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Who, What, When, Where, Why, and How of Coral Reef Communities • Who (organisms) can be found in a coral reef community? • What is a coral reef community? • When can you find a coral reef community (is it seasonal)? • Where are coral reef communities found? • Why are coral reef communities so important? • How do coral reef communities interact with other ecosystems in the ocean? Chapter 15 Coral Reef Communities Karleskint Turner Small • Let’s talk about primary productivity in the oceans - Photosynthesis by phytoplankton • Polar oceans are the most productive • Cold water holds more gases • Summer brings 24 hours of daylight for photosynthesis • Temperate waters are somewhat productive • Tropical waters are the least productive • That’s why there are nice clear, blue water • Coral reefs are like an oasis in the tropical water dessert Coral Reef Communities • Coral reefs are highly productive, but occur in nutrient-poor waters – This is made possible by the symbiotic relationship between coral animals and zooxanthellae – These symbionts + algae form the basis of the community; other reef animals depend on these organisms Organisms That Build Coral Reefs • Stony (true) corals are the primary organisms that deposit massive amounts of CaCO3 that compose most of the structure of coral reefs • Hermatypic: coral species that produce reefs, found in shallow, tropical waters and harbor zooxanthellae Organisms That Build Coral Reefs • Ahermatypic: corals that do not build reefs, which can grow in deeper water from the tropics to polar seas – most do not harbor zooxanthellae • Coralline algae and organisms such as fire coral, deposit lesser amounts of calcium carbonate on reefs – fire corals important in Caribbean reefs – coralline algae important in structure of Pacific reefs Organisms That Build Coral Reefs • Coral colonies – large colonies of small coral polyps, each of which secretes a corallite (cup of calcium carbonate) – the coral larva called a planula larva settles and attaches – a polyp develops, and reproduces by budding to form a growing colony – polyps’ gastrovascular cavities remain interconnected – a thin, usually colorful epidermis overlies the colony surface Organisms That Build Coral Reefs • Coral nutrition – corals have evolved several strategies for obtaining food – symbiotic zooxanthellae • supply 90% of nutritional needs of stony coral • zooxanthella provide glucose, glycerol and amino acids • coral polyp provides a suitable habitat and nutrients, absorbed directly through the animal’s tissues • zooxanthellae remove CO2 and produce O2 • need of zooxanthellae for sunlight limits depths to which stony corals can grow Organisms That Build Coral Reefs • Coral nutrition (continued) – corals as predators • tiny zooplankton or other small animals paralyzed by the cnidocytes (stinging cells in tentacles) are passed into the digestive cavity Organisms That Build Coral Reefs • Coral nutrition (continued) – other sources of nutrition • mesenteric filaments (coiled tubes attached to the gut wall) can be extruded from the mouth to digest and absorb food outside the body • corals can feed off bacteria living in their tissues, which feed on dissolved organic matter directly from the water Organisms That Build Coral Reefs • Reproduction in corals – Reproduction by fragmentation • in addition to budding, corals can also reproduce asexually by fragmentation • some branching corals are fragile and tend to break during storms • if they survive the storm, fragments can attach and grow into new colonies Organisms That Build Coral Reefs • Reproduction in corals (continued) – Sexual reproduction in coral • Many species of coral are hermaphroditic, some have separate sexes • mostly broadcast spawners—release both sperm and eggs into the surrounding seawater • some are brooders—broadcast sperm, but retain eggs in the gastrovascular cavity • spawning is usually synchronous among Pacific reef species, but nonsynchronous among Caribbean species Reef Formation • Involves both constructive and destructive phases • Bioerosion: the destructive phase of reef formation – boring clams or sponges attack exposed surfaces on the undersides of large corals – the coral stand weakens, then topples in a storm or ocean surge – accumulating debris smothers boring organisms, cracks are filled with CaCO2 sediments, and coralline algae cement it together • Types of reefs: • • • • Fringing Barrier Atoll Patch Types of Coral Reefs • Fringing reefs – develop along shores of tropical/subtropical islands or continental landmasses – Of all reef types, most affected by human activities because of their proximity to land, develop right next to the land Types of Coral Reefs • Barrier reefs – similar to fringing reefs but separated from the landmass and fringing reef by lagoons or deepwater channels – Great Barrier Reef is the world’s largest barrier reef Types of Coral Reefs • Atolls – usually elliptical, arise out of deep water and have a centrally-located lagoon – Often eroded dead volcano • Darwin’s theory of atoll formation: – corals colonize shallow areas around newlyformed volcanic islands to form a fringing reef – the island sinks and erodes, and a barrier reef is formed about the island – the island sinks completely, leaving an atoll Land Sea Reef Fringing reef Patch Lagoon reef Land Reef Sea Barrier reef Lagoon Sea Sea Reef Atoll Patch reef Reef Stepped Art Fig. 15-11, p. 422 Types of Coral Reefs • In addition, patch reefs can occur within lagoons associated with atolls and barrier reefs Reef Structure • Different reef types share common characteristics • Reef front or forereef: portion of the reef that rises from the lower depths of the ocean to a level just at or just below the surface of the water, on the seaward side – drop-off: a steep reef-front