Selective feeding by the echinoid,Evechinus chloroticus,and the removal of plants from subtidal algal stands in Northern New Zealand

Summary Feeding choice of the echinoid Evechinus chloroticus was examined for six fucoid and one laminarian species of algae. Three experiments were conducted to determine the algal choice by echinoids under controlled conditions. In the first experiment, the seven algal species were presented to echinoids in laboratory conditions. The second experiment had replicates of the algal species placed randomly on a subtidal rocky reef where echinoids were abundant and randomly dispersed. For the third experiment, which was also field-based, replicates of one highly-ranked species, Ecklonia radiata, were presented to naturally dispersed Evechinus. In addition, a series of controlled observations was used to examine the order in which echinoids removed algae from mixed species stands on subtidal boulders and to determine if this was related to the experimentally demonstrated choices of algal species.The results of the first two experiments showed that there were differences between algal species in the amount of material grazed by echinoids. Rankings of algal species from the field experiment were not correlated with rankings from the laboratory experiment. The order of removal of algal species from natural stands was correlated with the laboratory-based experimental rankings of algal species, but not with the rankings from the field-based experiment or with algal species availability. There were differences between algal species in their vulnerability to grazing by echinoids, as measured by regression analyses on the amount of material grazed from algal replicates vs. the number of attached echinoids. Within each species, echinoid numbers exerted a non-linear effect on the removal of algal material. In the third experiment, where only one species of algae was presented, the echinoids still distributed themselves non-randomly amongst replicates, aggregating on some samples.Data on the finer scale distribution of algal species over the entire subtidal reef on which these experiments and observations were conducted indicate that Evechinus are not often presented with a choice of adult plants of several different species in natural stands.The evidence from this study supports the conclusion that feeding preferences by echinoids are labile and do not clearly exert the major influence on the removal of plants from natural stands. Preference, as determined from experimental rankings of algal species, is only one of a number of factors which may affect the removal of algae by echinoids. Other important factors are the density of echinoids present, algal susceptibility to removal, and the distribution and abundances of the various algal species and echinoids relative to each other. It is suggested that algal life history characteristics may be unaffected by echinoids and that coevolutionary arguments are not appropriate for describing echinoid-algal interactions.     

Patterns of distribution and abundance of large brown algae and invertebrate herbivores in subtidal regions of northern New Zealand

The distribution and abundance of subtidal laminarian and fucoid algae, echinoids, and four species of herbivorous gastropods were assessed. Line transects and stratified random quadrats were used, encompassing different depths and seven localities ranging over 1000km in northern New Zealand. This sampling was carried out to form a quantitative basis on which to pose hypotheses for later experimental work. Several patterns were apparent, both within and between localities. Fucoid algae were characteristic of depths between 0–6 m, while laminarian algae had peaks of abundance at 3–6 m and from 10–17 m. Intermediate depths were characterized by areas bare of large brown algae, with high densities of echinoids and gastropods. This pattern of a bimodal depth distribution of algae and abundant grazers at intermediate depth was a consistent feature of the northern coastal sites and the semi-exposed offshore islands. Size—frequency distributions for algae at these sites showed that the largest fucoids generally occurred in very shallow water, while the dominant laminarian alga tended to have larger individuals at depths of 10–14 m. In all cases, the biomass distribution with depth of the algal species mirrored their numerical abundance.

Differences between localities were associated both with environmental factors and the unique algal floras of some offshore islands. Echinoids and gastropods were rare at the two most exposed localities, which were also at the extremes of the geographic range. At these sites, depths between 6–10 m were occupied by large brown algae. Deep-water fucoid algae occurred at two offshore localities but the predominant species at one of them was peculiar to that island group.

Because of the general consistency between localities, and the few exceptions to these patterns, hypotheses are considered which could be tested on a local scale to account for echinoid and algal abundances and depth distributions. Our results indicate that the chief questions concerning the structuring of subtidal algal communities are demographic in nature and that an assessment of the status of individual organisms, rather than biomass or productivity, is the appropriate approach to resolve these questions.     

