Herbivory on coral reefs: algal susceptibility to herbivorous fishes

Summary The susceptibility of several tropical algal species to fish grazing was studied on the Belizean barrier reef off the Caribbean coast of Central America. Short-term transplant experiments indicate that plant species vary markedly in their rates of biomass loss to grazing by a shallow-water guild of herbivorous fishes. Algal species transplanted from habitats with low grazing pressure are highly susceptible to grazing, while species occurring in habitats with high herbivore densities are highly resistant to grazing. Algal species show differential susceptibility to grazing by two major components of the tropical herbivore guild, Acanthurus (surgeonfishes) and Sparisoma (parrotfishes).Variability in plant susceptibility to grazing by herbivorous fishes was not clearly correlated with morphological or chemical characteristics that have been previously suggested as plant defenses against herbivory. Plants found to be highly resistant to fish grazing, such as Halimeda, exhibit both morphological characteristics and secondary chemical compounds which do appear to reduce herbivory. In contrast, species of Caulerpa, Sargassum, Turbinaria, and Padina, which also possess alleged morphological and/or chemical defenses, are nevertheless highly susceptible to fish grazing.     

Niche partitioning of feeding microhabitats produces a unique function for herbivorous rabbitfishes (Perciformes, Siganidae) on coral reefs

Niche theory predicts that coexisting species minimise competition by evolving morphological or behavioural specialisations that allow them to spread out along resource axes such as space, diet and temporal activity. These specialisations define how a species interacts with its environment and, by extension, determine its functional role. Here, we examine the feeding niche of three species of coral reef-dwelling rabbitfishes (Siganidae, Siganus). By comparing aspects of their feeding behaviour (bite location, bite rate, foraging distance) with that of representative species from two other abundant herbivorous fish families, the parrotfishes (Labridae, Scarus) and surgeonfishes (Acanthuridae, Acanthurus), we examine whether rabbitfishes have a feeding niche distinct from other members of the herbivore guild. Measurements of the penetration of the fishes’ snouts and bodies into reef concavities when feeding revealed that rabbitfish fed to a greater degree from reef crevices and interstices than other herbivores. There was just a 40 % overlap in the penetration-depth niche between rabbitfish and surgeonfish and a 45 % overlap between rabbitfish and parrotfish, compared with the almost complete niche overlap (95 %) recorded for parrotfish and surgeonfish along this spatial niche axis. Aspects of the morphology of rabbitfish which may contribute to this niche segregation include a comparatively longer, narrower snout and narrower head. Our results suggest that sympatric coexistence of rabbitfish and other reef herbivores is facilitated by segregation along a spatial (and potentially dietary) axis. This segregation results in a unique functional role for rabbitfishes among roving herbivores that of “crevice-browser”: a group that specifically feeds on crevice-dwelling algal or benthic organisms. This functional trait may have implications for reef ecosystem processes in terms of controlling the successional development of crevice-based algal communities, reducing their potential to trigger macroalgal outbreaks.     

Territorial behavior of threespot damselfish (Eupomacentrus planifrons) increases reef algal biomass and productivity

Synopsis Algal growth and damselfish (Eupomacentrus planifrons) territories were studied in two reef habitats at Discovery Bay, Jamaica. Damselfish territories were contiguous in the reef flat (0 to 2.5 m), where the algal composition and biomass varied from territory to territory. In contrast, on the lower reef terrace (22 m), damselfish territories were often spatially segregated. While the algal composition of the territories was more uniform on the reef terrace, the total algal biomass was lower than in the territories on the reef flat. Damselfish are largely herbivorous, and they defend their territories against most intruding fish, including a number of herbivorous species. Areas of the reef terrace outside of damselfish territories were heavily grazed by herbivorous fishes and contained only small quantities of non-crustose algae.The reef terrace territories were characterized by a multispecific turf of algae (greens, blue-greens, and reds) covering the Acropora cervicornis framework and by the leafy, brown alga, Lobophora variegata. A rapid reduction in the biomass of brown algae and filamentous algae was noted when damselfish were permanently removed from their territories. Only calcified, encrusting algae — plants apparently somewhat undesirable as fish food sources — would be common on the terrace zone of this reef if damselfish territories were absent. Damselfish territoriality may significantly influence the dynamics of some reefs by increasing the biomass of the algal turf thereby increasing; reef productivity. Since blue-green algae, potential nitrogen fixers, occur in these algal turfs, the fish may also be indirectly affecting reef nutrition.     

