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.     

Marine macroalgae as foods for fishes: an evaluation of potential food quality

Synopsis A revitalized view of feeding by herbivorous marine fishes is sought through two questions. First, What characteristics of major taxa of algae identify them as predictably high or low quality foods? Second, are marine algae valuable foods for fishes which do not mechanically disrupt cell walls and do not harbor specialized enzymes or microbes capable of lysing cell walls? Energy, ash and nutrient content of 16 species of marine algae were employed to assess food quality of fleshy red, green, brown and calcareous red algae. On the basis of ash, calories, total protein and total lipid content, fleshy algae should be superior to calcareous algae as foods for fishes; in addition, green algae should be superior to brown algae and brown algae superior to red algae. When the probable digestibility of storage and extracellular carbohydrates is considered, green and red algae are predicted superior to brown algae as food. Two species of damselfishes (Pomacentridae) from the Gulf of California,Eupomacentrus rectifraenum andMicrospathodon dorsalis, eat red and green algae and ignore brown and calcareous algae. They feed, therefore, in a fashion consistent with predictions based only on algal chemistry. These fishes absorb at least 20–24% of the biomass, 57–67% of the protein, 46–56% of the lipid and 37–44% of the carbohydrate contained in algae eaten in the wild. Since these damselfishes do not masticate their food, it appears that herbivorous fishes can digest major fractions of algal nutrients without mechanical destruction of algal cells.     

Community structure and diet of roving herbivorous reef fishes in the Abrolhos Archipelago, south-western Atlantic

Community structure and diet of roving herbivorous reef fishes were analysed in 13 study sites around the five islands of the Abrolhos Archipelago, north-eastern Brazil (17°58' S; 38°42' W). This area has been part of the Abrolhos Marine National Park since 1983. Abundances and diets of fishes within the families Scaridae, Acanthuridae and Kyphosidae were compared among groups of sites differing in benthic community structure and exposure regime. The abundance of roving herbivorous fishes was higher in shallower sites than in deeper sites. At all study sites, observations of total herbivorous reef fish community structure revealed that 64% of fishes were acanthurids, 33% were scarids and only 4% were kyphosids. This pattern was predominant in all study sites. The majority of fishes examined in this study had the bulk of their diet based on both algae (35–90%) and detritus (35–65%). Among groups of algae, filamentous algae were the most commonly consumed as the majority of roving herbivorous fishes in Abrolhos feed as scrapers and excavators. The exceptions were the kyphosids, which had a diet consisting primarily of macroalgae (mainly Phaeophyta), and Acanthurus coeruleus that consumed mainly turf algae and very little detritus. The data collected indicate that roving herbivorous fishes in the Abrolhos Archipelago are more abundant at calm sites where detritus and delicate algae, their major food resources, are more abundant. The results presented here, along with underwater observations, suggest that the abundance and diet of these three fish families are associated with substratum cover.     

Activation of chemical defenses in the tropical green algae Halimeda spp.

Halimeda spp. are among the most common seaweeds on tropical reefs where herbivory is intense. These calcified seaweeds produce diterpenoid feeding deterrents; the major metabolites are halimedatetraacetate and halimedatrial. We found that most species of Halimeda on Guam immediately convert the less-deterrent secondary metabolite halimedatetraacetate to the more potent feeding deterrent halimedatrial when plants are injured by grinding or crushing. This conversion would therefore occur when fishes bite or chew Halimeda plants. We term this process of rapid conversion “activation”. Extracts from injured plants contained higher amounts of halimedatrial and were more deterrent toward herbivorous fishes than extracts from control plants. Herbivore-activated defenses are common in many families of terrestrial plants: however, this is the first example of an activated defense in a marine plant.     

