Herbivory and patch dynamics on rocky reefs in temperate Australasia: The roles of fish and sea urchins

Sea urchins are widely considered to be the major grazers in temperate subtidal systems, with herbivorous fish being browsers of minor importance. This paper reviews spatial and temporal patterns in these herbivores on rocky reels in temperate Australasia, with the aim of assessing their relative impacts on patch structure and dynamics. Herbivorous fishes are widespread and make up a significant numerical component the reel fish fauna. Sea urchins are also abundant, but not all geographic locations support actively grazing species. Both fish and sea urchins exhibit distinct patterns of distribution among depth strata. Within depth strata, all herbivores are restricted to (sea urchins) or forage preferentially in (fish) particular habitat patches, causing a mosaic of different feeding activities. These patches are either related to specific features of the habitat (e.g. Kelp patches, topography) or behavioural interactions. Foraging by sea urchins and demersal-nesting damselfishes is intense and persistent, whereas in the kelp-feeding fish Odax cyanomelas, foraging reaches greatest intensity at predictable locations during a few months of every year. Many fish and sea urchins consume some algae in preference to others. However, feeding preferences may determine the nature of the impact only in fishes. For sea urchins, preference may occasionally determine the order in which algae are consumed, but at high densities they consume all available macroalgae. Impacts of both types of herbivore on the abundance of algae have been recorded. Some sea urchins (e.g. Evechinus chloroticus, Centrostephanus rodgersii) appear to severely modify biogenic habitat structure by maintaining ‘barrens’ (areas devoid of macroalgae) over long periods. In contrast to this, the effects of fishes may be more transitory (e.g. seasonal impact of Odax cyanomelas on brown algae) or occur at smaller spatial scales (e.g. nest sites maintained by male Parma victoriae) Herbivorous and other fishes appear to respond to spatial patterns in algal distributions, rallier than having it major impact upon them. The relative effects of fish and sea urchins on the long-term dynamics of kelp forests are unknown, hut temporal patterns in herbivore abundance and behaviour, and algal demography arc urgent targets for research.     

Do native herbivores provide resistance to Mediterranean marine bioinvasions? A seaweed example

Generalist herbivores in marine ecosystems are poorly examined for their potential to serve as a source of biotic resistance against algal invasion. We assessed how one of the main generalist herbivores in Mediterranean rocky reefs (the sea urchin Paracentrotus lividus) affects Lophocladia lallemandii and Caulerpa racemosa, two algal invaders with strong detrimental effects on native benthic communities. In a comparison of sea urchin gut contents to algal community composition, strong preferences were exhibited, leading to no relationship between consumption and availability. Both C. racemosa and L. lallemandi were abundant in algal assemblages (>60% occurrence), but C. racemosa (20% of diet) was consumed more than L. lallemandi (3.5%). Experimental enclosures of sea urchins (12 sea urchins * m⁻²) were carried out in locations where L. lallemandii was already established and C. racemosa was rare (new invasion) or abundant (established invasion). C. racemosa was negatively affected by sea urchins only when it was rare, and no effect was detected when the alga was already abundant. Results for L. lallemandi were exactly opposite: urchins limited seasonal increases in L. lallemandi in highly-invaded areas. Because of the small amount of direct consumption of L. lallemandi, its decrease in abundance may be related to the grazing of native algae where L. lallemandii is attached. Overall, our results show that high densities of native herbivores may reduce invasive algae at low densities, due to a combination of direct and indirect effects, but it has no significant effect in highly-invaded areas.     

Patterns of distribution and movement of fishes, <Emphasis Type="Italic">Ophthalmolepis lineolatus</Emphasis> and <Emphasis Type="Italic">Hypoplectrodes maccullochi</Emphasis>, on temperate rocky reefs of south eastern Australia