Effects of algal canopy clearance on plant, fish and macroinvertebrate communities on eastern Tasmanian reefs

Changes in assemblages of plants, macroinvertebrates and fishes on three eastern Tasmanian reefs were monitored over 12 months in replicated control blocks and adjacent 10×12-m blocks cleared of fucoid, laminarian and dictyotalean algae. Removal of canopy-forming plants produced less change to biotic assemblages than reported in studies elsewhere, with the magnitude of change for fish and invertebrate taxa lower than variation between sites and comparable to variation between months.The introduced annual kelp Undaria pinnatifida exhibited the only pronounced response to canopy removal amongst algal taxa, with a fivefold increase in cleared blocks compared to control blocks. Marine reserves are suggested to assist reef communities resist invasion by U. pinnatifida, through an indirect mechanism involving increased predation pressure on sea urchins and reduced formation of urchin barrens that are amenable to U. pinnatifida propagation.Large invertebrates were more associated with turfing algae or the reef substratum than the macroalgal canopy. The herbivorous sea urchin Heliocidaris erythrogramma and abalone Haliotis ruber showed the strongest response to clearing amongst common macroinvertebrate species, with a halving of population numbers. Observed densities of the common monacanthid fish Acanthaluteres vittiger also declined by about 50%. The relatively high level of resistance shown by eastern Tasmanian reef biota to patch disturbance was attributed largely to high diversity and biomass of turfing macroalgae damping effects of canopy clearance.     

Influence of Diadema antillarum populations (Echinodermata: Diadematidae) on algal community structure in Jardines de la Reina, Cuba

The 1983-1984 mass mortality of Diadema antillarum produced severe damages on Caribbean reefs contributing to substantial changes in community structure that still persist. Despite the importance of Diadema grazing in structuring coral reefs, available information on current abundances and algal-urchin interactions in Cuba is scarce. We analyzed spatial variations in Diadema abundance and its influence on algal community structure in 22 reef sites in Jardines de la Reina, in June/2004 and April/2005. Urchins were counted in five 30 x 2m transects per site, and algal coverage was estimated in randomly located 0.25m side quadrats (15 per site). Abundances of Diadema were higher at reef crests (0.013-1.553 ind/m2), while reef slope populations showed values up to three orders of magnitude lower and were overgrown by macroalgae (up to 87%, local values). Algal community structure at reef slopes were dominated by macroalgae, especially Dictyota, Lobophora and Halimeda while the most abundant macroalgae at reef crests were Halimeda and Amphiroa. Urchin densities were negatively and positively correlated with mean coverage of macroalgae and crustose coralline algae, respectively, when analyzing data pooled across all sites, but not with data from separate habitats (specially reef crest), suggesting, along with historical fish biomass, that shallow reef community structure is being shaped by the synergistic action of other factors (e.g. fish grazing) rather than the influence of Diadema alone. However, we observed clear signs of Diadema grazing at reef crests and decreased macroalgal cover according to 2001 data, what suggest that grazing intensity at this habitat increased at the same time that Diadema recruitment began to be noticeable. Furthermore, the excessive abundance of macroalgae at reef slopes and the scarcity of crustose coralline algae seems to be due by the almost complete absence of D. antillarum at mid depth reefs, where local densities of this urchin were predominantly low.     

The structure of a nearshore fish community of Western Australia: diel patterns and the habitat role of limestone reefs

