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     

Sponge-mediated coral reef growth and rejuvenation

Sponges mediate consolidation of Porites furcata rubble on shallow Caribbean reefs by quickly adhering to rubble and stabilizing it until carbonate secreting organisms can grow and consolidate it to the reef. Experimental investigations demonstrate that the entire cycle from (1) temporary binding of rubble by sponges, through (2) rubble consolidation by encrusting coralline algae, to (3) colonization of consolidated rubble by corals, can be completed within 10 months. Bound rubble both adds to vertical reef growth and also provides stable substrata for colonization by corals. Corals that colonize stabilized rubble are damaged less and survive better than on unstable rubble. Rubble that is not temporarily stabilized by sponges does not become bound to the reef, because continuous movement disturbs the consolidation process, and does not provide suitable substrata for settlement and growth of corals. Sponge-mediated consolidation of rubble may increase rates of reef growth and enhance reef recovery after damage. This new role for sponges in reef growth is not obvious from examination of the internal fabric of a reef frame. Spongemediated consolidation may help to explain geographic and temporal differences in growth and morphology among shallow reefs of ramose corals.     

The Acropora inheritance: A reinterpretation of the development of fringing reefs in Barbados,West Indies

The discovery of the widespread occurrence of the remains of the reef coral Acropora palmata within the fabric of the fringing reefs on the west coast of Barbados requires a new interpretation of their Holocene development. Radiocarbon dating of the A. palmata framework suggests that reef construction by this species began as early as 2,300 years B.P. A. palmata probably flourished in Barbados into the present century but has now declined. The present fringing reefs are characterized by a core and base of A. palmata upon which subsequent colonization took place, especially by Montastrea annularis, Porites porites and coralline algae.     

Context-dependent corallivory by parrotfishes in a Caribbean reef ecosystem

This study assesses the patterns of corallivory by parrotfishes across reefs of the Florida Keys, USA. These reefs represent a relatively unique combination within the wider Caribbean of low coral cover and high parrotfish abundance suggesting that predation pressure could be intense. Surveys across eight shallow forereefs documented the abundance of corals, corallivorous parrotfishes, and predation scars on corals. The corals Porites porites and Porites astreoides were preyed on most frequently with the rates of predation an order of magnitude greater than has been documented for other areas of the Caribbean. In fact, parrotfish bite density on these preferred corals was up to 34 times greater than reported for corals on other reefs worldwide. On reefs where coral cover was low and corals such as Montastraea faveolata, often preferred prey for parrotfishes, were rare, predation rates on P. porites and P. astreoides, and other less common corals, intensified further. The intensity of parrotfish predation increased significantly as coral cover decreased. However, parrotfish abundance showed only a marginal positive relationship with predation pressure on corals, likely because corallivorous parrotfish were abundant across all reefs. Parrotfishes often have significant positive impacts on coral cover by facilitating coral recruitment, survival, and growth via their grazing of algae. However, abundant corallivorous parrotfishes combined with low coral cover may result in higher predation on corals and intensify the negative impact that parrotfishes have on remaining corals.     

Recruitment of scleractinians onto the skeletons of corals killed by black band disease

To determine what happens to scleractinian corals that have been killed by black band disease (BBD), massive corals with BBD were monitored for 11 years on a shallow reef (<10 m depth) in St. John, US Virgin Islands. Small quadrats (0.039 m²) were used to compare the rates of scleractinian recruitment to the skeletons of corals killed by either BBD or physical disturbance (Hurricane Hugo 1989). Coral recruitment was also quantified on the adjacent fringing reef using larger quadrats (0.25 m²) to detect possible biases associated with using small, permanent quadrats to assess recruitment to BBD-killed corals. Of 28 tagged colonies with BBD in 1988, 43% were lost to Hurricane Hugo in 1989, 7% were lost to unknown causes between 1991 and 1992, and 14 were monitored annually for 11 years; of these, 71% were dead and still in their original growth position in 1998. Between 1988 and 1997, corals recruited to the BBD-killed surfaces at a rate of 1.1 ± 0.3 recruits · 0.039 m⁻² · decade⁻¹ (mean ± SE, n = 14), although mortality reduced the density to 0.3 ± 0.2 recruits · 0.039 m⁻² by 1997. The rate of recruitment and the taxonomic composition of the coral recruits to BBD-killed corals were indistinguishable statistically from those to corals killed by Hurricane Hugo. This demonstrates that BBD creates space that is functionally the same as other dead coral surfaces in providing a substratum for coral recruitment. However, because coral recruits are dispersed widely, clumped in distribution and temporally variable in density on the fringing reef as a whole, it is unlikely that they will be found on monitored coral colonies that have been killed by BBD. While this hypothesis is consistent with the higher density of recruits on the fringing reef compared with BBD-killed corals, further studies are required to investigate alternative explanations such as the role of substratum age in favoring recruitment to surfaces other than those killed recently by BBD. Accepted: 26 August 1999     