Current ecological models predict that reef fish assemblages will be strongly influenced by habitat type. Here we test hypotheses about habitat types and abundance patterns of temperate reef fishes from broad spatial scales (100 s of km) to small spatial scales of metres to tens of metres. Habitat preferences are also described over long periods of time (22 years) for two abundant taxa. Patterns of distribution and abundance varied over ～ eight degrees of latitude (29.9–37.5°S) along the coast of New South Wales, Australia. Ophthalmolepis lineolatus (Labridae) preferred kelp and Barrens habitats and juveniles were most abundant in habitats rich in algae. This species also increased in abundance from North to South. In contrast, Hypoplectrodes maccullochi (Serranidae) were usually only found in the Barrens habitat and great variation was found among locations. Both taxa were most abundant on urchin grazed deep reefs (over 10 m deep). Habitat preferences of O. lineolatus and H. maccullochi appeared resistant to major environmental perturbations that included large El Niño events in 1991, 1998 and 2002. Home ranges of O. lineolatus varied from 52 m² to 1,660 m² and often overlapped; fish of all sizes were most abundant in algal dominated habitat. Limited movements and small home ranges (2.1–11.6 m²) combined with a strong affiliation for shelter indicated that most H. maccullochi are strongly site-attached. Habitat type is important to these taxonomically different fishes, but to varying degrees where H. maccullochi was more of a habitat specialist than O. lineolatus and would be more vulnerable to perturbations that alter Barrens. Changes in reef habitats will have a great influence on fish assemblages and this should also be considered in coastal planning (e.g. for Marine Protected Areas, MPAs) and the assessments of resistance and resilience of fishes to climate change.     

Predation and the Control of the Sea Urchin Echinometra viridisand Fleshy Algae in the Patch Reefs of Glovers Reef, Belize

The massive reduction in sea urchin Diadema antillarum populations since the mid-1980s has been associated with large increases in the abundance of fleshy algae on many Caribbean reefs despite the availability of other sea urchin and finfish grazers. This study examined the ecology of a grazer living sympatrically with D. antillarum, the common and abundant sea urchin Echinometra viridis. I examined the role that finfish and invertebrate predators play in controlling the distribution of E. viridis as well as the ability of this sea urchin to control exposed fleshy algae on the patch reefs of the Glovers Reef Atoll lagoon. I found that the major predators of this sea urchin were Calamus bajonado (jolthead porgy), Balistes vetula and Canthidermis sufflamen (queen and ocean triggerfish), Lachnolaimus maximus (hogfish), and a gastropod, probably Cassis madagascariensis. The abundance of E. viridis is constrained by predation, which restricts E. viridis to cryptic locations, such as crevices. Sea urchins bit a smaller percentage of experimental algal assays than finfish. Finfish herbivory was associated positively with patch reef topographic complexity. Unexpectedly, E. viridis abundance was positively correlated with fleshy algal abundance, but negatively correlated with the frequency of finfish bites. Predators restrict E. viridis to crevices and therefore reduce their influence on exposed fleshy algae, even at moderately high population densities (up to 10 per square meter). Since net benthic primary production of coral reefs is most strongly associated with herbivory on exposed surfaces, it would appear that E. viridis is unable to maintain the same production as reefs dominated by D. antillarum. Received 5 November 1998; accepted 2 June 1999.     

Spatial dynamics of benthic competition on coral reefs

The community structure of sedentary organisms is largely controlled by the outcome of direct competition for space. Understanding factors defining competitive outcomes among neighbors is thus critical for predicting large-scale changes, such as transitions to alternate states within coral reefs. Using a spatially explicit model, we explored the importance of variation in two spatial properties in benthic dynamics on coral reefs: (1) patterns of herbivory are spatially distinct between fishes and sea urchins and (2) there is wide variation in the areal extent into which different coral species can expand. We reveal that the size-specific, competitive asymmetry of corals versus fleshy algae highlights the significance of spatial patterning of herbivory and of coral growth. Spatial dynamics that alter the demographic importance of coral recruitment and maturation have profound effects on the emergent structure of the reef benthic community. Spatially constrained herbivory (as by sea urchins) is more effective than spatially unconstrained herbivory (as by many fish) at opening space for the time needed for corals to settle and to recruit to the adult population. Further, spatially unconstrained coral growth (as by many branching coral species) reduces the number of recruitment events needed to fill a habitat with coral relative to more spatially constrained growth (as by many massive species). Our model predicts that widespread mortality of branching corals (e.g., Acropora spp) and herbivorous sea urchins (particularly Diadema antillarum) in the Caribbean has greatly reduced the potential for restoration across the region.     