Synopsis Extensive limestone reefs are a characteristic feature of much of the coastline of Western Australia, and potentially represent a major habitat feature influencing the structure of the coastal fish community. The structure and temporal dynamics of the fish fauna and its relationships to nearshore patch reefs and surrounding habitat near Dongara, Western Australia, were examined using (1) diel gill-netting and (2) quantitative rotenone sampling of enclosed areas of substratum. Long-term and day-to-day variability of the fauna was low. Dominant species of gill-net collections were either associated with reefs or occurred in similar abundances at both reefs and surrounding sand/seagrass flats. The overall abundance, number of species and biomass of netted fishes was higher around reefs. Rotenone collections of the more sedentary species showed a similar pattern, but suggested, however, that a simple reef versus surrounding sand and seagrass habitat comparison is complicated by the canopy-forming seagrass Amphibolis that occurs on reef tops. Time of day had an important effect on overall fish abundance and number of species, with peaks occurring at crepuscular periods. This reflected dusk and dawn activity peaks of a dominant species rather than overlapping activities of many diurnal and nocturnal species. Diel switches between reef-edge habitat and surrounding sand/seagrass flats were uncommon despite expectations (based on literature examples) that patch reefs would function primarily as sheltering habitats and surrounding non-reef areas act as foraging habitat. High catches at reef-edge sites suggest that the majority of fishes forage on or near limestone patch reefs. Fish densities of around 0.8 individuals per m^-2 of bottom on these Western Australian reefs are relatively high in comparison to visual census estimates obtained for temperate reef systems in South Australia and New Zealand, but similar to those obtained using comparable netting methods in temperate Australian seagrass systems.     

The role of the reef flat in coral reef trophodynamics: Past,present, and future

The reef flat is one of the largest and most distinctive habitats on coral reefs, yet its role in reef trophodynamics is poorly understood. Evolutionary evidence suggests that reef flat colonization by grazing fishes was a major innovation that permitted the exploitation of new space and trophic resources. However, the reef flat is hydrodynamically challenging, subject to high predation risks and covered with sediments that inhibit feeding by grazers. To explore these opposing influences, we examine the Great Barrier Reef (GBR) as a model system. We focus on grazing herbivores that directly access algal primary productivity in the epilithic algal matrix (EAM). By assessing abundance, biomass, and potential fish productivity, we explore the potential of the reef flat to support key ecosystem processes and its ability to maintain fisheries yields. On the GBR, the reef flat is, by far, the most important habitat for turf‐grazing fishes, supporting an estimated 79% of individuals and 58% of the total biomass of grazing surgeonfishes, parrotfishes, and rabbitfishes. Approximately 59% of all (reef‐wide) turf algal productivity is removed by reef flat grazers. The flat also supports approximately 75% of all grazer biomass growth. Our results highlight the evolutionary and ecological benefits of occupying shallow‐water habitats (permitting a ninefold population increase). The acquisition of key locomotor and feeding traits has enabled fishes to access the trophic benefits of the reef flat, outweighing the costs imposed by water movement, predation, and sediments. Benthic assemblages on reefs in the future may increasingly resemble those seen on reef flats today, with low coral cover, limited topographic complexity, and extensive EAM. Reef flat grazing fishes may therefore play an increasingly important role in key ecosystem processes and in sustaining future fisheries yields.     

The role of herbivorous fishes and urchins in coral reef communities

Synopsis Herbivorous fishes and invertebrates are conspicious elements of coral reef communities where they predominate both in numbers and biomass. Herbivores and the coral reef algae on which they feed represent a co-evolved system of defense and counter-defense. Algal species have developed toxic, structural, spatial and temporal defense or escape mechanisms, while the herbivores employ strategies that involve anatomical, physiological and behavioral adaptations. Current research demonstrates that many reef fishes are highly selective in the algae they consume. Food selection in these fishes may be correlated with their morphological and digestive capabilities to rupture algal cell walls. Sea urchins select more in accordance with relative abundance, although certain algal species are clearly avoided.The determinants of community structure on coral reefs have yet to be established but evidence indicates a strong influence by herbivores. Reef herbivores may reduce the abundance of certain competitively superior algae, thus allowing corals and cementing coralline algae to survive. We discuss how the foraging activities of tropical marine herbivores affect the distribution and abundance of algae and how these activities contribute to the development of coral reef structure and the fish assemblages which are intimately associated with reef structure.This paper forms a part of the proceedings of a mini-symposium convened at Cornell University, Ithaca, N.Y., 18–19 May 1976, entitled Patterns of Community Structure in Fishes (G. S. Helfman, ed.).     

