Influence of sexual selection and feeding functional morphology on diversification rate of parrotfishes (Scaridae)

Scaridae (parrotfishes) is a prominent clade of 96 species that shape coral reef communities worldwide through their actions as grazing herbivores. Phylogenetically nested within Labridae, the profound ecological impact and high species richness of parrotfishes suggest that their diversification and ecological success may be linked. Here, we ask whether parrotfish evolution is characterized by a significant burst of lineage diversification and whether parrotfish diversity is shaped more strongly by sexual selection or modifications of the feeding mechanism. We first examined scarid diversification within the greater context of labrid diversity. We used a supermatrix approach for 252 species to propose the most extensive phylogenetic hypothesis of Labridae to date, and time-calibrated the phylogeny with fossil and biogeographical data. Using divergence date estimates, we find that several parrotfish clades exhibit the highest diversification rates among all labrid lineages. Furthermore, we pinpoint a rate shift at the shared ancestor of Scarus and Chlorurus, a scarid subclade characterized by territorial behaviour and strong sexual dichromatism, suggesting that sexual selection was a major factor in parrotfish diversification. Modifications of the pharyngeal and oral jaws that happened earlier in parrotfish evolution may have contributed to this diversity by establishing parrotfishes as uniquely capable reef herbivores.     

Community structure and habitat associations of parrotfishes on Oahu,Hawaii

Despite their ecological importance as bioeroders and their economic importance in commercial, artisanal, and recreational fisheries, there have been relatively few studies on parrotfish (Scaridae) ecology in Hawaii. Belt transects were conducted around the island of Oahu to survey current parrotfish distributions, size structure, species composition and associated habitats. Scarid communities in this heavily fished region are dominated by smaller species and smaller individuals within all species. Specific habitat characteristics such as rugosity, substrate diversity, and percent live coral cover were positively correlated with scarid numerical abundance. Scarids, however, were patchily distributed and were often absent from preferable habitats, suggesting that intense fishing pressure may be an important factor preventing these fish from fully exploiting available habitats. This study is the first thorough, broad-scale investigation of scarid community structure in Hawaii, and provides important information that has management and conservation implications for parrotfish in Hawaii and throughout tropical coral reef ecosystems.     

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.     

Direct and indirect effects of nursery habitats on coral‐reef fish assemblages,grazing pressure and benthic dynamics

Migrating species are common within seascapes, but the potential for these movements to alter the populations and functional roles of non‐migrating species (e.g. by increasing predation) is rarely investigated. This study considers whether the presence of nursery habitats (mangroves and seagrass) simply enhances the abundance of nursery‐using parrotfishes and piscivores on nearby coral reefs, or also affects other parrotfishes. Data from 131 reef sites and multiple seascape configurations across 13 degrees of latitude were used to model correlations between biophysical variables, including nursery habitat connectivity, and the abundance and grazing pressure of both nursery‐using species and other parrotfishes and piscivore biomass. Connectivity to mangroves and dense seagrass was positively correlated with the biomass of nursery‐using species, but was also negatively correlated with non‐nursery parrotfish populations. This reduction may be caused indirectly by nursery habitats increasing confamilial competition and predation by nursery‐using piscivores, particularly affecting small parrotfishes settling directly onto reefs. As key reef grazers, parrotfishes affect coral demographics. Consequently, a spatial simulation model predicted the impacts after five years of changes in grazing pressure because of nursery habitat connectivity. The model demonstrated that high nursery connectivity was correlated to changes in grazing pressure on nearby reefs that could potentially lead to differences in coral cover of ～3–4% when compared to low connectivity reefs. However, the direction of this change depended on the seascapes’ characteristics. Historically, large‐bodied, nursery‐using parrotfish would have increased grazing in all nursery‐rich seascapes. Overfishing means that nursery availability may have spatially variable impacts on coral cover, influencing reserve design. This study suggests that nursery availability may directly and indirectly modify an ecological process, and alter an ecological cascade (migrating species increase predator and competitor abundances, affecting other grazers and consequently corals). Therefore, elucidating the multi‐species impacts of animal movements is required to better understand ecosystem functioning.     