Consumers of sea urchins,<Emphasis Type="Italic"> Paracentrotus lividus</Emphasis> and<Emphasis Type="Italic"> Arbacia lixula</Emphasis>, in shallow Mediterranean rocky reefs

Underwater observations on fish and asteroid consumers (i.e. predators and scavengers) of sea urchins, Paracentrotus lividus and Arbacia lixula, were carried out at several locations in shallow Mediterranean rocky reefs. Observations conducted in the marine reserve of Torre Guaceto (Adriatic Sea) revealed that sparid fishes, Diplodus sargus and D. vulgaris, are the main fish predators of small (<1 cm in test diameter) and medium (1–4 cm) sea urchins, whereas the labrids Coris julis and Thalassoma pavo preyed only upon small sea urchins. Large D. sargus were able to prey upon small and medium, and occasionally large (>4 cm) sea urchins, whereas medium and small Diplodus preyed mainly upon small sea urchins. The number of sea urchins preyed upon by fishes was negatively related to sea urchin size for both species. P. lividus appeared to be subject to higher predation levels than A. lixula. The scavenger guild comprised 11 fish species, with D. sargus, D. vulgaris, Coris julis and Chromis chromis accounting for about 80% of scavenger fishes. Observations performed at several locations in the Mediterranean on the predatory asteroid Marthasterias glacialis revealed that only 3% of the detected individuals were preying upon sea urchins. Due to the importance of sea urchins for assemblage structure and functioning of Mediterranean rocky reef ecosystems, these results may have also important implications for management of fishing activities.Communicated by H.-D. Franke     

Grazing intensity influences the strength of an associational refuge on temperate reefs

Recent studies have emphasized the role of positive interactions in ecological communities, but few have addressed how positive interactions are mediated by abiotic stress and biotic interactions. Here, I investigate the effect of a facilitator species on the abundance of macroalgae over a gradient of herbivory. Grazing by sea urchins can be intense on temperate reefs along the California coast, with benthic macroalgae growing exclusively in physical refuges and interspersed within colonies of the strawberry anemone, Corynactis californica. Field experiments indicated that the net effect of C. californica on turf algae was strongly nonlinear over a gradient in density of sea urchins. At low intensities of urchin grazing, the anemone and macroalgae competed for space, with algae capable of overgrowing C. californica. At intermediate grazing intensities, C. californica provided a refuge for turf algae but not for juvenile kelp. Neither turf algae nor kelp benefited from the presence of C. californica at the highest levels of grazing intensity, as sea urchins consumed nearly all macroalgae. The hump-shaped effect observed for C. californica contrasts with the prevailing view in ecological theory that positive interactions are more common in harsh environmental conditions. The results reported here qualify this view and underscore the need to evaluate positive interactions over a range of abiotic stress and consumer pressure.     

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.     

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.     

Trophic Cascades Induced by Lobster Fishing Are Not Ubiquitous in Southern California Kelp Forests

Fishing can trigger trophic cascades that alter community structure and dynamics and thus modify ecosystem attributes. We combined ecological data of sea urchin and macroalgal abundance with fishery data of spiny lobster (Panulirus interruptus) landings to evaluate whether: (1) patterns in the abundance and biomass among lobster (predator), sea urchins (grazer), and macroalgae (primary producer) in giant kelp forest communities indicated the presence of top-down control on urchins and macroalgae, and (2) lobster fishing triggers a trophic cascade leading to increased sea urchin densities and decreased macroalgal biomass. Eight years of data from eight rocky subtidal reefs known to support giant kelp forests near Santa Barbara, CA, USA, were analyzed in three-tiered least-squares regression models to evaluate the relationships between: (1) lobster abundance and sea urchin density, and (2) sea urchin density and macroalgal biomass. The models included reef physical structure and water depth. Results revealed a trend towards decreasing urchin density with increasing lobster abundance but little evidence that urchins control the biomass of macroalgae. Urchin density was highly correlated with habitat structure, although not water depth. To evaluate whether fishing triggered a trophic cascade we pooled data across all treatments to examine the extent to which sea urchin density and macroalgal biomass were related to the intensity of lobster fishing (as indicated by the density of traps pulled). We found that, with one exception, sea urchins remained more abundant at heavily fished sites, supporting the idea that fishing for lobsters releases top-down control on urchin grazers. Macroalgal biomass, however, was positively correlated with lobster fishing intensity, which contradicts the trophic cascade model. Collectively, our results suggest that factors other than urchin grazing play a major role in controlling macroalgal biomass in southern California kelp forests, and that lobster fishing does not always catalyze a top-down trophic cascade.     