that forms a vertical wall – spur-and-groove formation or buttress zone: finger-like projections of the reef front that protrude seaward; disperses wave energy and helps prevent damage Reef Structure • Reef crest: the highest point on the reef and the part that receives the full impact of wave energy – where wave impact is very strong, it may consist of an algal ridge of encrusting coralline algae, lacking most other organisms, and penetrated by surge channels, grooves of the buttress zone • Reef flat or back reef: portion behind the reef crest – reef flat of fringing reefs ends at the shoreline – reef flat of atolls and barrier reefs descends into the lagoon Reef Structure • Coral populations on reef front are massive domeshaped brain corals and columnar pillar corals on intermediate slopes, below this region coral species form plate-like formations • Higher up on reef where wave energy is greatest, branching species of coral are found, e.g., elkhorn coral in Caribbean • In protected areas behind reef front, in shallow calm waters, small species of coral occur, e.g., rose, flower and star corals Coral Reef Distribution • Major factors influencing distribution (corals are sensitive): • temperature – corals do best at 23-25o C • light availability – photosynthetic zooxanthellae need light, corals not found below 60 meters • sediment accumulation – can reduce light and clog feeding structures • salinity, corals absent from areas of massive freshwater outflow, e.g., the mouth of the Amazon • wave action – moderate wave action is beneficial, brings in oxygenated seawater, removes sediment that could smother coral polyps • heavy wave action during hurricanes can damage reef structure • duration of air exposure – can be deadly Comparison of Atlantic and Indo-Pacific Reefs • Pacific reefs are older and have a greater depth of reef carbonates • Buttress zone is deeper on Atlantic reefs and coral growth may extend to 100 m down – Pacific coral growth rarely exceeds 60 m • Proportion of reef covered by corals may approach 100% on some Pacific reefs, but usually less than 60% on Atlantic reefs • Algal ridges more common in the Pacific because of wind and waves Comparison of Atlantic and Indo-Pacific Reefs • Hydrozoan Millipora complanata (fire coral) is dominant on Atlantic reefs – similar species never dominate in the Pacific • Atlantic corals nocturnal (night); Pacific corals diurnal (day) Comparison of Atlantic and Indo-Pacific Reefs • Greater sponge biomass in the Atlantic • Pacific has giant clams and sea stars that prey on corals Coral Reef Ecology • Source of nutrients for coral reefs – land runoff for reefs close to land – That’s why too much runoff can bring unwanted nutrients – source of nutrients for atolls unclear (usually out in the middle of the ocean) • possible explanations: – nutrients accumulated over time are efficiently recycled – reef bacteria and filter feeders capitalize on nutrients from dissolved/particulate organic matter – nutrients are stored in the biomass of the community’s inhabitants Coral Reef Ecology • Photosynthesis on Reefs – photosynthetic organisms: zooxanthellae, benthic algae, turf algae, sand algae, phytoplankton, seagrasses – more dense than tropical ocean, with greater biomass than reef animals – associations of producers with other organisms assist in efficient recycling, e.g., zooxanthellae with corals, cyanobacteria with sponges Coral Reef Ecology • Reef productivity – ratio of primary production to community respiration = P-R ratio • P = gross photosynthesis • C = community respiration – P-R ratio used to measure state of development of a biological community Coral Reef Ecology • Reef productivity (continued) – P-R ratio > 1 = primary production exceeds respiratory needs • biomass increases, excess biomass available for growth or harvesting – P-R ratio = 1 = steady state (climax) • little biomass remains available for growth – P-R ratios for coral reefs are typically close to 1 • high productivity balanced by high respiration Coral Reef Ecology • Reef productivity – increases in productivity are often the result of eutrophication • eutrophication: nutrient enrichment – eutrophication typically manifested as a dramatic proliferation of algae • if grazing doesn’t increase, algae can grow over and smother corals • This will happen with too much sewage and fertilizer runoff The Coral Reef Community • Competition among corals and other reef organisms – fast-growing, branching corals grow over slowergrowing, encrusting or massive corals and deny them light – slower-growing corals extend stinging mesenterial filaments from their digestive cavity to kill faster-growing corals – fast-growing corals can also sting and kill using long sweeper tentacles with powerful nematocysts The Coral Reef Community • Competition among corals and other reef organisms (continued) – Slower growing corals are more aggressive than fast growing corals – Massive corals are generally more shade tolerant and are able to survive at greater depths – as a result… • fast-growing, branching corals on many reefs dominate upper, shallower portions • larger, slower-growing corals dominate deeper portions The Coral Reef Community • Competition among reef fishes – High diversity • coral reefs - marine habitats with greatest diversity/abundance of fishes • seems to defy competitive exclusion principle, which suggests that no 2 species can occupy the same niche – 60-70% of reef fishes are general carnivores – about 15% are coral algae grazers or omnivorous The Coral Reef Community • Competition among reef fishes (continued) – hypotheses proposed to explain this: • competition model: factors such as time of day or night, size of prey, position in the water column, etc. provide each species with a unique niche (hence, no competition) • predation disturbance model: assumes