A standard unit for monitoring recruitment of fishes to coral reef rubble

We developed a benthic standard monitoring unit for recruitment of fishes (SMURF) to sample fishes that settle in coral rubble and tested it on three fringing reefs on the west coast of Barbados, West Indies. These rubble SMURFs are inexpensive to construct and permit newly settled fishes, including cryptic, secretive and small species, to be quickly and fairly completely (> 83%) removed by divers without returning the unit to the surface. Over a 3-month period involving 9 collections, 32 units on 3 reefs yielded 948 newly settled fishes belonging to 28 taxa (3.3 ± 0.2 S.E. recruits per sample, n = 287) in addition to numerous crustaceans. Most specimens were Sparisoma spp. (Scaridae) (41.8%) and Stegastes partitus (Pomacentridae) (13.6%), with moderate numbers of Scarus spp. (Scaridae) (8.6%), Lythrypnus spp. (Gobiidae) (7.4%), and Pseudogramma gregoryi (Serranidae) (7.2%). Sampling SMURFs at 1-day rather than 11-day intervals yielded approximately twice as many fish, although some taxa were not affected by sampling frequency, indicating taxon-specific differences in post-settlement loss rates. Netting, intended to reduce predation on settlers, did not affect settlement estimates unless algae grew on it, suggesting that the SMURF design minimizes fish predation even in the absence of the netting. Over time, however, the netting resulted in substantial algal growth that was associated with an increase in abundance of Sparisoma spp. and a decrease in the abundance of S. partitus. This suggests that settler estimates are more affected by microhabitat changes from algal growth than by predation. During low settlement periods, SMURFs performed better than light-traps in assessing the daily input of new settlers of two abundant taxa. Eight SMURFs per reef were sufficient to demonstrate differences in settlement rates and temporal correlations in settlement within and among reefs. This study underscores the potential of benthic SMURFs to measure settlement of coral reef fishes at a range of spatio-temporal scales with moderate research cost and effort, and to facilitate the study of the settlement stage of several taxa.     

Trophic significance of herbivorous macroinvertebrates on the central Great Barrier Reef

The common herbivorous macroinvertebrates on reef flats of the central Great Barrier Reef (GBR) were, in order of abundance: gastropod molluscs (Turbo and Trochus spp.); pagurid crabs; and the starfish, Linckia laevigata. The density of macroinvertebrates on Davies Reef was lowest in the thick-turf habitats at the windward reef-crest (0.6–0.9 m^–2) compared with 3.1 to 5.2 m^–2 elsewhere on the reef flat. Invertebrate grazer densities were similar on mid-shelf reef flats (mean: 2.3–3.6 m^–2) and significantly lower on outer-shelf reefs (0.3–1.0 m^–2). The ingestion rate of Turbo chrysostomus, the most abundant macroinvertebrate species, was derived from (a) faecal production and food absorption efficiency, (b) comparison of algal biomass on grazed and ungrazed natural substrata and (c) gut-filling rate and feeding periodicity in field populations. The ingestion rate of Trochus pyramis, the most common trochid and an abundant component of the macroinvertebrate fauna, was also estimated using (a). This gastropod fed continuously, whereas T. chrysostomus showed a distinct nocturnal feeding periodicity. T. chrysostomus and T. pyramis ingested daily means of 35 and 54 mgC animal^-1, respectively. Total gastropod grazing rates (mgC m^–2d^-1 in the field ranged from 11 in a thick-turf, reef-crest habitat to 144 on the open-grazed main flat. Grazing by gastropods accounted for between 0.3 and 8% of the net production of benthic algal food resources, depending on location on the reef flat. Across the whole reef flat the mean (areally-weighted) gastropod grazing rate was 6% of net production. A comparison of the relative roles of different types of grazers led to the conclusion that fishes are likely to have the greatest overall trophic impact on reefs of the central GBR. Even where macroinvertebrates are most abundant on reef flats, the yield from benthic algal communities to macroinvertebrates is estimated to be only one third of that due to fishes.Contribution no. 471 from the Australian Institute of Marine Science     