Is there a link between the type of habitat and the patterns of abundance of holothurians in shallow rocky reefs?

The presence of a mosaic of habitats, largely determined by sea urchin grazing, across shallow rocky reefs may potentially influence in differences in the distribution patterns of invertebrates. The aim of this paper was to assess, using a correlative approach, whether the type of habitat influences the abundance patterns of holothurians in the eastern Atlantic. We hypothesized that abundances of large (> 10 cm) holothurians varied among four types of habitat (3 vegetated habitats with low abundances of the sea urchin D. antillarum vs. ‘barrens’ with hyperabundances of sea urchins), and that these differences were consistent at a hierarchy of spatial scales, including two islands and several replicated sites within each type of habitat and island. Three species of large holothurians were found, accounting for a total of 300 specimens. We found remarkable differences in abundances of holothurians between the ‘barrens’ and the three vegetated habitats. This pattern was strongest for the numerically dominant species, Holothuria sanctorii. Total abundances of holothurians were between 5 and 46 times more abundant in ‘barrens’ compared with the vegetated habitats. Inter-habitat differences were species-specific with some inconsistent patterns from one island to the other. The total abundances of holothurians tended to increase with the abundance of sea urchins within ‘barrens’. Our study suggests that there may be a link, at least for the dominant species Holothuria sanctorii, between the distribution and abundances of large holothurians and the habitat across shallow-waters of the eastern Atlantic.     

Distributions and habitat associations of the bridled monocle bream Scolopsis bilineatus (Nemipteridae): a demographic approach

This study focussed on the demography and ecology of Scolopsis bilineatus at three locations on the Great Barrier Reef: the Lizard Island Group, Orpheus Island and One Tree Island. Scolopsis bilineatus lived for up to 16 years and had four distinct life‐history stages, which varied in their distribution patterns, habitat use and reproductive behaviour. Pre‐maturational sex change occurred whereby all males were derived from immature females, and males grew faster and larger than females. Small females and larger males generally formed pairs, which influenced their spatial distributions at small scales. Distributions of S. bilineatus were influenced by depth and exposure within reefs, particularly for juveniles, and most fish were found in shallow, sheltered habitats. Abundance was influenced by benthic cover, and was higher in areas of high coral cover and low where algae were abundant. Habitat associations were stronger at the microhabitat scale, and shelter sites were important for adults. Ontogenetic changes in microhabitat associations were found: juveniles occupied sand and rubble, and adults occupied shelters such as caves and overhangs. Adults showed site fidelity for shelter sites over a period of 4 days and returned to specific shelter sites repeatedly. These findings illustrate the importance of understanding the spatial ecology and habitat use of coral reef fishes, particularly with reference to size‐based changes within species.     

Echinoderms display morphological and behavioural phenotypic plasticity in response to their trophic environment

The trophic interactions of sea urchins are known to be the agents of phase shifts in benthic marine habitats such as tropical and temperate reefs. In temperate reefs, the grazing activity of sea urchins has been responsible for the destruction of kelp forests and the formation of 'urchin barrens', a rocky habitat dominated by crustose algae and encrusting invertebrates. Once formed, these urchin barrens can persist for decades. Trophic plasticity in the sea urchin may contribute to the stability and resilience of this alternate stable state by increasing diet breadth in sea urchins. This plasticity promotes ecological connectivity and weakens species interactions and so increases ecosystem stability. We test the hypothesis that sea urchins exhibit trophic plasticity using an approach that controls for other typically confounding environmental and genetic factors. To do this, we exposed a genetically homogenous population of sea urchins to two very different trophic environments over a period of two years. The sea urchins exhibited a wide degree of phenotypic trophic plasticity when exposed to contrasting trophic environments. The two populations developed differences in their gross morphology and the test microstructure. In addition, when challenged with unfamiliar prey, the response of each group was different. We show that sea urchins exhibit significant morphological and behavioural phenotypic plasticity independent of their environment or their nutritional status.     

