Coral habitat in the Aleutian Islands of Alaska: depth distribution, fine-scale species associations, and fisheries interactions

The first in situ exploration of Aleutian Island coral habitat was completed in 2002 to determine the distribution of corals, to examine fine-scale associations between targeted fish species and corals, and to investigate the interaction between the areas’ diverse fisheries and coral habitat. Corals, mostly gorgonians and hydrocorals, were present on all 25 seafloor transects and at depths between 27 and 363 m, but were most abundant between 100 and 200 m depth. Mean coral abundance (1.23 colonies m⁻²) far exceeded that reported for other high-latitude ecosystems and high-density coral gardens (3.85 colonies m⁻²) were observed at seven locations. Slope and offshore pinnacle habitats characterized by exposed bedrock, boulders, and cobbles generally supported the highest abundances of coral and fish. Overall, 85% of the economically important fish species observed on transects were associated with corals and other emergent epifauna. Disturbance to the seafloor from bottom-contact fishing gear was evident on 88% of the transects, and approximately 39% of the total area of the seafloor observed had been disturbed. Since cold-water corals appear to be a ubiquitous feature of seafloor habitats in the Aleutian Islands, fisheries managers face clear challenges integrating coral conservation into an ecosystem approach to fisheries management.     

Microbial consortia of gorgonian corals from the Aleutian islands

Gorgonians make up the majority of corals in the Aleutian archipelago and provide critical fish habitat in areas of economically important fisheries. The microbial ecology of the deep-sea gorgonian corals Paragorgea arborea, Plumarella superba, and Cryogorgia koolsae was examined with culture-based and 16S rRNA gene-based techniques. Six coral colonies (two per species) were collected. Samples from all corals were cultured, and clone libraries were constructed from P. superba and C. koolsae. Cultured bacteria were dominated by the Gammaproteobacteria, especially Vibrionaceae, with other phyla comprising <6% of the isolates. The clone libraries showed dramatically different bacterial communities between corals of the same species collected at different sites, with no clear pattern of conserved bacterial consortia. Two of the clone libraries (one from each coral species) were dominated by Tenericutes, with Alphaproteobacteria dominating the remaining sequences. The other libraries were more diverse and had a more even distribution of bacterial phyla, showing more similarity between genera than within coral species. Here we report the first microbiological characterization of P. arborea, P. superba, and C. koolsae.     

Comparing total body lipid content of young‐of‐the‐year Steller sea lions among regions of contrasting population trends

Steller sea lion (Eumetopias jubatus) young‐of‐the‐year (YOY) are nutritionally dependent upon their dam through the majority of their first year. Several indices of body condition were measured in YOY 1.5–9 mo of age captured in Southeast Alaska (n = 122), the Gulf of Alaska (n = 182), and the Aleutian Islands (n = 38) to test the hypothesis that nutritional stress impacted the ability of adult female Steller sea lions to adequately nourish their late gestation YOY in the central Aleutian Islands in the early 2000s. Body mass (kg) and percent total body lipid content (%TBL) increased with age in all three regions of Alaska that were sampled (P < 0.05). Young‐of‐the‐year 7–9 mo of age were leaner in Southeast Alaska (27.6% ± 1.0%) and Gulf of Alaska (29.5% ± 0.8%) than in the Aleutian Islands (35.7% ± 1.2%, P < 0.001). Condition indices calculated from morphometric measures did not strongly predict the %TBL measured by isotope dilution. The trend for Aleutian Island YOY to have larger body mass and larger body fat reserves are counter to what would be expected if dams were unable to adequately provision their late lactation YOY due to inadequate food availability in the central Aleutian Islands.     

Within-colony feeding selectivity by a corallivorous reef fish: foraging to maximize reward?