competition, but suggests that the effect of predation or other causes of death keep populations low enough to prevent competitive exclusion The Coral Reef Community • Competition among reef fishes (continued) – hypotheses proposed to explain this: • lottery model: assumes competition occurs, but suggests that chance determines which species of larvae settling from the plankton colonize a particular area of the reef • resource limitation model: suggest that available larvae are limited and that limitation prevents fish population from ever reaching the carrying capacity of the habitat The Coral Reef Community • Effect of grazing – reef is a mosaic of microhabitats with different levels of grazing and different algal communities – grazing of larger, fleshier seaweeds permits competitively inferior filamentous forms or coralline algae to persist – herbivory decreases with depth – damselfish form territories where they exclude grazers and permit abundant algal growth • provides habitat for small invertebrates • overgrows corals; fast-growing, branching corals are most successful near damselfish The Coral Reef Community • Effect of predation – predation of sponges, soft corals and gorgonians provides space for competitively inferior reef corals – small invertebrates are almost all well hidden or camouflaged, indicating the prevalence of predation in the reef The Coral Reef Community • Symbiotic relationships on coral reefs – cleaning symbioses • cleaner wrasses, gobies, etc. feed on parasites of larger fishes • cleaning organisms set up a cleaning station – Other symbiotic relationships • • • • clownfishes and anemones conchfish and the queen conch gobies and snapping shrimp crustaceans and anemones Evolutionary Adaptations of Reef Dwellers • Adaptive behaviors to avoid predation – invertebrates hide during the day and forage at night – producing a poisonous coating of mucus – burying the body in sand to hide – inflating to appear larger – hiding at night when nocturnal predators are active Evolutionary Adaptations of Reef Dwellers • Structural adaptations for feeding – cnidocytes (stinging cells) of cnidarians aid in prey capture – radioles (hair-like) appendages of Christmas tree worms are used to capture phytoplankton – non-bivalve mollusks use radula to graze algae – mantis shrimp have extremely sharp forward appendages – snapping shrimp use sound to defend territory and stun prey – crinoids (feathers stars) use basket of mucus to feed Evolutionary Adaptations of Reef Dwellers • Protective body covering – tough, defensive exteriors help animals avoid predation, but can limit mobility and growth • Role of color in reef organisms – color for concealment and protection – Many invertebrates have colors and stripes that allow them to blend in with the environment Evolutionary Adaptations of Reef Dwellers • Role of color in reef organisms (continued) – brilliant color of many fish actually helps them to blend in with colorful background of the reef – other types of camouflage • body shape – warning coloration • e.g., lionfish – other roles of color • defending territories • mating rituals Threats to Coral Reef Communities • Effect of physical changes on the health of coral reefs – hurricanes and typhoons topple and remove coral formations – El Niño Southern Oscillation (ENSO) • changes winds, ocean currents, temperatures, rainfall and atmospheric pressure over large areas of tropical and subtropical areas • can cause massive storms Threats to Coral Reef Communities • Why are coral reefs important? – protect coast from high surf conditions – remove large amounts of carbon dioxide from water and air – provide habitat for a huge diversity of invertebrates and fish – economical value, many people earn living by collecting and processing reef products – important place of recreation – have potential for harvesting pharmaceutical products Threats to Coral Reef Communities • Effects of human activities – Destructive fishing practices • overfishing, i.e., eliminating grazers, allows algae to overgrow reefs • poisonous chemicals used to capture fish also poison corals • explosives used to stun and capture fish can cause massive destruction to coral • bottom trawling for fish also destroys coral structures Threats to Coral Reef Communities • Effects of human activities (continued) – Coastal development • produces runoff containing nutrients, pesticides, toxic wastes • increases sedimentation and changes patterns of water flow Threats to Coral Reef Communities • Effects of human activities (continued) – Other human activities • coral mined for use as bricks, road-fill, cement component • removed to make jewelry • inexperienced snorkelers and boaters damage reefs Threats to Coral Reef Communities • Effects of human activities (continued) – effects of human-induced climate change in atmosphere • increased levels of carbon dioxide from burning of fossils fuels primary cause of ocean warming • causes corals to become stressed and more susceptible to coral bleaching and disease Threats to Coral Reef Communities • Effects of human activities (continued) – coral bleaching • a phenomenon by which corals expel their symbiotic zooxanthellae • most often associated with warming of the ocean water by ENSO or global warming • if the stress is not too severe, corals may regain zooxanthellae and recover • if the stress is prolonged, corals may fail to regain zooxanthellae and die Threats to Coral Reef Communities • Effects of human activities (continued) – coral diseases • black band disease: a distinct dark band of bacteria migrates across the living coral tissue, leaving behind a bare white skeleton • white pox: characterized by white lesions and caused by Serratia marcescens – other coral diseases: • white band disease • white plague • CYBD (Caribbean yellow band disease) or yellow blotch disease