Escapes from herbivory in relation to the structure of mangrove island macroalgal communities

Summary Some shallow habitats that surround mangrove islands exhibit abruptly discontinuous macrophyte boundaries; in other regions, plant distributional patterns are less defined. Where distinct boundaries do occur, fleshy algae predominate on the roots of the red mangrove, Rhizophora mangle, which do not contact the bottom sediments (hanging roots), while calcifying algae dominate on the substratum-penetrating roots and banks (=embedded-root habitat) surrounding the mangrove thickets. Considerable natural-history and floristic information reveals that the fleshy hanging-root species are not specialists, for that type of habitat. Experimental transplants showed that on banks and embedded roots where there typically are abundant macroherbivores (particularly sea urchins), most fleshy algae are eliminated.The dominants of the hanging-root habitat (e.g, Acanthophora spicifera, Spyridia filamentosa, Caulerpa racemosa var. Occidentalis) are 6–20 times more susceptible to herbivores than the dominants of the embedded-root habitat (e.g., Halimeda opuntia f. triloba, H. monile). Consequently, we suggest the former are relegated to the spatial refugia from herbivores (=non-coexistence escapes) provided by the hanging roots. Factors associated with these palatability differences include higher average calorific values (6.5 times) of the fleshy hanging-root dominants, greater proportions of organic content (2.6 times) and the general absence of calcification. The dominants of the embedded-root habitat show reduced edibility as a probable consequence of low calorific values, heavy calcification and potential herbivore-detering secondary metabolites. Correlative evidence and preliminary experimental results tentatively indicate that, in the absence of macroherbivores, the hanging-root dominants, which exhibit production rates 4.7 times greater than the dominants of the embedded-root habitat, are better competitors for space.We suggest that variations in herbivory are responsible, in part, for maintaining greater algal diversity in mangrove systems. At a study site with abundant sea urchins, five algal species were found only in the embedded-root habitat three species were confined to the hanging roots, while three others occurred in both. At an urchin-free site, no macrophytes were found only on embedded-root substrata, while one (in trace amounts) was found only on hanging-root habitat and eight occurred in both. We predict that in the absence of herbivores, the species assemblage characteristic of the hanging-roots would exclude many of the dominants from the embedded-root habitat.     

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.     

Global patterns in the effects of predator declines on sea urchins

Latitudinal gradients in the strength of biotic interactions have long been proposed, but empirical evidence for the expectation of more intense predation, herbivory and competition at low latitudes has been mixed. Here, we use a meta‐analysis to test the prediction that predation pressure on sea urchins, a group of consumers with a particularly strong influence on community structure in the world's oceans, is strongest in the tropics. We then examine which biotic and abiotic factors best correlate with biogeographic and within habitat patterns in sea urchin responses to predation. Consistent with expectations, predator impacts on sea urchins were highest in tropical coral reefs and decreased towards the poles in rocky reef habitats (> 25° absolute latitude). However, latitude and temperature were weakly correlated with effect sizes, and the strongest predictor of predator impacts was sea urchin species. This suggests an important role of prey identity (i.e. traits including behaviour, physical, and chemical defences) rather than large scale abiotic factors in determining variation in interaction strengths. Ecosystem‐shaping sea urchins such as Tripneustes gratilla, Diadema savignyi and Centrostephanus rodgersii were strongly impacted by consumers, indicating a tight coupling between predators of these species and their boom and bust prey. Anthropogenic activities such as over‐fishing, climate change and habitat destruction are causing rapid environmental change, and understanding how predation pressure varies with temperature, across habitats and among prey species, will aid in predicting the likelihood of ecosystem wide effects (via trophic cascades).     