Diet,food preference,and algal availability for fishes and crabs on intertidal reef communities in southern California

Synopsis Herbivory by wide-ranging fishes is common over tropical reefs, but rare in temperate latitudes where the effects of herbivorous fishes are thought to be minimal. Along the west coast of North America, herbivory by fishes on nearshore reefs is largely restricted to a few members of the Kyphosidae, distributed south of Pt. Conception. This paper presents information on natural diets and results from feeding choice experiments for two abundant kyphosids from intertidal habitats in San Diego, California —Girella nigricans andHermosilla azurea, and similar data for the lined shore crab,Pachygrapsus crassipes, which also forages over intertidal reefs. These results are compared with the availability of algae in intertidal habitats measured during summer and winter, on both disturbed and undisturbed habitats. The diets of juveniles ofG. nigricans andH. azurea collected from nearshore habitats were dominated by animal prey (mainly amphipods), but adults of these fishes, andP. crassipes, consumed algae nearly exclusively, with 26, 10, and 14 taxa of algae identified fromG. nigricans, H. azurea, andP. crassipes, respectively. Algae with sheet-like morphologies (e.g.Ulva sp.,Enteromorpha sp., members of the Delesseriaceae) were the principal algae in the diets of the fishes, and calcareous algae (e.g.Corallina sp.,Lithothrix aspergillum) and sheet-like algae (Enteromorpha sp.) comprised the greatest identifiable portion of the shore crab's diet. Feeding choice experiments indicated that the fishes preferred filamentous algae (e.g.Centroceras clavulatum, Polysiphonia sp.,Chondria californica) and sheet-like algae (e.g.Enteromorpha sp.,Ulva sp.,Cryptopleura crispa) over other algal morphologies, whereas the shore crab chose jointed calcareous algae (e.g.Lithothrix aspergillum, Corallina vancouveriensis, Jania sp.) most frequently. The diets and preferences for algae by the fishes were generally most similar to the assemblage of algae available in early successional (disturbed) habitats during summer when sheet-like and filamentous algae are abundant. The shore crab exhibited the opposite trend with a diet more similar to late successional (undisturbed) habitats.     

Large mobile versus small sedentary herbivores and their resistance to seaweed chemical defenses

Summary Small, relatively sedentary herbivores like amphipods and polychaetes (mesograzers) often live on the plants they consume and should therefore view plants as both foods and living sites. Large, relatively mobile herbivores like fishes commonly move among, and feed from, many plants; they should view plants primarily as foods and rarely as potential living sites. In marine communities, fishes that consume plants are also important predators on mesograzers. Since seaweeds avoided by fishes should represent safer living sites for small herbivores, mesograzers living on and consuming seaweeds that are not eaten by fishes should have higher fitness than mesograzers living on plants preferred by fishes. In previous work, we demonstrated that seaweed secondary metabolites that deterred feeding by a fish and sea urchin had no effect on feeding by a common amphipod (Hay et al. 1987a). We then hypothesized that mesograzers would, in general, be less affected by seaweed chemical defenses than larger, more mobile herbivores like fishes. In this investigation, we evaluate the generality of this hypothesis by comparing the feeding of an omnivorous fish (Lagodon rhomboides) with that of an omnivorous, tube-building polychaete (Platynereis dumerilii) to see if the mesograzer prefers seaweeds avoided by the fish and if it is less affected by seaweed chemical defense. Platynereis dumerilii fed almost exclusively on Dictyota dichotoma, the seaweed eaten least by Lagodon rhomboides. The diterpene alcohols (dictyol-E and pachydictyol-A) produced by Dictyota significantly deterred feeding by Lagodon but did not affect, or at one concentration stimulated, feeding by Platynereis. Our data support the hypothesis that small, relatively sedentary herbivores that live on plants are more resistant to chemical defenses than are large, relatively mobile herbivores that move among many plants.     