Direct evaluation of macroalgal removal by herbivorous coral reef fishes

Few studies have examined the relative functional impacts of individual herbivorous fish species on coral reef ecosystem processes in the Indo-Pacific. This study assessed the potential grazing impact of individual species within an inshore herbivorous reef fish assemblage on the central Great Barrier Reef (GBR), by determining which fish species were able to remove particular macroalgal species. Transplanted multiple-choice algal assays and remote stationary underwater digital video cameras were used to quantify the impact of local herbivorous reef fish species on 12 species of macroalgae. Macroalgal removal by the fishes was rapid. Within 3 h of exposure to herbivorous reef fishes there was significant evidence of intense grazing. After 12 h of exposure, 10 of the 12 macroalgal species had decreased to less than 15% of their original mass. Chlorodesmis fastigiata (Chlorophyta) and Galaxaura sp. (Rhodophyta) showed significantly less susceptibility to herbivorous reef fish grazing than all other macroalgae, even after 24 h exposure. Six herbivorous and/or nominally herbivorous reef fish species were identified as the dominant grazers of macroalgae: Siganus doliatus, Siganus canaliculatus, Chlorurus microrhinos, Hipposcarus longiceps, Scarus rivulatus and Pomacanthus sexstriatus. The siganid S. doliatus fed heavily on Hypnea sp., while S. canaliculatus fed intensively on Sargassum sp. Variation in macroalgal susceptibility was not clearly correlated with morphological and/or chemical defenses that have been previously suggested as deterrents against herbivory. Nevertheless, the results stress the potential importance of individual herbivorous reef fish species in removing macroalgae from coral reefs.     

Effects of fish predation and seaweed competition on the survival and growth of corals

On Caribbean coral reefs, high rates of grazing by herbivorous fishes are thought to benefit corals because fishes consume competing seaweeds. We conducted field experiments in the Florida Keys, USA, to examine the effects of grazing fishes on coral/seaweed competition. Initially, fragments of Porites divaracata from an inshore habitat were transplanted into full-cage, half-cage, and no-cage treatments on a fore-reef. Within 48 h, 56% of the unprotected corals in half-cage and no-cage treatments (62 of 111) were completely consumed. Stoplight parrotfish (Sparisoma viride) were the major coral predators, with redband parrotfish (S. aurofrenatum) also commonly attacking this coral. Next, we transplanted fragments of P. porites collected from the fore-reef habitat where our caging experiments were being conducted into the three cage treatments, half in the presence of transplanted seaweeds, and half onto initially clean substrates. The corals were allowed to grow in these conditions, with concurrent development of competing seaweeds, for 14 weeks. Although seaweed cover and biomass were both significantly greater in the full-cage treatment, coral growth did not differ significantly between cage treatments even though corals placed with pre-planted seaweeds grew significantly less than corals placed on initially clean substrate. This surprising result occurred because parrotfishes not only grazed algae from accessible treatments, but also fed directly on our coral transplants. Parrotfish feeding scars were significantly more abundant on P. porites from the half and no-cage treatments than on corals in the full cages. On this Florida reef, direct fish predation on some coral species (P. divaracata) can exclude them from fore-reef areas, as has previously been shown for certain seaweeds and sponges. For other corals that live on the fore-reef (P. porites), the benefits of fishes removing seaweeds can be counterbalanced by the detrimental effects of fishes directly consuming corals. Received: 31 May 1997 / Accepted: 2 September 1997     