The influence of predation and conspecific adults on the abundance of juvenile Evechinus chloroticus (Echinoidea:Echinometridae)

Summary Reefs dominated by red algae, associated with high echinoid densities, are consistent features of the shallow subtidal around northeastern New Zealand. Factors determining the abundance of juvenile Evechinus chloroticus were investigated in such a habitat. Using a factorial design, a field experiment was used to assess the influence of predators and adult E. chloroticus on juvenile abundance. The use of 2 m² exclusion cages enhanced juvenile E. chloroticus abundance over a 16 month period, an effect independent of conspecific adults. We attributed this effect to the exclusion of benthic-feeding, predatory fish. Several species forage over the study area at high densities and are known from gut content analysis to prey on juvenile E. chloroticus in the field. Invertebrate predators are at very low densities in the area. The possibility of caging and site artefacts confounding this interpretation is discussed.Adult E. chloroticus did not directly affect conspecific juvenile densities during the experimental period. However their removal produced a significant change in community structure toward one dominated by macroscopic brown algae. Echinoid removal led to rapid recruitment of laminarian and fucoid algae, predominantly Ecklonia radiata and Sargassum sinclairii. In addition, densities of herbivorous gastropods, particularly the limpet Cellana stellifera decreased in the echinoid exclusion area, as did the feeding rates of predatory fish. The consequences of E. chloroticus removal may be dependent upon the size of the area from which they are excluded.Despite the high densities of predatory fish, a low though consistent number of juvenile E. chloroticus escape predation. We suggest that these represent sufficient input into the adult grazing population to maintain the habitat. This interpretation argues against a key role for predators in structuring shallow water reef communities in northeastern New Zealand.     

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.     

Arctic rhodolith beds and their environmental controls (Spitsbergen, Norway)

Coralline algae (Corallinales, Rhodophyta) that form rhodoliths are important ecosystem engineers and carbonate producers in many polar coastal habitats. This study deals with rhodolith communities from Floskjeret (78°18′N), Krossfjorden (79°08′N), and Mosselbukta (79°53′N), off Spitsbergen Island, Svalbard Archipelago, Norway. Strong seasonal variations in temperature, salinity, light regime, sea-ice coverage, and turbidity characterize these localities. The coralline algal flora consists of Lithothamnion glaciale and Phymatolithon tenue. Well-developed rhodoliths were recorded between 27 and 47 m water depth, while coralline algal encrustations on lithoclastic cobbles were detected down to 77 m water depth. At all sites, ambient waters were saturated with respect to both aragonite and calcite, and the rhodolith beds were located predominately at dysphotic water depths. The rhodolith-associated macrobenthic fauna included grazing organisms such as chitons and echinoids. With decreasing water depth, the rhodolith pavements were regularly overgrown by non-calcareous Polysiphonia-like red algae. The corallines are thriving and are highly specialized in their adaptations to the physical environment as well as in their interaction with the associated benthic fauna, which is similar to other polar rhodolith communities. The marine environment of Spitsbergen is already affected by a climate-driven ecological regime shift and will lead to an increased borealization in the near future, with presently unpredictable consequences for coralline red algal communities.     

Large-scale associations between macroalgal cover and grazer biomass on mid-depth reefs in the Caribbean

Since the 1970s, macroalgae have become considerably more abundant on many Caribbean reefs and overfishing of grazing fishes has been implicated as a contributory factor. We explored relationships between algal cover and grazers (biomass of herbivorous fishes and abundance of the sea-urchin Diadema antillarum) on mid-depth reefs (12-15 m) in 19 areas at seven locations in Jamaica, Barbados, Belize, Grand Cayman and Cuba, between April 1997 and April 1998. Diadema antillarum density was never >0.01 m^-2, while herbivorous fish biomass (acanthurids and scarids ⁸ cm total length) varied from 2-5 g m^-2 in Jamaica to 17.1 g m^-2 in Barbados, and was strongly correlated, negatively with macroalgal cover and positively with 'cropped' substratum (sum of 'bare', turf and crustose-coralline substrata) cover. However, overfishing of herbivorous fishes alone cannot explain the widespread abundance of macroalgae, as even on lightly fished reefs, macroalgal cover was mostly >20%. Herbivorous fish populations on those reefs were apparently only able to maintain approximately 40-60% of reef substratum in cropped states, but due to low space-occupation by coral and other invertebrates, 70-90% of substratum was available to algae. The abundance of macroalgae on lightly fished reefs may therefore be a symptom of low coral cover in combination with the continuing absence of Diadema antillarum.     