Discriminating causes from consequences of persistent parrotfish corallivory

A detailed understanding of the dual role of parrotfish as both key herbivores and potentially important corallivores is essential to the study of coral health and reef trophodynamics. Some Caribbean parrotfish regularly consume live coral, and discriminate both among coral species and among colonies within a particular species. While they prefer Montastraea spp. corals, which are dominant Caribbean reef builders, causes of selective and persistent grazing of certain colonies remain unknown. We manipulated coral exposure to parrotfish grazing through a long-term cage exclusion experiment in Belize, comparing initially grazed vs. intact (non-grazed) Montastraea spp. colonies. We measured nutrition-related characteristics (C:N ratio, %C, and %N) as well as defensive characteristics (nematocyst density and skeletal hardness) to determine if any of these variables accurately predicted parrotfish grazing. There were substantial reductions in coral nutritional quality (C:N) associated with parrotfish grazing, although these changes appear to be a consequence rather than a cause of parrotfish selectivity. Likewise, nematocyst densities were suppressed in grazed corals, also likely a result of chronic grazing stress. We found no intraspecific differences in skeletal hardness related to grazing. These results provide further demonstration of the physiological consequences of grazing, but the cause of preferential grazing by parrotfishes on certain Montastraea spp. colonies still requires further investigation.     

Parrotfish predation on massive Porites on the Great Barrier Reef

Parrotfish grazing scars on coral colonies were quantified across four reef zones at Lizard Island, Northern Great Barrier Reef (GBR). The abundance of parrotfish grazing scars was highest on reef flat and crest, with massive Porites spp. colonies having more parrotfish grazing scars than all other coral species combined. Massive Porites was the only coral type positively selected for grazing by parrotfishes in all four reef zones. The density of parrotfish grazing scars on massive Porites spp., and the rate of new scar formation, was highest on the reef crest and flat, reflecting the lower massive Porites cover and higher parrotfish abundance in these habitats. Overall, it appears that parrotfish predation pressure on corals could affect the abundance of preferred coral species, especially massive Porites spp, across the reef gradient. Parrotfish predation on corals may have a more important role on the GBR reefs than previously thought.     

Quantifying production rates and size fractions of parrotfish‐derived sediment: A key functional role on Maldivian coral reefs

Coral reef fish perform numerous important functional roles on coral reefs. Of these, carbonate sediment production, as a by‐product of parrotfish feeding, is especially important for contributing to reef framework construction and reef‐associated landform development. However, only limited data exist on: (i) how production rates vary among reef habitats as a function of parrotfish assemblages, (ii) the relative importance of sediment produced from eroded, reworked, and endogenous sources, or (iii) the size fractions of sediment generated by different parrotfish species and size classes. These parameters influence not only overall reef‐derived sediment supply, but also influence the transport potential and depositional fate of this sedimentary material. Here, we show that parrotfish sediment production varies significantly between reef‐platform habitats on an atoll‐margin Maldivian reef. Highest rates of production (over 0.8 kg m⁻² year⁻¹) were calculated in three of the eight platform habitats; a rubble‐dominated zone, an Acropora spp. dominated zone, and a patch reef zone. Habitat spatial extent and differences in associated parrotfish assemblages strongly influenced the total quantities of sediment generated within each habitat. Nearly half of total parrotfish sediment production occurred in the rubble habitat, which comprised only 8% of the total platform area. Over 90% of this sedimentary material originated from eroded reef framework as opposed to being reworked existing or endogenously produced sediment, and comprised predominantly coral sands (predominantly 125–1000 µm in diameter). This is comparable to the dominant sand types and size fractions found on Maldivian reef islands. By contrast, nearly half of the sediment egested by parrotfish in the Acropora spp. dominated and patch reef habitats resulted from reworked existing sediments. These differences between habitats are a result of the different parrotfish assemblages supported. Endogenous carbonate production was found to be insignificant compared to the quantity of eroded and reworked material. Our findings have important implications for identifying key habitats and species which act as major sources of sediment for reef‐island systems.     