Spatial pattern of coral diversity in Luhuitou fringing reef, Sanya, China

84 quadrats from 5 vertical transects of Luhuitou fringing reef are investigated in detail by using video-quadrat and indoor-interpretation methods. The results show that (1) the reef consists of 69 species of hermatypic corals belonging to 24 genera and 13 families which are unevenly distributed in abundance. (2) Among all the corals, Porites lutea is the most dominant species with importance value percentage up to 36.62%; Porites and Acropora are dominant genera with importance value percentages 43.85% and 22.88%, respectively. (3) There exist distinct spatial differences in coral communities. Both the coral covers and coral diversity indices on the northeastern transects are higher than those on the central and southern transects. (4) Coral communities also show remarkable zonal characteristics with less coral species occurring on reef flat than on reef slope. The importance value percentage of the sole dominant coral genus, Porites, is over 50%, while on the reef slope, the importance value percentages are 28.33% for the first dominant genus Acropora and 26.71% for the second dominant genus Porites. Our further analysis suggests that the spatial and zonal differences of coral diversity pattern are correlated with both natural environmental changes and human activities. The shallow water reef flat is frequently exposed at low tide and it receives more anthropogenic influences (including dredging and trampling) than the deep water reef slope. Thus, the coral community on the reef flat is not as well developed as that on reef slope. The relatively poor coral covers and coral diversity indices on the central and southern transects are closely related to heavy human activities around these sites such as aquaculture, fishing and coastal sewage drainage. Therefore, the impact of human activities must be taken into account in developing strategies for the protection of this coral reef.     

Spatial pattern of coral diversity in Luhuitou fringing reef, Sanya, China

84 quadrats from 5 vertical transects of Luhuitou fringing reef are investigated in detail by using video-quadrat and indoor-interpretation methods. The results show that (1) the reef consists of 69 species of hermatypic corals belonging to 24 genera and 13 families which are unevenly distributed in abundance. (2) Among all the corals, Porites lutea is the most dominant species with importance value percentage up to 36.62%; Porites and Acropora are dominant genera with importance value percentages 43.85% and 22.88%, respectively. (3) There exist distinct spatial differences in coral communities. Both the coral covers and coral diversity indices on the northeastern transects are higher than those on the central and southern transects. (4) Coral communities also show remarkable zonal characteristics with less coral species occurring on reef flat than on reef slope. The importance value percentage of the sole dominant coral genus, Porites, is over 50%, while on the reef slope, the importance value percentages are 28.33% for the first dominant genus Acropora and 26.71% for the second dominant genus Porites. Our further analysis suggests that the spatial and zonal differences of coral diversity pattern are correlated with both natural environmental changes and human activities. The shallow water reef flat is frequently exposed at low tide and it receives more anthropogenic influences (including dredging and trampling) than the deep water reef slope. Thus, the coral community on the reef flat is not as well developed as that on reef slope. The relatively poor coral covers and coral diversity indices on the central and southern transects are closely related to heavy human activities around these sites such as aquaculture, fishing and coastal sewage drainage. Therefore, the impact of human activities must be taken into account in developing strategies for the protection of this coral reef.     

The effect of natural variation in substrate architecture on the survival of juvenile bicolor damselfish

