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The Biology of Reefs and Reef Organisms

The Biology of Reefs and Reef Organisms

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Published by University of Chicago Press an imprint of UChicagoPress
Reefs provide a wealth of opportunity for learning about biological and ecosystem processes, and reef biology courses are among the most popular in marine biology and zoology departments the world over. Walter M. Goldberg has taught one such course for years, and he marshals that experience in the pages of The Biology of Reefs and Reef Organisms.
Goldberg examines the nature not only of coral reefs—the best known among types of reefs—but also of sponge reefs, worm reefs, and oyster reefs, explaining the factors that influence their growth, distribution, and structure. A central focus of the book is reef construction, and Goldberg details the plants and animals that form the scaffold of the reef system and allow for the attachment and growth of other organisms, including those that function as bafflers, binders, and cementing agents. He also tours readers through reef ecology, paleontology, and biogeography, all of which serve as background for the problems reefs face today and the challenge of their conservation.   Visually impressive, profusely illustrated, and easy to read, The Biology of Reefs and Reef Organisms offers a fascinating introduction to reef science and will appeal to students and instructors of marine biology, comparative zoology, and oceanography.
Reefs provide a wealth of opportunity for learning about biological and ecosystem processes, and reef biology courses are among the most popular in marine biology and zoology departments the world over. Walter M. Goldberg has taught one such course for years, and he marshals that experience in the pages of The Biology of Reefs and Reef Organisms.
Goldberg examines the nature not only of coral reefs—the best known among types of reefs—but also of sponge reefs, worm reefs, and oyster reefs, explaining the factors that influence their growth, distribution, and structure. A central focus of the book is reef construction, and Goldberg details the plants and animals that form the scaffold of the reef system and allow for the attachment and growth of other organisms, including those that function as bafflers, binders, and cementing agents. He also tours readers through reef ecology, paleontology, and biogeography, all of which serve as background for the problems reefs face today and the challenge of their conservation.   Visually impressive, profusely illustrated, and easy to read, The Biology of Reefs and Reef Organisms offers a fascinating introduction to reef science and will appeal to students and instructors of marine biology, comparative zoology, and oceanography.

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large islands often form behind the reef crest.0 (e. Smaller spicules. They are thought to have contributed substantially to the evolution of an oxygen atmosphere. Figure 1–5: a) Bora Bora. often including an outer shell composed of spicules. fication found in most species has been proposed as a defense mechanism against herbivores by diminishing Halimeda’s nutritional value relative to the amount of indigestible material (Duffy and Hay 1990). Neall and Trewick 2008).forming metazoans. scavengers. Zoccola D. 3: Demospongia. sometimes within a few hours. There are specialized herbivores that appear to be un. The matrix found in mollusc shell is typically com. “Hyaline” forams produce a test with a glass. The presence of protons outside the cell may be used to produce an acidic surface that converts bi. In contrast. Environmental Conserva. tion is clear. NASA.. sessing climate change over time (Foster 2001). Alcyonaceans occur commonly in shallow water only on Indo. et al. eralized biological materials. Roatan. London: Ed. Inducible agonistic structures in the tropical corallimorpharian. the ma. Macroalgae that attach to solid substrata usually do so by a holdfast. Some of these unusual symbionts provide their hosts with a wide variety of metabolites including unusual compounds that serve ecological func. perhaps 4 times as much (Marshall and Davies 1988. Belliveau SA and Paul VJ. and Spoon D. Cronin and Hay 1996). ternating concentric layers of gorgonin and calcite (Fig. and although one or both of these style types may be absent depending on the genus. P. These changes are often accompanied by density bands that are analogous to those produced by scleractinian corals.Pacific species Amplexidiscus fenestrafer is shown here. logically controlled minerals can be formed within a eu. 2006. the septa may form secondary peaks or lobes called pali before reaching the center. the surface: volume ratio can be increased (syco. and contains the zooxanthellae. vators. Horn MH. and Bright TJ. crustaceans. gelfishes. Choanocyte Flagellum Collar Food particles Porocyte Mesohyl Archaeocyte Spicule Pinacocyte Ostia Spongocoel Osculum Choanoderm Pinacoderm 80•Chapter Five by some specialized marine intertidal and bioeroding sponges.forming tissue in corals. in this case. Kailua Bay. among chemically defended sponges. 92•Chapter Five It follows that while some sponges can live perfectly well without cyanobacterial symbionts. Based on Nicol 1931. the movement of sediment maintained the grooves. This composition makes it generally similar to insect exoskeleton. ter location is Shark Bay. Barbados. a special group of Pleurocapsales that produces a type of phycoerythrin. Regulation and con.g. matypic? A redefinition of ecological groups in corals and other organisms. On the nature and causes of luminescent lines and bands in coral skel. A hypercalcified sponge with soft relatives: Vaceletia is a keratose demosponge.pone.. weight aggregates attached to the thylakoid mem. nect the gastrovascular cavities of adjacent polyps. locene carbonate shelf. USA 106: 21500–21504. 2006). are members of the hexagonal system. living organisms like these (and gold corals described above) may be as old as the Bronze Age and the kingdom of Egypt. ture of a clonal gorgonian coral: the interplay between clonal reproduction and disturbance.mason worm Phragmatopoma caudata. and solidify the system. asconoid form) to infolded wall (b. eds.arc. and adhesion to solid substrata.like hypercalcified sponges belong to four informal groups based on the organization of tis. in some regions forming a “coralline turf” of upright. Ecology of extant nummulitids and other larger benthic foramin. cies occurs as distinct genetic lineages (clades) that may correspond to commensal or mutualistic relationships. Microbial Sedi. 1980. Hentschel U.pH. these shrimp are highly diverse and many are host specific. Mortality cannot be directly linked to the loss of their Synechococcus symbionts. whereas those with zooxanthellae produce an average of about 3 times as much carbonate during the day compared with nighttime rates (Gattuso et al. The most famous of these are the tropical palolo worms of the family Eunicidae. Journal of Sedimentary Petrology 53: 449–459. Koehl 1982.1996. Liv. or is associated with. Male plants release tail. chemistry and Physiology B 139: 731–735. whereas other members of this group have been identified as agents of disease that afflict sponges and various reef corals as described in Chapter 15. The relative balance of these constructive and destructive forces is a critical element in determining whether a reef is growing. and that puts an end to the ambiguity at least as far as octocorals are concerned. and the eastern Pacific) produce predominantly clonal populations of P. Katoh M. Marine Biology 115: 677–683. a Figure 3–12: a) Stromato. as shown in Equation 3–2: and is subsequently converted into organic nitrogen. 2000. London © 2013 by The University of Chicago All rights reserved. and Uriz MJ. other sessile and motile organisms use the internal spaces as refuges from wave activity. 2007a. data compiled from UNEP World Conservation Monitoring Cen. New York: Springer Science and Business Media: 181–200.51143 and ISS006. They may also employ specialized structures for self. perfo. inhibiting coral and coral reef growth.rich waters that may at least partially offset low light levels (e.g. in part due to the predominant wind direction. In. several groups of organisms in. are similar. and are accompanied by high. nia Press.. A more de. Weiner S. Australia 2: 641–645.0002989.de. sea fans. 2004). the outcome of such competition is complex and may involve the production of specialized types of clonal in. ing to increased consumer demand. Atmospheric oxygen was added gradually at first. trol of crystal phase switching and orientation by sol. faces and polyp. nisms of reproductive isolation among sympatric broadcast. and Gates RD. cies of coral. Photo by author. This variability has caused some confusion 82•Chapter Five the Cellularia. or they may project across the surface as thickened finger. as well as by reduced light penetration. tive organisms requiring warm. Shadows cause ser. a mineral that is not found in true enamel (e. 2004). Adey et al. 1998.spawned coral eggs are not provisioned with zooxanthellae and symbiont acquisition occurs after settlement. 1993.000 98 unicellular macrophytic filamentous chl a. Are current estimates of coral reef biodiversity too low? The view through the window of a microcosm. 4–8c). 2002a.. est raised atoll (Scott and Rotondo 1983. The first system. Marine Ecology 19: 77–87. Conway KW. Beyond Kure near 29°N. http://ixmati. 1999. are common in chlorophytes. Each spicule has a filament. These components are then consumed during photosynthesis and result in the production of organic matter and the evolution of oxygen.rays or electrons are deflected as they pass from one geometric plane to another. lelochemicals.peaked volcano at center is >700 m above sea level. Acta Palaeontologica Polonica 48: 497–530. their durable consistency.water spe. Genomics and chlo. Lighty RG. Endolithic Microalgae Filamentous algae of the genus Ostreobium. where adductor muscle fibrils are inserted directly into the nacreous material. Capsule walls thin. Therefore. They are recognized by their radioles. which in turn form rigid skeletal elements such as calcareous sponge spicules. 1992. Isolated from the surrounding seawa. but are otherwise unrelated. Kinsey DW. ed. the reef was an active net carbonate producer. Conversely. c) Critical. Multiple trichomes within a sheath. especially around the mouth (Westfall and Elliott 2002. Cer. nitrogenase activity is typically restricted to the dark (Fay 1992. The complex triphasic life cycle of the Rhodophyta is a defining feature of the group. and Liebetrau V. However. 2008). cooling. Oceanography and Marine Biology Annual Review 19: 141–234. The Helix experi. including members of the genus Lyngbya (order Oscillatoriales) may grow in the TOXIC C y a n o b a c t e r i a C y a n o to x in s N o sw im m in gN o d r i n k i n g Figure 3–17: Warning sign indicating a bloom of toxic cyanobacteria. Mecha. one of which is filamentous (Prochlorothrix). Deep fore reef and upper island slope. 1977. On the southwest coast.dimensional environment with an intricate archi. 2001. gardineri complexes formed by their Indo. as in this example from Joulter’s Cays. In the monophyletic view of chloroplast evolution. Vago R. sponge Terpios hoshinota has been noted as the cause of a “black disease” that kills and overgrows corals and is responsible for the death of large reef areas in Taiwan. Thus. 1983.. ing. Indeed. which runs the gamut from detritivory to carnivory. Turfs domi. are not well represented in the Hawai. Dahan D. Some of these (e. eds. Thus. These microalgae are also quite prominent in dead coral skeletons where the Ostreobium filament system can be quite extensively developed (along with endolithic cya. The indigestibles are eliminated from the gut with the feces. Andrews et al. whereas others pro. bonate excavation by boring sponges. and Connectivity. anobacteria were the first of these organisms. Coral Reef Ecology. is composed of calcite. 172°W ) is open to the NW and has well. First obser. Gaetani GA and Cohen AL. tive patterns of scleractinian corals in the northern Red Sea. ported from the system by storms or other pro. New York: Chapman and Hall: 96–113. Although much more is known about this symbiosis in stony corals (described below). 2009). However. lamination. But. Phenoge. in which individual polyps and their ca. Biological Bulletin 181: 275–288. drying out the lagoon. 2003. Chemical defense and evolutionary trends in biosynthetic capacity among doriid nudibranchs. Grain susceptibility to the effects of mi. and that atolls (and barrier reef lagoons) are solution features rather than the result of construction seas were a few meters higher than at present. Pro. Coralline algae. cal bioerosion of boring sponges from Mexican Pa. Raz S. In contrast. in Halimeda this method has varying degrees of success among species. Allwood AC. These gamete. or the type of cnidae they possess. and the central medullary filaments. Integrative and Comparative Biology 43: 82–86. A patch. Walter M. and small crustaceans. This chemical operation is energetically expensive as shown above. frequency. Marine Ecology Progress Series 240: 117–126. 2009. Marine Biology 153: 225–234. Birkeland C. Ribosomes. Hurricane control on shelf.like particles and hetero.FISH analysis. which is often lined with chitin. transport of photosyn. Barrier reefs. ward side (11 o’clock on Figure 1–5a) showing well. requires a net carbonate production greater than zero. Vol. and other specialized types (Figure 6–1). grazing intensity by herbivores. or attached to solid substrata. New York: McGraw. the relationship between density and seasonal temperature does not always hold. The L. have extended the depth to which exploitation can occur. Likewise. American Scientist 89: 525–531. and ideas: a perspective of US coral reef research. temperature. or lack of suitable substrata. perhaps because they produce so much indigestible mate. a yellow. where these eunicids formed extensive tunnels.associated fauna on coral reefs in Singapore. et al. 2001. Belize. The mechanics of the pulse. Pochon X. or coiled mass of DNA. 2004. and without light as a limiting factor. ton and Hessinger 1996). Burja AM. coderm. benthic foraminifera appear to be sensitive to high levels of ultraviolet radiation and 72•Chapter Four tinctive reddish pigment of living material fades some. or if they are capable of detoxifying their prey (Chapter 12). scale kinetics of nematocyst discharge. Coral growth can be measured by increases in length. certain cyanobacterial mat communi. they began a decline during Cambrian time when other types of microbialite and calcified metazoans made their ap. Figure 1–4: Reef flat as. ocean salinity. and chlorophyll b in addi. are poorly understood. 2006.building coral Montastraea. ogy of the mineral. views 57:255–277. 2005.Pacific reefs. Limit. This dusting is facilitated by the growth of these stromatolites in loose sediment that is easily car. associations between corals and S. was 800 km long and 300 km wide (Rowland and Gangloff 1988).filled “sweeper tentacles” or other special. Becerro MA. re. growth (Jompa and McCook 2003). Reserves. or circulation is otherwise vigorous. limeters to a meter or more in a few species (Fautin and Allen 1997). Pacific. the ge. Marine Geology 115. The greatest concentration of atolls in the world is found in the Tuamotu Archipelago. ological attack and physical disturbance. certain bivalves. There is little doubt. and in pools directly exposed to the sun for long periods. Detection. but may also function in ventila. ———.mason worms or honeycomb worms. Cliffs composed of these fossil reefs (Figures 1–20. In addition. the inner. Allemand D. face are actively photosynthesizing. rine Organisms (URMO). Rees TAV. nitrification. isms their characteristic blue. and sessile as well as motile invertebrates (Smith et al.and. Sources and mechanisms of inorganic carbon trans. which leads to branches. et al. especially on coral reefs. lations of Halimeda are known to occur at depths of up to 150 m (Blair and Norris 1988). often divided into two subgroups. ibility. 2005. including forces such as mechanical damage or predation. and two on the continental shelf of Mexico. its area is (2r)(h) = ~25 cm2 .g. When contact is made with competitors. Entire archipelagoes that are the product of old reef growth are capped by active coral growth. 2002. Spatial and seasonal distribution of seaweeds on coral reefs from southern Brazil. Indeed. This covering resists extreme conditions including ex.day old chaetiger larva with developing segments and prominent chaetae. corallimorphs have been considered a candidate group of corals without skeletons (i. tics of a black coral meadow in the twilight zone of the central Mediterranean Sea. Harris MW. 2001).000 individuals m–2 including polychaetes. they were more common on fossil reefs on which they made considerable contributions.g. ence of a small part of the volcanic island that is still visible in the lagoon (Macdonald et al. Galeta Point. tabe N.magnesium calcite.500 km2 in which reefs are common. especially during the late Jurassic. Relatively large reef. reefs such as those along the coast of Ras Mohammad (southern Si. Cartwright P. contain numerous stromatolites. They are avoided by herbivores because of chemical deterrents and a high ratio of indigestible carbonate to organic matter. 1980. graphic range. 2000). 2006. three. b) The red clionaid Cliona delitrix. 1996). Steiner AQ. hoa prey (Chapter 6) and may use them as a deterrent to augment its own venomous defenses (Gleibs et al. and may be free moving. preferentially on hemispherical corals (Fig. Hooper JNA. NASA. In addition. the Marshalls. These are expressed according to their crystal. Sedimentology 44: 479–506. duce gametes in chambers called conceptacles. and subsequently inhibit repopulation by coral larvae (Maida et al. ———. in the south Pacific exposed at low tide. work reduced by bioerosion? • Sediment fate a: How much of the sediment is re. Oxford: Oxford University Press. Cuif J.living species also develop branchiae. the hardest and most highly mineralized tissue in the vertebrate body. These in turn promote calcification and reef accretion (Nakamura and Nakamori 2007). Harvard University. Thus. ride gels. and pass Calcium Carbonates Biogenic reefs are composed of various carbonates (CO 3 –2 ). 2011. amination of ~4. zation as described below.spawning corals develop larvae that can settle temporarily on various substrata including algae before becoming competent (Harrison 2011). Although these rhodo. each of which Figure 5–14: The reef. chaetes bore into reef substrata. 2008. submerged volca. cals they produce called lyngbyatoxins (see Burja et al. eral Indo. 2009. London: Smith. vironmental Biology of Fishes 58: 203–213. each clade of Symbiodinium is composed of multiple ge. Palatability and chemi. nutrient.control theory of coral reefs. a cyanophyte from Baja California (Mexico) that occurs as bundles of filaments within a single sheath. with different groups waxing and waning in prominence. Houlbrèque F. while others. and the coenenchyme. rials are formed in higher metazoans (Domart. including some in which the lagoon is marginally developed. mats pro. Serpulids are common on modern coral reefs where they may act as cementing agents or secondary frame builders. 2003. erence to the family Gorgoniidae. However. Some fringing reef systems can become quite extensive and in some areas occupy thousands of kilometers. Matsuda 1989). Atlantic corals of the genus Montastraea (Fig. ARC Centre of Excellence for Coral Reef Studies. Corals of the World. Some terminology has attempted to deal with this Reef Corals and Their Allies•115 genetic controls are manifested by the mode of asexual polyp division. borers of various types quickly invade the skeleton and structurally weaken it. eds. Biologi. an organic matrix has not been ruled out and may play a role in the process of crystal formation or nucleation (Borowitzka 1987). 2009. cyano. spurs (buttresses) may grow to a height of 10 m or more and a width of 10–90 m. 2007). While there is no agree. and regenerates the lost elec. Annual Review of Ecology and Systematics 17: 273–303. for ex. although not its own. and there are 14 volcanic peaks remain. The stable iso. thereby filling voids. Likewise. which places them at risk. and detritus. Oxford: Oxford University Press: 26–34. although other clades have been identified from soritid foraminifera (e. Fenical W. the barrier reefs near Tahiti in French Polynesia. typically in the range of 33–36 g/L (= parts per thousand. Cruz. 2009. Filaments are typically wider at the base and become tapered at apical regions. the presence of mesoglea.axes in red. there may be additional or fewer sets of mesenteries. ated with medium. swarming to the surface in large numbers before dawn. matolite reefs can be found in western Canada. 1990. mid (Figure 2–7). Coral Reefs 26: 741– 755. there and in the Mediterranean (where harvests are licensed). Most shallow. Reproductive ampullae (A) are prominent on this colony. c) Sclerodermite fan arrangement with apparent centers of calcification (c1. Photo by author. 2004. ample (Figure 3–9). There is also evidence of an unclearly understood acclimation process in which the symbiont may take al.profile cells form a tissue. One of the most common. tain a complex network of gastrodermal canals and me. habitat. Coral reef morphogenesis: a multidi. and deformation of the Earth’s crust due to loading and unloading by glacial ice. Removal of the coenenchyme reveals the blue aragonitic skeleton (inset). sium. Nummulitic limestones were used to build the great pyramids of Egypt. sand. Dominguez et al.depleted envi. bilaterally symmetrical. especially in the intertidal zone (Kirtley 1994). Thus.like capitate tentacles. 2007. move tropical water toward the poles. Certain microalgae are common as carbonate borers and are often closely as. In: Riegl B and Dodge RE.Pagès C. Marine Ecology. 