Changes in gastropod assemblages in freshwater habitats in the vicinity of Basel (Switzerland) over 87 years

Net pen fish farms generally enrich the surrounding waters and the underlying sediments with nutrients and organic matter, and these loadings can cause a variety of environmental problems, such as algal blooms and sediment anoxia. In this study we test the potential of biofiltration by artificial reefs for reducing the negative environmental impacts surrounding fish farms in the Gulf of Aqaba, Red Sea. Two triangular-shaped artificial reefs (reef volume 8.2 m³) constructed from porous durable polyethylene were deployed at 20 m; one below a commercial fish farm and the other 500 m west of this farm in order to monitor the colonization of these reefs by the local fauna and to determine whether the reef community can remove fish farm effluents from the water. Both reefs became rapidly colonized by a wide variety of organisms with potential for the removal of compounds released from the farms. Within the first year of this study fish abundances and the number of species reached 518–1185 individuals per reef and 25–42 species per reef. Moreover, numerous benthic algae; small sessile invertebrates (bryozoa, tunicates, bivalves, polychaetes, sponges, anemones) and large motile macrofauna (crustaceans, sea urchins, gastropods) settled on the reef surfaces. Depletion of chlorophyll a was measured in the water traversing the artificial reefs in order to assess the biofiltration capacity of the associated fauna. Chlorophyll a was significantly reduced to a level 15–35% lower than ambient concentrations. This reduction was greatest at intermediate current speeds (3–10 cm s^–1), but was not influenced by current direction. The reef structures served as a successful base for colonization by natural fauna and flora, thereby boosting the local benthic biodiversity, and also served as effective biofilters of phytoplankton.     

Community structure and biogeography of shore fishes in the Gulf of Aqaba,Red Sea

Shore fish community structure off the Jordanian Red Sea coast was determined on fringing coral reefs and in a seagrass-dominated bay at 6 m and 12 m depths. A total of 198 fish species belonging to 121 genera and 43 families was recorded. Labridae and Pomacentridae dominated the ichthyofauna in terms of species richness and Pomacentridae were most abundant. Neither diversity nor species richness was correlated to depth. The abundance of fishes was higher at the deep reef slope, due to schooling planktivorous fishes. At 12 m depth abundance of fishes at the seagrass-dominated site was higher than on the coral reefs. Multivariate analysis demonstrated a strong influence on the fish assemblages by depth and benthic habitat. Fish species richness was positively correlated with hard substrate cover and habitat diversity. Abundance of corallivores was positively linked with live hard coral cover. The assemblages of fishes were different on the shallow reef slope, deep reef slope and seagrass meadows. An analysis of the fish fauna showed that the Gulf of Aqaba harbours a higher species richness than previously reported. The comparison with fish communities on other reefs around the Arabian Peninsula and Indian Ocean supported the recognition of an Arabian subprovince within the Indian Ocean. The affinity of the Arabian Gulf ichthyofauna to the Red Sea is not clear. Received in revised form: 2 November 2001 Electronic Publication     

Consumptive and non-consumptive effects of an invasive marine predator on native coral-reef herbivores

Invasive predators typically have larger effects on native prey populations than native predators, yet the potential roles of their consumptive versus non-consumptive effects (CEs vs. NCEs) in structuring invaded systems remains unclear. Invasive lionfish (Pterois volitans) may have ecosystem-level effects by altering native fish grazing on benthic algae that could otherwise displace corals. Lionfish could reduce grazing by decreasing the abundance of herbivorous fishes (CEs), and/or the predation risk posed by lionfish could alter grazing behavior of fishes (NCEs). To test for these CEs, we manipulated lionfish densities on large reefs in The Bahamas and surveyed fish populations throughout June 2009–2011. In July 2011, NCEs of lionfish were measured by observing fish grazing behavior on algal-covered substrata placed in microhabitats varying in lionfish presence at different spatial scales, and quantifying any resulting algal loss. Lionfish reduced small herbivorous fish density by the end of the 2010 summer recruitment season. Grazing by small and large fishes was reduced on high-lionfish-density reefs, and small fish grazing further decreased when in the immediate presence of lionfish within-reefs. Lionfish had a negative indirect effect on algal loss, with 66–80 % less algae removed from substrata in high-lionfish-density reefs. Parrotfishes were likely driving the response of herbivorous fishes to both CEs and NCEs of lionfish. These results demonstrate the importance of considering NCEs in addition to CEs of invasive predators when assessing the effects of invasions.     