Foraging theory predicts that individuals should choose a prey that maximizes energy rewards relative to the energy expended to access, capture, and consume the prey. However, the relative roles of differences in the nutritive value of foods and costs associated with differences in prey accessibility are not always clear. Coral‐feeding fishes are known to be highly selective feeders on particular coral genera or species and even different parts of individual coral colonies. The absence of strong correlations between the nutritional value of corals and prey preferences suggests other factors such as polyp accessibility may be important. Here, we investigated within‐colony feeding selectivity by the corallivorous filefish, Oxymonacanthus longirostris, and if prey accessibility determines foraging patterns. After confirming that this fish primarily feeds on coral polyps, we examined whether fish show a preference for different parts of a common branching coral, Acropora nobilis, both in the field and in the laboratory experiments with simulated corals. We then experimentally tested whether nonuniform patterns of feeding on preferred coral species reflect structural differences between polyps. We found that O. longirostris exhibits nonuniform patterns of foraging in the field, selectively feeding midway along branches. On simulated corals, fish replicated this pattern when food accessibility was equal along the branch. However, when food access varied, fish consistently modified their foraging behavior, preferring to feed where food was most accessible. When foraging patterns were compared with coral morphology, fish preferred larger polyps and less skeletal protection. Our results highlight that patterns of interspecific and intraspecific selectivity can reflect coral morphology, with fish preferring corals or parts of coral colonies with structural characteristics that increase prey accessibility.     

Coral reefs in the Southern Barents Sea: habitat description and the effects of bottom fishing

Fifteen Lophelia reefs from offshore to coastal areas off northern Norway were studied using video. Health status of the coral habitat (degree of physical impact, % cover of living tissue, colony size), occurrence of trawl marks and lost fishing gear, height of coral colonies and associated fauna were analysed from 44 video-lines. Fishing impact was more frequent on the offshore reefs (36.5% of the observed areas) than those in the coastal reefs (0.6%). The most visible effects of fishing were broken and displaced coral colonies. At some sites only small scattered fragments of live corals were observed, indicating recent impact. The mean colony height of Lophelia and gorgonian corals at impacted sites was around half the size of those at non-impacted sites. Both species richness and abundance was higher at non-impacted coral habitats compared to impacted. The actinarian Protanthea simplex and unidentified brittlestars were the only taxa with higher abundance on impacted compared with non-impacted habitats. The reefs at the offshore location were protected against bottom trawling in 2009 through the establishment of a marine protected area (MPA), but a general ban against trawling on known coral reefs had already been implemented in 1999. In the MPA, signs of regrowth were observed. Most of the observed damage probably occurred over 10 years earlier. Results show that live and non-impacted cold water coral reefs have an important ecological function by enhancing the local biodiversity and fish abundance. Preventing further damage to impacted reefs may lead to full recovery within a few decades.     

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.     

Patterns of chemical defense among Caribbean gorgonian corals: a preliminary survey

Ship-board assays employing the common Caribbean wrasse Thalassoma bifasdatum (Bloch) were undertaken to determine the palatability of food pellets coated with freshly-extracted, lipid-soluble metabolites of 37 types of Caribbean gorgonian corals representing at least 19 species from 11 genera. Extracts of 19 types (51%) were highly unpalatable (zero or one of five pellets eaten), four types (11%) were moderately unpalatable (two or three of five pellets eaten) and 14 (38%) were palatable (four or five of five pellets eaten) to fish in feeding assays. Gorgonians of the genera Pterogorgia (three types) and Eunicea (nine types) were consistently highly unpalatable, those of the genus Plexaurella (four types) were palatable and those of the genus Plexaura were most frequently palatable (six of eight types). Further assays of serial dilutions of extracts from seven representative, unpalatable types revealed that extracts inhibited fish feeding at pellet concentrations near or below the concentrations that metabolites occur in the gorgonian soft tissue. Extracts of Erythropodium caribaeorum (Duchassaing and Michelotti) and Pseudopterogorgia rigida (Bielschowsky) deterred fish feeding at pellet concentrations less than an order of magnitude lower than those found in the soft tissues of the corresponding gorgonians. Thin layer chromatographic analyses of extracts revealed the presence of lipid-soluble, secondary metabolites in a majority of the highly unpalatable extracts, although secondary metabolites were also present in a smaller percentage of palatable extracts. These data support the hypothesis that the soft tissues of many gorgonian corals contain lipid-soluble feeding deterrents which act as a defense against predation.     

