast of Nordland and further north, sea urchins are infected by nematodes, resulting in a low, but significant increase in their mortality. No re-growth of kelp beds has been found in the most infected areas. In the late 1960s and the early 1970s, a high occurrence of echinoderm larvae was observed in deeper waters. This was a period with cold water, which may have caused high recruitment of sea urchins. The bet-hedging life strategy of sea urchins may account for the sudden increase in the size of the populations. In the present paper I propose the hypothesis that higher individual growth rates and higher mortality rates in the south than in the north may explain the decrease in the populations, which may in turn account for the re-growth of kelp in the southern areas.     

Latitudinal variation in intertidal algal community structure: the influence of grazing and vegetative propagation

Summary The hypothesis that sea urchin grazing and interactions with turf-forming red algae prevent large brown algae from forming an extensive canopy in the low intertidal zone of southern California was tested with field experiments at two study sites. Experimental removal of sea urchins resulted in rapid algal recruitment. Crustose coralline algae which typically dominate the substratum in areas with dense urchin populations were quickly overgrown by several species of short-lived green, brown and red algae. The removal of urchins also significantly increased the recruitment of two long-lived species of large brown algae (Egregia laevigata and Cystoseira osmundacea at one study site and E. laevigata and Halidrys dioica at the other). The experimental plots at both sites were eventually dominated by perennial red algae.A two-factorial experiment demonstrated that sea urchin grazing and preemption of space by red algae in areas where urchins are less abundant are responsible for the rarity of large brown algae in the low intertidal of southern California. The three dominant perennial red algae, Gigartina canaliculata, Laurencia pacifica and Gastroclonium coulteri, recruit seasonally from settled spores but can rapidly fill open space with vigorous vegetative growth throughout the year. These species encroach laterally into space created by the deaths of large brown algae or by other disturbances. Once extensive turfs of these red algae are established further invasion is inhibited. This interaction of algae which proliferate vegetatively with algae which recruit only from settled spores is analogous to those which occur between solitary and colonial marine invertebrates and between solitary and cloning terrestrial plants.It is suggested that a north-south gradient in the abundance of vegetatively propagating species, in grazing intensity and in the frequency of space-clearing disturbances, may account for latitudinal variation in intertidal algal community structure along the Pacific coast of North America.     

Biological interactions maintain the boundaries between kelp forests and urchin barrens in the Aleutian Archipelago

This study used benthic surveys and manipulative experiments to examine (1) if boundaries between kelp forests and urchin barrens exist at multiple locations spanning the Aleutian Archipelago, (2) if these boundaries are spatially stable, and (3) how changes in algal density within the kelp forests influence the ability of urchins to invade them. Our results demonstrate that sharply punctuated kelp forest-urchin barren boundaries occur throughout the Archipelago, and they are spatially stable for at least 2 years. Further, when all macroalgae were experimentally removed from the kelp forest side of the boundaries, urchins rapidly moved into these clearings and excluded macroalgae for up to 2 years. However, these movements were not observed where 75% or less of the macroalgae was removed (leaving 25% or more in place), suggesting that even low macroalgal abundances can prevent urchins from invading the kelp forests. Further, urchin densities were negatively related to kelp density, again indicating that kelp can reduce urchin densities. While the ability of urchins to overgraze kelp forests is widely known, our results indicate that kelp can inhibit urchins, that these inhibitory influences are a widely recurrent phenomenon, and that this interaction is important to maintaining kelp forests across the Aleutians.     

Effects of removing sea urchins (Strongylocentrotus droebachiensis): Stability of the barren state and succession of kelp forest recovery in the east Atlantic