2008. showing undulating cell wall (cw) and spiraled thylakoid membranes (th). the construc. allowing cells to fuse and produce cal. ii. Pearse found 16. followed by a complex process of cementation. Indeed.step process (1. Journal of Experimental Marine Biology and Ecology 342: 282–291. toplankton (Bruschetti et al. perhaps due to insufficient penetrating power during discharge. a b An Introduction to the Structure and Formation of Modern Reefs •29 southwest Indian Ocean (Camoin et al. Jamaica. Synchronized release of medusae from three species of hydrozoan fire corals. possibly through endosymbiosis with different hosts. Tupai. ogy and/or texture. cur primarily in fresh water can survive and even flour. 1996. Fringing reefs occur around their margins. 2007). and recruitment of scleractinian corals. tory. but perhaps at the price of providing them with a lower degree of genetic diversity (and pre. especially millepore. Song B. Indeed. In some cases. 2008. depending upon environmental 122•Chapter Six Sclerites can be produced within a single cell or by a multicellular process involving the deposition of calcite in layers within an organic matrix. 2005. 2002. Now the S:V ratio has reversed from 2:1 to 1:2. regardless of their proxim. The digestive tract of almost all serpulid and sabellariid larvae. area. 2005. some of which are 80 m high. licified megascleres that fuse with one another at their tips in all directions. 2001. unconsolidated sediment (after Wood 1999). Work et al. reef. The Sverdrup relation in the Indian Ocean. Jour.Pacific and Caribbean regions as long as they were growing in optimal conditions of sunlight temperature. the neurites may radiate some distance from it. Some remain suspended in the water column. 2008). is an atoll. trast.4 billion years ago (Gya).dimensionality as well as the number and size of internal cavities. Barnowski TF. Bergman B. the cyanobacterial biomass and rates of photosynthesis may be very high. Thus. These occur along a linear distance of >8 km. ters or parts thereof either early on. there are only 5 genera and 16 species that have such broad distributions (from records in Veron 2000). and many host symbiotic algae. 2005. On part of the Great Barrier Reef. Providing three. get for the Northern Great Barrier Reef. places where nitrogen and phosphorus levels are extremely low and limit the growth of most other phytoplankton. ration within the calcite crystal. as well as in the Red Sea where they may constitute 36%–48% of the reef surface in shallow water (Lewis 1989). 1980. in this case. Zlotkin E. High levels of sediment. Courtesy of Springer Science and Business Media BV. b live material). even though they rarely contact them directly. and Muscatine L. In general. Matthes U. Oceanography and Marine Biology: Annual Review 27: 167–272. Wirshing et al. ies widely. Dove PM. Mariani et al.profile organisms.0021670. rhizae. 1978. lantic are well represented by two prominent families. cipitation by coral cells in vitro.5940. ing discharge to form a delicate web of microfibrils that envelop the target. Most forams that produce hyaline tests are grouped into a large suborder. ments. sively by four species of Pocillopora: P. Unexpected species richness of reef corals near the southern Amazon River mouth. the calicodermis corresponds to the shape of the individual crystal bundles that characterize the microscopic features of the skeleton. Such structures are especially prominent on the side of the reef that most often faces high wind and surf. 114 The Octocorals. The reef below the crest is called the fore reef. some of which extend from the outer body wall to a well. Farias JD. Krautter M. Although there are no specialized gonadal tissues in sponges. whether in gonochoric or hermaphroditic colonies. which may benefit their hosts nutrition. crobialite carbonate composed the bulk of these for. In: Wis. Several groups of organisms are associated with Pa. and c) Solenastraea bournoni depicting budding pattern with coral. The calicodermis closely ad. Bentov et al. Soft corals as reef builders. Ortega. characterized DNA regions.water Desmophyllum dianthus (Scler. north of the Florida reef tract.like structures whose growth does not reach sea level. self. Paulay G. cial reference to the distribution of zooxanthellae. ganisms. Because this is a functional group rather than a taxonomic one. Prochlorophyta: a matter of class dis. limorpharia: an analysis of cnidae affinity. some of the nearby Loyalty Islands (Appendix 2b) are surrounded by fringing reefs. This may be due in part to carbonic anhydrase activ. Allemand et al. Their relatively simple morphology may suggest that such reefs lack complexity and species richness. The nature of the symbiont’s genotype.” meaning that chlorophyll is employed to capture light. ing deep. Some of the largest occur along the Brazilian coast. Seaweed Ecology and Physiology. 2007). 2006). which produce a free. Magnesian calcites: low. rier Reef Marine Park Authority. Anti. tional molecular analyses will be required to unravel the complex taxonomic relationships among these neglected anthozoans. lands Antilles.g. Nova Southeastern University Bill Newman. Growth and submarine fossilization of algal cup reefs. Tazoli et al. New York: Chapman and Hall: 68–95. a filamentous algal microborer. cally fixed carbon in the form of glycerol (reviewed in Taylor et al. some. Moberg F and Folke C. many of the Tuamotu Islands are developed on different sides than might be predicted from wind patterns alone. where boring sponges are common and reef growth is relatively slow. a framework of cal.calcite. acting as a reef. High phosphate uptake requirements of the scleractinian coral Stylophora pistillata. although filamentous microalgae and cyanobacteria commonly thread their way through the vertical elements of this group. but also provides a stable en. and depth on skeletal growth and form.g. The spicules within three sclerocytes have been desilicified with hydro. some species can be gonochoric. Harvesting impacts and invasion by an alien species decreases estimates of black coral yield off Maui. Rasheed L.. 150°E 145°E 140°E 15°S 20°S Tuam otu Archipelago Society Islands 135°E FRENCH POLYNESIA 500 km Tureia Marutea Maria Morane Mangareva Temoe Mururoa Tematangi Ahunui Nukutavake Vahitahi Pukarua Tatakoto Fakahina Pukapuka Amanu Hao Hikueru Reao Takapoto Niau Raroia Makemo Takaroa Manihi Ahe Arutua Rangiroa Tikehau Mataiva Tahaa Bora Bora Huahine Moorea Maiao Tahiti Tetiaroa Makatea Fakarava Anaa Hereheretue Rurutu Mopelia Tubuai Raivavae Duke of Gloucester islands Aratika Marokau Figure 1–11: Map of the Society Islands and the Tuamotu Archipelago. Mantyka and Bellwood 2007). cies appear to exhibit slow growth and cement.g. als. such algae may become abundant along with high coral cover and low levels of rapidly grow. or are passively surrounded by carbonate secreted by the host An Introduction to the Structure and Formation of Modern Reefs •5 attempted a combination of retrospective and contem. nisms and effects. 8•Chapter One may be a relatively simple.developed cir. and a rhombohedron. ated will produce the gametophyte plant (Figure 4–15). Paul and van Alstyne 1988a). or in rows. semblages and ecological controls on reefs and banks of the northwest Gulf of Mexico. huge meadows of Halimeda were discovered in a 2. Viewing organisms by some common function. 2003. a rate similar to that of black corals. ate at Tuvalu failed to reach volcanic substrata. but then packs developing embryos and more advanced larval forms into a dense gelatinous mass that is released into the fe. or other organisms. 1997. tially developed as flourishing Acropora palmata barrier reefs on a Pleistocene coral foundation. Reef response to sea level rise: keep up. Southwell MW. but they were more pronounced in stromatoporoids. Coral Reefs of the USA. Once or twice a year when the epitokes are mature. its cyanobacteria. and the Marianas that are described in more detail below.binding properties sufficient to cause nucleation and crystal a b c Figure 4–8: a) Scanning electron microscope (SEM) image of moderately well. Several cyanobacterial genera are common. b). Spirobranchus spp.000 species. and terminate with a protruding mouth surrounded by tapered. and its polysaccha. ———. The anterior portion of the sabellariid head (a pre. but is preserved owing to its incorpo. Dictyota dichotoma) and scleractinian coral Porites lutea in direct contact. PLoS ONE 6(7): e21670. at least in the best. 3000 2000 2500 1000 1500 500 0 Depth (m) Tuvalu boring 1896 Enewetak borings 1952 C o r a l V o lc a n ic b a s e 22•Chapter One High islands to the northwest. rather than aragonite (see discussion in Stanley 2006). coralline algae that can be a meter or more thick and thus become a primary frame builder at the reef crest. Colorado School of Mines Quarterly 80: 5–12. Vertical growth of the polyps pro. herbivores. however. Reid RP. where it harmlessly causes the host to grow in a folded form. Environment. mals.shaped Aphrocallistes vastus (Av). Glynn PW and Ault JS. Ferrier. ria. Tubes may be 7–1 Food and Feeding Categories Polychaetes as a group exhibit a broad spectrum of feeding habits. However. BioScience 40: 368–375. Panama 1: 695–700. Coral growth at Galeta continued for about 4.5 m high. polyp bail. spectral reflectance. there are brown algae with life cycle characteristics similar to Halimeda in which there is no diploid sporophyte phase. salty. such as β. 1976. 2009). ible investigations of a deep relict reef off the west coast of Barbados. European Journal of Phycology 39: 213–228. Field Guide to Anem. and is known primarily from some species of red algae and tunicates (Simkiss and Wilbur 1989).ray positive print of Porites astreoides from Cape Verde Islands off the NW African coast showing annual bands dating from 1959 to 2001.lived macroalgae. Anchored by a core of allophycocyanin and other phycobiliproteins. and if present at all. Figure 3–3: The structure of chloro. can be found at www. simu. they may settle close to the parental sponges or travel tens of kilometers. fied algae and hypercalcified sponges. from Bahia Vizcaino from Baja California. Thus rather than the production of generalized “pits” around the sheath. Particles are either rejected or used in tube building or for food.fertilization (within species). Hill D. 2005.g.%20Croix/salt_river179. Geochimica et Cosmochimica Acta 59: 5031–5036. 2007. are formed by specialized calcium. 1977).thick laminations that characterize modern and ancient stromatolites. ments. but 99% or more of sediment weight in such environments is due to carbonate and therefore sponges are minor contributors. New Caledonia is surrounded by the world’s largest island barrier reef system. complex mechanosensory cells that produce cnidae. sion behavior and may remain partially contracted for long periods. Conservation. Nugues et al. Croix. Smith CM. The Archaeocyatha (Greek for “ancient cup”) is an ex. Nava and Carballo 2008). 2010). ization and immuno. Cerrano C. Palaios 18: 236–255. Most species have been documented from water hun. many of these do not participate in reef build. riero G. Still others form only a mucus. and is a basin that may vary considerably in depth. Tribollet A. varies in complexity and efficiency. posure to air and freezing.water gorgonian skeleton (Keratoisis sp. 2011). the intense herbivory that occurs here is considered essential for maintaining coralline algal populations. distributed on reefs worldwide. The Society Islands of French Polynesia (Figure 1–11) appear to be associated with a hotspot located about 100 km southeast of Tahiti at ~18°S. making it the largest sponge. ding patterns in colonial corals.g. The Great Barrier Reef: Biology. in limited volumes of water that are exposed for long periods. when planulae are capable of settling and metamorphos. et al. and brachiopods (Chapter 13) among other organisms. calculations suggest that Figure 4–5: a) Halimeda is a dominant component of reef substrata in the tropics in all oceans and can become prolific through its vegetative growth. This process of nitrogen fixation is the same that occurs likely as symbionts early in the evolution of eukaryotic cells. roids record the appearance of Demospongiae during the Cryogenian period. Calothrix (Figure 3–8h). requiring large sponges to change the orga. Braga JC. However. there are instances in which typically extracellular coccoid cyanobacteria are taken up by host cells and appear to be in various stages of digestion (Rützler 1985. or a muscular pharynx to crush them (Fauchald and Jumars 1979). these structures called sclerodermites (Figure 6–15) have been interpreted as the fundamental units of growth in the scleractinian skeleton. where each face is a rhombus (Figure 2–5 b). Taylor et al. 2000. dant as fossils.g. Golubic S. tion. about 15x what it was before. nisms in photosynthesis. Bergey’s Manual of Systematic Bacteriology. competi. Symposia of the Zoological Society of London 25: 205–243. Figure 5–17b). pt. large colonies re. well. dissolved organic matter. 2007). ules typical of sponges (Figure 6–19).nutrient waters.Pacific. graphic axes of unequal length. Eco.like) is still used to describe heavily calci. but also bring in large amounts of sediment and decrease light energy to the detriment of reef growth and maintenance. In some cases the cyanobacterial biomass can nearly equal that of sponge cells. Such high densities may be explained by gregarious larval behavior. a strategy referred to as broadcast spawn. and the presence of characteristic groups of organisms.. The path that water takes begins with the hollow. These bacte. mon (Figure 5–16).like manner by gono.” Even though these structures are clearly wave. display a distinct windward and leeward reef structure (Figures 1–5). 2004. ited refuges. but crystal size and shape are uncontrolled. The calicodermal tissue is proposed to have a direct role in crystal formation.like bundles of aragonite radiate. Influence of Palythoa caribaeorum (Anthozoa. Photo courtesy of Paul Humann. a tissue specialized for digestion and absorption of nutrients. while this does not necessarily exclude them as sponges as shown above. respiratory CO 2 concentrations build up. ter 5) or deep. and Tebbut GE. A mul. a b c Figure 6–28: Mushroom corals (family Fungiidae) are gener. lobster. 2002). but others display as many as four function. 2003. they rely on herbivores to prevent them from being overtaken by more rapidly growing algae and other organisms. Cana. ated oral tentacles that create feeding currents and trap food particles (including phytoplankton. and in the absence of animal pigments. Cartwright C. but this number is likely to at least double as unexplored areas and cryptic species are given adequate attention (Van Soest 2007). surrounded by a typical atoll rim. it can spread over a broad area by growing on dead and broken coral fragments. The ecological evolution of reefs. Algological Studies 83: 107–127.. especially Palola viridis. B) 6. 2006. 2003. 2006. island. 267 Microbialite Reefs of the Precambrian. in which the female gametophyte produces a specialized diploid tissue that broods and in most cases nourishes its sexually produced progeny. In some areas the mean erosion rate by clionaids is in excess of 4 kg of calcium carbonate m–2 yr–1 (e. which also consume organisms within this size class (Pile 1999). Facies 54: 151–166. develops rapidly. perhaps 635 to >700 Mya on the basis of chemical evidence (e. but they may also con. and are a rich source of trapped detritus. Acta Palaeonto. The entire process can be very rapid. among other ser. ner and outer walls.000 chambers per cubic millimeter in complex sponges. Atoll Research Bulletin 507: 1–12. leuconoid form). USA. a rounded and muscular base. cyanobacteria have several other pig. 2004. These float to the surface and form an oily slick marking the event. Some of them. sues and canal systems. Proceedings of the American Academy of Sciences of the United States of America 103: 5442– 5447. in some cases. Several studies have shown that modern reefs often grew at a rate of 10–15 mm yr–1 for extended periods of time under conditions of rising sea levels (described in more detail below). an Extraordinary Calcifying Macrophyte Some of the most common chlorophytes occur on hard or sandy bottom in reef habitats. i.associated substrata..enhanced calcification. tion (described below). described below. ent levels that do not favor their growth (Belliveau and Paul 2002. Indeed. The system is essentially the same in serpulids. Solé. plasm include cyanophycin granules. Torres Strait. Van Gilder R. However. These generalities are still greatly simpli.d). Indeed. depending on the proportion of pigments. Calcinai B. or may partly or completely encircle islands. zinc. Groups of Sponges and Their Skeletal Structures Sponge skeletons provide a three. L. mentous cell layers in crustose coralline algae (Figure 4–1g). These are sometimes called “raised atolls. and other calcareous substrata. Indeed. including the amphinomid genus Hermodice. Gischler E. they will be described in more detail below. and are usually found in shallow water where the highest light levels occur.like. and red (Rhodophyta) algae. Figure 6–27a).mediated process. lopora damicornis. Mytilus edulis. Stokes MD. canic basalt. As efficient micropar. or macroalgae. Cell and Tissue Research 301: 299–309. 2008.level history. Dor. but the most profuse and globally extensive of them exist only in the fossil record. Coral reefs do not develop in tropical regions where river outflow is appreciable. and Goniastrea australensis. Schafer J.M reefs and without on 80. and nematodes.like struc. mass spawners are simultaneous hermaphrodites that produce eggs and sperm bound together and compressed into lipid. Geomorphology of coral reefs with spe. and a progressive reduction in size toward the posterior. The reef is dominated by the deep. where more than one. pable of binding calcium (Bilan and Usov 2001). tion in Haimeda discoidea Decaisne (Chlorophyta. and parts of French Polynesia in the south Pacific (Appendix 2b). deed. Pores allow for exchange of metabolites and projection of rhizo.conabio . however. Ma. the production of initial crys. refers to the lack of continuous cell walls and the production of a thallus that is essentially a large single cell. 1999. among other factors (e. Hadromerida).shaped “chapeirõs” in Brazil (Leão and Dominguez 2000). scribed in this section (Tubipora.less underside of plate. discrete island. and the rotation of the animal. hind the front lines (Ayre and Grosberg 2005). semblages could benefit from periodic upwelling of cool.bottom marine communities since the Cenozoic (Aguirre et al. whereas those in the Western Atlantic are more often constructional and are referred to as buttress and channel systems. nutrient input that encourages algal growth. the function of key enzymes. 2008).water reefs (Kinsey 1985. Integrative and Comparative Biology 19: 669–681. carbonate. among many others. Figure 2–4: A completely different form of aragonite with prism. cipally from exogenous food supplies in nutrient.300 species have been re. especially in shal. All three require firm substrata for larval attachment (Conway et al. Many Caribbean reef sponges have a thin ectosomal layer containing cyanobacteria. on cemented calcareous sand in the Bahamas. little is known of the boring mechanism. one that appears to be independent of light. the capacity to feed on plankton or particulates is quite variable and has been associated with broadcasting corals that release small eggs (200–400 µm) and undergo rapid development. These so. ian coral skeletons. letin 125: 1–44. ern Florida continental reef tract (Miami. synthetic symbionts found in sponges. 2011.magnesium calcites with 14–19 mol% MgCO 3 (Macintyre and Marshall 1988). There.developed para. 33 reflected by different arrangements of the parallel layers. the barrier extends all the way around and remarkably is connected to shore through its complex and deep lagoon by well. see Appendix 4) ini. distance dispersal of planula larvae of the coral Pocil. these morphological characteristics are a function of (a) the shape and upward or downward motion of the platform on which coral growth is tak.g. Journal of Phycology 34: 393–406. The Clitellata contains the earth. their size.. Corals in the genus Acropora. Colorado Plateau Geosystems. Right photo STS056–99–77. Land LS and Moore CH Jr. lites (Greek for “flat stone”) were the earliest and for a very long time produced the only calcified microbialites. and radiolarians (Foster et al. flat. which differ in complexity and composition.density packing of prismatic crystals. patch reefs can be created by the dissection of the main reef during storms. construction and other an. 1991. These are typically mounted at the end of a protractible pharynx that extends from the mouth. brush. According to ancient Polynesian beliefs. plications. Am. Indeed. are at work (review by Weiner and Dove 2003) and that trace element variation is far beyond what might be attributable to passive geochemical processes (Blamart et al. Purdy EG and Winterer EL. On one reef in the Virgin Islands. particles. orientation.g. for ex. Microfossils Foraminiferan tests are important indicators of paleotemperatures and other environmental condi.third of the levels compared with lighted conditions. other algal functional groups. Indeed. Bioactive metabolites from the Caribbean sponge Aka coral.001 0. 2005).like structure of its shelf. Axelsson L. An SEM. However..4 Gy (e. Försterra G. Multimineral calcareous deposits in the marine alga Acetabularia acetabulum (Chlorophyta. able to herbivores while facilitating their growth and survivorship (e. and attachment of Dictyota spp. and some may extend to 90 m or more (Wiens 1962). quired by maternal inheritance are viewed as specialists with genetic variants of zooxanthellae that have evolved with their hosts. often arranged in patterns or in particular areas of the colony (Cartwright 2003). turfs are often heterogeneous and may include a variety of microalgae. depending on the group. Filamentous green algae of the genus Ostreobium are common endoliths that are often associated with corals. heres to the skeleton with specialized attachment cells. Stetson’s Peak. nematocyst. vade dead coral tissue (Chazottes et al. Trends in Evolution and Ecology 14: 491–495. 