The dynamics of architectural complexity on coral reefs under climate change

One striking feature of coral reef ecosystems is the complex benthic architecture which supports diverse and abundant fauna, particularly of reef fish. Reef‐building corals are in decline worldwide, with a corresponding loss of live coral cover resulting in a loss of architectural complexity. Understanding the dynamics of the reef architecture is therefore important to envision the ability of corals to maintain functional habitats in an era of climate change. Here, we develop a mechanistic model of reef topographical complexity for contemporary Caribbean reefs. The model describes the dynamics of corals and other benthic taxa under climate‐driven disturbances (hurricanes and coral bleaching). Corals have a simplified shape with explicit diameter and height, allowing species‐specific calculation of their colony surface and volume. Growth and the mechanical (hurricanes) and biological erosion (parrotfish) of carbonate skeletons are important in driving the pace of extension/reduction in the upper reef surface, the net outcome being quantified by a simple surface roughness index (reef rugosity). The model accurately simulated the decadal changes of coral cover observed in Cozumel (Mexico) between 1984 and 2008, and provided a realistic hindcast of coral colony‐scale (1–10 m) changing rugosity over the same period. We then projected future changes of Caribbean reef rugosity in response to global warming. Under severe and frequent thermal stress, the model predicted a dramatic loss of rugosity over the next two or three decades. Critically, reefs with managed parrotfish populations were able to delay the general loss of architectural complexity, as the benefits of grazing in maintaining living coral outweighed the bioerosion of dead coral skeletons. Overall, this model provides the first explicit projections of reef rugosity in a warming climate, and highlights the need of combining local (protecting and restoring high grazing) to global (mitigation of greenhouse gas emissions) interventions for the persistence of functional reef habitats.     

Herbivorous fish assemblages and herbivory pressure on Laamu Atoll, Republic of Maldives

While herbivory is recognized as a fundamental process structuring coral reef communities, herbivore assemblages and processes are poorly described for reefs in the Indian Ocean region. We quantified herbivorous fish assemblage structure (abundance and diversity) in Laamu Atoll, Republic of Maldives, in four reef habitat types: faro reef flats, faro reef slopes, inner and outer atoll reef slopes (20 sites in total). Herbivorous fish assemblages, representing a total of 30 species, grouped strongly by habitat type, with the highest absolute abundance observed on faro reef flats and lowest abundance on inside atoll rim reef slopes. Removal of Thalassia seagrass blades by ambient herbivore assemblages was used in a bioassay to assess relative herbivory pressure among four habitat types (eight sites). Also, at one site a choice herbivory assay was performed to assess herbivore preference among four benthic plants across three depth zones. Relative herbivory, as indicated by Thalassia assays, was highest on inside atoll rim reef slopes and lowest on outside atoll rim reef slopes. Thalassia consumption did not correspond to overall herbivorous fish abundance, but corresponded more closely with parrotfish abundance. In the choice assays, herbivores showed strong preferences among plant types and consumption of most plant types was higher at mid-depth than in the shallow reef flat or deep reef knoll zones.     

Different Surrounding Landscapes may Result in Different Fish Assemblages in East African Seagrass Beds

Few studies have considered how seagrass fish assemblages are influenced by surrounding habitats. This information is needed for a better understanding of the connectivity between tropical coastal ecosystems. To study the effects of surrounding habitats on the composition, diversity and densities of coral reef fish species on seagrass beds, underwater visual census surveys were carried out in two seagrass habitat types at various locations along the coast of Zanzibar (Tanzania) in the western Indian Ocean. Fish assemblages of seagrass beds in a marine embayment with large areas of mangroves (bay seagrasses) situated 9 km away from coral reefs were compared with those of seagrass beds situated on the continental shelf adjacent to coral reefs (reef seagrasses). No differences in total fish density, total species richness or total juvenile fish density and species richness were observed between the two seagrass habitat types. However, at species level, nine species showed significantly higher densities in bay seagrasses, while eight other species showed significantly higher densities in reef seagrasses. Another four species were exclusively observed in bay seagrasses. Since seagrass complexity could not be related to these differences, it is suggested that the arrangement of seagrass beds in the surrounding landscape (i.e. the arrangement on the continental shelf adjacent to the coral reef, or the arrangement in an embayment with mangroves situated away from reefs) has a possible effect on the occurrence of various reef-associated fish species on seagrass beds. Fish migration from or to the seagrass beds and recruitment and settlement patterns of larvae possibly explain these observations. Juvenile fish densities were similar in the two types of seagrass habitats indicating that seagrass beds adjacent to coral reefs also function as important juvenile habitats, even though they may be subject to higher levels of predation. On the contrary, the density and species richness of adult fish was significantly higher on reef seagrasses than on bay seagrasses, indicating that proximity to the coral reef increases density of adult fish on reef seagrasses, and/or that ontogenetic shifts to the reef may reduce adult density on bay seagrasses.     