Parasites of forage fishes in the vicinity of Steller sea lion (Eumetopias jubatus) habitat in Alaska

Fish serve as intermediate hosts for a number of larval parasites that have the potential of maturing in marine mammals such as Steller sea lions (Eumetopias jubatus). We examined the prevalence of parasites from 229 fish collected between March and July 2002 near two islands used by Steller sea lions in Southeast Alaska and island habitats in the Aleutian Islands. Sea lion populations have remained steady in Southeast Alaska but have been declining over the last 30 yr in the Aleutian Islands. Even though the fish samples near the Southeast Alaska haul-outs were composed of numerous small species of fish and the Aleutian Islands catch was dominated by juveniles of commercially harvested species, the parasite fauna was similar at all locations. Eleven of the 20 parasite taxa identified were in their larval stage in the fish hosts, several of which have been described from mammalian final hosts. Four species of parasite were more prevalent in Southeast Alaska fish samples, and seven parasite species, including several larval forms capable of infecting marine mammals, were more prevalent in fish from the Aleutian Islands. Nevertheless, parasites available to Steller sea lions from common fish prey are not likely to be a major factor in the decline of this marine mammal species.     

Fine-scale population genetic structure in Alaskan Pacific halibut (<Emphasis Type="Italic">Hippoglossus stenolepis</Emphasis>)

Pacific halibut collected in the Aleutian Islands, Bering Sea and Gulf of Alaska were used to test the hypothesis of genetic panmixia for this species in Alaskan marine waters. Nine microsatellite loci and sequence data from the mitochondrial (mtDNA) control region were analyzed. Eighteen unique mtDNA haplotypes were found with no evidence of geographic population structure. Using nine microsatellite loci, significant heterogeneity was detected between Aleutian Island Pacific halibut and fish from the other two regions (F _ST range = 0.007–0.008). Significant F _ST values represent the first genetic evidence of divergent groups of halibut in the central and western Aleutian Archipelago. No significant genetic differences were found between Pacific halibut in the Gulf of Alaska and the Bering Sea leading to questions about factors contributing to separation of Aleutian halibut. Previous studies have reported Aleutian oceanographic conditions at deep inter-island passes leading to ecological discontinuity and unique community structure east and west of Aleutian passes. Aleutian Pacific halibut genetic structure may result from oceanographic transport mechanisms acting as partial barriers to gene flow with fish from other Alaskan waters.     

Benthic Composition of a Healthy Subtropical Reef: Baseline Species-Level Cover,with an Emphasis on Algae,in the Northwestern Hawaiian Islands

The Northwestern Hawaiian Islands (NWHI) are considered to be among the most pristine coral reef ecosystems remaining on the planet. These reefs naturally contain a high percent cover of algal functional groups with relatively low coral abundance and exhibit thriving fish communities dominated by top predators. Despite their highly protected status, these reefs are at risk from both direct and indirect anthropogenic sources. This study provides the first comprehensive data on percent coverage of algae, coral, and non-coral invertebrates at the species level, and investigates spatial diversity patterns across the archipelago to document benthic communities before further environmental changes occur in response to global warming and ocean acidification. Monitoring studies show that non-calcified macroalgae cover a greater percentage of substrate than corals on many high latitude reef sites. Forereef habitats in atoll systems often contain high abundances of the green macroalga Microdictyon setchellianum and the brown macroalga Lobophora variegata, yet these organisms were uncommon in forereefs of non-atoll systems. Species of the brown macroalgal genera Padina, Sargassum, and Stypopodium and the red macroalgal genus Laurencia became increasingly common in the two northernmost atolls of the island chain but were uncommon components of more southerly islands. Conversely, the scleractinian coral Porites lobata was common on forereefs at southern islands but less common at northern islands. Currently accepted paradigms of what constitutes a “healthy” reef may not apply to the subtropical NWHI, and metrics used to gauge reef health (e.g., high coral cover) need to be reevaluated.     