Evaluation of algal regulation by herbivorous fishes on Caribbean coral reefs

The role of herbivorous fishes in maintaining low macroalgal cover was evaluated on coral reefs on several reef sites from Guadeloupe, either protected or not. Grazing by herbivorous fishes was assessed on different algal facies using fish-bite counts. Algal consumption by fish was estimated as well as algal production. Bite counts revealed that herbivorous fishes feed preferentially on algal turf and avoid brown macroalgae. The algal consumption varied between 0.4 and 2.8 g m⁻² days⁻¹ and was higher inside marine protected areas than outside. Comparison with algal production revealed that herbivorous fishes did not succeed in regulating algal growth. The insufficient number of grazers may lead to the dominance of stable assemblages of macroalgae on coral reefs, preventing the recovery of reef into previous coral-dominated ecosystems.     

Branching coral as a macroalgal refuge in a marginal coral reef system

Marginal coral reef systems may provide valuable insights into the nature of ecosystem processes in systems on the trajectory towards a phase shift to an alternate ecosystem state. This study investigates the process of herbivory in a marginal coral reef system in the Keppel Islands at the southern end of the Great Barrier Reef. Branching Acropora coral and the brown macroalga Lobophora variegata occupied up to 95% of the reef crest substratum at the three surveyed reefs. Feeding rates of herbivorous fishes and removal rates of Lobophora were directly quantified within areas of branching Acropora and on planar surfaces. Feeding rates by herbivorous fishes were habitat dependent with the highest bite rates being found in planar habitats for both Lobophora and the epilithic algal matrix (EAM) by 1–2 orders of magnitude, respectively. Feeding rates on Lobophora were, however, much lower than rates on the EAM. The low rates of Lobophora removal and significantly lower rates of herbivory in branching habitats were consistent with the high biomass of this brown alga throughout the Keppel Islands and with its distribution on reef crests, where Lobophora biomass was 20 times greater in branching than in planar habitats. This lack of feeding by herbivorous fishes within branching coral habitats in the Keppel Islands contrasts with the typical role of coral and topographic complexity on herbivores on coral reefs and highlights the potential for complex interactions between algae, corals and fishes on coral reefs. On marginal systems, herbivory may modify algal distributions but may be unable to contain the proliferation of algae such as Lobophora.     

Effects of herbivores, nutrient enrichment, and their interactions on macroalgal proliferation and coral growth

We conducted a 20-week manipulative field experiment on shallow forereefs of the Florida Keys to assess the separate and interactive effects of herbivory and nutrient enrichment on the development of macroalgal communities and the fitness of the corals Porites porites and Siderastrea siderea. Excluding large herbivorous fishes produced macrophyte blooms both with and without nutrient enrichment. In contrast, there were no direct effects of nutrient enrichment. There were, however, small, but significant, interactive effects of herbivory and enrichment on macroalgal cover. Following nutrient enrichment, total macroalgae and the common seaweeds Dictyota spp. were suppressed in the presence, but not in the absence, of large herbivorous fishes—suggesting that fishes were selectively feeding on nutrient-enriched macrophytes. Access by large herbivores prevented algal overgrowth of corals, but these large fishes also directly grazed both corals. Excluding fishes did not alter survivorship of either coral species, but did decrease parrotfish grazing scars on both corals and increased the net growth of P. porites. Nutrient additions had no direct effects on the survivorship of corals, but there was a trend (P = 0.097) for nutrients to stimulate the growth of P. porites. The preponderance of experiments available to date indicates that loss of key herbivores is a major factor driving macroalgal blooms on coral reefs; anthropogenic nutrient pollution generally plays a more minor role.     