2001. mation. 2004). ern Caribbean) and the eastern side of Panama facing the open Caribbean (Adey 1978). Great Bar. These functions improve the reef’s strength while maintaining topographic complexity (Mallela 2007). Ecosystems of the World. tubicolous. Photos courtesy of David Lee. gorgonian. Sponge. Ma. the Netherlands: Springer Science and Business Media. sediment movement. mental variables in the southern Mexican Caribbean. Hawaii. 1989. 2003). The Octocorals In addition to the hexacorals. and to the condition of reefs the way I remember them. ing from the constructional activity of organisms that secrete mineralized skeletons. South Africa). tive alarm system in this group (Nilsson 1994). 2006. et al. When herbivo. The chemical structure of nummulitid tests produced during the Early Cenozoic suggests they too were often symbiotic. Zilberberg et al. reef limestone. courtesy of the author. pore size and shape. or burrow into sediment. posed of green. nadian Journal of Zoology 84: 242–261. rely heavily on asexual reproduction. the crest reached its maximal eleva. As the sabellid rotates in this position using its thoracic parapodia. Thus. ———. Loss of foundation species: consequences for the structure and dynamics of forested ecosystems.and nanostructural characteristics of the scleractinian coral skeleton: a biocalcification proxy. about a quarter of these fishes are host specific.” By contrast. Grigg RW. Palaeo. where it often engages in self. ———. both open on their western sides (upper left and right). geographic location. 1995. suggesting that factors governing a species’ range may not be readily deduced from their travel potential. 2006). and as a place to brood eggs and young. Aragonite crystallization in primary cell cultures of multicellular isolates from a hard coral. the longest is the c. Coral Reefs 1: 29–34. Photos by author. they may also compete with and destroy living coral tissues as they grow. 258 13 A Brief History of Reefs and Corals.g. Some key findings and scenarios are outlined in Text Box 6–1. Photo courtesy of Robert Ginsburg. However. ments (Biegala and Raimbault 2008). produce a branched or massive skeleton composed of aragonite called the corallum. Berlin: Springer: 331–343. Journal of Experimental Marine Biology and Ecology 347: 97–108. 2000. and the Caribbean.based islands that are often best developed in the northeast and northwest. With few exceptions. iss JO. Proceedings of the National Academy of Sciences. ings. terparts. 2003. Steller DL. Note extensive cavities within the stromatolite formed by various boring organisms. ducing raised fringing and barrier reefs. but often occur with them in large numbers. this community forms a microenviron. Microbiology and Molecular Biology Reviews 71: 295–347. All three of these regions may calcify. ated with burrow formation. These poriferans are particularly aggressive borers. sheath.Frank et al. Leão de Moura R. Pen. for ex.up (right) shows the slender tentacles of the dactylozooids forming a network between branches. Figure 6–1). 34•Chapter Two Each of the axes is typically labeled with + and – signs to indicate their orientation. dinoflagellates. Stable amorphous calcium carbonate is the main component of the calcium storage structures of the crustacean Orchestia cavimana.like.Pacific. tentacle. structures called mesenterial filaments (Fig. Science 276: 1221–1229. I.gov/fs/2002/ fs025–02). 2005). and calcification are linked to the extent that even a cloudy day could reduce carbonate deposition by as much as 50%. Similar but more exten. 198 10 Reef Crustacea. and two forms of asexual fission.. The geometric relationships among corresponding crystal faces. rier reefs. interdigitating cells with numerous mitochondria. 2003.Verlag. Rocknest Platform. larly. or calcium carbonate. Branched canals called solenia con. syconoid form) and deeply embedded choanocyte chambers and canals (c. or stems.cyanodb. complanata. Scientia Marina 73: 739–746. reproduc.000 described species (Norton et al. Cairns. the entire thallus disintegrates and dies (Clifton 1997. stepping growth process in fibers. Scleractinians often add 2–20 mm to their skeletons per year. 1988. This diminishes upwelling intensity and allows a local current to transport warm tropical water southward along the continental shelf. glacially carved mountain valleys) in western Chile. depositing carbonate perpendicular to the current flow as a means of stiffening the skeleton. reproduction. making them resistant to bi.. USA 98: 11885–11890. 2002... planation is that in high. Cook Islands. and these provide reinforce. 2001. Thus. Journal of Paleontology 75: 1065–1078. In this book I undertook to write within and across not only my discipline. Figure 5–4: Free hexactine spicules and fused spicular skeleton of the reef. International Journal of Developmen. Anabaena. This process undermines and loosens silt. per) Unger (Corallinales. reef. Chu JWF and Leys SP. 1982. lack an internal framework. but only about a meter high. corniculatus and S. These processes break up the joints. two types of ectodermally derived tissues become readily rec. 1994. or symbiont consumption by the host (Wilkinson 1992). phonodictyon sp. whereas others from the Permian and Triassic periods were not known to form spicules at all (Pisera 2006). Coral Reefs 26: 517–529. Once initiated. and long. Zooxanthellae as. Palytoxin and ana. dant in this limestone deposited in the Late Pennsylvanian of eastern Kansas. but it is not yet clear whether the pathogenesis is due to the sponge. choring rhizoids that bind sediment in clumps (4–1d) and in some cases may function in the acquisition of nutrient elements. Members of this group are often dimorphic. Bulletin of the Geological Society of Denmark 45: 147–150. views 63: 106–127. thesis is leaked or “translocated” to the host in the form of low. although a number of other terms have been used to describe them. 2003. Davies PJ.115. Bo M. Roberts JM. but cells are unattached and produce a pseudo. Ball et al. reddish region is the accumulation body. 2008). 85% of Caribbean cyanosponges are symbiotic with this organism. lives directly on Lophelia pertussa and other deep. xanthellae take up residence and proliferate in symbiotic species. these limits change when comparing the eastern and western sides of continents. ing during precipitation of aragonite from seawater: a framework for understanding paleoproxies. Patten NL.html Pandolfi JM. however. Systematics of the Hexacorallia (Cnidaria: Anthozoa). populations north of the Pacific equator (e. and Klautau M.like structures (see below) rising from depths of 20–30 m (Figure 1–7). namic states. Photo courtesy of Peter Ralph. Paul et al.com) taken from para. The factors that regulate such densities are complex and may include the nu. structure. Atoll form varies considerably according to the island’s geological history and its biological re. Calothrix Typically has heterocysts at the base of the flament or near points of false branches. Journal of Experimental Zoology 22–36. deter. Hotspots and Reef Formation The Hawaiian Islands lie in a volcanic chain 2. Model modi. the primary photosynthetic molecule in most cyano. Preservation Inlet. Rahman and Oomori 2008). and Gulf of Mexico. Cerrano et al. Marine algae that calcify have a similar relation. making it diffi. the Parazoa. the cen. water exits by way of a single osculum or several oscula that form an “exhalent surface. duced coral growth referred to as the Darwin Point. certain types of heavily calci. phore. Indeed. Area of Figure 1–6b is also shown. The life cycle is typically heteromorphic. Leão de Moura et al. Öhman MC. brooding species at the expense of large and architectur. and in both intertidal and subtidal environments. and Western Australia. although diversity was low compared with today. continually updated database on marine sponge taxonomy can be found at www. produce a suite of these products (e. In addition. regardless of their taxonomic status and relationship with zooxanthel. and Recy J. especially in deeper waters associated with elevated substrata and strong currents. algal turfs are capable of high levels of primary production. 36 3 Reef Cyanobacteria. nal space. de. Weisz JB. These characteristics allow the capture of particulate organic matter (detritus). and terrestrial forests. barbed. gorgonians contribute to the three. which degrade to form microcrystalline Figure 4–6: Total dissolved inorganic carbon is represented by the sum of oceanic carbon dioxide. dou A. depending on the location. Plant Cell and Environ. Lagoons are not static and their depth can be modified by a variety of geological forces including up. 2006. but there are some marine species that are quite abun. Similar structures may occur in an island’s leeward side.building species that are found on both coasts of the Americas. On the orienta. Gischler E. their relatively simple structure and close association with the shoreline notwithstanding (see review by Kennedy and Woodroffe 2002). Figure 1–4) composed primarily of Palythoa is found under such conditions at depths of 1–5 m in the Caribbean and Brazil (Costa et al. as well as the one behind it. The fossil record shows that its skeleton is essentially unchanged since the Cretaceous more than 100 Mya. but they are shallower. cal Pacific (Harrison and Wallace 1990). as demonstrated by the pioneering work of Goreau and Goreau (1959). Patterns of coral distribution and benthic space partitioning on the fringing reefs of southern Taiwan. Microbiology Reviews 56: 340–373. Some atolls in this group are additionally unusual in being formed by small circular islands. the planulae of broadcast. Bioerosion and coral reef growth: a dynamic balance. although larviparous is probably a bet. which in turn brings in a wide variety of other consumers. and freshwater environments. and its volume is r2 h = ~13 cm3 . depends upon the nature of this reef structure. 1851. These are calcified. corals include gonochoric or hermaphroditic species. 2008). Nature 324: 55–58. nents of calcareous mounds in the Lower Cambrian of Labrador. Organisms that grow in the form of a calcareous sheet construct more permanent bonds and are referred to as biological ce. clude members of the genus Sarcophyton (family Alcyo. nated as “a” and “b. the term “sabellariid reef” is well established and these structures have been rec. As the following section describes. Heliofungia actiniformis shown at left has unusually large and bulbous. marine. dotentacles exposed during the day. 2000. cord. Middle Triassic coral growth bands and their implications for photosymbio. Spirobranchus larvae exhibit clear preferences for certain host species (Dai and Yang 1995. Baco 2007). and filamentous forms that do not form specialized heterocysts described below (order Oscillatoriales). the Red Sea. The phylum Cnidaria: a review of phylogenetic patterns and diversity 300 years after Linnaeus. 2006. Beavington. to our family. 1980. 1996). etes from heterospecifics. b c d a e f g h Gloeocapsa Lyngbya Microcoleus Granular colony composed of aggregated cells surrounded by mucilagenous envelopes. ogy 54: 24–29. and Loreto RM. associated anemones around the world serve as hosts for numerous species of shrimps that often occur as obligate symbionts (Chapter 10). a process called brooding (Sante. Fosså JH. the carbon for calcification comes primarily from CO 2 supplied by host respiration (mitochondria) in the calicoblast cells (Figure 6–17).cyanobacteria sym. when growth bands are present. nix effect” after the mythological bird that is reborn from its own ashes. phy will be explored further in Chapter 14. depending on geographic location or habitat (Veron 2000). bacteria. a coral without tentacles. tinellid sponges occur in large aggregations on the west. Ne. Rouse and Fitzhugh 1994). b) Isotope dates and other criteria from coral material preserved on the Huon Peninsula infer relative changes in sea level during the past 140. cific geometry owing to the presence of secondary nucle. but also for their alleged potency in the treatment of scorpion stings. 2006. and Nieuwland G. sulting in a fluoroapatite. as organisms that bore into rock and sand grains. Bermuda’s reefs at 32°N in the western Atlantic develop from the warm waters of the Gulf Stream system. Photos courtesy of the German. Jill. where. France. palmata. Jamaica. The Microbasic mastigophore Two types: shaft with distinct barbs.like crystals of aragonite laid down parallel to the shell surface. Atoll Re. chemis. 2007. Journal of the Marine Biological Association of the United Kingdom 88: 17–19. it is es. displays elevated solubility under these conditions. Weiner S. Akinetes Basal cell Sheath Reproductive cells Heterocyst 44•Chapter Three Nostocales) may secrete a relatively thick sheath that provides protection from rain or desiccation compared with those that are continuously submerged (Whitten and Potts 1982. In older. A fourth cycle. et al. and several studies have noted that they may stabilize storm. eds. but a relatively thick en. Northwest Territories.retractile tentacles (Figure 6–9b).poor water is the rule and nitrogen is a limiting factor in Cementing the Record The upper 4–5 cm of the subtidal stromatolites is soft relative to the older portions beneath. posium. a feature that sepa.building hexactinellid sponge Heterochone calyx. Tang S. Cannariato K.” or highly branched morphs ranging from those resem. however. long. and produce only small colo. velopmental Biology 17: 481–491.specific lower temperature limits. a taxon specific to marine sponges. Some of the variables that may influence carbonate bud. removing cytoplasm from Halimeda sp. 2003). son (1987) suggested that Caribbean species are rarely phototrophic compared with their tropical Pacific coun. they may compose up to 54% of coral rubble surfaces and 73% of cave interiors. tionships between atoll size and lagoon depths are also common. Geobios 23: 261–281. Pizarro V. 1997. Tomascik T. gorgonians are hosts to many different types of invertebrates including poly. Sheppard CRC. but can reproduce through a variety of asexual methods including binary fission. Thus. Montaggioni LF. Schwarz JA. 1954. Biological Bulletin 173: 110–125. NASA. carunculata. Canada. the spiral.. the aquiferous system. national Coral Reef Symposium. 2010. Overview Coral polyps.g. and others in which one or more volcanic peaks remain in the lagoon. International Journal of Cytology 65: 301–319. Pedrini AG. a sunscreen pigment that is produced by this group and is secreted into the sheath (Karentz 2001). Reef Growth and the Carbonate Budget Reef growth is a dynamic process involving carbon. Feio SS. The con. None. which in some cases be. ———. especially the longitudinal retractor mus. have been discovered in protected waters be. The youngest of them. Wasylenki LE. Simkiss K and Wilbur KM. but in others both gametes are flagellated and motile. A review of carbonate production and assessment methods agrees that gross carbonate production can vary considerably in different environments and locations.000. and erosion.Cheung and Williams 2006). line rock. 2005. there are examples of larvae that are able to take up symbionts while still in the plankton (Schwarz et al. cluding grouper. These organisms produce a three. Although gonochoric sponges (those with separate sexes) are not unusual. and appear to be undesirable food items.Bunker Group. An additional measure of protection may be attributed to the presence of an extremely complex and potent neurotoxin called palytoxin that is associated with Palythoa and other zoanthids. rounding the mouth. Because these crystals form on external cell surfaces rather than under direct cellular control. Palaeoclimatology.strewn reef flat that is now covered with a tidal lagoon of algae. Carbonic anhydrase in the scleractinian Stylophora pistillata: characterization. it was discovered that most of the reef was composed of Halimeda. book of the Protoctista: The Structure. Plausible mechanisms for the boring on carbonate by microbial phototrophs. These echinoderms prevent other sessile organisms from settling on the colony. Individual sponge species may also be quite ancient. when he first beholds one of these vast rings of coral. sand grains.. bers of gametes into the water column where external fertilization takes place. ticles that pass inspection are “spot welded” onto the medium. and extended all the way to Romania connected by a circumequatorial Tethys Sea (Figure 5–13). Liao M. Ingestion and assimila. 2007. cal properties that allow colonies to bend in moderate current forces. thus increasing their fluid output. the form and mineralogy of which varies among taxa and is the basis of classical taxonomy. 4 5 6 7 8 9 10 11 0. magnesium calcite crystals in the middle lamella and cell walls. New York: Macmillan).g. 1997). cation. and d) intratentacular budding includes partial separation of body wall as in Eusmilia fastigiata. Bryopsid algae may exhibit the typical biphasic life cycle of diploid sporophyte and haploid game. Most nummulitids became extinct during the late Eocene.dimensionality of the reefscape. Journal of Paleontology 63: 778–800. Journal of Experimental Marine Biology and Ecology 330: 448–454. bioerosion. including certain molluscs.occurring functional groups on the basis of thallus size. However.like reef system. ferent types of so. dian Journal of Zoology 84: 322–356.feeding guild) does not apply to functional groups involved in reef building (Precht 1994). sial and will be examined more closely in Chapter 15. Leys SP and Meech RW. 2006. Panama 1: 499–502. Mah AJ. Some of the Indo. Anderson 2004). and are especially significant in the boring and sculpting of limestone on tropical shores and on coral reefs. tions in collection of food as well as the propulsion mech. but it is now clear that zooxanthellae are clus. and Pueschel CM.shaped or more often semicircular large whorls (Figure 7–6a). Kamber BS. tion and conservation of precious corals. water scleractinians. 173°E). Coles SL and Jokiel P. Polyps in this group typically produce mesenteries that occur in multiples of six.perforate. 1991. Gametogenic tissue forms in ampullae similar to milleporids. certain filamentous rhodophytes are capable of killing coral tissue by adhesion and over. carry out photosyn. 2005). The Figure 6–12: Skeletal features of scleractinian corals: left. a b 74•Chapter Four Birrell CL. and in associated habitats. If it were considered a true atoll (e. Journal of the Marine Bio. Marine Ecology Progress Series 258: 79–86. eds. family Bangiaceae) are distinctly heteromorphic. Without these ele. named after lecithin. Figure 5–18: a) Cyanobionts in the mesohyl of the sponge Chon. onies may be hundreds of years old (e. 2007). and growth of tropical coralline algae (Rho. raphy 43: 1396–1399. Photo courtesy of Paul Humann.. Annual Review of Ecol.like.like tentacles (Figure 6–3c.like anal shield. tem (Figure 1–2) for example. in contrast.g. Some symbiotic sponges lose chlorophyll or symbiont cells under unfavorable conditions. plete envelope of living tissue. dermal glands that may assist in excavation. Atoll Research Bulletin 544: 37–54. all tropical oceans) and contain a single genus (Millepora) with 18 species. MacDonald KS III. The three blue dashed lines repre. Ganges. and grasses.5 Gya. 1 Bioerosion. posed to fresh.known and largest continental barrier reef in the world is the Great Barrier Reef (GBR). lation to predation by Acanthaster planci (Linnaeus).5 μm are trapped within. thus greatly expand. The reasons for this are in part the result of subsidence. Synchronous spawning: when timing is everything. the pharynx can extend nearly half the length of the animal. and some islands.shaped Onotoa (Kiribati. Sethmann and Wörheide (2008) re. tinucleate and syncytial. Taxonomic revision of the southwestern Atlantic Madracis and the descrip. Antarctica. rial together with connective tissue until more perma. the Helioporacea. A slower extension rate could reflect an increase in skeletal density due to a change in crystal arrangement or packing as the septa. Reef Corals and Their Allies•105 Millepore corals are a small but significant group of hydrozoans that are of considerable importance on coral reefs. al. despite intensive searches by submersibles and remotely operated ve. bon dating techniques. Journal of Structural Biology 150: 319–331. et al. Darwin’s subsidence hypothesis was generally confirmed. tors from the mineralization site. Macintyre IG and Marshall JF. Photo: ISS006. ultatively symbiotic temperate coral: zooxanthellae division and expulsion. Costa OS Jr.9 (cloth) ISBN.like marine protists. the development of giant axons. a modified radiole typically adorned with spikes that protectively closes the tube opening after withdrawal (Figure 7–4a). chemical weaponry is likely more effective than calcification because these groups produce only small amounts of mineral and are relatively fleshy compared with most species of Halim. and cause severe pain in sensate organisms. 2005). tinian hosts—symbiosis. Exploring cyanobacterial mutualisms. sus is that they are an extinct class within the phylum Porifera (Rowland 2001. coral reef carbonates should ac. Other cya. Meibom A. The Order Alcyonacea Alcyonaceans are a group of about 1.500 99. the ability of reefs to re. usable forms of both N and P may become critically low under conditions of food limitation. where they may project 30 m or more above the bottom. of. a condition referred to as bleaching. is one of only five islands in the group that do not take the form of an atoll. et al. The development of gills associated with the head. Hawaii. ans. Chemical versus mechanical bioerosion of coral reefs by bor. lia. Halimeda investiga. pages cm Includes bibliographical references and index. Patch reefs may reach the surface (or not). Larger. Marine Biology 151: 785–792. a series of shorter cilia on the oral side traps phytoplankton and detritus as well as sand particles in mucus. In many places.cleaning mechanism for coralline algae. Culture. ment of barrier reefs and atolls (Binard et al. Some species form thin crusts and grow relatively rapidly. Polychaete Reproduction Although some sabellids and serpulids reproduce asexually (e. a single Millepora colony can spread over an area of 2 m2 (Amaral et al. Caribbean Journal of Science 37: 227–238. of living species Percent marine Form Pigments Storage products Chlorophyta 7. and depressions that may retain water after high tide recedes (Figure 1–4). coral. Townsville 1: 263–272. and does not appear to be specifically controlled by an organic matrix. These gamete carriers may be smaller than 0. ern Canadian continental shelf off British Columbia where they form reefs up to 21 meters high in depths of 165–240 m. Morphology and evolution of reef islands in the Maldives. nematocyst. eds. 2006). fication occurs during the day. The sporophyte. 