A new species of Polymastia (Porifera, Hadromerida, Polymastiidae) from the Aleutian Islands, Alaska, USA

Polymastia fluegeli n. sp. is described from deep water off the Aleutian Islands (Alaska, USA). P. fluegeli is disc-shaped and lives partly buried in the sediment, with only the papillae protruding above the surface. This new species has a basal layer of agglutinated sediment particles occurring between the choanosome and the ectosomal lower layer. This conspicuous sediment layer is not described for other species of Polymastia. Occurring spicule types and sizes are different from known species of the area.     

Dead bleached coral provides new surfaces for dinoflagellates implicated in ciguatera fish poisonings

Synopsis Dead sections of bleached corals in the United States Virgin Islands and British Virgin Islands were found to be colonized by filamentous algae harboring epiphytic dinoflagellates implicated in ciguatera fish poisonings. The dinoflagellates Ostreopsis lenticularis, Prorocentrum concavum, and P. lima were found in association with filamentous algae growing on dead sections of bleached Montastrea annularis and Acropora cervicornis. Several fish species from the families Acanthuridae, Pomacentridae, and Scaridae were observed to readily consume this filamentous algae/epiphytic dinoflagellate food source. Such fishes are common prey for large piscivores inhabiting tropical reefs. An increased incidence of ciguatoxic fishes may occur on reefs where bleaching events have caused significant coral mortality.     

Diversity and antibacterial activity of culturable actinobacteria isolated from five species of the South China Sea gorgonian corals

This study describes the diversity and antibacterial activity of culturable actinobacteria isolated from five species of gorgonian corals (Echinogorgia aurantiaca, Melitodes squamata, Muricella flexuosa, Subergorgia suberosa, and Verrucella umbraculum) collected in shallow water of the South China Sea. A total of 123 actinobacterial isolates were recovered using ten different isolation media, and assigned to 11 genera, including Streptomyces and Micromonospora as the dominant genera, followed by Nocardia, Verrucosispora, Nocardiopsis, Rhodococcus, Pseudonocardia, Agrococcus, Saccharomonospora, Saccharopolyspora and Dietzia. Comparable analysis indicated that the numbers of actinobacterial genera and isolates from the five gorgonian coral species varied significantly. It was found that 72 isolates displayed antibacterial activity against at least one indicator bacterium, and the antibacterial strains isolated from different gorgonians had almost the same proportion (~50 %). These results provide direct evidence for the hypotheses that gorgonian coral species contain large and diverse communities of actinobacteria, and suggest that many gorgonian-associated actinobacteria could produce some antibacterial agents to protect their hosts against pathogens. To our knowledge, this is the first report about the diversity of culturable actinobacteria isolated from gorgonian corals.     

Coral and sea urchin assemblage structure and interrelationships in Kenyan reef lagoons

Patterns of hard coral and sea urchin assemblage structure (species richness, diversity, and abundance) were studied in Kenyan coral reef lagoons which experienced different types of human resource use. Two protected reefs (Malindi and Watamu Marine National Parks) were protected from fishing and coral collection, but exposed to heavy tourist use. One reef (Mombasa MNP) received protection from fishermen for one year and was exploited for fish and corals prior to protection and was defined as a transitional reef. Three reefs (Vipingo, Kanamai, and Diani) were unprotected and experienced heavy fishing and some coral collection. Protected and unprotected reefs were distinct in terms of their assemblage structure with the transitional reef grouping with unprotected reefs based on relative and absolute abundance of coral genera. Protected reefs had slightly higher (p<0.01) coral cover (23.6 ± 8.3 % ± S.D.) than unprotected reefs (16.7 ± 8.5), but the transitional reef had the highest coral cover (30.8 ± 6.4) which increased by 250% since measured in 1987: largely attributable to a large increase inPorites nigrescens cover. Protected reefs had higher coral species richness and diversity and a greater relative abundance ofAcropora, Montipora andGalaxea than unprotected reefs. The transitional reef had high species richness, but lower diversity due to the high dominance ofPorites. Sea urchins showed the opposite pattern with highest diversity in most unprotected reefs. Coral cover, species richness, and diversity were negatively associated with sea urchin abundance, but the relative abundance ofPorites increased with sea urchin abundance to the point wherePorites composed >90% of the coral cover at sites with the highest sea urchin abundance. Effects of coral overcollection was only likely for the genusAcropora (staghorn corals). A combination of direct and indirect effects of human resource use may reduce diversity, species richness, and abundance of corals while increasing the absolute abundance of sea urchins and the relative cover ofPorites.     