Reef fish structure and distribution in a south-western Atlantic Ocean tropical island

The community structure of the reef fish fauna of Trindade Island, a volcanic oceanic island located 1160 km off the coast of Brazil, is described based on intensive visual censuses. Seventy-six species were encountered in 252 censuses, with mean ± S.E. of 99 ± 3 individuals and 15.7 ± 0.3 species 40 m(-2) transect. The average fish biomass, calculated from length-class estimation, was 22.1 kg 40 m(-2) transect. The species contributing most to biomass were, in decreasing order, Melichthys niger, Cephalopholis fulva, Kyphosus spp., Holocentrus adscensionis, Sparisoma amplum, Sparisoma axillare, Acanthurus bahianus and Epinephelus adscensionis. Carnivorous fishes were the largest trophic group in terms of biomass, followed by omnivores and roving herbivores. The two predominant types of reef habitat, fringing reefs built by coralline algae and rocky reefs made of volcanic boulders, showed significant differences in the biomass and the abundance of the trophic guilds. Within each habitat type, significant differences in species richness, density and biomass were detected among crest, slope and interface zones. Although similar in overall species composition to coastal reefs in Brazil, the fish fauna of Trindade Island shares certain characteristics, such as a high abundance of planktivores, with other Brazilian oceanic islands. Despite comparatively high fish biomass, including the macro-carnivorous species habitually targeted by fisheries, signs of overfishing were evident. These findings highlight the urgency for a conservation initiative for this isolated, unique and vulnerable reef system.     

A Paleocene coral—algal—sponge reef from southwestern Alabama and the ecology of Early Tertiary reefs

The Late Paleocene Salt Mountain Limestone from southwestern Alabama is a coral-algal-sponge buildup which further characterizes the faunal makeup of early post-Cretaceous reefs. Thin sectioning has disclosed a variety of lithologies, including large foram-algal packstone, algal bindstone, and sponge bafflestone. A low-diversity fauna of massive scleractinian corals caps the sequence, but may be developed intermittently throughout the section as well. The constructional importance of coralline algae and the low diversity of scleractinian corals are characteristic of Paleocene reefs in general. Sponges, however, are virtually unknown in earliest Tertiary sediments. Their abundance in the Salt Mountain demonstrates not only their local contribution to Early Tertiary reefs, but may also reflect an opportunistic response of sponges as reef constructors following the extinction of oligotrophic, rudist-coral reef communities of the Late Cretaceous. □ Paleocene, reef, paleoecology, sponges, extinction.     

Transformation of algal turf by echinoids and scarid fishes on French Polynesian coral reefs

The respective roles of regular echinoids and scarid fishes in the transformation of turf algae, the main food resource for reef herbivores, were investigated on French Polynesian coral reefs. The role of one species of parrotfish (Scarus sordidus) was compared with that of four species of echinoids. The degree and ways of degradation of the algal matter were determined by the organic matter percentage, the composition of the sugar fraction, and the concentration and composition of chlorophylltype pigments as assayed by HPLC analysis. Chemical analyses were performed on anterior and posterior intestines for scarids, intestinal contents and faeces for echinoids, and on fresh algal turf as a control of initial food quality. A decrease in mean percentage of organic matter in gut content was observed from intestine (9.7%) to faeces (7%) in sea urchins, but not in parrotfishes. The total sugar fraction decreased from fresh algal turf (32% of total organic matter) to echinoid (28%) to scarid (18%) gut contents. The ratio of insoluble to soluble sugars (I/S ratios) was higher in echinoids (2.6) than in scarid gut contents (1.0). A decrease in the total pigment concentration was measured from fresh algal turf to echinoid and it was found to be even lower in scarid gut contents. Chromatograms showed that the composition of chlorophyll-type pigments in scarid intestines was very similar to fresh algal turf, with a dominance of native forms, mainly chlorophyll a and b. On the contrary, degraded pigment forms dominated in echinoids. The main degraded products were pheophorbides in sea urchins, and chlorophyllides in parrotfishes. These results provided evidence for differentiation in digestive processes occurring in the two types of grazers. Echinoids released higher degraded algal material than did scarids. Thus, these two types of grazers play different roles in the recycling of organic matter on coral reefs.     