Coral reefs offer settling fish larvae a spatial mosaic of microhabitats that differ not only in structural complexity but also in the abundance and diversity of predators. This paper provides evidence that interactions between predators and prey are causally linked to variation in relevant architectural characteristics of natural substrates. Juvenile bicolor damselfish, Stegastes partitus, experienced greater mortality on Montastrea annularis boulder coral than on piles of Porites porites coral rubble. This pattern was consistent on both back and fore reef habitats. Architectural differences, variable encounter rates with predators, and access to different food sources all contributed to higher mortality. Spatial and structural differences among refuges provided by the two substrates were most important in affecting survival. Porites rubble contained almost three times the number of crevices (39.9 vs. 14.3 crevices m^–2AM²), had a much smaller mean crevice size (15.9 vs. 48.5 cm²), and had a more complex internal structure than Montastrea coral. Survival of juvenile fish living on Montastrea coral was positively correlated with crevice density (r² = 0.41) whereas survival of fish living on Porites rubble was negatively correlated with crevice size (r² = 0.43). These patterns were evident on fore reef habitats only, whereas on back reef habitats no clear patterns emerged. The effect of these natural differences in architecture on mortality rates of juvenile S. partitus was experimentally tested in the field using a combination of natural and artificial substrates. As the number of large shelters was increased I found that the density of potential predators increased but the survival rates of juvenile bicolor damselfish declined. These results highlight the importance of structural architecture among common reef substrates in affecting predator-prey relationships and in determining the survivorship and small-scale distribution patterns of juvenile reef fish.     

An energy budget for Porites porites (Scleractinia), growing in a stressed environment

An energy budget was determined for the coral Porites porites living in a stressed environment for comparison of the energy inputs and expenditure with those of the same species living in an adjacent clear water fore reef environment. The stressed site was characterised by higher sedimentation and lower irradiances than at the fore reef site. Zooplankton ingestion was found to be an insignificant component of the energy intake: the coral is fully autotrophic under stress conditions. The integrated 24 h rate of photosynthetic energy production on a clear sunny day was 20% higher for stressed corals compared to fore reef corals. This was largely the result of photoadaptation which resulted in increased values for and decreased values for I_k in the hyperbolic tangent function equation for the photosynthesis versus irradiance curve. The energy investment in growth of animal tissue was lower in stressed corals. The percentage translocation of photosynthase to the animal tissue remained at about 78%, but the respiration rate of the animal tissue was reduced by 3 fold. These data combined with the high rate of photosynthetic production predict a net daily energy surplus of 67% in stressed corals compared with the 45% surplus in unstressed corals. Scope for growth is reduced under stress conditions.     

Complex interactions in the control of coral zonation on a Caribbean reef flat

Summary This study uses short-term assays and long-term transplant experiments to document the potential importance of fish predation and herbivory to the distribution and abundance of reef-building corals in a Caribbean back-reef system. Experimental manipulations of fish access reveal that the zonal patterns of the two reef-building corals Porites astreoides and P. porites f. furcata, dominant on shallow back-reef habitats, are strongly associated with the feeding intensity of parrotfishes. Differential palatability of the two corals to parrotfishes, the proximity of protective cover for large grazers and the availability of small refugia to harbor a cryptic grazer fauna are suggested as major features contributing to the observed patterns. A model predicting the interactions of various algivore/corallivore guilds on the relative dominance of Porites and algal populations is presented.     

Chronic and catastrophic natural mortality of three common Caribbean reef corals

Compared to catastrophic impacts from storms, disease epidemics and bleaching events, little is known about the effects of more routine chronic mortality in reef corals. To monitor this ongoing mortality, monthly visual assessments of the cause of tissue damage were related to mortality rates (changes in planar surface area) of tagged colonies of three common reef corals: Montastrea annularis, Porites astreoides and Diploria strigosa at Buck Island Reef National Monument, St. Croix, US Virgin Islands. During the study Hurricane Hugo, the most powerful cyclone to affect the area in at least 60 y, made a direct impact on the site. Effects of the hurricane were extremely localized, with certain exposed sites being almost completely razed while others showed no detectable changes in community structure. Mortality caused both by the hurricane and by other factors during the 26 month study varied between species and also between site locations around the island. Differences in susceptibility were not dependent solely on gross morphology, because two robust, massive species showed opposite responses to hurricane damage and chronic mortality. Diploria strigosa was virtually unaffected by chronic factors, but was heavily damaged at exposed sites during the hurricane. In contrast, mortality from predation and tissue necrosis was high in Montastrea annularis, but it largely escaped damage from the hurricane because it was absent from the most severely scoured locations. Porites astreoides, with populations dominated by much smaller colonies, was affected by both chronic and hurricanerelated mortality. Differences in susceptibility to the various types of natural disturbance among species, coupled with high spatial and temporal variability in the effects of such disturbances, may be critical to the maintenance of species diversity on the reef.     

The Effect of Parrotfish (Scaridae) on Coral in Barbados,W. I

Among the five species of parrotfish studied, only largeSparisoma viride (and rarely large Scarus vetula) graze on live coral. Of the several species of corals grazed, Montastrea annularis is grazed most extensively, and variations in its form may be the result of grazing scars. Parrotfish feeding on algae at the bases of the corals Madracis mirabilis and Porites porites break off the corals. This attack causes recession of the stands of M. mirabilis at an average rate of 1 cm/month.     