Eggs and late-stage embryos of Allocareproctus unangas (family Liparidae) from the Aleutian Islands

Four unusual masses of adhesive fish eggs surrounding limbs of the octocoral Primnoa sp. were collected in Seguam Pass in the Aleutian Island Archipelago at a depth of 397 m. The eggs contained embryos in the flexion stage of development. When dissected and cleared and stained, counts of dorsal and anal fin rays and vertebrae allowed identification of the eggs as those of Allocareproctus unangas Orr and Busby, a newly described snailfish species. This is first report of snailfishes of the genus Allocareproctus, and of liparids other than those in the genus Liparis, spawning outside of lithodid crabs.     

Grenadier abundance examined at varying spatial scales in deep waters off Newfoundland, Canada, with special focus on the influence of corals

There is a growing body of research examining the effects of corals on fish communities, species abundances, and biodiversity. Yet, few studies have quantitatively examined what factors are influencing the distribution of individual fish species. In general, many researchers believe they know what influences the distribution of grenadiers on large spatial scales, but numerous studies have shown the distributions of organisms are often determined by various factors that change in relative importance when viewed at differing scales. Our study used video collected from three deep canyons off Newfoundland, Canada (North west Atlantic) to examine how the factors apparently influencing the distribution of four grenadiers (Macrouridae: Coryphaenoides rupestris, Coryphaenoides carapinus, Nezumia bairdii, and Macrourus berglax) change when assessed at varying spatial scales. We paid special attention to the influence of deep-water corals found in the study area (large gorgonians/antipatharians, small gorgonians, sea pens, soft corals, and cup corals). The factors that influenced grenadier presence and/or abundance (and the magnitude of this effect) varied as different sampling resolutions were examined. We found C. rupestris abundance was positively related to cup coral abundance in transects longer than 10 m, likely as a result of similar habitat preferences between both taxa. When significant relationships between depth and C. rupestris presence and/or abundance were found, they were always negative. Depth was a significant predictor of C. carapinus abundance in transects longer than 10 m. Very few predictors of M. berglax abundance or presence could be found. Depth and the number of small gorgonians were consistent predictors of N. bairdii abundance.     

Heavy metals distribution in the coral reef ecosystems of the Northern Red Sea

Concentrations of seven heavy metals (Cu, Zn, Pb, Cd, Ni, Co and Fe) were measured in the seawater, sediments, common scleractinian reef-building corals and soft corals (Octocorallia : Alcyonacea) at seven reef sites in the Northern Red Sea: I (Hurghada), II (Ras Za’farana), III (El-Ain Al-Sukhna), IV (El-Tur), V (Sha’b Rashdan), VI (Sharm El-Sheikh) and VII (Dahab). Levels of heavy metals were considerably elevated in seawater, sediments and corals collected from reef sites exposed to increased environmental contamination, as a result of diversified natural and anthropogenic inputs. Soft corals of genera Lithophyton, Sarcophyton and Sinularia showed higher concentrations of Zn, Pb, Cd and Ni than hard coral genera Acropora and Stylophora. Soft coral Sarcophyton trocheliophorum collected from El Ain Al-Suhkna (Gulf of Suez) had greater concentration of Cu, followed by hard corals Acropora pharaonis and Acropora hemprichi. The elevated levels of Zn, Cd and Ni were reported in the dry tissue of soft coral Sinularia spp. On the other hand, the soft coral Lithophyton arboreum displayed the highest concentration of Pb at Sha’b Rashdan (Gulf of Suez) and elevated concentration of Zn at Sharm El-Sheikh. Sediments showed significantly higher concentration of Fe than corals. The higher levels of Fe in hard corals than soft corals reflected the incorporation of Fe into the aragonite and the chelation with the organic matrix of the skeleton. The greater abundance of soft corals in metal-contaminated reef sites and the elevated levels of metals in their tissue suggesting that the soft corals could develop a tolerance mechanism to relatively high concentrations of metals. Although the effects of heavy metals on reef corals were not isolated from the possible effects of other stresses, the percentage cover of dead corals were significantly higher as the concentrations of heavy metals increased.     