The role of turtles as coral reef macroherbivores

Herbivory is widely accepted as a vital function on coral reefs. To date, the majority of studies examining herbivory in coral reef environments have focused on the roles of fishes and/or urchins, with relatively few studies considering the potential role of macroherbivores in reef processes. Here, we introduce evidence that highlights the potential role of marine turtles as herbivores on coral reefs. While conducting experimental habitat manipulations to assess the roles of herbivorous reef fishes we observed green turtles (Chelonia mydas) and hawksbill turtles (Eretmochelys imbricata) showing responses that were remarkably similar to those of herbivorous fishes. Reducing the sediment load of the epilithic algal matrix on a coral reef resulted in a forty-fold increase in grazing by green turtles. Hawksbill turtles were also observed to browse transplanted thalli of the macroalga Sargassum swartzii in a coral reef environment. These responses not only show strong parallels to herbivorous reef fishes, but also highlight that marine turtles actively, and intentionally, remove algae from coral reefs. When considering the size and potential historical abundance of marine turtles we suggest that these potentially valuable herbivores may have been lost from many coral reefs before their true importance was understood.     

Grazing pressure of herbivorous coral reef fishes on low coral-cover reefs

The impact of grazing by herbivorous fishes (Acanthuridae, Scaridae, and Pomacentridae) on low coral-cover reefs was assessed by measuring rates of benthic algal production and consumption on inshore and offshore reefs in the upper Florida Keys. Algal production rates, determined in situ with caged and uncaged experimental plates, were low (mean 1.05 g C m⁻² day⁻¹) and similar among reef types. Algal consumption rates were estimated using two different models, a detailed model incorporating fish bite rates and algal yield-per-bite for one species extrapolated to a guild-wide value, and a general regression relating fish biomass to algal consumption. Algal consumption differed among reef types: a majority of algal production was consumed on offshore reefs (55–100%), whereas consumption on inshore patch reefs was 31–51%. Spatial variation in algal consumption was driven by differences in herbivorous fish species composition, density, and size-structure among reef types. Algal consumption rates also varied temporally due to seasonal declines in bite rates and intermittent presence of large-bodied, vagile, schooling species. Spatial coherence of benthic community structure and temporal stability of algal turf over 3 years suggests that grazing intensity is currently sufficient to limit further spread of macroalgal cover on these low coral-cover reefs, but not to exclude it from the system.     

A preliminary study of the biology and ecology of the blue starfish Linckia laevigata (L.) on the Australian Great Barrier Reef and an interpretation of its role in the coral reef ecosystem

The blue starfish Linckia laevigata grazes coralline algae. The starfish populations studied were composed entirely of adults. Spawning takes place in October at the southern end of the Great Barrier Reef. On reefs which were unaffected by Acanthaster planci, L. laevigata was confined to algae covered reef tops and rubble banks. On reefs affected by A. planci, L. laevigata had extended its range and was feeding on and among the coralline algae covered dead hard coral skeletons on the reef perimeter. Coral regrowth, followingattack by A. planci , was found to be slower on reefs with populations of L. laevigata living on the reef perimeter than on reefs where they were absent from this region. It is suggested that grazing by L. laevigata destroys small coral colonies and newly settled larvae thus slowing down the rate of coral regeneration. The consequences of this reduced rate of recolonisation is also discussed.