Facultative mutualism between an herbivorous crab and a coralline alga: advantages of eating noxious seaweeds

Because encrusting coralline algae rely on herbivory or low light levels to prevent being overgrown by competitively superior fleshy algae, corallines are relatively rare in shallow areas with low rates of herbivory. In contrast to this general trend, the branching coralline alga Neogoniolithon strictum occurs primarily in shallow seagrass beds and along the margins of shallow reef flats where herbivory on macrophytes is low. This alga apparently persists in these habitats by providing refuge to the herbivorous crab Mithrax sculptus at mean densities of 1 crab per 75 g of algal wet mass. When crabs were removed from some host corallines, hosts without crabs supported 9 times the epiphytic growth of hosts with crabs after only 30 days. Crabs without access to a coralline alga were rapidly consumed by reef fishes, while most of those tethered near a host alga survived. These results suggest that the crabs clean their algal host of fouling seaweeds and associate with the host to minimize predation. However, to effectively clean the host, the crab must consume the wide array of macroalgae that commonly co-occur with coralline algae in these habitats, including chemically defended species in the genera Halimeda, Dictyota, and Laurencia. Crabs did readily consume these seaweeds, which were avoided by, and are chemically defended from, herbivorous fishes. Even though crabs readily consumed both Halimeda and Dictyota in whole-plant feeding assays, chemical extracts from these species significantly reduced crab feeding, suggesting that factors other than secondary chemistry (e.g., food value, protein, energy content), may determine whole-plant palatability. Having the ability to use a wide variety of foods, and choosing the most profitable rather than the least defended foods, would diminish foraging time, increase site fidelity, and allow the crab to function mutualistically with the host alga. Despite the obvious benefit of associating with N. strictum, M. sculptus did not prefer it over other habitats offering a structurally similar refuge, suggesting that these crabs are not N. strictum specialists, but rather occupy multiple habitats that provide protection from predators. Structurally complex organisms like N. strictum may commonly suppress competitors by harboring protective symbionts like M. sculptus. It is possible that diffuse coevolution has occurred between these two groups; however, this seems unlikely because both herbivore and host appear to respond most strongly to selective pressures from predators and competitors outside this association.     

Potential Economic seaweeds of Hengchun peninsula,Taiwan

There are 25 genera and 76 species of economically important marine algae found in the Hengchun Peninsula. The greatest number of useful species is found in the Rhodophyta. Sargassum is the most common genus of the Phaeophyta and has the largest biomass and the widest distribution, but fewer useful species. The local people harvest seaweeds from natural beds for food, fodder, fish bait, fertilizer and medical purposes. However, only a few of the available seaweeds are used and in small quantities. Most of the species are seasonal and more abundant in spring and winter than in the summer. Four genera,Sargassum,Enteromorpha,Ulva andCodium have large standing crops and are widely distributed in the coastal water. Production of the other genera is lower. There are many other useful seaweeds present in this area, some of which are also used as landscape plants in aquaria and as fertilizer in horticulture. The potential commercial uses forHalymenia microcarpa are also briefly discussed.     

Mesoherbivores reduce net growth and induce chemical resistance in natural seaweed populations

Herbivory on marine macroalgae (seaweeds) in temperate areas is often dominated by relatively small gastropods and crustaceans (mesoherbivores). The effects of these herbivores on the performance of adult seaweeds have so far been almost exclusively investigated under artificial laboratory conditions. Furthermore, several recent laboratory studies with mesoherbivores indicate that inducible chemical resistance may be as common in seaweeds as in vascular plants. However, in order to further explore and test the possible ecological significance of induced chemical resistance in temperate seaweeds, data are needed that address this issue in natural populations. We investigated the effect of grazing by littorinid herbivorous snails (Littorina spp.) on the individual net growth of the brown seaweed Ascophyllum nodosum in natural field populations. Furthermore, the capacity for induced resistance in the seaweeds was assessed by removing herbivores and assaying for relaxation of defences. We found that ambient densities of gastropod herbivores significantly reduced net growth by 45% in natural field populations of A. nodosum. Seaweeds previously exposed to grazing in the field were less consumed by gastropod herbivores in feeding bioassays. Furthermore, the concentration of phlorotannins (polyphenolics), which have been shown to deter gastropod herbivores, was higher in the seaweeds that were exposed to gastropod herbivores in the field. This field study corroborates earlier laboratory experiments and demonstrates that it is important to make sure that the lack of experimental field data on marine mesoherbivory does not lead to rash conclusions about the lack of significant effects of these herbivores on seaweed performance. The results strongly suggest that gastropods exert a significant selection pressure on the evolution of defensive traits in the seaweeds, and that brown seaweeds can respond to attacks by natural densities of these herbivores through increased chemical resistance to further grazing.     