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.     

BICARBONATE STIMULATION OF CALCIFICATION AND PHOTOSYNTHESIS IN TWO HERMATYPIC CORALS1

A wide range of bicarbonate concentrations was used to monitor the kinetics of bicarbonate (HCO₃^?) use in both photosynthesis and calcification in two reef‐building corals, Porites porites and Acropora sp. Experiments carried out close to the P. porites collection site in Barbados showed that additions of NaHCO₃ to synthetic seawater proportionally increased the calcification rate of this coral until the concentration exceeded three times that of seawater (6 mM). Photosynthetic rates were also stimulated by HCO₃^? addition, but these became saturated at a lower concentration (4 mM). Similar experiments on aquarium‐acclimated colonies of Indo‐Pacific Acropora sp. showed that calcification and photosynthesis in this coral were enhanced to an even greater extent than P. porites, with calcification continuing to increase above 8 mM HCO₃^?, and photosynthesis saturating at 6 mM. Calcification rates of Acropora sp. were also monitored in the dark, and, although these were lower than in the light for a given HCO₃^? concentration, they still increased dramatically with HCO₃^? addition, showing that calcification in this coral is light stimulated but not light dependent.     

Indicators of UV Exposure in Corals and Their Relevance to Global Climate Change and Coral Bleaching

A compelling aspect of the deterioration of coral reefs is the phenomenon of coral bleaching. Through interactions with other factors such as sedimentation, pollution, and bacterial infection, bleaching can impact large areas of a reef with limited recovery, and it might be induced by a variety of stressors including temperature and salinity extremes, and ultraviolet light. Under conditions of ocean warming, often associated with calm and stratified waters, photobleaching of UV-absorb-ing chromophoric dissolved organic matter (CDOM) is increased, and penetration of both UV-B and UV-A is greatly enhanced. Indices of UV-specific effects in coral tissue are needed to test whether UV increases, associated with global climate change, are harmful to corals. To address this challenge, we have evaluated UV-specific effects in corals and have characterized factors that alter penetration of UV radiation over coral reefs. An immunoblotting assay was developed to examine UV-specific lesions (thymine dimers) in coral and zooxanthellae DNA. We observed dose-dependent increases of thymine dimers in coral (Porites porites var porites) exposed to artificial solar irradiance in a solar simulator, although effects were not strictly proportional. UV measurements were made in July 1999 at Eastern Sambo reef and nearby sites, including profiling along transects from reef to shore. Results of these analyses indicate that the coral at Eastern Sambo reef (at 3-4 meters) were receiving UV-B radiation that was equivalent to 25 to 30% of surface UV irradiance. However, the water just inside the reef in Hawk Channel (located closer to land) was considerably more opaque to UV. This water photobleached with loss of UV absorbance and fluorescence when it was exposed to simulated solar radiation. These results indicate that photobleaching of the DOM and transport of near-shore water out over the reefs might play a key role in controlling UV penetration to the reef surface.     

PHOTOSYNTHESIS IN SOME ALCYONACEAN CORALS

Burkholder , Paul R., and Lillian M. Burkholder . (Brooklyn Botanic Garden, Brooklyn, New York.) Photosynthesis in some Alcyonacean corals. Amer. Jour. Bot. 47(10) : 866–872. Illus. 1960.—Quantitative production of oxygen in bottles, and amounts of chlorophyll and carotenoid pigments were studied in selected species of Alcyonacean corals, the stony coral Porites, porites, and turtle grass, Thalassia testudinum, in Puerto Rico. Photosynthetic production exceeds the consumption of oxygen in combined respiration of the corals and their contained algal zooxanthellae, both in short-term light experiments and over 24 hr. of natural, alternating day and night. Observations of oxygen production in 8 species of corals, exposed to full sunlight in 1 ft. of water, gave assimilation values of 0.9 mg. O₂ (equivalent to 0.27 mg. C fixed) per mg. chlorophyll a per hour. A maximum assimilation value of 1.56 was observed in Antillogorgia turgida. From data on the increased oxygen it was calculated that from 0.33 to 1.65 mg. residual fixed carbon per g. of fresh coral were accumulated in Porites porites and Antillogorgia turgida during a day and night period of 24 hr. Photosynthesis of turtle grass in the same environment was more than 3 times the rate of these corals. The zooxanthellae contain carotenoids in abundance, along with chlorophylls a and c. On a unit of chlorophyll a basis, the photosynthetic systems of corals appear to be less active in reduction of carbon dioxide in bottles than some other photosynthetic systems of the sea, such as turtle grass and phytoplankton observed under optimum conditions.     