Variation in fish density,assemblage composition and relative rates of predation among mangrove,seagrass and coral reef habitats

We tested the hypothesis for several Caribbean reef fish species that there is no difference in nursery function among mangrove, seagrass and shallow reef habitat as measured by: (a) patterns of juvenile and adult density, (b) assemblage composition, and (c) relative predation rates. Results indicated that although some mangrove and seagrass sites showed characteristics of nursery habitats, this pattern was weak. While almost half of our mangrove and seagrass sites appeared to hold higher proportions of juvenile fish (all species pooled) than did reef sites, this pattern was significant in only two cases. In addition, only four of the six most abundant and commercially important species (Haemulon flavolineatum, Haemulon sciurus, Lutjanus apodus, Lutjanus mahogoni, Scarus iserti, and Sparisoma aurofrenatum) showed patterns of higher proportions of juvenile fish in mangrove and/or seagrass habitat(s) relative to coral reefs, and were limited to four of nine sites. Faunal similarity between reef and either mangrove or seagrass habitats was low, suggesting little, if any exchange between them. Finally, although relative risk of predation was lower in mangrove/seagrass than in reef habitats, variance in rates was substantial suggesting that not all mangrove/seagrass habitats function equivalently. Specifically, relative risk varied between morning and afternoon, and between sites of similar habitat, yet varied little, in some cases, between habitats (mangrove/seagrass vs. coral reefs). Consequently, our results caution against generalizations that all mangrove and seagrass habitats have nursery function.     

Post-larval French grunts (Haemulon flavolineatum) distinguish between seagrass, mangrove and coral reef water: Implications for recognition of potential nursery habitats

Environmental cues like sound, magnetic field, oceanic currents, water chemistry or habitat structure are believed to play an important role in the orientation of reef fish towards their settlement habitat. Some species of coral reef fish are known to use seagrass beds and mangroves as juvenile habitats. Once oceanic larvae of these fish have located a coral reef from the open ocean, they still have to find embayments or lagoons harbouring these juvenile habitats. The sensory mechanisms that are used for this are still unknown. In the present study, experiments were conducted to investigate if recruits of the French grunt (Haemulon flavolineatum) respond to habitat differences in water type, as mangrove/seagrass water may differ in biotic and abiotic compounds from coral reef water. Our results show that post-larvae of a reef fish that is highly associated with mangroves and seagrass beds during its juvenile life stage, choose significantly more often for water from mangroves and seagrass beds than for water from the coral reef. These results provide a more detailed insight in the mechanisms that play a role in the detection of these juvenile habitats.     

Kenyan coral reef lagoon fish: effects of fishing,substrate complexity,and sea urchins

Population density, number of species, diversity, and species-area relationships of fish species in eight common coral reef-associated families were studied in three marine parks receiving total protection from fishing, four sites with unregulated fishing, and one reef which recently received protection from fishing (referred to as a transition reef). Data on coral cover, reef topographic complexity, and sea urchin abundance were collected and correlated with fish abundance and species richness. The most striking result of this survey is a consistent and large reduction in the population density and species richness of 5 families (surgeonfish, triggerfish, butterflyfish, angelfish, and parrotfish). Poor recovery of parrotfish in the transition reef, relative to other fish families, is interpreted as evidence for competitive exclusion of parrotfish by sea urchins. Reef substrate complexity is significantly associated with fish abundance and diversity, but data suggest different responses for protected versus fished reefs, protected reefs having higher species richness and numbers of individuals than unprotected reefs for the same reef complexity. Sea urchin abundance is negatively associated with numbers of fish and fish species but the interrelationship between sea urchins, substrate complexity, coral cover, and management make it difficult to attribute a set percent of variance to each factor-although fishing versus no fishing appears to be the strongest variable in predicting numbers of individuals and species of fish, and their community similarity. Localized species extirpation is evident for many species on fished reefs (for the sampled area of 1.0 ha). Fifty-two of 110 species found on protected reefs were not found on unprotected reefs.     