Stability properties of the barren state of a kelp forest-sea urchin system were studied in northern Norway. The ability of the sea urchin Strongylocentrotus droebachiensis to maintain high population densities and recover from perturbations, and the succession of kelp forest revegetation, were studied experimentally by reducing the sea urchin density on a barren skerry. Additional information was obtained from community changes following a natural, but patchy, sea urchin mortality that varied between sites. On the barren grounds, high sea urchin densities (30 50 per m²) is maintained by annual recruitment. Severe reductions of sea urchin densities initiated luxuriant kelp growth, while more moderate reductions allowed establishment of opportunistic algae (during spring and early summer), but no kelps. Succession of algal growth, after the severe decline in sea urchin densities, followed a predictable pattern. At first the substrate was colonized by filamentous algae, but within few weeks they were outcompeted by the fast growing kelp Laminaria saccharina. After 3–4 years of the removal experiment, the slower-growing, long-lived kelp L. hyperborea became increasingly dominant. Increased food availability after reduction in sea urchin density led to increased individual growth of the remaining sea urchins. However, the population density did not increase, neither from recruitment nor immigration from adjacent areas with high sea urchin densities. Possibly, early establishment of a dense kelp stand, may represent a breakpoint in the ability of sea urchins to reestablish a barren state. The ability of L. saccharina quickly to invade and monopolize an area may have both positive and negative effects on the succession towards the climax L. hyperborea kelp forest. Competitive interactions may slow the process, but development of a dense stand of L. saccharina will also reduce grazing risk on scattered recruits of the more slowly growing L. hyperborea.     

A COMPARISON OF DRAGON KELP,EUALARIA FISTULOSA, (PHAEOPHYCEAE) FECUNDITY IN URCHIN BARRENS AND NEARBY KELP BEDS THROUGHOUT THE ALEUTIAN ARCHIPELAGO1

The Aleutian Archipelago coastal ecosystem has undergone a dramatic change in community composition during the past two decades. Following the removal of ～99% of the sea otters, Enhydra lutris, from the ecosystem, changes to the benthic communities resulted in widespread losses to most of the region’s kelp beds and corresponding increases in the prevalence of urchin barrens. Within the urchin barrens, the few kelps that have remained are exposed to elevated light, nutrients and currents, all of which may enhance their physiological condition and thus result in greater fecundity. To explore this further, we examined patterns of sporophyte fecundity in the dominant canopy‐forming kelp, Eualaria fistulosa, in both urchin barrens and in nearby kelp beds at seven Aleutian Islands spanning a range of 800 km. We found that the average weight of E. fistulosa sporophyll bundles was significantly greater on sporophytes occurring in the urchin barrens than in the nearby kelp beds. Furthermore, the average number of zoospores released per cm² of sporophyll area was also significantly greater in individuals from the urchin barrens than the nearby kelp beds. When these two metrics were combined, our results suggest that individual E. fistulosa sporophytes occurring in the urchin barrens may produce as many as three times more zoospores than individual E. fistulosa sporophytes occurring in the nearby kelp beds, and thus they may contribute disproportionately to the following year’s sporophyte recruitment in both urchin barrens and the adjacent kelp beds.     

Organization of macroalgal assemblages in the Northeast Pacific: the assumption of homogeneity and the illusion of generality

Evidence for the geographic generality of the causes of intertidal zonation and the indirect effects of a keystone predator, the sea otter, on subtidal kelp assemblages was examined. Most research on intertidal algal assemblages has been done at a few protected sites where zonation is distinct. Surveys of wave-exposed intertidal sites in central and northern California show that assemblage structure is highly variable. This indicates that our present understanding of assemblage organization, including the effects of mussel-algal interactions, may not be widely applicable. Surveys of kelp forest habitat along the entire coast of California suggest that deforestation by sea urchins is uncommon in the absence of sea otters. These examples indicate that the generality of commonly accepted causes of algal assemblage structure in the Northeast Pacific may be an illusion based on assumptions of environmental homogeneity.     

The influence of herbivores on primary producers can vary spatially and interact with disturbance

Identifying the major drivers of ecosystem change remains a central area of ecological research. Although top–down drivers of change have received particular focus, we still have little understanding of how consistently these factors control an ecosystem's shift in both directions, between different ecosystem states. Using a crossed experiment in a shallow embayment in southeastern Australia, we investigated the roles of disturbance (kelp removal) and sea urchin herbivory (via increased density) to determine their contributions to shifts away from a kelp‐dominated community. In a second experiment, done in urchin barren areas at two sites, we tested whether reductions in ambient sea urchin densities allowed an algal shift in the reverse direction. In both experiments, we observed that high densities of sea urchins could negatively influence kelp and macroalgal abundance. However, in the kelp bed, a moderate or severe disturbance resulted in a comparable algal response, irrespective of urchin density. The influence of sea urchins also varied dramatically between the two urchin barren sites. Here, reducing urchin densities resulted in algal recovery at one site, but at the other site, substantial colonisation of barren areas by canopy‐forming brown algae and Ulvales occurred across all (low, medium, and high) urchin density treatments. Our findings illustrate multiple pathways of urchin barren creation and algal recovery, and reveal that shifts both to and from an urchin barren state can occur irrespective of herbivore pressure. These alternate pathways can operate over short spatial distances or with different regimes of disturbance.     