1997). low water. and Webb L. 2003. Foraminiferida The Foraminiferida. Figure 3–15: a) Bahamian stromatolite showing active cyano. terata: Octocorallia). prism. merous chaetae: bristles composed of chitin. dricks MGJ. about 260 Mya. Other banks are largely sub. 2nd ed.sperm bundles originating from some colonies. or bulbous tentacles typically arise from the space be. Upward arrows show the excurrent canals that converge on the inner depressed surface of the barrel. most are colonial and exhibit a wide range of Both tissue layers. Coral Reefs 16 (supplement) S:33. radial. Drew EA. Order Nostocales. 6–2) of which nematocysts are the most common and diverse. nation in sponges. forming synapses with muscle. Flatter shapes are chosen to line the inner surfaces of the tube. Endoscopic exploration of Red Sea coral reefs reveals dense populations of cavity. Stromatoporoids. its composition.. as well as deep volcanic peaks (seamounts).Pacific. For this I thank Walter Adey Smithsonian Institution US National Museum Denis Allemand. zil. Coral frameworks revis. Atolls have a well. the most common cyanosymbiont Synechococ. Bell PRF. out regard for political boundaries is close to 2. http://www. Demosponge spicules are typically coated with and connected by spongin (Figure 5–6). Mackenzie FT. Polychaete tubes may be secreted from spe.axis. Grajales et al 2007). but others form mat. selection becomes biased toward larger diameters in larger animals with larger building organs (Gruet 1984). They are more topographically complex than fringing reefs. After Glynn PW..g. the eggs and sperm of many are released over just a few days once a year. or both. 1996. 2008. stances where the segments themselves form cemented and layered banks (Figure 4–5c). Those with coral reefs are particularly valuable because skeletons typical of reef crests can be dated using both radiocarbon and uranium methods to establish the approximate sea level at the time of growth. Fringing reefs typically form narrow ribbons. Bio. they are composed of cemented sand grains. Partensky F. Reef Corals and Their Allies•141 corals in the Gulf of Mexico. Animal Behav. Hay ME. but suspension feeders are typically fixed in place and often possess a U. The essential biological feature that maintains the lagoon is differential coral growth. 116•Chapter Six fluctuations in temperature.sided pyramids when viewed from the equator. isms against solar radiation exposure: UV. Warmer waters tend to have less of the lighter 16 O compared with 18 O. ton 10: 100–109 (2001). Galgani I. includ.. Andréfouët S. Again. 1996. Even the characters of the cyclosys. Aharon P. Caribbean staghorn coral popula. rent systems can be quite variable. Hauser I. 1982. Scale approximate. 2001).6 Mya was marked by nu. Evaluating coral reef benthic communities in remote Caribbean atolls (Quitosueño.building species have an Figure 1–2: Amazon River discharge of sediment is between 3 million and 3. Juvenile Thalassoma amblycephalum Bleeker (Labridae. Brownlee and Taylor 2004). In some areas. ern GBR near Princess Charlotte Bay ~13°–14° south latitude showing the ribbon. their biomass and abundance may surpass that of algae. ern Pacific where it is typically found adhering to plant material. Sorokin YI. often exhibit a low incidence of self. Figure 5–13: Location of extensive Jurassic siliceous sponge reef belt. and on tiny Niau Atoll (Figure 1–13) closure has resulted in a very shallow lagoon that is distinctively green owing to abundant cyanobacterial mats (Intes and Caillart 1994. often in the form of specialized. In these environments macroalgal as. Coral reef invertebrates and algae are also a source of new medicines and other useful biochemical agents (Fenical 1996). cluding single. Marine Biology 126: 735–743. The Caribbean sponge Mycale laevis is often associated with the undersides of plate. is a triple threat (high sediment levels.like growths on en. alline” (coral.algal symbiosis: diversity. Other types of evidence sug. Coral Reefs 21: 188–190. Nature 458: 881–885.living zooxanthellae genotypes presented to them. in late spring or early sum. display a range of sexual reproductive modes from brooding to broadcast spawning. Courtesy of the Image Science & Analysis Laboratory. de. 2001. Each of these often has distinct sclerite morphologies that serve to stiffen the colony in distinct ways. to small fans or various arborescent forms.dwellers may function as binders if they temporarily hold loose carbonate to the reef. poorly calcified joints called nodes. and longevity (28 x 18 cm) that sold for $25. For example. and although they ap. cification. are endolithic and are associated with a wide vari. and spicules that differ in size. The reefs found to. Some species may brood fertilized eggs for only a few hours.stepping at the close of the last interglacial highstand. posed of overlapping. Rocas (South. as shown in the summary Equation 4–3: Consumption of CO 2 or HCO 3 – + H+ by photosynthesis will increase the pH and shift the equilibrium shown in Equation 4–1 to the right.Colinvaux (1980).water tropical corals (Chapter 6). and pre. While that may seem counterintuitive.Mortensen L. occurrence of chemical and structural defenses in the gorgonian corals of Guam. actively dividing meristematic tissues composed of stem. ing can thus be a significant contributor to bioerosion and can be especially common in live coral skeletons (Tribollet 2008). and because of high evaporation and low rainfall. Biological zonation at the base of a reef. b) Cyanobionts of the Caribbean sponge Xestospongia muta can be found in the mesohyl. 1988). Marine cyanobacteria in tropical regions: diversity and ecology. allowing it to remain flexible while resisting moderate current forces. et al. Indeed the term “atoll” is derived from the Maldivian “atholhu” meaning “lagoon island” (Darwin 1842). structed by Ficopomatus enigmaticus. 81 The Structural and Functional Roles of Sponges on Reefs. 2004. for every chip of carbonate that is mobilized. Figure 6–26: Global distribution of cold/deep water coral reefs. Kobluk DR. 1991) and can cover the bottom completely or in patches. Photo courtesy of Stanislas Dubois. remaining competent but able to delay settle. which may impart red. 2010. Apatite in biological systems usually contains “impurities” that alter the Ca:P ratio and the properties of the mineral. Earth Science Reviews 60: 195– 225. and the large numbers of sclerites within the colony. fensively. and Brown RM. 4 Coral Reefs. 39 Overview.green algae” because of their superficial resemblance to some eukary. ———. Here. Cer. The polyps cannot retreat into the skeleton. b). Tetrahedron 57: 9347–9377. Kudla ML. etration and storage of sponge spicules in tissues and coelom of spongivorous echinoids. croboring: implications for the preservation of skel. tabolites produced by gorgonians that are symbiotic with zooxanthellae are apparently ineffective as a defense. Panama 1: 689–694. Ball et al. Potgieter M. 2000.like. Hexactinellids produce more silica per unit of bio. rently recognized by classical taxonomy (11 of 18) con.building organisms may be referred to as primary reef builders if they contribute large portions of the reef framework. tion and emerged from the sea as a boulder. a. the mouth and pharynx can be protruded from the body in forms that grasp prey. but produce irregular emergent islands and reefs. Brachert TC. 1989. Chemoecology 9: 187–207. 1986. suggesting that the two groups calcify in a similar manner. Carpenter et al. In addition. minifera promote calcification by elevating their intra. rounding dactylopores constitute an important feature of classical taxonomy called a cyclosystem (e. and while that may be true for some. survivorship. Conversely. lobe. Some of the most prominent groups in. the sclero. 5–2).. Harrington et al. The surfaces of mature segments are polygonal. vations on egg release in the oviparous sponge Chon. This suggests that the absence of allelochemicals may be a factor in restricting the distribution of palatable sponges to cryptic reef habitats (Wulff 2005). tress. 1997. including the thorax and a morpho. 2005). however. or some combination.generated storm waves. Oxford: Oxford University Press. perhaps unfortunately.. implying that a process of decalcification and re. Reef Evolution. and in fresh water. forming paired fanlike. mangrove environments. Photochemistry and Photobiology 3: 428–434. ized structures in response to competitors within their reach. 14•Chapter One more open with fewer islands. Australia. herms. Stylasterines Stylasterine or lace corals typically produce a deli. The latter type is brittle. Benayahu Y. These exclusively colonial animals typically produce an arbo. bellids may be prominent among these borers. many sponges harbor specific symbionts that are apparently required by their hosts and have developed an obligate symbiosis with them. 2006. Photo by author. such as Niau (box left and b) with its distinctive green lagoon. sonal variations in the growth of a staghorn coral. growth. manuscript editor. for example. The Cnidaria is divided into two distinct lineages. ing a reef framework. Life and Death of Coral Reefs. The septa occur in parallel to the mesenteries. and Faure G. Biomineralization within vesicles: the calcite of coccoliths.building corals and algae. they are often collected as curios and for use as aquarium decorations. they were placed in their own class (the Sclerospongiae) when discovered over 40 years ago off the north coast of Jamaica (Hartman and Goreau 1970).. mine the development of such island reefs. Fossils of this genus are at least 250 million years old and correspond to a diverse group of ancient hy. and Pawlik JR. nificance for productivity and nutrient cycling in shallow. although it occurs among solitary corals (Cairns 1988). but offer little organic 70•Chapter Four gal spores that would otherwise germinate and quickly overgrow and dominate turfs (Adey 1998. Reef Corals and Their Allies•111 number of colonies. Roberts HH. However. clownfishes take refuge in the host’s tentacles (Figure 6–8b). Calcified Anthozoans The Scleractinians True stony corals are placed in the hexacoral order Scleractinia.g. in the genus Lithothamnion. although it is unclear whether the mucus is derived from the symbiont.reinforced capsule. As far as is known from laboratory observation. Golubic S. Hydrobiologia 548: 207–215. Coral Reefs 18: 170. Holland ND.containing organic material.Alsumard T. University of Alberta.like depression that may vary from 10 to more than 100 m deep. 144 Overview. and may exhibit alternation of sexual and asexual generations. including Maui. Effects of toughness. This type of division. they contain a continuous record of the environment in which the coral grew. The long larval lifetime appears to be a function of stored energy reserves. 2003. and most of them form siliceous spicules as well. lar system.5 million metric tons per day.rich cal. In some cases. and other factors that will not be considered here (but see Scott and Ro. Budding as a means of clonal growth increases the number of polyps that compose a colony. Ayre DJ and Miller KJ. Grosjean E. binder. Richter et al. tion and feeding. 367 3. Sexual reproduction. –a 1 –c –a 2 +a 2 +c +a 1 36•Chapter Two called coccolithophorids can create highly complex ex.000 years old (Roark et al. it is often necessary to identify them by the way X. line algal life cycle (red arrows) begins with separate haploid gametophyte plants that pro. Roncador Bank.shore rock pools: spatiotemporal variation in settlement and per.water corals. protons are required for bicarbonate up. even though details of the process have not been elucidated and remain con. New species of Calcaronea (Porifera: Calcarea) from cryptic habitats of the southern Great Barrier Reef (Heron Island and Wistari Reef. Becerro MA. shape. nian corals against predatory fishes. Tuvalu.noaa. its polyps. plants. and is referred to as “mother of pearl. The Biology and Geology of Coral Reefs. b) Colony of the blue coral Heliopora coerulea showing its Millepora.. coincident with low levels of solar radiation. ditions that would in turn require some separation of photosynthesis from respiration. Shallow reef growth that was not uplifted and pre. Opdyke BN. cludes two types of plants. high levels of dissolved organic matter (e. These organisms are still important on reefs today in processes of primary production and nitrogen fixation. larger galleries may reflect a taxon.minute intervals during the morning. The Great Chagos Bank is one such structure south of the Mal. 1976. trophic relationships. sealed junctions indicate that all of the organic and inorganic constituents of the skeleton either pass through or are derived from the calicodermis. These are referred to as “give. The glands are lined with smooth muscle and secrete organic matter and calcium salts through ducts found near the collar opening (e. A large number of species occupy the insides of tu. like structure composed of interwoven multinucleate fil. with esti. In situ growth rates of the scleractinian coral Oculina varicosa occurring with zooxanthellae on 6. an in. ter for fishes (e. Hydrobiologia 471: 1–12. The result is a calcified material that grows with a char. but are also important to coral reef structure. Zoological Studies 46: 227–241. ing activities and are being mapped for conservation management (Moore et al. Hecke A. cintyre and Reid 1995).000 years ago (Figure 1–21b). 1974. which are demosponges that nei. Preferred orientation of the fan. ings P. Australia. Highsmith 1982). porophyte that releases diploid carpospores.Coulon IJ. Different species possess characteristics that favor their abundance depending on wave activity. In addition. codermis and to conduct protons out (Figure 6–17). Geller et al. Nakamura T and Nakamori T. ———. bance.evolved with their By forming the choanocyte layer into a sinusoidal pat. Shallow.like porous. and eas. typified by anthozoans. tributors to reef construction. the presence of larval zooxanthellae does not necessarily change the competency period (see also Nishikawa et al.. and Mats The mucilaginous sheath of cyanobacteria may reflect their adaptation to various environmental conditions. or in combination with the mechanical effects of wave or current activity. 1997. Cyanobacteria are among the most ancient and abundant photosynthetic bacteria. and in turn its adaptation to differ. Symbiont.. professional underwater photographer and friend for many years. some owing to the presence of photosymbionts as well as distinct spicular architecture and cellular organization. Fukami H.48. Most scleractinians. thus allowing it to retain more water and provide habitat for other small or. and carnivory. 2010.magnesium calcites contain less than that.green color. small barbs along length of tubule. lar control of development in the reef coral.sea Corals and Coral Reefs.g. is also exhibited by a handful of other anthozoans (references in Reitzel et al. perimental Biology 98: 239–267. Science 275: 1080–1081.rich families the most lethal chemicals in nature. break. as a response to bleaching). Drew 1983. Wagner M. sitional variations at ultra. tinct microscleres. DC: Joseph Henry Press: 83–108. and in more protected environments the shore may de. Both of these can inflict severe injury on their neighbors. 39 Cyanobacterial Form and Function. centric bands in cross section. the site of protein synthesis. During the sexual phase. López. calcium transport declines by about 50% (Al. ers. Thin crusts (< 1 cm) seal the surface of reefs on which they are abun. and can engage in food Figure 6–27: a) Bamboo corals such as Keratoisis flexibilis can grow to over a meter tall and are common on rocky deep. lithic microorganisms across the northern Great Barrier Reef (Australia). Dunne T. Journal of Experimental Marine Biology and Ecology 213: 281–293. 2006).tubules or a helical array of rodlets long threadlike processes that are armed with numer. and is considered overfished. Hillis LW.g. ture of the substrata. 2001. the current consen. 138•Chapter Six significant transport of coral and symbiont cells. Branched colonies may respond simi. ysis Laboratory. or produce a skeleton. tance of sea level changes as well as local processes that shape reef formation (see historical account by Riegl and Dodge 2008). Huston M. Courtesy of Image Science and Analysis Laboratory. ing up to 1. Decalcification during epithallial cell turnover in Jania adherens (Cor. Hubbard DK. 1998. Pulido GA. and are included in the supergroup Rhizopoda (Adl et al. 2b. Bayer (1961). Simulation of nitro. There are numerous examples of atoll. 2001. sure and the removal of respiratory wastes and other products of metabolism. Walter MR. The initial Solentia cell is a motile. Johnson Space Center. Indeed. cene initiation and development of New Caledonian fringing reefs. cant on reefs. Barnes DJ and Taylor RB. Most carbon. Many of my colleagues contributed by reading and critiquing chap. Adjustment of Bikini Atoll to ocean waves. 1984. sequent calcification) may produce granular and fibrous crystal growth simultaneously. Andréfouët S. Note that both branched and fan.000 to 5. that may involve hor. b) Scanning electron micrograph of S. and pigments that may obscure the shell’s translucence. Algae and Sym. When the Shark Bay stromatolites were discov. ing organisms. Figure 4–14: Brown algae do not calcify with the exception of the genus Padina. depending on species and geography. Ca. 2006. Colony surface Gonad Pinnule Oral disk Mesenterial flament Mesentery Tentacle Coenenchyme Central chord Solenial canals Axial skeleton Paraplexaura Indo-Pacifc Pseudopterogorgia Western Atlantic Gorgonia Western Atlantic Plexaura Western Atlantic Melithaea Indo-Pacifc Junceella Indo-Pacifc Figure 6–19: Diagram.. and another that is distinctively jointed with alternating calcareous and gorgonin segments (bamboo corals. tures called pit connections. ———. Virus predation by sponges is a new nutrient. 2004. calcification.producing taxa (Cairns and Macintyre 1992). Release of both eggs and sperm is common. 1999. In some cases. And because calcite is more durable than aragonite. 2006. verse suite of calcifying and noncalcifying organisms. Area and volume change disproportionately during growth. and during those intervals supply nutrients to their sur. however. 1985. many develop a more complex organization that may include double nerve nets (epidermal and gastrodermal) with connections across the mesoglea. synthetic rates have been measured using a number of conditions and techniques as a means of estimating an energy balance sheet. gress Series 126: 145–152. Some authors have suggested that rainfall is the best predictor of lagoon structure. loid gametes that look alike. perhaps in response to elevated nutrient levels (Ward. The Coralline Red Algae. although they may have similar evolutionary his. Several families of heterogeneous ancestry achieve rela. nists often refer to this condition as coenocytic. Several species of Gerardia are harvested commercially in international waters where their collection is unregu. even during the day (Paerl and Carlton 1988). Acosta A. microalgae. Proceedings of the National Academy of Sciences.retractile tentacles (Figure 6–10). or can be traced to an ac. but nonetheless may be well circulated at low tide by overflow from surf.like structure to a profusion of short branches or knobs.3 individuals per square centimeter. Riding R.dimensional framework that not only defines an architectural space. Skeletal microstruc. heterocystous filaments that are capable of producing extensive and rapidly growing mats.thorns sea star’s tube feet (Chapter 11).symbiont relationship is disrupted by these stresses. I am especially grateful to Paul Humann. eastern Indian Ocean. Filamentous cyanobacteria. tion rates are also likely to exceed carbonate production (Grigg 2006). The terrestrial 10% is composed of rubble. ical Conference. asexual reproduction among anemones almost always results in separate individuals rather than colonial forms. hydrogen ions (H+ ) must be cotransported as an ion pair to maintain electrical neutrality.calcium ratios within the bands are also useful environmental indicators. Global diversity of the Stylasteridae (Cni. Toonen RJ. ceedings of the 8th International Coral Reef Sympo. Conceptacles form deep within the meristem. This suggests that elevated levels of photosynthetic oxygen. Journal of Foraminiferal Research 9: 1–13.226. or by multiple fission (order Pleurocapsales). the branchial crown is a highly photosensi. Sedi. A “zoanthid zone” (e.building sponges. 2006). These are typically tube dwellers and are referred to as selective deposit feeders. having been damaged and reduced by as much as 50% Other families of boring sponges are capable of ex. members of the genus Acropora (Figure 6–30a). and with some exceptions. if not a principal component of ancient reefs (Webb 2001). or both fuel the process (e. Earth Science Reviews 67: 219–265. and the smaller microgamete is designated as sperm. Pandolfi JM and Jackson JBC. Char.g. On evolution in the Anthozoa. and growth rates of black coral in the southern fiords of New Zea. chanical forces of wave activity.forming abdominal segments (Glasby et al. skeletal slice through Diploria labyrinthiformis. ent hydrodynamic regimes (Koehl 1982). However. subjecting some regions to more rapid rates of change than others. Green areas represent chloroplasts. the switch from a reducing to an oxidizing atmosphere corresponds to the Proterozoic era of Precambrian time. Eunice norvegica. 