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What is a ‘ reef’? • complex structures formed by reef-building , , mollusks, and I. Introduction to coral reefs • result from a balance II. Characteristics of coral reef fishes between constructive and destructive processes III. Maintenance of diversity • external framework provides , refuge from predators, and food • distribution is limited by light, temperature, and hard substrate for initial colonization (between 30o N and 30o S latitude)

Physical structure of coral skeletons (cracks and crevices) provides refuge from predators

Distribution Diversity of corals and fishes highest in central Indo-Pacific • worldwide in tropical • lower on western margins continents, where occurs (cooler & murkier)

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1 Reef Zones Coral Morphology – variety of colony skeletal structures • reef flat (& ) • reef crest • reef slope

Primary is very high on coral reefs Dynamics: coral reefs change over time 1. Physical disturbances • storms hurricanes, cyclones Production (kg Carbon per m2 per year) (affect fast-growing branching corals most) Average Oceanic areas 0.1 kg • sedimentation Rainforest 2 kg • heating () 2 kg Coral Reef 1.5-5 kg 2. Biological disturbances • disease • waters where coral occur are very nutrient poor (“oligotrophic”) • high productivity possible because of tight recycling of nutrients, photosynthetic fixation of carbon (by corals and algae) and nitrogen (by blue-green algae)

II. Coral reef fishes A. : dominated by perciforms • in , for example, 51 of 103 Families are Perciforms Summary of coral reef attributes • 17 of the 20 most speciose families Perciforms B. Morphology (general characteristics): • high structural complexity provides for fishes Generally: • laterally compressed • high productivity provides food • physoclistous swim bladders • many different habitat types (e.g., zones) • highly modified jaw and pharyngeal apparatus (provides • dynamic (though not as much as kelp !) powerful and efficient feeding mechanisms) • pectoral fin locomotion common (MPF rowing) but very high morphological diversity (though dominated by families with small ) C. Diversity: extremely high • approximately 1/3rd to 1/2 of all fishes live on coral reefs (i.e., 10,000+ species)

2 D. Vision especially important (apparent in diversity of coloration) E. strong Habitat Associations • many species associate with particular features of the reef • may remain tightly associated with particular site for entire life

F. Life-history characteristics Major Families of Coral Reef Fishes • mostly egg layers (oviparous) -- broadcast or demersal spawners 1. The Labroids • 95%+ have pelagic larval stage & sedentary adult stage a. Labridae (wrasses) (bipartite life cycle)

1. The Labroids 2. Pomacentridae (damselfishes) a. Labridae (wrasses) b. Scaridae (parrotfishes)

3 3. The Acanthuroids (mostly ) 3. The Acanthuroids a. Acanthuridae (surgeonfishes) a. Acanthuridae b. Siganidae (rabbitfishes ) -- Indo-Pacific only

3. The Acanthuroids 4. The Chaetodontoids a. Acanthuridae a. Chaetodontidae () b. Siganidae c. Zanclidae (moorish idol) -- Indo-Pacific only

4. The Chaetodontoids 5. Haemulidae (Grunts) a. Chaetodontidae b. Pomacanthidae (angelfishes)

4 6. Gobiidae (gobies) 7. Blenniidae (blennies)

8. Synodontidae (lizardfishes) 9. Mullidae (goatfishes)

Nocturnal Families Nocturnal Families 1. Apogonidae (cardinalfishes) 1. Apogonidae 2. Holocentridae (squirrelfishes)

5 Nocturnal Families The highly evolved: 1. Apogonidae 1. Ostraciidae (boxfishes) 2. Holocentridae 3. Muraenidae (moray eels)

The highly evolved: The highly evolved: 1. Ostraciidae 1. Ostraciidae 2. Tetraodontidae (pufferfishes) 2. Tetraodontidae 3. Diodontidae (pocupinefishes) 3. Diodontidae 4. Balistidae ()

The large predators: The large predators: 1. Serranidae (groupers) 1. Serranidae 2. Lutjanidae (snappers)

6 The large predators: The large predators: 1. Serranidae 1. Serranidae 2. Lutjanidae 2. Lutjanidae 3. Lethrinidae (emperors) 3. Lethrinidae 4. Carangidae (jacks)

Hypotheses to explain high diversity: III. Diversity A. Resource partitioning -- What allows so many species to coexist without some • reef fishes have obvious trophic specializations: specialized being outcompeted? feeding behavior, territoriality (may be defense of food), and (e.g., thousands of species on Philippine reefs) modified mouths

• basic ecological theory: species that share resources should compete and eventually a competitive dominant excludes all other species, causing low

Hypotheses to explain high diversity: B. Does predation keep populations below ? A. Resource partitioning • evidence that predation is potent force affecting reef • reef fishes have obvious trophic specializations: specialized assemblages: feeding behavior, territoriality (may be defense of food), and modified mouths – high abundance of piscivores; tremendous losses in many fishes just after settlement • but is there evidence of current ? – defensive structures, colors, & toxins – most species have overlapping resource use – behavioral defenses: schooling, twilight changeover behavior – experimental studies give mixed results

7 C. Is settlement of larvae too low to reach carrying capacity Summary of evidence for mechanisms that allow species ( limitation hypothesis) coexistence in high diversity communities

Evidence: • removal of adults does not Competition may not cause competitive exclusion because… increase recruitment rate • species use resources in different ways • after total removal of adults, • predators keep densities low can take several years to • limited return of planktonic larvae to reef keeps densities low return to original numbers • recruitment pulses are sporadic & unrelated to reproductive output

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