Relationships between benthic cover, current strength, herbivory, and a fisheries closure in Glovers Reef Atoll, Belize

Benthic cover, current strengths, and fish abundance and diversity were examined on 150 lagoonal patch reefs and mapped to determine their distribution, inter-relationships, and relationship to the fisheries closure in Glovers Reef Atoll. Current strength was highest at both the northern and southern ends of the atoll and largely controlled by local wind and weakly by tidal forcing. Benthic functional group distributions varied throughout the atoll and had distinct areas of dominance. In contrast, dominance of coral species was weaker, reflecting the lost cover and zonation of Acropora, Porites, and Montastraea that were reported in the 1970s. Hard and soft corals dominated the windward rim, while the central and leeward lagoon had lower current strengths and sea grass and fleshy green algae were relatively more abundant. Brown erect algae were relatively more common in the north and calcifying green and red algae the southern ends of the atoll. Only Montastraea-Agaricia agaricites distributions were similar to reports from the 1970s with high relative dominance in the southern and northeast atoll. The central-northern zone, which was described as an Acropora zone in the 1970s, was not recognizable, and Porites porites, P. astreoides, Millepora alcicornis, and Favia fragum were the most abundant species during this survey. Hard and soft coral cover abundance declined away from the reef rim and tidal channels and was associated with fast seawater turnover and high surgeonfish abundance. Consequently, the windward rim area has retained the most original and persistent hard-soft coral and surgeonfish community and is considered a priority for future management, if the goal is to protect coral from fishing impacts.     

Diversity of symbiotic algae of the genus Symbiodinium in scleractinian corals of the Xisha Islands in the South China Sea

Symbiotic algae （Symbiodinium sp.） in scleractinian corals are important in understanding how coral reefs will respond to global climate change. The present paper reports on the diversity of Symbiodinium sp. in 48 scleractinian coral species from 25 genera and 10 families sampled from the Xisha Islands in the South China Sea, which were identified with the use of restriction fragment length polymorphism （RFLP） of the nuclear ribosomal DNA large subunit gene （rDNA）. The results showed that： （i） Symbiodinium Clade C was the dominant zooxanthellae in scleractinian corals in the Xisha Islands; （ii） Symbiodinium Clade D was found in the corals Montipora aequituberculata, Galaxea fascicularis, and Plerogyra sinuosa; and （iii） both Symbiodinium Clades C and D were found simultaneously in Montipora digitata, Psammocora contigua, and Galaxeafascicularis. A poor capacity for symbiosis polymorphism, as uncovered by RFLP, in the Xisha Islands indicates that the scleractinian corals have low adaptability to environmental changes. Further studies are needed to investigate zooxanthellae diversity using other molecular markers.     

Predator release of the gastropod <Emphasis Type="Italic">Cyphoma gibbosum</Emphasis> increases predation on gorgonian corals

When large, predatory fishes and invertebrates were excluded from areas of a coral reef in the Florida Keys, USA, densities of the normally rare gastropod Cyphoma gibbosum, a principal predator of gorgonian corals, increased 19-fold. Gorgonians in predator exclosures were grazed more frequently and extensively by C. gibbosum than were gorgonians in uncaged areas. In exclosures, 14% of all gorgonians showed recent predation by C. gibbosum, with 62% of the entire colony surface being removed from these attacked individuals. In areas where predators of C. gibbosum were not excluded, only 5% of gorgonians exhibited recent damage, with only 26% of the colony surface being removed from these few damaged individuals. Thus, the increases in both frequency and extent of attack combined to produce an 8× increase in gorgonian damage following removal of large predators. These patterns suggest that predators typically suppress C. gibbosum populations, that overfishing of these predators could release C. gibbosum from top-down control, and that this release will allow increased damage to gorgonian corals.