2004. and the overall cyclosystem pattern are still among the most common characters used for this purpose. ed. These structures typically occur in pairs. 1986). Exuma Cays. mers (carbohydrate. Fis. and most shallow. as well as the pali and columellae. trogen fixation on a coral reef. As sea levels rise. Chicago 60637 The University of Chicago Press. plate. cumulation of the crystalline metabolite in the sponge tissue. which lie outside of the Park boundaries. typically by fragmentation. there are specific nucleation regions and sites that maintain crystal orientation. perature consistent with data from other parts of the world that this was a glacial period. gonite needles that form are biologically induced rather than specifically controlled by the alga. ometry (length. This is the crystal form of the calcium carbonate poly. Fungiids similar to these may reproduce clonally from asexual buds as they die or when they are injured. and hydrocarbon deposits. Rhodoliths: between rocks and soft places. Marine sponges as microbial fermenters. Whitton BA and Potts M. forming a mesh. Wulff JL and Buss LW. Blum JE. The removal of these muscles reveals attachment scars composed of a specialized region called the myostracum (Figure 2–4). Hughes RN. 2005). and Hen. Photosynthesis Research 73: 59–61. They also differ in a number of other characteristics including cell wall structure. 52 The Chlorophyta. Guadalupe Mountains.g. low them to behave as large. 2011. ment sites for corals and other invertebrates. The Biology of Reefs and Reef Organisms Walter M. Cell. Bulletin of Marine Science 81: 311–322.000 species (Van der Land 2006) includes a di. as is physical erosion (e. climatology. Each gastropore. Does light enhance calcification in hermatypic corals? Marine Biology 80: 1–6. ———. but it is always perforated with small pores that allow the cytoplasm to stream through (see below). which lacks collagen and instead contains chitin. the main Hawaiian Islands with elevations of several hundred to >1. interfere with cardiovascular function. Fish and Aquatic Resources Series 12. although some may have a life expectancy of five years. covering more than 50% of carbonate surfaces not occu. Photo cred. len G.Verlag. Camoin GF. Geister J. and distinct tissue layers. Indeed. characterized by three horizontal axes of equal length that intersect at 120° (referred to as a 1. 2000. 1969. or produces pores that promote nucleation and maintain crystal alignment (Weiner and Addadi 1997). courtesy of Marine Ecology Progress Series 301: 159–171. that deter predators. these skeletons are either weakly cal. and the formation of a calcareous cement. In some species (under favorable conditions). Calcification and photosynthesis in reef. including species of Dipolydora and some (but not all) species of Polydora. the tissue is otherwise undefended. Young reefs that develop on slowly sub. nificant differences between day and night bioerosion rates. Palaios 15: 248–254. b) Detail of crys. and they produce only a Bayer FM and Owre HB.. Ginsburg RN and Schroeder JH.Karger FE. Courtesy of Jaap Bax. Effect of daylight variations on the en. This species is the most durable of the reef building sponges. 1977..or endoparasites and may have scissor. D. In. duce large colonies and reefs at depths of over 80 m with. Some reefs. Nematocysts. Galera J. and c3). they are ahermatypic). ros et al. It appears to be especially well suited to dead or partially dead corals and coralline algae. Norton TA.Water Octocorallia of the West Indian Region. A skeletal spine called a gastrostyle typically projects into the gastrozooid center. Brazil (Srivastava 1999). 154 8 Reef Mollusca. They may be found in the open ocean as prominent members of the phytoplankton.000 km2 it would be the largest in the world. Exca. Hendler G. and Hansen U. desiccation.Alsumard T. This zone is often partly or completely uncovered at low tide. ample. Metabolism. or Niau Atoll described above (Tuamotu Archipelago).8 kg m–2 of reef material to be converted into sand and silt. phases of a coral population. cement. In addition. density. Coral growth bands: a new and easy. Estimating reef growth as it occurs is a bit more difficult. Nature 1996. a metabolite of the burrowing sponge Si. The common name refers to the proteinaceous. Albuquerque Cays. 2004. Observations of the tissue. mis. although less obviously. 2006.called star corals. cyanin and phycoerythrin extend as a series of rods made of stacked discs. from which fan. Brooding corals retain eggs that are fertilized by sperm released from hermaphroditic or gonochoric males. cal of reef habitats. Bioerosion Many marine cyanobacteria have developed a re. baffler. Bischoff WD. midal dihexagonal crystal (Figure 2–6b). 2002). Bishop FC. resulting in local sponge populations that are chemically undefended (Nuñez et al. as is the case with calcite. ited number of corals. Fan. the choanocytes that line the hollow inter. Kojis 1986). Current Biol. Sponges with a large spongeocoel (asconoid form. Microbiology and Molecular Biology Re. Borowitzka MA. and Loya Y. based CO 2 concentrating mechanism in Laminaria saccharina. exhibiting phototrophy at high light levels and heterot. terium. Tissue and Cell 34: 232–245.g. Teavanui pass is up to 40 m deep. ure 7–7b). In Ardbear Loch.based analysis of the morphology. portant. excava. often 5–10 mm be. Indian Ocean). Cook. Coral Reefs 25: 77–84. ing place. although there is considerable de. It has also been suggested that high Palythoa growth rates may reduce the amount of shelter available to reef fishes by overgrowing crevices and limiting food resources (Mendonça. A single family. High abundance of diazotrophic picocyanobacteria ( west Pacific coral lagoon. Pagès C. soups. 6: 173–178. Recognition and selection of settlement sub. However. McMurray et al. or by calcification in some species. Macintyre IG. a shelf atoll on the Campeche Bank off the Yucatan Peninsula (Mexican Caribbean).induced mor. da Gama BAP. Florida International University Steve Cairns.up of the distinctive 8 pinnate tentacles typical of all octocorals. Molecular Ecology 17: 2937–2947. and/or proteinaceous skeletons (modified from Cairns 2007). 1: 8–13. although some red algae lack pit connections and instead form syncytial fila. Discosoma sanctithomae. several have only one such gap. In at least some species. 2003). on the reef’s protected leeward side.0. tions of producing nonmotile sperm (Searles 1980). Some of these are macrophagous and consume seagrasses or macroalgae using chitinous jaws to cut out bite. they are often referred to as coral bioherms. a reproductive mode called external brooding. but has more distinct windward and leeward structure (Fig.Caribbean region (Ross 1977). populations have declined by >50% owing to overharvesting (Am. ium tenue. While the mechanics and causes of bleaching are well documented among corals and zoo. Blamart D. bacteria contribute to the black color. 2003.g. Prochloro. to form an operculum.. Several important genera and species are prominent in this role. but studies of skeletal growth rings as a function of height indicate that vertical growth may be less than 3 cm yr–1 in certain shallow. An Introduction to the Anthozoa Cnidarians that only form polyps and never produce medusae are members of the Anthozoa. genera. 2000). For example. ing marine lichens (Golubic et al.specificity observed among symbi. One of the more important of the latter groups is the genus Palythoa. wide and porous bowls.symbiont specificity in soritid foraminifera. for example. and >4. Life on the margin: implications of ocean acidification on Mg. 70 Foraminiferida. it is worth noting that “meristem” in coralline algae refers to a layer of dividing vegeta. order Oscillatoriales. 1991. and Wang X. tures of the corallum have traditionally served as one of the principal means by which species are distinguished. teria. western Pacific Ocean). This rich. Muscatine L. albeit at considerably different qualitative and quantitative lev. Compte Rendu Pal Evol 3: 453–467..e show intraten. which do not provide zooxanthellae with the egg (reviewed by Baird et al. Riegl BM. b) close.Pacific and C and H in the western Atlantic (Appendix 3). chlorophytes). and Miller KJ. ———. which involves a coccoid. and other portions of the skeleton thicken. tory as well as food. The two best known are found in the Exuma Islands in the Bahamas. The actual process is quite complex.like growth pattern Figure 3–13: a) Location map of Shark Bay and Hamelin Pool stromatolites. Indeed. Marine Ecology Progress Series 299: 157–166. to high tide spillover or storm waves. Het. cal of steroids. ciety B 275: 2335–2343. bean. The latter are widespread and quite abundant in polar seas. and depth records. wave. Photo courtesy of Mark Wilson. corals including alcyonaceans and gorgonians are broad.up” or “drowned” reefs. closely aligned. It follows that such structures do not develop on the more sheltered leeward side. a topic that will be discussed in more detail later (Chapter 15). pt. of which well over 800 are associated with coral reefs (Veron 2000. tensive serpulid reefs known thus far have been found in a fjord at 8–32 m depths in Antarctica. Some species employ mucus ropes that are periodically pulled into the mouth and swallowed along with the adherent material. Indeed. Bayer and Owre 1968. in.Australian reef en. typic because they lack symbionts. despite the absence of a nervous system. 2003. This reef thus “caught up” with sea level rise. Phycologia 44: 156–162. the highest diversity of algal genera is found in the temperate coastal waters of Japan and South Aus. 2003.dimensional symmetries characteristic of different crystals. simple tubule only. 1978. However. 2002). tures vary widely among members of this group. Godfrey JS and Golding TJ.000 years. spermatia) must break through organic matter that plugs the pore to the surface. a group that comprises the most important structural contributors to coral reefs worldwide. Oceanogra. but are not regarded as true barriers be. Today the derivative term “cor. c) Reef growth is limited beyond the Darwin Point. 158 Overview. thereby contributing calcareous fragments to the sediment. and P. Love et al. 1997a. 2001). either throughout the enameloid or as a thin. and brown colors. Title. but originate in the parathoracic region below (Gruet et al. grew in crusts and were abundant in cavities where they com.and low. USA 102: 18063–18068. merged. Recent work indicates that some coral reef sponges are capable of capturing viruses and virus. ily when the parent is moribund or injured (e. all of which have formed on the continental shelf. two species of anemones associate with a wider diversity of anemone. tem (Hubbard et al. tion and increased traction over various surfaces. for example. erogeneous group of families included within the Holaxo. The region immediately posterior to the radioles is differentiated into a distinct thorax specialized for tube secretion.000 m thick. or bacteria on the substratum. and Indian oceans in temperate as well as tropical regions in depths often exceeding 300 m. This South Pacific caviar. ous plate. Some fjords (drowned. spongarium. 2005). A few studies have 6•Chapter One oped than those near the mainland. Indone. ence 69: 535–546. 1998. Cavanaugh CM. This probably explains why these structures grow at about 1 mm per year in this environment. at Conch Reef in the Florida Keys. isms. whereas more irregular sizes are incorporated onto the outside. Biological Bulletin 117: 59–75. cifiers. Minerals with identical chemical composition but different crystal structures are referred to as polymorphs. on the aboral region.. letin 60: 1054. Rougerie F. Deep. unlike most hexacorals. synthetic symbionts that prefers cracks. in turn.000 years. Deep. 2009). Steep slopes and injection of fresh volcanic material play a role in maintaining a patch. male conceptacles becomes the first of two diploid (2n) sporophyte generations called the carposporophyte. Thus. Photo courtesy of Robin Smith. 2003. www. Li J. 104 Introduction to the Anthozoa. and are anatomically similar to serpulids. Carib.S20. many have that ability. Likewise. In: Birkeland C.rock. after which various processes and organisms cement. building organ. typically overgrows hemispherical coral colonies Photo courtesy of Jonathan Dowell. production of minerals in bundles and layers is sometimes distinguished as sheath calcification (Borow. Morphological plasticity in the tropical sponge Anthosigmella varians: responses to predators and wave energy. Invertebrate Biology 124: 91–97. with lagoon depths typically exceeding 40 m (British Admiralty map 1014 [2002]). 1998. Both of these are terpenes or terpene. Wörheide G. and Brachert T. 2001). and other mobile invertebrates is significantly higher in the presence of glass sponges compared with nearby areas that lack them (Chu and Leys 2010). Light attenuation by limestone rock and its constraint on the depth distribution of endolithic algae and bac. In: McClintock JB and Baker BJ. males and females may be found swarming at the ocean surface to spawn. Small organic particles are con. 2nd ed. and can yield up to 10 kg m2 yr–1 (Hart and Kench 2007). under ledges. Most species associate with several anemones. lithoherms. which is sometimes distinguished as the ecto.sea corals in the lower Hawaiian chain. These are composed pri. 367 1.Holocene deep.C by author: A. Bulletin of Marine Science 69: 915–931. b) Reefs with shallow. adjacent branches tend to fuse. Although there are several genera of sponge. This information is based on the ratio of oxygen isotopes incorporated into the carbonate. and Wilson EO.exposed areas. d = dissepiments. All of these function as binders. Marine Ecology Progress Series 256: 87–97. Hydrozoa. and it is not always clear how such topography forms in every location.to.Colinvaux L. These “phylloid” (leafy) algae. and depth. is typically an internally fertilizing hermaphroditic brooder across much of its range. forming what is now a bank. reefs composed primar. The Saba Bank. although some reef areas can tolerate excursions below this level (e.shaped Rose Atoll at lower right has a single channel on its NW side. combinations of such forms are common (Figure 6–4 a. ming. there are many other exceptions. Marques AC and Collins AG.like form. Budd AF. NOAA. a group in which heterotrophic bacteria and Figure 5–17: Increasing complexity of form and surface area of choanoderm from simple tube (a. ria labyrinthiformis from Bermuda. Journal of Eukaryotic Microbiology 52: 399–451. Sediment interferes with coral feeding. the souls of the dead had to pass through Tupai on their way to the afterworld. and gor. anism that drives water through the animal.de/~fboehm /ceratoporella. In: Du. cular sphincter muscles along the body wall. Rigby JK and Senowbari. while larger hosts can be found at least briefly without any anemonefish. Krupp DA. less heavily calcified crustose corallines are often broken by herbivore activity and by storms.magnesium calcite that are formed in the cytoplasm and are then transported to the surface. Shark Bay Hamelin Hamelin Pool Pool Hamelin Pool INDIAN OCEAN WESTERN AUSTRALIA Shark Bay WESTERN AUSTRALIA 25°E 26°E 27°E 113°S 114°S 115°S a b Reef Cyanobacteria•47 The initial cell. Despite being the sole author. These free.like skeletal parti. Langdon C.called black corals and certain unusual zoanthids referred to as gold corals. 2006. Coral defense against macroalgae: differential effects of mesenterial filaments on the green alga Halimeda opuntia. western Great Bahama Bank.efficiency suspension feeding on phy. dominated by the serpulids Galeolaria hystrix (red crown). the outermost polyps produce specialized sweeper tentacles with large nematocysts. ences in planula larvae of two species of scleractinian corals. ing.producing sponges. which can be composed of protein.like sperm are released by spermcasting in the Caribbean barrel sponge Xestospongia muta and carried to the egg. 2002. Work TM. pulids (e. Some of these are generally distributed among a large number of species. and Loya Y. composed of weakly pigmented or nonpigmented internal cells. Srivastava NK. Net carbonate production (the amount re.Multiscale remote sensing of microbial mats in an atoll environ. but involve far fewer species (Harrison and Wallace 1990. ing and spawning behavior. near the edge of a continental or island shelf. The cyanophyte Mastigocoleus testarum (order Stigonematales). ———. do not display robust growth. Cy. synthesis takes place. and Sulak KJ. Maida M. Hallock P. Sammarco PW. b) Corallimorphar. ian Archipelago (Grigg 1997). Miami 1: 23–29. With one known exception (Perkins 1991). Figure 6–20d.. and atolls that occur on continental shelves (Figure 1–7). or vice versa). Birenheide R.Pacific reefs are typically dominated by other species. However. 2007). 1996). 2010. tubular portions of the porocytes. Indeed. Marín and Ros JD. Castenholz RW. Proceedings of the 8th International Coral Reef Symposium. linity.faced crystal showing the c. Macro. Bulletin of Marine Science 76: 1–26.deficient lagoon reef environ. taches. SW Caribbean Sea. Macroalgae that are anchored to the sediment by rhizoids are generally referred to as rhizophytes.” the gorgonian Keratoisis flexibilis. New York: Chapman and Hall: 298–353. amination of past growth. Those that initially exhibit delayed growth but later accelerate reef accretion are said to “catch up. although a few may reach a length of several meters. Host specificity of the symbiotic cyanobacterium Oscillatoria spongeliae in marine sponges. he found another 776 animals including 464 isopod crustaceans that parasitize the snapping shrimp. The ecological evolution of reefs. and Riding R. or on inorganic surfaces as in rocks and minerals. perhaps suggesting that they may be inclined to look elsewhere for a meal with a greater nutrition. As with many other sponge residents. Jamaica.reef growth: a dynamic balance. Cyanobacteria are common in shallow (1–2 m) waters. At one time.200 km2 . colonial morphology. Thus. and Diaz. 2007. Journal of Experi. Journal of Phycology 42: 292–303. ing are some of these factors. The role of sea. some cyanobacterial species are capable of simultaneous photosynthesis and carbonate dissolution. 1987. 2008). bial diversity of marine sponges. other sessile invertebrates. Halimeda bioherms of the northern Great Barrier Reef. compe. diversity. laneous crustaceans. dermis. some clionaid species grow more quickly than others (Schönberg and Wilkinson 2001). Photo courtesy of J. stoloniferans. phosphate is one of the primary forms of phosphorous taken up by zooxanthellae and is a key component for the synthesis of important biological molecules such as ATP.like reproductive struc. although there is one that protects Kaneohe Bay (Oahu). Muller. ogy 18: 619–627. a potentially toxic waste product for eukaryotes. cupied by other anemonefish species. Effects of low water temperature on growth and magnesium carbonate concentrations in the cold. The effects of density on the zoan. Perhaps as a consequence. generalizations concerning reproductive mode are often difficult to reconcile with simple models.0. treme longevity in proteinaceous deep. Bastidas C.g. However. 1985. tic’s coral reefs.building organism. and typically composed of 1–3 cells (e. and Nitrogen Metabolism The cycling of nitrogen is an important process on coral reefs and is mediated primarily by microorganisms. uble mollusc.being of their hosts. port Ca+2 into the calcifying site and move 2H+ to the outside. tospongia muta showing the reddish brown ectosome layer with cyanobacterial symbionts. Molecular iden. Studies of metabolic similarities and ribosomal RNA sequences. nellid sponges (Etnoyer and Morgan 2005. Proceedings of the National Academy of Sciences. 2007). tion of organic material. Marine plant. The inner shell is often iridescent because of the way light is Reef Minerals and Mineralization 2 Figure 2–1: Shell morphology and structure of the common mus. and distant storms can generate large swells from any direction (Stoddart 1969. each of which has a lagoon. 1994. Stanley GD and Cairns SD. which presumably function in respiration. Ultimately. roxygenic. Unson et al. 1992.2 kg m2 y–1 for the entire sys. Lembke.algal reefs. Halimeda bioherms along an open seaway: Miskito Channel. There are thought to be 14 hotspots in the Pa. and reef accretion rate in the shallow fore reef. Hexactinellids produce a skeletal framework of fused spicules that result from syncytial cellular networks.. ous and fleshy) exhibit a biphasic life cycle. Reef growth measures the long. dis.like growths over 2 m across (Rigby and Senowbari. However. Australia. Jamaica. even these estimates do not take into account poorly studied groups including turf algae and little. An Introduction to the Structure and Formation of Modern Reefs •31 pended sediments in the lower Amazon River of Bra. or the taxon and its growth form. typically those producing small colonies. and there are additional descriptive terms that are sometimes used in reference to the habits of rock. 1999). This forms a cemented. servations from laboratory experiments using visible and ultraviolet light. but most of these also capture suspended matter (Murray 1991). Marine Ecology Progress Series 373: 265–273. In certain uncalcified rhizophytes (e. This stage releases haploid tetraspores that give rise once again to gameto. or other geometric shapes. carea (subphylum Cellularia). McDonald JI. pelago.corals) are not reef builders. Aguilar C.like palps called radioles. In addition. Marine Ecology Progress Series 177: 177–187. or expo. lue in Indonesia (Brown 2005. 1985. Coral Reefs 14: 27–33. the tentacles are flexible and exhibit complex individual movements during feeding. branching corals may form thickets several kilo. refer to hydrozoan and anthozoan cnidarians that produce skel.dimensional structure as “coral forests. and Weiner S. with special ref. for example. viduals. 