Effects of seaweed extracts and secondary metabolites on feeding by the herbivorous surgeonfish Naso lituratus

We examined 22 species of algae and two species of seagrasses from coral reef habitats around Guam to determine if they possessed chemical defenses against the acanthurid Naso lituratus. Whole plants (18 species) were offered to determine whether they were preferred or avoided by N. lituratus in the laboratory. Organic extracts of 15 algae and one seagrass were applied to palatable seaweeds and offered to N. lituratus in the laboratory to determine if the seaweeds were chemically defended. Extracts that deterred feeding were further fractionated if sufficient amounts were available, and the fractions and associated pure compounds were tested in similar feeding assays. N. lituratus was significantly deterred from feeding by crude extracts from five different species of algae: Avrainvillea obscura, Bryopsis pennata, grazed Halimeda macroloba, Neomeris annulata, and Portieria (=Desmia) hornemannii. The pure compounds avrainvilleol from A. obseura, ochtodene from P. hornemannii, one fraction and one brominated sesquiterpene from N. annulata, and two fractions from T. expeditionis also deterred feeding. These results, together with previous work, suggest that tropical herbivorous fishes differ in their responses to plant chemistry, and this variability precludes broad generalization about the effects of marine plant secondary metabolites on herbivorous fishes.     

Herbivory facilitates growth of a key reef‐building Caribbean coral

The decline of reef‐building corals in conjunction with shifts to short‐lived opportunistic species has prompted concerns that Caribbean reef framework‐building capacity has substantially diminished. Restoring herbivore populations may be a potential driver of coral recovery; however, the impact of herbivores on coral calcification has been little studied. We performed an exclusion experiment to evaluate the impact of herbivory on Orbicella faveolata coral growth over 14 months. The experiment consisted of three treatments: full exclusion cages; half cage procedural controls; and uncaged control plates, each with small O. faveolata colonies. We found that herbivorous fish exclusion had a substantial impact on both macroalgal cover and coral growth. Fleshy macroalgae reached 50% cover within some exclusion cages, but were almost absent from uncaged control plates. Critically, O. faveolata calcification rates were suppressed by almost half within exclusion cages, with monthly coral growth negatively related to overgrowth by fleshy macroalgae. These findings highlight the importance of herbivorous fishes for coral growth and the detrimental impact of macroalgal proliferation in the Caribbean. Policy makers and local managers should consider measures to protect herbivorous fishes and reduce macroalgal proliferation to enable coral communities to continue to grow and function.     

Predation Risk,Resource Quality,and Reef Structural Complexity Shape Territoriality in a Coral Reef Herbivore

For many species securing territories is important for feeding and reproduction. Factors such as competition, habitat availability, and male characteristics can influence an individual’s ability to establish and maintain a territory. The risk of predation can have an important influence on feeding and reproduction; however, few have studied its effect on territoriality. We investigated territoriality in a haremic, polygynous species of coral reef herbivore, Sparisoma aurofrenatum (redband parrotfish), across eight reefs in the Florida Keys National Marine Sanctuary that were either protected or unprotected from fishing of piscivorous fishes. We examined how territory size and quality varied with reef protection status, competition, predation risk, and male size. We then determined how territory size and quality influenced harem size and female size to understand the effect of territoriality on reproductive potential. We found that protected reefs trended towards having more large predatory fishes and that territories there were smaller but had greater algal nutritional quality relative to unprotected reefs. Our data suggest that even though males in protected sites have smaller territories, which support fewer females, they may improve their reproductive potential by choosing nutritionally rich areas, which support larger females. Thus, reef protection appears to shape the trade-off that herbivorous fishes make between territory size and quality. Furthermore, we provide evidence that males in unprotected sites, which are generally less complex than protected sites, choose territories with higher structural complexity, suggesting the importance of this type of habitat for feeding and reproduction in S. aurofrenatum. Our work argues that the loss of corals and the resulting decline in structural complexity, as well as management efforts to protect reefs, could alter the territory dynamics and reproductive potential of important herbivorous fish species.     

Impact of Herbivore Identity on Algal Succession and Coral Growth on a Caribbean Reef

Background

Herbivory is an important top-down force on coral reefs that regulates macroalgal abundance, mediates competitive interactions between macroalgae and corals, and provides resilience following disturbances such as hurricanes and coral bleaching. However, reductions in herbivore diversity and abundance via disease or over-fishing may harm corals directly and may indirectly increase coral susceptibility to other disturbances.