Algal resistance to herbivory on a Caribbean barrier reef

Field and laboratory research at Carrie Bow Cay, Belize showed that macroalgae, grouped in functional-form units resisted fish and urchin herbivory in the following order (from high to low resistance): Crustose-Group, Jointed Calcareous-Group, Thick Leathery-Group, Coarsely Branched-Group, Filamentous-Group and Sheet-Group; thereby supporting the hypothesis that crustose, calcareous and thick algae have evolved antipredator defenses and should show the greatest resistance to herbivory with a gradation of increasing palatability towards filaments and sheets. Of the 21 species examined, several (e.g.,Dictyota cervicornis on grids,Laurencia obtusa andStypopodium zonale) had exceptionally low losses to fish grazing, probably due to chemical defences. The sea urchin,Diadema antillarum, was more inclined to feed on algae with known toxic secondary metabolites than were herbivorous fishes; hypothetically related to the differences in mobility and concomitant modes of feeding. Tough leathery forms such asSargassum polyceratium andTurbinaria turbinata resisted grazing by bottom feeding parrotfishes (Scaridae) and surgeonfishes (Acanthuridae) but were susceptible when suspended midway in the water column, possibly due to the presence of rudderfishes (Kyphosidae) which readily consume drift Sargassaceae. The overall tendencies support our predicted relationship between grazer-resistance and algal morphology. In conjunction with our previously reported findings concerning primary productivity, toughness and calorimetry for many of the same species, these results lend credence to generalizations relating form with function in marine macroalgae.     

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     

Chemical defenses and the susceptibility of tropical marine brown algae to herbivores

Summary I assayed phenolic and tannin concentrations in a number of species of temperate and tropical brown algae of the genera Sargassum and Turbinaria. Tropical species in both genera contained consistently low levels of phenolics and tannins (species means ranged between 0 and 1.6% [measured as % dry weight of the thallus]). Levels of phenolics in temperate species of Sargassum were variable and consistently much higher than in tropical species (species means ranged between 3 and 12% by dry weight). This pattern of latitudinal variation in phenolic levels in Sargassum conflicts with previous predictions for latitudinal variation in the chemical defenses of marine organisms. The low levels of phenolics present in the tropical species that I analyzed may also explain recent results (Hay 1984; Lewis 1985) demonstrating that tropical Sargassum and Turbinaria are often preferentially consumed by herbivorous fishes and echinoids.     

Marine-terrestrial contrasts in the ecology of plant chemical defenses against herbivores

Small marine herbivores that live on the plants they consume often selectively eat seaweeds that are chemically defended from fishes. Their feeding is unaffected or stimulated by the plant metabolites that deter fishes, and these small herbivores dramatically reduce their susceptibility to predation by associating with host plants that are noxious to fishes. Ecological similarities between these small marine herbivores and numerous terrestrial insects suggest that herbivorous insects also may have evolved a preference for toxic plants because this diminishes their losses to predators, parasites and pathogens. Although marine and terrestrial plants and herbivores evolved in strikingly different environments, the ease of experimentation in some marine systems makes them ideal for addressing certain questions of fundamental importance to both terrestrial and marine workers.     