Age-dependent zonation of the periwinkle Littorina littorea (L.) in the Wadden Sea

On sedimentary tidal flats near the island of Sylt (German Bight, North Sea) abundance and size distribution of periwinkles, Littorina littorea L., were studied in low intertidal and in shallow and deep subtidal mussel beds (Mytilus edulis L.). In low intertidal mussel beds, surveys revealed that high densities (1,369±571 m^–2) of juvenile snails (≤13 mm) were positively correlated with strong barnacle epigrowth (Semibalanus balanoides L. and Balanus crenatus Bruguière) on mussels. A subsequent field experiment showed that recruitment of L. littorea was restricted to the intertidal zone. Abundances of periwinkles (213±114 m^–2) and barnacles abruptly decreased in the adjacent shallow subtidal zone, which served as a habitat for older snails (>13 mm). L. littorea was completely absent from disjunct deep (5 m) subtidal mussel beds. Snail abundance varied seasonally with maxima of >4,000 m^–2 in low intertidal mussel beds in October and minima in July, just before the onset of new recruitment. I suggest that the presence of cracks and crevices among the dense barnacle overgrowth in intertidal mussel beds favoured recruitment and survival of juvenile snails. Larger (older) specimens are assumed to actively migrate to the less favourable adjacent subtidal. Therefore, intertidal mussel beds are considered as nurseries for the population of L. littorea in the Wadden Sea. Received in revised form: 25 September 2000 Electronic Publication     

Physical disturbance and community organization on a subtidal cobble bed

This study examines the role of physical disturbance in regulating the development and maintenance of macroalgal community structure in a subtidal cobble bed on the Atlantic coast of Nova Scotia, Canada, after a mass mortality of sea urchins. Displacement by hydrodynamic forces of cobbles with and without attached macroalgae was measured in mark-recovery experiments. Cobbles without macroalgae experienced low levels of overturning or displacement by waves. In contrast, cobbles with large fleshy algae (such as kelps and fucoids) were displaced by drag forces along the seabed during periods of moderately strong wave action. Displacement of these “drifters” occurred once the attached alga reached a critical mass (relative to the weight of the anchoring cobble), which varied among species of differing morphology. The rate of displacement of drifters was an exponential function of the ratio of attached algal biomass to rock weight. The loss of larger erect forms from the cobble bed results in a patchwork of low-lying turfs. Clusters of small boulders, experimentally transplanted to the cobble bed, accumulated a greater biomass of large fleshy algae than surrounding cobbles after one year, likely because of the greater stability of boulders. Algal-mediated hydrodynamic displacement of cobbles appears to be an important form of physical disturbance in the cobble habitat, restricting the types and sizes of algae that can grow on cobbles following an urchin die-off.     

Alien marine fishes deplete algal biomass in the Eastern Mediterranean

One of the most degraded states of the Mediterranean rocky infralittoral ecosystem is a barren composed solely of bare rock and patches of crustose coralline algae. Barrens are typically created by the grazing action of large sea urchin populations. In 2008 we observed extensive areas almost devoid of erect algae, where sea urchins were rare, on the Mediterranean coast of Turkey. To determine the origin of those urchin-less 'barrens', we conducted a fish exclusion experiment. We found that, in the absence of fish grazing, a well-developed algal assemblage grew within three months. Underwater fish censuses and observations suggest that two alien herbivorous fish from the Red Sea (Siganus luridus and S. rivulatus) are responsible for the creation and maintenance of these benthic communities with extremely low biomass. The shift from well-developed native algal assemblages to 'barrens' implies a dramatic decline in biogenic habitat complexity, biodiversity and biomass. A targeted Siganus fishery could help restore the macroalgal beds of the rocky infralittoral on the Turkish coast.     

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.