Methodology and Principal Findings

In two experiments over two years, we enclosed equivalent densities and masses of either single-species or mixed-species of herbivorous fishes in replicate, 4 m² cages at a depth of 17 m on a reef in the Florida Keys, USA to evaluate the effects of herbivore identity and species richness on colonization and development of macroalgal communities and the cascading effects of algae on coral growth. In Year 1, we used the redband parrotfish (Sparisoma aurofrenatum) and the ocean surgeonfish (Acanthurus bahianus); in Year 2, we used the redband parrotfish and the princess parrotfish (Scarus taeniopterus). On new substrates, rapid grazing by ocean surgeonfish and princess parrotfish kept communities in an early successional stage dominated by short, filamentous algae and crustose coralline algae that did not suppress coral growth. In contrast, feeding by redband parrotfish allowed an accumulation of tall filaments and later successional macroalgae that suppressed coral growth. These patterns contrast with patterns from established communities not undergoing primary succession; on established substrates redband parrotfish significantly reduced upright macroalgal cover while ocean surgeonfish and princess parrotfish allowed significant increases in late successional macroalgae.

Significance

This study further highlights the importance of biodiversity in affecting ecosystem function in that different species of herbivorous fishes had very different impacts on reef communities depending on the developmental stage of the community. The species-specific effects of herbivorous fishes suggest that a species-rich herbivore fauna can be critical in providing the resilience that reefs need for recovery from common disturbances such as coral bleaching and storm damage.     

Parrotfish abundance and selective corallivory on a Belizean coral reef

Parrotfish are important members of coral reef communities because they consume macroalgae that would otherwise outcompete reef-building corals for space. However, some Caribbean parrotfish species also feed directly on live corals, and thus have the potential to negatively impact coral fitness and survival. This study investigates selective grazing by parrotfish on particular coral species, differences in grazing incidence among reef habitats and intraspecific discrimination among colonies of several coral species. We also investigate spatial and temporal patterns of parrotfish species abundance across habitats on the Belize barrier reef, and examine correlations between parrotfish abundance and grazing intensity across reef habitats. We found that members of the Montastraea annularis species complex, major builders of Caribbean reefs, were preferred targets of parrotfish grazing across all reef habitats, while M. cavernosa, Agaricia agaricites, Diploria strigosa, Porites astreoides and Porites porites were not preferred; Siderastrea siderea was preferentially grazed only in the spur and groove habitats. Parrotfish grazing preferences varied across habitats; M. annularis was grazed most often in shallow habitats, whereas M. franksi was consumed more at depth. Although it was not possible to directly observe parrotfish grazing on corals, we did find a positive correlation between Sparisoma aurofrenatum abundance and M. franksi grazing incidence across habitats. Finally, when we compared our results to parrotfish abundances measured by a previous study, we found that Sparisoma viride and Sp. aurofrenatum, two species known to be corallivorous, had increased abundances between 1982 and 2004. In light of escalating threats on Caribbean reef corals, it would be important for future studies to evaluate the impact of parrotfish corallivory on coral survival.     

Seasonally changing habitat use patterns among roving herbivorous fishes in the southern Red Sea: the role of temperature and algal community structure