Scales of dispersal among hosts in a herbivorous marine amphipod

Abstract The spatial arrangement of plants may constrain the expression of herbivore preferences due to interaction between the scales of herbivore mobility and vegetation structure. I tested the hypothesis that the presence of the herbivorous marine amphipod Peramphithoe parmerong (Ampithoidae) on a low‐preference, poor‐quality host (the brown alga Padina crassa) was due to limited dispersal and the clumped nature of algal distributions. In contrast to limited dispersal in laboratory assays, amphipods rapidly colonized vacant algae in the field with natural densities being achieved within 1 day. These high rates of colonization did not decline with increasing distance from existing algal beds over the spatial scales separating Pa. crassa from the higher‐quality host algae (Sargassum spp.). Thus, the spatial arrangement of host algae did not interact with herbivore mobility to constrain host choices.     

The harvesting of macroalgae in New Zealand

Several species of algae have been commercially harvested in New Zealand, mainly for extraction of agar and alginates. In the past, the harvest was comprised mostly of shore-cast plants. There has been more recent interest, however, in harvesting attached plants of Pterocladia spp., Porphyra spp., Gracilaria sordida, Durvillaea spp., Macrocystis pyrifera, and Ecklonia radiata. The ecological effects of harvesting attached algae depend largely on the sizes of plants, the season of removal, the patch size of clearances, and the proximity and identity of mature plants. These have not been well-studied for seaweeds in New Zealand, but population and life history studies indicate that harvesting methods affect the continuity of algal resources, at least on a local scale, and are crucial factors in their management.     

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.     

Economically important seaweeds in Mar Piccolo,Taranto (southern Italy): a survey

A preliminary survey was carried out of the marine algae in the Mar Piccolo Basin, Taranto (southern Italy), in order to assess the degree of sewage pollution and to determine the presence of any economically important species that might be productively utilized. The following commercial species, belonging to a drift algal community, were found: Gracilaria bursapastoris, G. cf. verrucosa, Solieria sp. and S. filiformis. The species showed an aggregated distribution within the community and a clear cyclic annual pattern. Gracilaria bursa-pastoris exhibited the highest mean biomass values in May, whereas Solieria spp. were the most abundant in October. At the present time, only the harvesting of G. bursa-pastoris appears profitable. However, further detailed studies are required to more fully assess the standing crop of the Solieria spp. and G. cf. verrucosa. Considering the environmental situation in the basin, it might be more appropriate to cultivate, rather than harvest, these commercially important seaweeds. With the large number of sewage outlet plants available in the basin, the cultivation of seaweeds in a wastewater treatment-aquaculture system should be considered.     

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

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

Feeding complementarity versus redundancy among herbivorous fishes on a Caribbean reef

Herbivory is an important driver of community structure on coral reefs. Adequate understanding of herbivory will mandate better knowledge of how specific herbivores impact reef communities and the redundancy versus complementarity of their ecological roles. We used algal communities generated by herbivore manipulations to assess such roles among Caribbean herbivorous fishes. We created large enclosures on a 16- to 18-m-deep reef to create treatments grazed for 10 months by: (1) only Sparisoma aurofrenatum, (2) only Acanthurus bahianus, (3) no large herbivorous fishes, or (4) natural densities of all reef fishes. After 10 months, we removed cages and filmed how free-ranging reef fishes fed among these treatments that differed in algal community structure. In general, Acanthurus spp. and Scarus spp. rapidly grazed exclosure and Sparisoma-only treatments, while Sparisoma spp. preferentially grazed exclosure and Acanthurus-only treatments. These patterns suggest complementarity between Sparisoma spp. and both Acanthurus spp. and Scarus spp. but redundancy between Acanthurus spp. and Scarus spp. Despite these generalities, there was also within-genera variance in response to the different treatments. For example, large Scarus spp., such as Scarus guacamaia, fed more similarly to Sparisoma spp., particularly Sparisoma viride, than to other Scarus spp. Moreover, the three common Sparisoma species differed considerably in the macroalgae to which they exhibited positive or negative relationships. Thus, herbivorous reef fishes vary considerably in their response to different algal communities and exhibit complex patterns of compensatory feeding and functional redundancy that are poorly predicted by taxonomy alone.