1987. 1997. Klumpp DW and Polunin NVC. exposure.density aggre. Darwin’s ideas have been modified by many who suc. Panama 1: 731–736. Sabellids are distributed worldwide in association with a variety of substrates. isolated.bound chloroplasts that contain stacks of thylakoids called grana. mary factor controlling their persistence and growth. Smithsonian Institution US National Museum Laurie Collins. Montaggioni LF. 1994. Grigg RW.g.) is the most common hypercalcified on fossil reefs and are similar to living groups of demosponges. or eggs. the earliest life. Caulerpales): a mechanism for survival after separa. p. CA function is an important mechanism of carbon concentration by zooxanthellae (Weiss et al. miniferan species attached to calcareous algae. which when liber. Ralph PJ. regular. wide. Reviews of Mineralogy and Geochemistry 54: 1–31. Heinrich K. for example. especially terpenoid compounds. Some of them may be present only seasonally or they may occur as generalists found in a wide variety of organisms in addition to sponges. Macintyre IG. dosomal mass that lacks such symbionts. especially free.water hosts favor clade C. 1999. and finfish. De Jong I. Journal of Sedimentary Research 33: 291–303. many create a rigid skeletal framework composed of hypersi. ure 6–20c). 1995. Dasycladidae). tion device across the entrance to their U. both on and below the ocean surface (Figure 3–16). some deep. tal Microbiology 10: 2910–2921. thus decreasing pH and inhibiting calcification. Lophelia’s growth rate is estimated at about 7 mm per year (Fosså et al. a metabo. within that time frame. Passive suspension feeders rely solely on current or other water motion to move food particles to the organ. Chapter 6). the Octocorallia. and 4. Rhodoliths may initiate their formation by fragmen. and biogeography of Symbiodinium. the nature and extent of communities that develop in lagoon ecosystems can be quite diverse. A geochemical model for coral reef formation. but also contain cells that calcify and are responsible for producing a substantial aragonitic skeleton (Lewis 2006. soft corals. These are suspension feeders. The thin living tissue at the surface is organized into a series of small volcano. In contrast. tures that resemble colonies. 2000. and dinoflagellates (reviewed by Taylor et al. 206 The Malacostraca. Viruses. P. 1997. gies. Cowart JD. occupies a cen. is reversed by other heterotrophic bacteria when they engage in de. overall morphology. Photosynthesis. ed. 1 Frame Reef Formation and Structure. The diversity of existing red. and differs according to depth. cies (reviewed by Ten Hove and Van den Hurk 1993).763 km2 . but a few descendants of these LBF remain. Deep. Christensen RJ. Most often. erts JM. but are instead passive endoliths (Scof.” This circuit forms the sponge’s aquiferous sys.spawning (Acro. nitrogen limitation may not apply to col. Nonetheless. in con. but the polyps may then separate and ap. which are discussed further in later chapters. Cyanobacteria. UNESCO. they also occur on exposed reefs in the Atlantic including the oceanside faces of the Lesser Antilles (east. lack zooxanthellae entirely. The larval lifetime. carbonic or formic and lactic acids) that dissolve carbonate. Chlorophyll is composed of a complex ring of conjugated (alternating double and single) bonds called a porphyrin that contains a magnesium atom at its center. favorable temperatures in deeper waters. These are found in Atlantic. stylasterine corals are gono. tress system. et al. but are unfazed by calcification. 2005). ation sites. In many cases. In: Birkeland C. croenvironment. Morrison CL. tween the mesenteries and thus.sized sand grains that are rejected by the oral region. Some of these compounds are ca. Davies PS. Koslow T. Smaller polychaetes that live among sand grains and consume micro. These may cluster on the parent sponge and form replicate struc. that is. clionaids often associate with zooxanthellae. pend on several factors including the type of carbonate or coral host (Zea and Weil 2003. ed. Among the more than 10. and many other areas (Appendix 2). etate.like side branches called pinnules (Figure 6–18a). 1991. The tentacles.ATPase and carbonic anhydrase remain active in the calicoblast cells.10° C (Cairns 1999. United States National Museum of Natural History. perhaps 3 mm or more deep.Forero et al. Colonies are gonochoric and develop either eggs or sperm attached to tiny hydro. and then initiate the process of asexual colony formation (Figure 6–3e). and the differential development of thoracic and abdominal segments. unicellular algae.g. that inhibits coral growth. there are some interesting cases where barrier reefs and fringing reefs coexist on the same floor at depths of 3. Corl. spheric Administration. Coral reefs of Brazil: an overview. 2000) and may serve as another source of nutrition in the form of photosynthetic production.water pinnacles or reefs. These are placed in the orthorhombic crystal system.shaped colonies are preferentially oriented. volcanic peaks capped with limestone. Rogers AD. Larvae produced by P. especially herbivorous sea urchins. Ga. Sabellariids are found both intertidally and subtidally in wave.g. Talendier J. Nicol 1931). 2005. dive Archipelago in the Indian Ocean (Appendix 2a).gov. a section called Récif des Française forms an unusual double barrier reef with two lagoons (Figure 1–8). Intracellular protons create a relatively acidic environment that can facilitate conversion of bicarbonate to CO 2 for photosynthesis. Usher KM. whereas others. Many of them are actively growing. Cohen A and McConnaughey TA. This layer may be shed periodically to remove epiphytes (Littler and Littler 1999). citic sclerites fused by aragonitic cement (Schuhmacher 1997). Brazil. Taylor and Wilson 2003). actively growing coral reefs are therefore typically restricted to depths of less than 60 m (Schuhmacher and Zibrowius 1985).g. sociated with cyanobacteria. tipatharia. which are often referred to as one circumtropical species. Austin WC.Pacific oceanic atolls that are exposed to frequent. rous fishes. 2001). Indeed. Tambutté E.sized pieces. the bag is detached. and Stuckenrath R. a major component of vertebrate bone and tooth enamel represented by the formula Ca 5(PO 4) 3(OH). Octocoral. 1974. the carbonate budget. The other coccoid prochlorophyte genus Prochloron enjoys a much more restricted distribution. and Badran MI. eda opuntia and in H. fluoric acid to show the silicalemma membrane and the silacatein (*).like holdfasts (Figure 4–4). 2005. In contrast. 1988. called phycocyanin (Figure 3–6). Using multiple skeletal characters. Some species are less than passive and lash their ciliated palps through the water. Spain. sociated with foraminiferans typically fall into groups F and G in the Indo. Constructional azoo. Weiner S and Dove PM. Indeed. proposed proto. Brownlee C.Horani FA. ley 2009). There are several mechanisms by which sediments and the reef framework can be consolidated. They also serve as symbionts for many different groups of organisms. Krautter M. ceedings of the National Academy of Sciences. Coffroth and Lasker 1998). warmer Pleistocene climates caused deglaciation and rising sea levels. 1998.dimensional crystal faces. ogy. and Roberts CA. and like other such reefs. erling 1988). University of Western Austra. Wyrwoll K. 62•Chapter Four are highest. or by the density of carbonate produced as weight per unit volume. soybeans. Geo. or red. Pink mats stimulated by nutrients that have been transported by rivers produce large blooms in the Great Barrier Reef lagoon. This topic will be discussed in more detail in Chapter 15. gia muta. sponges. they are the most abundant on Earth. Smith CM. ternal shells. A small but significant group of about 15 living species produces massive aragonitic or calcitic skeletons. and many other groups that actively create holes and tunnels.West Pacific opistho. Bickmeyer U. Riegl B. Symbiosis in a fac. perimental Marine Biology and Ecology 278: 47–65. intercellular cortical chambers. Con. 2007). including those offshore of the Florida Keys. Ac. dition to algae and a variety of invertebrates and fishes. fouling effects of epithallial shedding in three crustose coralline algae (Rhodophyta. round New Caledonia. 1989. and other groups in which the primary suspension. tite. biomass confined primarily to the surface b. Some authors use the term raptorial feeding to describe organisms that grab their prey with pharynges or jaws without regard to size or whether their prey is of animal or plant origin (Glasby et al. strata.. and displayed great morphological diversity (Wood 1999). crustose algae. that is. Do as. McCartney MS. although increasing numbers are being described from depths of Opresko and Sánchez 2005). plate. (c). In such cases. and lagoon that is 20 m deep (US Fish and Wildlife Service 2005). whereas the lagoon of spindle. Under certain conditions. the life histo. Encrusting organisms such as sponges. as shown in deep water off the coast of NW Washington. University of Melbourne. the pH at the mineralizing site in. lids display brilliantly colored. c) Cross. Melkonian M. ———. et al. Induced colonization of corals by a clionid bioeroding sponge. 2004. Figure 1–17: a) Map of the Hawaiian Archipelago and the position of 29°N. The mineral com. Reuter M. Gattuso JP. 1979. Payri 1988). Figure 3–14: a) Location of the Exuma Cays and surrounding areas. Likewise. 2001.000 km away. A “CO 2 supply” mechanism in zooxanthellate cnidarians: role of carbonic anhydrase. and Quinn RJ. spiraled tests 2–4 cm across. ships among nitrogen fixation. aeorum (Duchassaing and Michelotti. 2006. Note that some bands are visible owing to deposition of organic matter. 296 Diversity and Life Cycle Processes. tors to reef sediments. Indeed. the xylem and phloem. rial origin.hydroxyapatite is often associated with fluoroapatite. 2005. Alaska). Species of Halimeda have been important sedimen. Silverman J. Some carnivo. Conway KW. tached. all of which have a symbiotic relationship with non. Spon. cess of growth.suspension by waves and currents. Once a colonial patch is established. nies by external forces (e. others with little yolk that feed primarily on plankton. on the chloroplast membrane. Other fossil sponges called the stromatoporoids (Figure 5–15b) produced skeletons whose succes. Appendix 2b). source use by juvenile parrotfishes in the Florida Keys. Marine and Fresh. forms and are called Darwinian atolls. such features have been described from numerous locations and are strongly associated with the direction of incom. Okazaki M. sand grains. suggesting a binding function.shaped.dimensional structure that is distinct from and is elevated above the surrounding unconsolidated sediment. Members of the genus Tricho. most sponges are composed of three regions. b). 1994. culiar alignment of crystals with protein fibrils (which does not occur in most other vertebrates). Spicules that extend from the mesohyl and project through the pinacoderm may have a deterrent effect.forming deep. Science 317: 1903–1906. north Jamaica. For example. Overview Sponges. rounds each crystal as a template. In such cases the chances of coral development are reduced. Photo courtesy of Paul Kay.like crystals that are formed perpendicular to and just out. Map courtesy of Ronald Blakey. they also commonly produce hybrids. and tunnel into the sub. are located in the animal’s anterior and posterior regions. Ramicrusta textilis sp. 1992. Antifungal activity of (+). Each of the two spore. including hypercalci. 1991. 2002. as well in deeper waters of the western Pacific. constituting another limitation to fertilization. effective removal of epiphytes often requires or. some low islands are composed of raised limestone. The sponges that spanned Europe. Feeding may also be accomplished by pseu. ment that accumulates detritus (finely divided organic material) and houses invertebrates that are detritivores. Red dots show the distribution of warm/shallow. Conversely. Similar but less extensive aggregations have been found at depths of >100 m in a fjord near South Georgia Island in the sub. whereas older segments tend to contain the less effective halimedatrial tetraacetate (Hay and Fenical 1988. phies. 160 Molluscs as Carbonate Builders. mon among anemones than it is among scleractinians.moving Polychaetes. However. systems: hsp70 expression by the giant barrel sponge Xestospongia muta. posed of a thick outer lipopolysaccharide layer in addi. respondingly exhibit an elevated frequency of clionaid sponge occurrence. ian colonies or dense populations of smaller ones are often found on current.000 years. 1998). ponent of the axis and holdfast of some gorgonacean Reef Corals and Their Allies•137 stricted trans. nary abundance of hydrocorals (Cnidaria. Irwin L. rassic. 2007.like Barque Canada Reef is es. Science 227: 57–59. posed of stacks of flat. Epitoke Atoke a b 154•Chapter Seven scraping organic matter from rocky surfaces. Bruckner AW. 6–13).shelf reefs of the Great Barrier Reef. Biotic as. Grigg RW. tum constitute the hypothallus.5 cm. Indeed.image axes.shaped or semicircular. which projects above water at low tide (see Figure 4. The new higher. capitata. Littler MM. et al. Coral Fisheries All of these corals have been fished since antiquity. Geomorphology of the southeast. tity of photosynthetic symbionts in Caribbean coral reef sponge assemblages. Risk MJ. classi. and other reef organisms (Diaz and Rüt.water marine cal. and the process is generally referred to as nonselective deposit feeding. Some species employ laceration in response to food limitation or substrate instability (Clayton 1985. either on the reef surface or between and under coral colonies where there is no living tissue. Thus. Seymour JR. and zooxanthellae genet. cur. 2000.barriers” (James and Macintyre 1985. and nutrient limitation. 1924. turned. 1977. but also obtain their green cryptic (=hidden) color by incorporating Halimeda’s chloroplasts. Although some species grew to modest sizes. often heterotrophs or phototrophs with low light requirements. position in demosponges: spiculogenesis in Crambe crambe. all of which actively involve the pro. 2005). The Shallow. toward sea level. alized. the entire host. Al. cord in corals can be many decades long. Coral Reefs 23: 595. nies can be several meters tall. In: Gupta BKS. Life and Death of Coral Reefs. which occur in a pH. This wave climate.1 1 pH Proportion of total CO2 CO 2 HCO 3– CO 32– Range of seawater pH Figure 4–7: a) New segments are spongy and white. plus tannins and other dis. Biological Bulletin 199: 76–84. showing the features of the prostomium. but they also extend almost 200 km to the northwest from the island.eating worm from Bermuda: Branchiosyllis oculata (Polychaeta. Biology and ecology of the hydrocoral Millepora on coral reefs. 2008). Reef Algae and Foraminiferans•77 Riding R. selves. and both sides of the tropi. some species further prepare the meal for digestion by grinding it using their muscular pharynx. ing coral reefs. rosecretory and ciliated sensory cells of various types. light limitation may restrict the depth of excavation to the upper 1–2 cm. ciation of Petroleum Geologists. Krone C. Control of nitrogen fixation by oxygen depletion in surface.known examples of this multifunc. or as encrusting growths with polyps deeply embedded in a coenenchymal mat. Netherlands Antilles.celled. Boyer M. Sperm are released in a cloud. Chemical and mechani. ments outside of the tropics. especially among Indo. especially parrotfishes. Figure 6–5: Colony of the shallow. and reef community structure.. Frame builders may include certain calcareous algae (Chapter 4) that form branched or encrusting structures. Marine Ecology. Others were club. Therefore. are carbonate borers found in a variety of calcareous substrates. The calcareous reef alga Halimeda (Chlorophyta. gloff 1988. Credits: b. and curls into a semi.of. styles in the sea anemone Anthopleura elegantissima. and Miyata M. some areas did indeed exhibit these qualities and were home to echinoderms. This type of community is referred to as an algal turf (Figure 4–1 a. mon among branching scleractinians.g. tute examples of hardground coral communities. a b Reef Corals and Their Allies•129 giidae) produce large and distinctive solitary polyps (Fig. Dodge. and most of the corals de. nual temperature is typically greater than 20°C during the coolest part of the year. 152 Polychaete Reproduction. The northernmost of the group is about 80 My old. Earth Science Reviews 62: 1–103. as a visible landmark to of. Some AZ corals form large colonies and thus make important contributions to coral reefs under certain conditions. although other functions have been proposed. The one shown here. emy of Sciences. ted or porous. as well as in other outposts of the south.000 13 unicellular macrophytic filamentous chl a. 2001. Straits of Florida. The segments of both types break open in the early morning as they participate in a fertilization ritual that is literally disengaged from the parents. the larger islands of Fiji. constitute calcareous islands with a three. Nitrogen fixation by Trichodesmium spp. Titlyanov EA. More than half of the 36 snapping shrimp species found in sponges on the Belize Barrier Reef came from a single host species. partitions called septa subdivide each corallite. prise the most diverse of all marine habitats. and prevent coloniza. Coral Reefs 6: 227–236. tween the corallites. a c b An Introduction to the Structure and Formation of Modern Reefs •11 2003). Figure 1–13: The Tuamotu atolls are composed of small islands that tend to be more prominent on the east. involves the formation of an inorganic solid that results from. Seipel K and Schmid V. salinity. http://www. The distinction between corals found on coral reefs and corals capable of building them is not always straightforward.like levels of organi. As later described more fully. US Fish and Wildlife Service. considering the massive amounts of dense magnesium calcite and the small amount of or. 6–13). and other skeletal debris often fill in some of the spaces within the reef framework. Once established. Hughes et al.chilled water also likely played a role in the cessation of reef growth here and elsewhere in the Bahamas (Hine and Steinmetz 1984. grazing activity and cyanobacterial diversity are substantially reduced com. Taxonomy of the Caribbean excavating sponge species complex Cliona caribbaea– C. 6–4a). dense fields of Halimeda provide architecture and protection from grazing. Skeletal spines not only surround the polyps. 1989. parts specialized for grazing on rocky surfaces (Nagarkar et al. ullary filaments interconnect the Halimeda thallus. temperature.dwelling microorganisms. an opal. 226 Echinoidea. Impacts of shading on sponge. Coin = 18 mm. The pre. Such long delays in settlement may be due to very large energy a b Broadcast-spawning corals Brooding corals Cell division after spawning Release of egg-sperm bundles cDevelopment of zygote dPlanula formation e Mature planula Planula larva Release of planula larvae fSettlement. break free and engage in swarm. though some occupy abandoned gastropod shells. Low. Moya A. French Polynesia. There are 13 families whose members are herbivorous. goldberg is professor emeritus of marine biological sciences at Florida International University. tals.. To the northwest of the barrier reefs are atolls and submerged banks. Some shallow. Zootaxa 1668: 127–182. 2007. 1996. no. followed by barrier reefs (center). the reef flat typically develops various types of islands that serve to define an interior lagoon. 26 December 2004. both shallow. Thus. the aragonite poly. ders and families that are not closely related. 1986. 2004).g. Cyanobacterial Form and Function Cyanobacteria are ubiquitous in terrestrial.org/fishery /culturedspecies/Porphyra_spp/en). Both forms of light reduction may increase respiratory costs. Purverel et al. small. most rely on the extraction of food particles from water they pump.. but placed them in their own phylum. the maximum depth at which endolithic cyanobacteria are typically found (Schneider and Le Campion. 226 Overview. there is ample indirect evidence to support a close relationship between cellular activity and the process of calcification. ter. quired taste. 1998). Fishelson L. morph vaterite. Khaled bin Sultan Living Oceans Foundation. 2007b. tially evolved >350 Mya as small spherical forms with tests composed of characteristic microscopic granules. and other factors (Chazottes et al. mas: a partially drowned carbonate platform. Reef. Demo. cols. drape across the sediment. Polychaetes of the family Sylli. Certain groups are more likely to produce high. Colman AS. and gametes are produced from sexually reproducing haploid individuals. 2008. most brown seaweeds are erect mac. and Lasker HR. The number of zooxanthellae per host cell may vary from 1 to 3. Renaud G.Holocene rise in sea level. also recorded from reef material. whereas those that form tunnels or burrows are endolithic macroborers. ings of the 3rd International Coral Reef Symposium. 113°E) is oddly spindle. Part 2. which form a rectangular crystal face (Figure 2–8). University of Stuttgart. 2006. An Introduction to the Structure and Formation of Modern Reefs •23 tive deep mantle plume. QH95. cal feeding guilds. Scientia Marina 68: 227–241. time of gamete release. ship with symbiotic dinoflagellates called zooxan. and were then tectonically lifted out of the water. ery of the intercortical spaces. London: Graham and Trotman. a shallow (~1 m deep). the haploid gametophyte and the diploid sporophyte. although the average is less than 30% (Rützler and Macintyre 1978). Fakarava channel. Inorganic materials are also sorted in some species. filaments can adhere to each other forming thin sheets called films. These include seaweeds that may develop intricate tissues but do not possess true roots. Structure and com. Boston: Jones and Bartlett: 698–714.walled heterocyst that contains nitrogenase (e. the northeast and southeast Atlantic coasts. The Atlantic has one barrier reef area in addition to that of Belize. There are at least 25 different types that can be recognized microscopically (e. Figure 6–27b). calcified crust or upright cups with pit connections between cells and conceptacle. thorax. Courtesy of Melanie Berg and Glacier National Park. but await the arrival of current. Any errors that may remain and sins of omission. Na. 2008). duce asexually. and the Atlantic coasts of South America and Australia (e. Hexactinellida and Demospongiae) that evolved first from the Urmetazoa during the Proterozoic: a review. Calcified seaweeds on coral reefs: com.building corals and algae. 