Mangrove Habitat Use by Juvenile Reef Fish: Meta-Analysis Reveals that Tidal Regime Matters More than Biogeographic Region

Identification of critical life-stage habitats is key to successful conservation efforts. Juveniles of some species show great flexibility in habitat use while other species rely heavily on a restricted number of juvenile habitats for protection and food. Considering the rapid degradation of coastal marine habitats worldwide, it is important to evaluate which species are more susceptible to loss of juvenile nursery habitats and how this differs across large biogeographic regions. Here we used a meta-analysis approach to investigate habitat use by juvenile reef fish species in tropical coastal ecosystems across the globe. Densities of juvenile fish species were compared among mangrove, seagrass and coral reef habitats. In the Caribbean, the majority of species showed significantly higher juvenile densities in mangroves as compared to seagrass beds and coral reefs, while for the Indo-Pacific region seagrass beds harbored the highest overall densities. Further analysis indicated that differences in tidal amplitude, irrespective of biogeographic region, appeared to be the major driver for this phenomenon. In addition, juvenile reef fish use of mangroves increased with increasing water salinity. In the Caribbean, species of specific families (e.g. Lutjanidae, Haemulidae) showed a higher reliance on mangroves or seagrass beds as juvenile habitats than other species, whereas in the Indo-Pacific family-specific trends of juvenile habitat utilization were less apparent. The findings of this study highlight the importance of incorporating region-specific tidal inundation regimes into marine spatial conservation planning and ecosystem based management. Furthermore, the significant role of water salinity and tidal access as drivers of mangrove fish habitat use implies that changes in seawater level and rainfall due to climate change may have important effects on how juvenile reef fish use nearshore seascapes in the future.     

Comparison between community structures of fishes in <Emphasis Type="Italic">Enhalus acoroides</Emphasis>- and <Emphasis Type="Italic">Thalassia hemprichii</Emphasis>-dominated seagrass beds on fringing coral reefs in the Ryukyu Islands,Japan

To clarify differences in community structures and habitat utilization patterns of fishes in Enhalus acoroides- and Thalassia hemprichii-dominated seagrass beds on fringing coral reefs, visual censuses were conducted at Iriomote and Ishigaki islands, southern Japan. The numbers of fish species and individuals were significantly higher in the E. acoroides bed than in the T. hemprichii bed, although the 15 most dominant fishes in each seagrass bed were similar. Cluster and ordination analyses based on the number of individuals of each fish species also demonstrated that fish community structures were similar in the two seagrass beds. Species and individual numbers of coral reef fishes which utilized the seagrass beds numbered less than about 15% of whole coral reef fish numbers, although they comprised about half of the seagrass bed fishes. Of the 15 most dominant species, 5 occurred only in the two seagrass beds, including seagrass feeders. Ten other species were reef species, their habitat utilization patterns not differing greatly between the two seagrass beds. Some reef species, such as Lethrinus atkinsoni and L. obsoletus, showed ontogenetic habitat shifts with growth, from the seagrass beds to the coral areas. These results indicate that community structures and habitat utilization patterns of fishes were similar between E. acoroides- and T. hemprichii-dominated seagrass beds, whereas many coral reef fishes hardly utilized the seagrass beds.     

Persistence and Change in Community Composition of Reef Corals through Present,Past, and Future Climates

The reduction in coral cover on many contemporary tropical reefs suggests a different set of coral community assemblages will dominate future reefs. To evaluate the capacity of reef corals to persist over various time scales, we examined coral community dynamics in contemporary, fossil, and simulated future coral reef ecosystems. Based on studies between 1987 and 2012 at two locations in the Caribbean, and between 1981 and 2013 at five locations in the Indo-Pacific, we show that many coral genera declined in abundance, some showed no change in abundance, and a few coral genera increased in abundance. Whether the abundance of a genus declined, increased, or was conserved, was independent of coral family. An analysis of fossil-reef communities in the Caribbean revealed changes in numerical dominance and relative abundances of coral genera, and demonstrated that neither dominance nor taxon was associated with persistence. As coral family was a poor predictor of performance on contemporary reefs, a trait-based, dynamic, multi-patch model was developed to explore the phenotypic basis of ecological performance in a warmer future. Sensitivity analyses revealed that upon exposure to thermal stress, thermal tolerance, growth rate, and longevity were the most important predictors of coral persistence. Together, our results underscore the high variation in the rates and direction of change in coral abundances on contemporary and fossil reefs. Given this variation, it remains possible that coral reefs will be populated by a subset of the present coral fauna in a future that is warmer than the recent past.     