Effects of eutrophication on bioeroding sponge communities with the description of new West Indian sponges, Cliona spp. (Porifera: Hadromerida: Clionidae)

Abstract. Pieces of coral rubble ( Porites porites ), collected from across 3 fringing reefs that lie along a eutrophication gradient, were examined for the presence of clionid sponges. A similarity analysis of species composition showed that reef zone had less effect on clionid community composition than did other factors affecting the reefs as a whole. Except on the back reef, the Zones, distances, and depths within the reefs had no significant influence on the number of clionid invasions. Reef comparisons demonstrated that clionid abundance increased with increasing eutrophication. Clionids were found in 41% of the pieces collected from the most eutrophic site vs. 24% from the least eutrophic. Because clionids are the principal bio-eroders of coral reefs, any increase in their abundance will likely result in greater bioerosion rates. The mean abundance of Type 3 corals (in which fragmentation is the primary mode of propagation) is positively related to the frequency of boring sponge invasion. suggesting that increased bioerosion may be partly responsible for community shifts toward Type 3 corals in polluted waters. Cliona cf. vastifica , found for the first time in Barbados, flourishes on the most eutrophic reef and may become an important bioeroder under the highly eutrophic conditions that have begun to plague West Indian reefs. Two new species of Cliona (Porifera: Hadro-merida: Clionidae) are described.     

Newly dominant benthic invertebrates reshape competitive networks on contemporary Caribbean reefs

Competition is a fundamental process structuring ecological communities. On coral reefs, space is a highly contested resource and the outcomes of spatial competition can dictate community composition. In the Caribbean, reefs are increasingly dominated by non-scleractinian species like sponges, gorgonians, and zoanthids, yet there is a paucity of data on interactions between these increasingly common organisms and historically dominant corals. Here, we investigated interactions among these groups of sessile benthic invertebrates to better understand the role of spatial competition in shaping benthic communities on Caribbean reefs. We coupled surveys of competitive interactions on the reef with a common garden competition experiment to determine the frequency and outcome of interference competition among eight focal species. We found that competitive interactions were pervasive on Florida reefs, with 60% of sessile benthic invertebrates interacting with at least one other invertebrate. Increasingly common non-scleractinian species were some of the most abundant taxa and consistently outcompeted the contemporarily common scleractinian species Porites porites and Siderastrea siderea. The encrusting gorgonian, Erythropodium caribaeorum, was the most aggressive species, reducing the live area of its competitors on average 42% ± 7.04 (SE) over the course of 5 months. Surprisingly, the most aggressive species declined in size when competing, while some less aggressive species were able to increase or maintain area, suggesting a trade-off between aggressiveness and growth. Our findings suggest that competition among sessile invertebrates is likely to remain an important process in structuring coral reefs, but that the optimal strategies for maintaining space on the benthos may change. Importantly, many non-scleractinian species that now dominate reefs appear to be superior competitors, potentially increasing the stress on corals on contemporary reefs.

     

Herbivory by crabs and the control of algal epibionts on Caribbean host corals

Summary A short-term experiment was conducted to examine the relationships among the branching coral Porites porites, algal epibionts, and a facultative crab associate Mithrax sculptus in Belize, Central America. Initial field observations suggested that coral colonies supporting resident crabs generally had lower algal cover than colonies without crabs. The hypothesis was tested that Mithrax significantly depresses host coral algal cover and thereby indirectly affects host survivorship and growth. Crab accessibility to an array of coral colonies, similarly covered with algal epibionts, was manipulated in three treatments. Results strongly support the hypothesis, with significant differences in algal cover (primarily Dictyota spp.) noted among treatments after only one month. Caged heads with crabs included and uncaged natural controls allowing crabs free access averaged less than 10% cover, whereas mean algal cover exceeded 75% where crabs were excluded. The uncaged treatment, in which crabs were allowed free access to Porites heads was not significantly different from the crab inclusion treatment. Collectively, these results demonstrate that under natural conditions, crabs can have pronounced effects on host corals by reducing fouling algal epibionts. Furthermore, these facultative coral associates may have more important, albeit localized effects on Caribbean corals than has been suggested previously.