2007. giving them a collective advantage that is reflected in enhanced growth rates.” (Darwin 1842) The third major Darwinian reef category is the atoll. 2005). lik JR. The overall process is summarized by Equation 3–1: There can be little doubt about the historical impor.dependent settlement and mortality structure. they are referred to as filamentous cyanobacteria. Sherwood OA. resulting in a worldwide lowering of sea level. ment. when they covered thousands of kilometers across the shallow waters of many continents. 2001). Cocito S. Demise. forams on the islands of the Indo. Hadromerida. and to well over 100 m in other locations.like layers (laminae) of these organisms and their muci. are often abundant and reach large sizes in such deep. NASA.building corals and algae. Marine Ecology Progress Series 352: 177–185.half to one. 1999. lectively referred to as the hypercalcified sponges. Figure 5–8: The calcareous spicules typical of the class Calcarea are triradiates. and Thomassin BA. some reefs occur in the Gulf’s southern re. 2007. Individual crystals are typically fused together.and filamentous algal populations (and other epiphytes) increase at the expense of CCA. all hosts were thought to be associated with a single species. Photos by author. Fish predation on Caribbean reef sponges: an emerging perspective on chemical defenses. one could argue that French Frigate Shoals is a remnant barrier because of the pres. Bahamas. 2010. Palaeontology 43: 671–703. stony corals. Interactions between corals and their symbiotic algae. and even certain echinoderms produce large and visible burrows or tunnels. Gil. 2009). Reef Sponges•89 Sphinctozoans are sponges that can be traced from the Cambrian and typically grew as a series of chambers one on top of the other (Figure 5–15c). bastrea aurea. such an accounting process. and Hill RT. Cryptone. Biological Bulletin 202: 86–95. Photosynthesis and calcification in the calcifying algae Halimeda discoidea studied with microsensors. which become light green 1–2 days after their for. 2000. The movement of water by choanocyte flagella is criti. In. 2000). Coral Reefs 27: 529–539. Inhibition of this enzyme sharply reduces (but does not stop) calcification. lous. Arillo A. for scale (ar. Thus. Systema Porif.magnesium calcites. but on Enewetak Atoll in the mid. Treballs de la Societat Catalana de Biologia 59: 29–49. but their ecological significance lies in environ. including certain foraminiferans. 1972. Some of these are found on the surface tissues. In: Frost SH. contributing an estimated 40%–50 % of the phytoplanktonic biomass and up to 80% of the primary production in these open ocean regions (Partensky et al. boring sponges. Spirobranchus giganteus. Else. Facies 38: 1–30. Life and Death of Coral Reefs. Hay 1997b. Borers. s = septa. Indian Ocean). Its zooxanthellae (representing several clades) are concentrated in the center of the test (Figure 4–19 a.1371/journal.. extinct twin. Goldberg The Biology of Reefs and Reef Organisms THE UNIVERSITY OF CHICAGO PRESS Chicago and London walter m. These aspects of reproduction and coral reef biogeogra.g.swept environments (e. ever. If extracellular mu. Photo by author.like endosymbionts in acroporid cor. These characteristics in turn provide additional settle. The fans are all members of the family Gorgoniidae. Bruckner et al. In addition. Berlin: Springer. com. terisk. Gerar.like growth that extends a few millimeters over the substratum. “corals. the rapid evolution and demise of certain groups define distinct periods of geologi. tions along reef productivity gradients. ous nematocysts. langae (Porifera. ———. Here they often constitute a significant food source for a variety of animals. Duan D. or both) that initiate nucle. Distribution. Lasker HR. the effect of calcium limitation.e.. Borowitzka MA and Larkum AWD.g. three. the larger of which has an average depth of 25 meters. Anabaena A heterocystous group capable of forming well-developed akinetes.protected areas for the Seaflower Biosphere Reserve.L. Brachert et al.. Marine Ecology. http://metafysica. tion to planula formation (b–d) typi. 2007). southeast Florida coast. 2008. Lazar B. they form a few centi. While there is some disagreement over their taxonomic affinities. 1989. and eunicids have been variously described as carnivores. but those within archaeocytes are often degenerate (*). perhaps by removing them as they age or by adjusting their density seasonally so that peak populations occur during the coolest part of the year.deficient environments (Usher et al.Aquatic Invertebrates/WG9. bioses: Plants. however.and epilithic cyano. quent swells from the NE trade winds. 2002. it increases its skeletal thickness and adds new corallites to the periphery of the colony. even regionally within a taxon. egories. Certain cyanobacteria produce chlorophyll a as well as chlorophyll b. although some produce thin crusts and can even overgrow neighboring corals (Pueschel and Saunders 2009). each of which is assembled within Golgi. These are shed continuously during the brief lifetimes of the thallus. when particulate or. These constitute the outermost layer of the new segment. The ability of one organism to “steal” defensive chemicals produced by another is referred to as kleptochemistry. Clausade M. Dissolution of corals by euendo. 1980.30167. even though the west coast of tropical Africa is at latitudes parallel to the Caribbean. The sabellid radioles are functionally identical and morphologically similar to serpulids.Sand terrace 20 m . these reefs are composed of individuals that exhibit slow growth rates and may have begun forming about 9. primarily at the expense of bicarbonate. author.. ———. ing from simple filaments to the development of more complex strata. Baird AH. Photo courtesy of Marie Kospartov and the Australian Research Coun. In this sense. Many symbiont. pro. 1961.dominated reef associated with a shipwreck on Palmyra Atoll. International Journal of Developmental Biology 50: 589–599. Viewed broadly. Note grains above are generally ovoid and separate.water coral reefs: out of sight—no longer out of mind. Macrobioeroders with activity on the surface: • grazing molluscs and fishes • excavating parrotfishes and certain sea urchins b. Hosts may include single. tion of gorgonin. specialized tentacles in some anemones may form flattened and branched pseudotentacles below those involved in feeding. 227 Asteroidea. This region of the spectrum is referred to as photosynthetically active radiation or PAR. Skeletal growth can be measured by changes in ge. Figure 7–8: Details of reefs constructed by Phragmatopoma caudata in Stuart.shaped. Rützler K and Macintyre IG. Figure 7–13). CCA play important physical and ecological roles on modern coral reefs. perhaps sug. Engel and Pawlik 2000. water corals (described below) form extensive reefs and are therefore hermatypic. lus if its growth is still active.226. forams more often form similar relationships with diatoms. Figure 6–4: a) This Millepora colony is encrusting and may have grown over another coral. Puglisi MP. Scott GAJ and Rotondo GM. PLoS ONE 3: e2429. tions. 168°W) is square. 2006). Richey JE. ent surface). et al. plications for biomineralization and paleo. ture of the substratum. especially on offshore islands (Spalding et al. the spreading of the ridge upon which they lie. Car. and Takemura A. red seaweeds hold the record for the deepest known algae at 268 m (880 ft) where irradiance levels are about 1/1000 of surface light (Littler et al. bonate and CO 2 as these carbon sources are transported 4–1 Some Relationships between Calcification and Photosynthesis Photosynthesis and calcification are complexly related processes. as well as high levels of carbonate and light. 1996. Spalding et al. chyme. Croix reefs. vironments. Earth Science Reviews 58: 163–231.brown. petitors before spreading over them with a sheet. Although described in more detail later (Chapter 8). ronments. Many occur with epizoic invertebrates that use them for support (Figure 6–11) and fishes that use them for shelter. ample. Leichter JJ. tion combined with aggressive expansion over the coral surface can quickly overwhelm their hosts (see Figure 5–9b). The lat. Hydrobiologia 530/531: 513–520. duced year round. forming a branched pseudofilament (unattached cells. high light intensity. and Furevic DM. Some reefs may have ini.” The typical test may look like a cluster of spheres or a miniature snail shell. However. Milanese M. thus allowing their hosts to survive in otherwise N. Calvert crystal with a more open structure and weaker bonds. ettii (Codiales. tenance of these communities by consuming macroal. tral GBR lies up to 260 km away.g. are more likely to occur in bur. mental information and has been used to reconstruct ancient climates including seawater temperatures. Continental Shelf Research 11: 771–790. 1997. Coral species diversity and environmen. surrounded by filamentous rhodophytes. a dense pavement of heavily calcified crustose Figure 4–16: a) Illustration of crustose coralline algae show. some within this family broadcast spawn. and Hillis L. Sclerites are typically sculpted with regularly arranged girdles of tubercles and warts that distinguish them from long. four of which are summarized here and further described below: • Maintenance of open space. Biomimetic ceramics and composites. Figure 5–20: Hypothetical and simplified diagram of relation. ———. Hydrobiologia 471: 125–131. Perhaps because of this. ———.” Fore reef Reef fat Sea level Increasing wave exposure Reef limestone Shore reef Fore Reeff fat Sea l e Reef crest Reef crest Coralline algal pavement Plate-like or encrusting Heavily branched Loosely branched Short. The vesicles actively concentrate specific ions and may assume distinct shapes that govern the morphol. indicat. hibit rapid vegetative growth by the addition of new segments attached vertically to older ones. cified or completely uncalcified. Octocorallia. Reiswig HM. and Ohno T. fect of seafloor habitat complexity on survival of juve. cellular pH. The crystals here are thin. Schlesinger et al. port for coral calcification and photosynthesis. more conspicuous species are often associated with living tissues of corals and demosponges. Its blade. long before the evo. Chazottes V. 1988. Simpson AGB. These sabellariids form colonies with less relief than S. Although some hexactinellids produce individual spicules.like structure that begins with intracellular vesicles of calcium and bicarbonate that are transported to the cell surface where they combine and mineralize on an extracellular organic matrix. Courtesy of Louis De Vos. sessment of the Andros Island reef system. with a total area of 18. Acroporid corals are prominent among these. ern Brazil. ments where they are densely packed and cover large areas of the bottom. Using radioactive calcium and carbon. Its branches begin with M. no. Biological Bul. In addition. 2006. which in turn damages their ability to grow.dimensional micro. Erwin PM and Thacker RW. and sloping. the archaeocyaths were relatively short. 1999. the asexually produced forms are the only type known (Grell 1979. gen mixtures. or become in. creased growth rate and frequency of asexual division (Holbrook and Schmitt 2005). Goldberg 2002a).3). Weiner et al. This area often develops its own distinctive coral community called a back reef. depending on the host (Erwin and Thacker 2008 a. 2008.1981. taxonomic. and in the Persian Gulf. Cairns SD. the Hexacorallia and the Octocorallia.like structures in fjords. Erlich H. some reefs. Some atolls are completely enclosed. and increases their vulner. the end segments of the abdomen curl into a J. Reproduction in the phylum Porifera: a synoptic overview. In.org/urmo. but they are most apparent when a colony is thinly sliced and photo. Courtesy of Andy Bruckner.swimming sperm (spermcasting). the entire plant is referred to as a thallus.sibling or parent. Dai 1993). Some are aggressive and will attack their hosts. and Telesnicki GJ. 2002). In: Lang JC. The contribution of these symbionts to host nutrition appears to be similar to anthozoan reef corals. sistency and is composed of small magnesium calcite crystals that reduce the porosity of reef limestone over time. 1996. Molecu. Deep water macroalgal communities adjacent to the Florida Keys reef tract. the fertilized eggs hatch into small larvae that drift with the plankton until settling on a coral reef as juve. pigmented cell layers called the cortex and one or more inner layers. Conversely. where seamounts rather than atolls occur. and the ori. six of the ten most common species on Australia’s Great Barrier Reef are essentially phototrophs and produce up to three times more oxygen through symbiont photosynthesis than they consume in respiration (Wilkinson 1983). Figure 4–3: A cross. Chemical defenses are also effective against many scavengers. meij and Sandin 2008). are locally outcrossed. and it contributes substantially to primary production on a global scale (Capone et al.. Shimizu K.e. ———. Canadian Society of Petroleum Geologists Mem.lived radioactive elements to cal. ISS004.100 km2 . several groups of organisms described in subsequent chapters employ a combination of physical and chemical processes. ment for four months or more. and Cubozoa (Marques and Collins 2004). This anemone is associated with 11 other clownfish species over its geographic range. 2005. forming shallow galleries in coral colonies. Trace element. composed of small. One hermit crab species uses abandoned sabellariid tubes as a home and is restricted to this habitat (Gherardi and Cassidy 1994). merous climate swings. A Caribbean species. ismal. the symbiosis can become permanently disrupted resulting in the death of the host (e. Mechanical or chemical processes or both are associ. Kure Atoll at 28. 2004). The specimen was found lying damaged on the seafloor at a depth of 150 m in Adak Strait (Aleutian Islands. Clifton KE and Clifton LM. lated. 50•Chapter Three munities on agents and rates of bioerosion (Reunion Island. Reef Sponges•87 3. 1994. Considerable amounts of Halimeda carbonate have been found even deeper (30–60 m) on bank reefs in the Bahamas where circum. polychaetes in the family Sabellidae produce an uncalci. high. but a single structure) in the Indian Ocean Republic of the Maldives. In some species. A model to explain the differences between Pacific plate island. 318 The Biology of Coral Bleaching and Disease. Oxygen relations of nitrogen fixation in cya.dwelling (tubicolous) species. The polyps typically occur on one side of the skeleton and. initial calcification appears not to interfere with subsequent mineral deposition.g. and then release a zygote rather than a complete larval form (e. whereas small corals that play a lesser role in reef construction are called ahermatypic. including chemical cues and availability of suit. Uni. rina Fabricius. Wilson PA. but mul. Bak RPM. and other nitrogen. duced zero. The structural complexity of the ner. Polychaete jaws are not new inventions and are common among Paleozoic fossils known as scolecodonts (Greek for “worm tooth”). Vol. tropical scleractinian coral families cur.like gorgo.reef growth: a dynamic balance. family Dictyotaceae) are common on hard substrate typically as algal turfs (Fig. iour 70: 97–110. temperate locations. all of which have abrasive or biting mouth. encrusting sheets of Archaeolitho. a subject that is described in more detail in Chapter 15. Hase C. The Ca. vidual units (ommatidia) composing the sabellid eye and well over 200 eyes in species that have been exam. and their effects on the diversity of tropi. Roark et al. Rodriguez. and may include coconut palms together with broadleaf woodland species. Kramer PA. As water is moved. Photo courtesy of Bill Rudman. as well as photosynthetic cyanobacteria. The stem cell concept in sponges (Porifera): Metazoan traits.lived. but the most prominent on tropical reefs is the Bryopsidales. damicornis and other coral species (see Carlon 1999) may have complex mat.. a b Reef Corals and Their Allies•107 hicles. 2004. In addition to these forms of microphagy. Bora Bora and its barrier reef is at center. percalcified sponges called the chaetetids (Figure 5–15a). Furthermore. calcareous algae. 2009. further details in Chapter 4). Several species are often found on sand or sand and mud and are referred to as psammophilic (sand. Some authors have identified par.g. and up to 12 mm long. dance (HMA) sponges (Hentschel et al 2003). Daly RA. 90•Chapter Five did not form reefs in the sense of physically complex. Some investigators have emphasized the importance of producing an alkaline environment by metabolic pro. compared with the deeper endosome. Such “barrier. Marine Biology 142: 643–648. Rees et al. ure 6–28) that grow to 50 cm in diameter and have the unusual habit of forming numerous asexual buds primar.forming stage. tively large sizes (>3 cm) and are included among an informal group called large benthic foraminifera (LBF). The most ex. locene reefs of St. Nicotine is an example of a simple alkaloid that can be seen throughout the more com. Photo courtesy of Walter Adey. seal. The ecology of Millepora: a review. Marine Biology 147: 761–774. While much is known about the general benefits of coral symbiosis. the epithallus. This rather high cost could make evolutionary sense if some symbiont groups performed better than others did under stress. These microphagous animals may use jaws or teeth to crack the diatom shells. Foster R. prepared with liquid nitrogen freeze.. shallow water in the Caribbean. Chemical defenses: from compounds to communities. et al. these reefs were found from North America to southern and Eastern Europe. may be up to 2 cm long and constitute the major supporting architecture. Y. Rhodolith bed diversity in the Gulf of California: the importance of rhodolith structure and consequences of disturbance. Estimates of brown algal origins are more variable. 1991.” Kiritimati in the Line Islands (central Pa.Deep reef slope 100+ m . Tambutté S. While the mechanics of this dark calcification are not entirely clear. Bahamas. morph is formed by many reef organisms. Where have all the carbonates gone? A model comparison of calcium carbonate budgets before and after the 1982–1983 El Niño at Uva Island in the eastern Pacific. mens of C.g. International Journal of Plant Sciences 162: 263–270. http:// www. Photo courtesy of Bernhard Riegl. Marine Ecology Progress Series 356: 123–138. Ireland.Pacific reefs that had become dissected by erosion (e. including flicking motions. Indeed. Hyella An endolithic cyanobacterium forming pseudoflaments. ed. et al. mentology 47: 915–921. This sedimentary species that define a community’s physical structure (e.transport of bicarbonate for photosynthesis (Adey 1998. Guille G. Crustose coralline algae (CCA) often rely on habitat exclusion of competitors as may occur in the wave. Benavides LM and Druffel EM. Levitan DR.energy environments often favor branching corals. Naked corals: skeleton loss in Scleractinia. which they superficially resemble as described below. both as a producer of fixed carbon and the cause of the erosion of carbonate. in some species. All boring sponges have specialized etching cells (Pomponi 1980) that extend pseudopodia as they bore into carbonate. Some syllids are carnivorous sponge specialists and may be found by the thousands in the canals of their hosts where they appear to consume its tissue (Pawlik 1983). come altered. est wave. degree of separation.building species (Zann and Bolton 1985). However. a topic that will be explored further in Chapters 3 and 13. The calicodermis is rich in mitochondria and generates high volumes of CO 2. stromatoporoids (b). erpa taxifolia (Chlorophyta). and Vollmer SV. bingers of global change? Micropaleontology 46: 95–104. represented by both fleshy and calcareous families and genera. by increasing the deposi. Randall JE and Fautin DG. fusion can produce formations that are far more extensive. However. and trap organic matter that is brought to the mouth Photo courtesy of Paul Humann. The role of endolithic cyanobacteria in the formation of lithified laminae in Bahamian stromatolites. Coral Reefs 14: 79–86. Fiore and Jutte 2010). tion by the encrusting excavating Caribbean sponge Cliona delitrix. vating sponge Cliona tenuis and corals influenced by higher. secretion of organic material for new wall production. Zooxanthellae Clade Distribution.usgs.. ation and crystal growth by binding to calcium or other ions (reviews by Simkiss and Wilbur 1989. However. Folke C. Indeed. and to implicate the presence of zooxanthellae early in the evolution of the Scleractinia >200 million years ago (e. anemone. The skel. Caribbean coral reefs. USA 106: 15374–15378. Sedimen. Indeed. Indeed. rine benthic algae. 1996. The best. Rogers et al. Photo courtesy of Paul Hu. For example. On El Capi. Photo courtesy of Allison Daley. making it resistant to hydrodynamic forces of extension and compression (e. they are called isogametes. Mechanisms of high colonial growth and rates of reproduction facilitate this domi. Schönberg CHL and Wilkinson CR. b). Kump LR. At this stage of their life history. dolfi and Jackson 2001).boring palolo worm Eunice viridis. Thereafter.” Organ. thropogenic enterprises. certain herbivorous gastropod molluscs (e. Mitochon. These had a similar massive. non. To put this into a paleoecological perspective. The global biodiversity of coral reefs: a comparison with rain forests. Van der Land J. vii 1 An Introduction to the Structure and Formation of Modern Reefs. Facies 44: 265–282. Vree. 1983. Based on Hyman 1940. and low. tridae): does it dictate spawning site preferences? En. from the Gulf of Alaska. 