Non‐reef habitats in a tropical seascape affect density and biomass of fishes on coral reefs

Nonreef habitats such as mangroves, seagrass, and macroalgal beds are important for foraging, spawning, and as nursery habitat for some coral reef fishes. The spatial configuration of nonreef habitats adjacent to coral reefs can therefore have a substantial influence on the distribution and composition of reef fish. We investigate how different habitats in a tropical seascape in the Philippines influence the presence, density, and biomass of coral reef fishes to understand the relative importance of different habitats across various spatial scales. A detailed seascape map generated from satellite imagery was combined with field surveys of fish and benthic habitat on coral reefs. We then compared the relative importance of local reef (within coral reef) and adjacent habitat (habitats in the surrounding seascape) variables for coral reef fishes. Overall, adjacent habitat variables were as important as local reef variables in explaining reef fish density and biomass, despite being fewer in number in final models. For adult and juvenile wrasses (Labridae), and juveniles of some parrotfish taxa (Chlorurus), adjacent habitat was more important in explaining fish density and biomass. Notably, wrasses were positively influenced by the amount of sand and macroalgae in the adjacent seascape. Adjacent habitat metrics with the highest relative importance were sand (positive), macroalgae (positive), and mangrove habitats (negative), and fish responses to these metrics were consistent across fish groups evaluated. The 500‐m spatial scale was selected most often in models for seascape variables. Local coral reef variables with the greatest importance were percent cover of live coral (positive), sand (negative), and macroalgae (mixed). Incorporating spatial metrics that describe the surrounding seascape will capture more holistic patterns of fish–habitat relationships on reefs. This is important in regions where protection of reef fish habitat is an integral part of fisheries management but where protection of nonreef habitats is often overlooked.     

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.     

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.     

Reciprocal facilitation and non‐linearity maintain habitat engineering on coral reefs

Ecosystem engineers that create habitats facilitate the coexistence of many interacting species. This biotic response to habitat engineering may result in non‐intuitive cascading interactions, potentially including feedbacks to the engineer. Such feedback mechanisms, either positive or negative, may be especially important for the maintenance of biogenic habitats and their community‐wide facilitation. Here, we describe the complex interactions and feedbacks that link marine habitat‐forming engineers, the reef‐building corals, and a group of herbivores, the parrotfishes; the latter preventing the overgrowth of macroalgae, a major competitor of corals. Using density data of eight parrotfish species on a Caribbean reef, we first describe the form of the response of parrotfish abundance to increasing topographic complexity generated by coral growth. Topographic complexity enhanced parrotfish abundance by promoting habitat suitability, but the shape (linear vs asymptotic) and strength of this response varied across species and size. Parrotfish grazing intensity, estimated from data on abundance and species‐, size‐ and life phase‐specific grazing rates also increased with topographic complexity despite an increase in the surface area over which parrotfish graze. Depending on fish species, this functional response was found to be linear or asymptotic. Using a simple analytical model we then explored the effects of topographic complexity and fishing pressure on coral‐algal competition, with particular emphasis on the implications of non‐linearities in the intensity of grazing. Simulations demonstrate that fishing and habitat degradation impair the performance of grazing, but that an asymptotic response of grazing intensity to topographic complexity increases the ecological resilience of coral reefs. Parrotfish and corals are mutually beneficial by creating a loop of positive, indirect feedbacks that maintain their own structure and function: coral growth promotes habitat suitability for parrotfish, concordantly enhancing grazing intensity, which in turn facilitates coral growth by reducing competitive exclusion by macroalgae. We conclude that the resilience of biogenic habitats is enhanced by non‐linear biotic responses to engineering and by the emergence of reciprocal facilitation linking habitat engineering and response organisms.