Habitat selection and juvenile persistence control the distribution of two closely related Caribbean damselfishes

Summary On many Caribbean fringing coral reefs, two closely related and ecologically similar damselfishes, the beaugregory (Stegastes leucostictus Müller and Troschel) and the cocoa damselfish (S. variabilis Castelnau), occupy nonoverlapping vertical distributions. In St. Croix (USVI), beaugregory are very abundant in shallow water back reef habitats (1–2 m depth) while cocoa damselfish are restricted to the base of the forereef (10–15 m depth).In this study, the roles of habitat selection at settlement and juvenile persistence were investigated to determine their influence on this pattern of zonation. Settlement events observed at intervals over a two-year period revealed that habitat selection occurred at settlement and was confined to habitats occupied by adults. In addition, differences in juvenile persistence (due to mortality and/or emigration) were found when species were translocated between depths. Over a period of 100-days, juvenile beaugregory moved from 1 m to 12 m depth suffered four-fold greater losses at the deeper sites than shallow water controls, while translocated cocoa damselfish suffered twice as many losses in shallow water than controls at 12 m depth. Despite these differences in persistence, growth rates of the two species were similar and independent of depth. These results indicate that preferential habitat selection at settlement, perhaps an evolutionary response to differential juvenile mortality, may play a deciding role in determining distributions of ecologically similar species of coral reef fishes.     

Predation of the sea urchin Heliocidaris erythrogramma by rock lobsters (Jasus edwardsii) in no-take marine reserves

The formation of sea urchin ‘barrens’ on shallow temperate rocky reefs is well documented. However there has been much conjecture about the underlying mechanisms leading to sea urchin barrens, and relatively little experimentation to test these ideas critically. We conducted a series of manipulative experiments to determine whether predation mortality is an important mechanism structuring populations of the sea urchin Heliocidaris erythrogramma in Tasmania. Tethered juvenile and adult sea urchins experienced much higher rates of mortality inside no-take marine reserves where sea urchin predators were abundant compared to adjacent fished areas where predators were fewer. Mortality of tagged (but not tethered) sea urchins was also notably higher in marine reserves than in adjacent areas open to fishing. When a range of sizes of sea urchins was exposed to three sizes of rock lobsters in a caging experiment, juvenile sea urchins were eaten more frequently than larger sea urchins by all sizes of rock lobster, but only the largest rock lobsters (> 120 mm CL) were able to consume large adult sea urchins. Tagging (but not tethering) juvenile and adult sea urchins in two separate marine reserves indicated that adult sea urchins experience higher predation mortality than juveniles, probably because juveniles can shelter in cryptic microhabitat more effectively. In a field experiment in which exposure of sea urchins to rock lobster (Jasus edwardsii) and demersal reef fish predators was manipulated, rock lobsters were shown to be more important than fish as predators of adult sea urchins in a marine reserve. We conclude that predators, and particularly rock lobsters, exert significant predation mortality on H. erythrogramma in Tasmanian marine reserves, and that adult sea urchins are more vulnerable than smaller cryptic individuals. Fishing of rock lobsters is likely to reduce an important component of mortality in H. erythrogramma populations.     

Seaweed succession on artificial reefs on different bottom substrata

Artificial rest reefs were set on sandy and rocky bottoms at 5–10 m depth along the coast of southern Japan. Mature thalli ofSargassum, Gelidium and other seaweds were transported from other coastal areas, packed in mesh bags and attached to the reefs to start the beds. After one year, the seaweed flora on the reef on a sandy bottom consisted of more than 20 species, includingSargassum spp. andGelidium amansii, which are important animal food species. Coralline algae were the dominants on the rocky bottom reefs. The lower biomass on reefs on the rocky bottom was due to grazing by urchins. The same number of species was present in the first and second years on reefs on sandy bottoms, but there were moreSargassum thalli the second year.Maximum algal biomass of the artificial reef in May of the second year was 9998 g wet wt m^–2 in sandy areas, 441 g wet wt m^–2 in boulder areas and 228 g wet wt m^–2 in rocky areas. Reefs on rocky bottoms continued to be covered by coralline algae and several species ofCodium andDictyota.