Coral reefs are characterized by intense herbivory. Spatial patterns in herbivory—particularly along the depth gradient—influence the distribution and abundance of algae. Depth gradients in herbivorous reef fishes are generally assumed to be temporally stable, but this assumption has rarely been questioned. Here, we use underwater visual census and herbivore exclusion experiments to study the community composition and temporal patterns in habitat use by roving herbivorous fishes in an environment characterized by profound seasonal changes in algal biomass and distribution and extreme summer temperatures. Among the 18 species of roving herbivores recorded, parrotfishes were dominant in species richness and biomass, while regional endemic species represented 77 % of the total biomass. During most of the year, roving herbivores aggregate in the shallow reef zones and their biomass declines with depth. The herbivore community on the reef flat is distinct from that in deeper zones. The former is characterized by Siganus rivulatus, Acanthurus gahhm and Hipposcarus harid, while the deeper reef zones are characterized by S. ferrugineus, Chlorurus sordidus and Ctenochaetus striatus. In summer, the distinct community structures among reef zones are lost as reef flat herbivores tend to exploit deeper reef zones and some reef crest species venture on to the reef flat. This summer change in herbivore distribution is also reflected in reduced turf biomass and increased yield to herbivores in the deeper reef zones. Habitat use is related to the feeding mode such that browsers dominate the reef flat and scrapers the reef crest, while the seasonal changes correspond to changes in availability of targeted algal resources. These seasonal changes appear to be driven by the extreme temperatures in summer, reaching 36 °C on the shallow reef flat.     

Habitat correlates of the distribution and biomass of Seychelles' reef fishes

Synopsis Relationships between quantitative measures of habitat type and the biomass of Chaetodon, Scarus and Parupeneus species were investigated across 35 reef sites in the Inner Seychelles Group. Multiple regression was used to determine the proportion of variance in biomass between sites which could be explained by depth, exposure, vertical relief, topographic complexity, live coral cover, coral rubble cover, rock cover, sand cover, underlying carbonate substrate, underlying sand substrate, underlying rock substrate and an index of fishing intensity. A significant proportion of the variance in biomass was explained by habitat variables and the index of fishing intensity for 7 of 12 Chaetodon species (23–52% of variance explained), 3 of 6 Parupeneus species (33–40%), and 10 of 13 Scarus species (14–46%). Within genera, different groups of habitat variables explained the variance in biomass for different species and, of the variables studied, only the proportion of underlying sand substrate failed to explain a significant proportion of the variance in biomass for any species. Quantitative relationships between the biomass of Chaetodon and habitat were often in accordance with those suggested by previous studies of their ecology, life-history and distribution at other Indo-Pacific locations. However, the habitat associations of the Parupeneus and some Scarus species have not been studied at other locations and clearly warrant further investigation. It was concluded that habitat was an important determinant of the distribution of many Seychelles reef fishes, but that the habitat variables examined were rarely the most important determinant of biomass. However, the inclusion of a procedure to collect habitat data provided a useful means by which to reduce the unexplained variance associated with visual census biomass estimates and therefore improves the possibility of elucidating the effects of other factors on the biomass of Seychelles reef fishes.     

Seaweeds preferred by herbivorous fishes

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

The relative impacts of grazing by caribbean coral reef fishes and Diadema: effects of habitat and surge

The effects of grazing on fleshy algal turfs by large herbivorous fishes and by the black spiny urchin, Diadema antillarum (Philippi), were investigated in three shallow (<3 m), moderately exposed, reef flat habitats in the San Blas Islands on the Caribbean coast of Panama. Grazing by Diadema was found to be a primary determinant of the biomass of fleshy algae in this shallow reef environment, and its impact was exposure-dependent. Feeding activity of Diadema was inhibited during the protracted periods of heavy wave action that characterize the dry season when strong north winds (24–27 kph) are typical. In the shallowest, most exposed elkhorn bluff habitats urchin grazing was minimal even during calm periods due to the effects of waves generated by unpredictable, light winds. In a relatively protected backreef habitat, water movement was inadequate to discourage grazing significantly, even during the turbulent dry season. As a result, algal biomass (decalcified dry weight) was typically three-fold greater in the elkhorn bluff habitat than on the ackreef. In habitats of intermediate exposure (elkhorn fields) algal biomass was reduced during calm periods, but increased to the level characteristic of the elkhorn bluff habitat during protracted rough periods that inhibited grazing by Diadema.A caging experiment in the backreef habitat demonstrated a relatively minor effect of grazing by herbivorous fishes in comparison to that of grazing by Diadema. Observations on the distributions and abundances of herbivorous fishes also indicated a relatively minor role for these grazers as determinants of algal biomass in the more exposed habitats.