150•Chapter Seven Sabellariids are morphologically complex. 2008. Lindquist N and Hay ME. Botanica Marina 45: 346–355. cal time. biofilms control other microbial processes including cell. and value as habi.g. in brackish lagoons. onts and is employed for heterotrophic digestion. 1988. both living and dead. Sherwood OA. Marine Ecology Progress Series 372: 77–86. Oahu. the Great Barrier Reef).e.Pacific (Tanner 1997). formed reef knolls up to 2..H. Some anemones deploy specialized inflated “fighting tentacles” that develop from those involved in feeding. dinoflagel. habitats. 2003). Conversely. Fig. Chaves. Van Soest R. and is considerably larger than any living Ca. 2003. and may travel distances of only a few hundred meters from the source (e. this project took on a life of its own and consumed more years than I care to admit. a large atoll in the northern Caribbean.Johnson Space Center. Thickest recorded Holocene reef section. ten by Darwin (1842) after his voyage on HMS Beagle from 1831 to 1836. These structures did not require a pre. In some regions of the Persian (Arabian) Gulf. rial. Interestingly. for example. feeding types. ing ~3. constitutes a repetitive impact for reefs along these coastlines and is thought to be a major factor in retarding their growth (Jokiel 2008). Halimeda appears to lack a sporophyte stage and produces gametes that are released massively and synchronously. This photo shows living colonies of both species in Bonaire. 2008). As further de. basal skeletal structure. 2003. cance of endoliths in dead. Beer S. binations of these. As they monopolize space.secreting organisms: The voids within a reef framework are typically filled by sediment. Shallower regions may develop extensive patch or fringing reefs if outer reef waters spill over the reef rim during high tides. However. Banaigs B. Panama 1: 743–748. likely by piercing the target organism with the bullet.de. populating reef environments after severe storms or after pattern in which there are no distinctly separate polyps (Figure 6–31a. Hayward DC. the open edges of the column quickly heal and regenerate missing parts. and carbonate deposition. This may be due in part to their ability to move nutrients to their shaded sides (Underwood 2006). ed. Journal of Natural Products 70: 504–509. Coral Reefs 21: 57–72. but only cyanobacteria. large popu. itats. 2004).900 individual coral reefs. shallow water. both ecologically and structurally. attract the epitokes with lanterns and scoop up large numbers of them with hand nets (Figure 7–11b). Calcification and photosynthesis in reef. Proceedings of the 8th Coral Reef Symposium. These are cases of facultative symbioses. Lawrence: University of Kansas Press: 166–190.divegallery. ogy of scleractinian corals. the Ro. gesting a new and unexplored carbon source for these organisms (Patten et al. Reef Evolution.. an elaborate and highly organized internal membrane system that is folded into onion. Science 209: 689–691. 1981. The age and origin of the Pacific islands: a geological overview. circulation is so poor that reef development is inhibited. ria) with no light requirements. In either case. Reviews in Mineralogy and Geochemistry 54: 1–29. If rim growth were able to keep pace with continued sinking. which may drop gradually or precipitously into deeper water. Sponges. Walters et al. ronment. once a host is colonized. cular tissues. they usually defend it from being oc. noa resedaeformis and Paragorgia arborea. New York: Elsevier: 209–252. gorgonians (Walker and Bull 1983. so.like ac. disease. 2006). Fan TY. come arched over. 1999. 130•Chapter Six sharing as well as other communal behaviors (Hughes 1989. 1983). the catalyst for this process may involve additional ac. Nanosecond. but is a common trait among many groups of anemones as it is in other anthozoans. Sometimes.g. Papers of the Tortugas Laboratory. Marine Biology 146: 301–313. may be dense and sink to the bottom. roplast evolution: what did cyanobacteria do for plants? Genome Biology 4: 209. Huvadhoo. Rates of active carbonate accretion vary considerably by geography. The beat of elongated lateral cilia creates a feed. although some groups have more than two or none at all.000 years ago. In general. perhaps reflecting chemical attraction and the selection of corals that do not employ aggressive behaviors toward them.124 m (Macintyre et al. Sexual reproduction in octocorals. and precipitate calcium carbonate in place. cium from one end of the filament to the other (Garcia. although in many places the benthos is com. 326 Direct and Indirect Effects of Human Activities on Reefs. and key differences between algal calcification and that of scleractinians. ous other organisms as parasites.18 in Chapter 4). organic sheaths (Figure 4–13). 108•Chapter Six Anthozoans are divided into two subclasses. vations of some common antipatharian corals in the marine park of Bunaken (North Sulawesi. troversial.level classification of eukaryotes with emphasis on the taxonomy of protists. Australia). winter. grooves (channels) may exhibit similar variability. Fautin DG and Allen GR. Figure 3–8e). ships between reef growth and sea level in the Great 30•Chapter One Gradstein FM and Ogg IG. alveolata. American Zoologist 39: 160–183. Russia. bicarbonate. Report on the Scientific Results of the Voyage of the HMS Challenger during the Years 1873–1776. it was re. d). representing both uncalcified genera as well as the calcified groups considered here. 2000. letin 141: 568–591. indicating that these organisms evolved 40 My be. 2009. totrophy and predation in the hermatypic coral Sty.like structures called bioherms. but even these are quite narrow.Morales BO. 2005.weight compounds including certain amino and fatty acids. but most annelid species (>8. Richmond RH and Hunter CL. each of which communicates with the external world by a nar. and four identical rectangular faces that make up the sides (Figure 2–5a). These house the oral tentacles as well as a horseshoe. 52 levels of anatomical complexity among seaweeds rang. Others may be able to supplement their diet with nutrient. Savalia savaglia. sular osmotic pressure and a powerful molecular spring mechanism (Nüchter et al. 2004. with a 100 µm–long megasclere forming within 40 hours (Uriz et al. Science 309: 1027–1028. not simply smaller versions of the megascleres. Multer HG and Clavijo I. whereas more complex molecules obtained by prey capture appear to be incorporated into the struc. certain sea urchins) readily consume such calcified materials (Solandt and Camp. Mariners. Plusses and Minuses The amount of carbonate produced on reefs is typi. These structures contain a hollow tubule that may be distinguished further by an enlarged base (shaft) that is often armed by numerous spine.like front composed of platy corals. especially below 10 m. Proceedings of the Royal Society B 278: 1323–1328. Marine Ecology Pro. Microbialites are typically very difficult to recognize in the fossil re. and creates a very unsuitable environment for settlement of coral larvae. the majority of corals are broadcast. A few unusual demosponges. one of the principal species of reef. ease called botulism. it is possible to con. and may serve as break. 2004. From the send. Most of the shallow. locomotion. Aragonite is precipi. are very common. 1988. 2007.. Scott DB. and have more of an undulating surface with a signifi.g. ment on terminology.like. 1988.. and other chlorophylls depending on the group. and Dai CF. and STS093. Some species actively transport bicarbonate across the cell membrane and convert it to CO 2 internally. in which about half of the approximately 20 species produce aragonite Brown Algae The division Phaeophyta is almost entirely marine and is composed of 900–1. but also are employed as offensive weapons against other sessile organisms including other sponges that may compete for space (e. 1971. Developmental insights into the origin of complex colonial hydrozoans. A sheathed. chemically controlled calcification process (reviewed by Cohen and McConnaughey 2003). 2006). stylasterines. suggest that the two groups calcify in a similar manner (Lee and Carpenter 2001).5 million metric tons of sediment across the Brazilian continental shelf each day (Meade et al. In hemispherical or plate. Introduction to Mineralogy. some of which mediate their ability to reduce predation pres. Grazing animals on reefs remove epilithic algae or organic matter. Koehl. Annelids and Sipunculans•149 capped. Reef coral survival and mortality at low temperatures in the Arabian Gulf: new species. Indian Ocean).dependent coral and coralline algae to rising sea levels. and light levels over time. Washington. havior. alcyonaceans.. Dominguez LL. or may become part of it. goon? Perhaps. most of the >3800 species of Hydrozoa (Van der Land 2006) form polyps. Olympic Coast National Marine Sanctuary. Friedmann EI. ductions from the gross amount in order to derive net carbonate production. Those shown here have a canopy of iridescent Dictyota sp. is composed of massive coral lobes that project vertically as much as 25 m from their bases.7.like or encrusting colonies in which the basal layer forms a distinct carpet of unfused sclerites. 1c. 2002. Arvedlund M. these are differentiated into one or more outer. Journal of Experi. consistent with that of many unbranched shallow. Reef Sponges•83 of silicic acid on a protein template (Cha et al. De Beer and Larkum 2001). the biology and chemistry of mollusc shell formation is complex and incompletely understood. the matrix sur. litrix and other clionaids are increasing on reefs. 46•Chapter Three of the former. Rhodophyta). eschel and Wegeberg 2005). nucleic acids. tats. deserts. tion. The coelenteron is a simple tube that lacks the tissue specializations found in anthozoans (described below). ———. gella that pull water through the porocytes (the inhal.and microgametes from Halimeda fuse producing a zygote that enlarges and elongates into a stage called a protosphere. ure 6–5). The branches are bro.harvesting antennae.000 meters thick was found as a cap on vol. In: Pitcher TJ. James NP and Macintyre IG. NOAA Photo Library. In other groups. are often 50 m deep. and these are sometimes distinguished as active or pas. Reef Zonation Reefs are typically divided into distinct zones as a function of depth. and certain herbivorous fishes (Capper et al. including a thin surface veneer. Contrast between solitary and clonal life. Subsequently. late xanthophylls and carotenes may impart a golden brown color to the host’s tissues (Figure 6–16). the Mal.like structures. reef.Colinvaux 1980). Hermatypic and Precious Octocorals Three groups of unrelated octocorals are considered together here because their skeletons are formed from fused sclerites. albeit at variable concentrations.. Whatever the mechanism. but banding prominence may differ among differ. and Santos AM. 1983.Kudla 1997). Aquatic Botany 92: 157–160. Science 152: 343–344. The influence of wave exposure on the ecological zonation of Caribbean coral reefs.energy. ontology 70: 347–355. with time the shells became progressively larger. Marshall CP. rica (Grotzinger 1989. After Glynn PW. the abundance of fish. >40% of all de.and deposit. Thus. portant coral reef species. ure 7–6b).Fonnegra and Zea 2010). Kahng SE. and Kitazato H. Figure 6–3d). It is thought that the source of this material is likely to be bacterial (Ramos and Vasconcelos 2010). transloca.* This high velocity discharge process involves a complex cascade of events that stem from high intracap. and a large concentration of xanthophylls in the chloroplasts that account for their characteristic color (Table 4–1). ally critical because most reef. The timing of reproduction var. poorly circulated lagoons are particularly susceptible to dramatic physical changes. Fautin DG. ate (Figure 7–11a). The data from these remarkably well. caves. Fission is typical of single. rounded struc. Cohen A. saline marine environment of Shark Bay. Among poorly calcified spe. Pro. 2006. While more information is be.Morales et al. 2007. In some cases. Some polychaetes feed on detritus and microorganisms that are suspended in the water by allowing them to settle passively onto ciliated feeding appendages associated with the mouth. and form starch as a grain. rather than reefs. bers of the benthic community whose skeletons can pro. ated for size. stalked. Dill RF. Brown algae do not figure prominently in the produc. Müller WEG. Reef Corals and Their Allies•135 corals including P.g. Granato AC. Its segmented branches. in turn. Coccoid cyanobacteria are more common than filamentous types. In Halim.000 years and nearly reached the surface in less than 2. well. nies whose polyps are embedded in a coenenchyme riddled with gastrovascular canals. In: Freiwald A and Rob. ment appears to have been weak. Thus. 2000).g. et al. tors. 2007. which becomes extremely common on intertidal and subtidal reef zones. eton reflects this organization because each tube is con. rial incorporated into the skeleton can indicate the pe. Such reefs in.g.mx/institucion/cooperacion_internacional/Taller NDF/Links. Sponges that brood embryos to this extent are sometimes referred to as “viviparous. They are also aggressive.Verlag: 31–50. tain blue. Reefs of the deep: the biology and geology of cold. LeGeros RZ. In more favorable cir. Marine Ecology Progress Series 363: 139–147. 2007). daria. protective outer cap. and Nielsen LE. circular. cheilus longicauda. From seawater. a basin. Journal of Phycology 41: 1010–1024. blue. These in turn bear 1–3 rows of specialized. Vigorous circulation on and around the shallow reef crest gives rise to elevated dissolved oxygen. ing coral tissue. Wood 1999).and gravel. Most taxa produce more modest sand. Several explanations have been proposed to explain this capability. Seminars in Cell and De. the utricle tips fuse or become closely appressed over the surface of the segment (Figures 4–7b. 2006 and Fukami et al. Marine Ecology Progress Series 397: 319–332. Marine Ecology Progress Series 12: 137–143. Decline in surface temperature and salinity in the western tropical Pacific Ocean in the Holocene epoch. similar to the adaptation described for the filamentous O. mak. Honolulu: University of Hawaii Press. Because Monaxons Polyaxons Microscleres Tetraxons Figure 5–5: Demosponge spicules may be present in a variety of forms.) are absent in this group.Neto JP. or thin sheets that lie on or within the substratum and are referred to as microalgae. is a zone in which further growth is often limited by its proximity to sea level. Lang 1974). and then translocated once again for use by their hosts. Feeding polyps are often bottle or vase. However. well as certain green algae (Chapter 4) and fungi that constitute a “mature” endolith in the roots of leguminous plants like alfalfa. The hydrocoral genus Millepora in Indonesia. However. Bald et al. Harrison PL and Wallace CC. In northeastern Brazil and southwestern India. Baird AH. liths in the massive corals Cyphastrea serailia. Tsunami im. 2008.celled coccoid and filamentous types. The proportion of individual spicules and spongin often dictates the relative degree of flexibility and stiffness of the sponge body. and resist desiccation during low tide (Shick 1991). 2007.. Sydney. the family Coralliidae. Miake Y. ern Caribbean fringing reef. left from Wikipedia. Various endo. Within a few hours or up to 2 days after spawning. Indeed. Thus.. The Biology of Cyanobacte. Schierwater B. Littler MM and Littler DS. solu. The spaces between the cortical filaments calcify first. and produces a basket. 1985.. Vol. and thick. Protected lagoons may support mangrove communities or seagrass and algae ing the surface at their crests during low tide and rising from deeper water farther offshore than fringing reefs. and solid solution behavior.. Atmospheric dinitrogen fixation by benthic com. However. giganteus complex. 318 Climate Change. courtesy of the journal. Tai Y. mately 10 m sec–1 ) within a period as short as 700 nano. they were all but extinct. On the basis of relative deterrence. large antipathar. sometimes in rather spectacular fashion. an invasive species presumed to be native to Australia that has also success. Coral Reefs 4: 19–26. Inset: Details of branching structure. 1998..4 mm yr–1 . and Fabrizio MC.500 valid species. although distinct from algae. gin of the oral disc or may be numerous and arranged in radiating rows around the mouth. but the volume is now >200 cm3 . similar to those favored by crustose coral.000 years ago the reef was about 20 m below sea level. Panama 1: 701–706. and Paul VJ. 255 Fishes That Perch or Live on Corals. but most of their biomass is confined to irregular channels and galleries within the substratum. The eggs pro. some filamentous forms produce a thick. regardless of the source. In contrast. Although much remains to be learned concerning the specifics of mineral formation (and demineralization). Lar. Halimeda species are among the most abundant tropical algae. Pomponi S. organic compounds. As Trichodesmium blooms increase in size and frequency.like openings often built up by sclerites that are visible on the colony surface (Figure 6–20a). Marine Ecology Progress Series 186:119–126. a province of Indonesia. Some algae maintain acidic regions outside the plasma membrane that shift the equilibrium shown in Equation 4–1 to the left (Mercado et al. Shashar N. Kump 2008). Durand I. mates of 1 million–9 million species (Paulay 1997. The primary tube secreted by the juvenile is composed of Figure 7–4: a) The serpulid Spirobranchus giganteus expanded on a coral host showing the radioles and spiny operculum. Tubicolous and Burrowing Polychaetes Freely mobile polychaetes include a number of or.3 cm yr–1 . in this case exuded as toxic mucus from the boring sponge Aka coral. reef develop. The aboral region may form an adhesive pedal disc. mounds. which produces ge. 2010). scrape. In contrast. Bayer FM. ing them with >70% of their carbon requirements. Likewise. the idea that glaciation and erosion is responsible for many of the historical features of reef formation. Calcaxonian gorgonians. The flexible axial skeleton with its growth layers and central chord indicates that this is a holaxonian gorgonian. 1988. and other large organisms. some species (e. Others have suggested that such changes indicate that physiological processes. but are far more common on islands. Feeding behavior. and Braekman J. building corals and grows with their hosts.like structures called acrorhagi that develop on the upper portion of the col. Turf algae are typically dominant on undisturbed reef surfaces. Berlin: Springer. fluoride may be Figure 2–6: More complex forms of the hexagonal system: a) a dihexagonal. ers. and Thiel V. Competition among marine mac. 120•Chapter Six 6–1 A Hypothesis Linking Coral Calcification and Photosynthesis: A Tale of Two Enzymes Calcification in zooxanthellate corals is not as much dependent upon light as it is enhanced by it. trate a pool of silicic acid. fied organisms. fill can be influenced by the presence of erect but flexible organisms such as algae and octocorals (Chapter 6) that reduce moderate current velocities along reef surfaces. drilla nucula (Demospongiae. the US Virgin Islands. typically forming cycles of six. I: system level studies. although in some cases aragonite or amor. abun. biology of endolithic microorganisms in living coral skeletons. 1983. Some species produce identi. or other well. Wood R. Schuhmacher H and Zibrowius H.binding and cement. the reef crest. Oxford: Oxford Univer. algae lack true roots. Figure 1–20: Major reef terraces of the Huon Pen. The large Caribbean barrel sponge Xestospongia muta is an example of a gonochoric.. Birkedal et al. and Bavestrello G. carbonate sediments may be imported or exported from reefs. The apatite crystal is composed of two six. and the develop. Some species have been described as car. Foraminifera: a biological overview.Jones 1988). Uncalcified Reef Anthozoans Sea Anemones With about 1200 species in 46 families (Daly et al. 2003. Shashar N. Francis L. However. Inhabitants of certain sponges at Dry Tortugas. bacteria are also responsible for the production of unique compounds found in sponges. 2007).secreting organisms and thereby functions as a surface cement or binding agent. Annual Review of Ecology and Systematics 19: 19–111. when fossil cells and the first multi. they generally are not prominent con.like mounds found in Cambrian and Precambrian times.water fishing practices and are further threatened by climate change. Rollion. 2008. i. In some cases. among others. Figure 4–7a) and during the warmer months. Many of them do not attain large sizes and are not recognized as reef builders (i.West Pa. 2002. the GBR has a boomerang shape. The family Sabellariidae (“sahbel are eeidee”) is constituted by 107 species called sand. markable ability to bore into and erode carbonate sub. tangle themselves around Halimeda or other Dictyota species (Lirman and Biber 2000. wise. 1998. Broadcast spawners are typically her. Crystals are oriented and aligned within the cell walls rather than scattered randomly within intracellular spaces as in Halimeda. and Erez J. they are known to interfere with the early life stages of some species (Ver. larviparous spermcaster (Figure 5–3). Coral reef fishes constitute almost a third of all known fish species. phorous may be limited. The hydrozoan coral Millepora dichotoma: speciation or phenotypic plasticity? Marine Biology 143: 1175–1183. or they may browse with one or more paired jaws that protrude from a muscular pharynx. Indeed. cyanobacteria form a natural group of organisms and are a monophyletic taxon.shaped Tarawa (Kiribati. which are stalked. are described further in Chapter 15. and are estimated to be between 6 thousand and 10 thou. tal spines.” More generally. cellular cyanobionts in open ocean dinoflagellates. construct bioherms. Figure 1–3).producing polyps do not divide and may be resis. one Fishes and Their Sea Anemones. Either of these will decrease calcification levels during the day (Häder et al. Palaeoecology 166: 89–100. some parrotfishes prefer older. Fiesler L. Bruckner AW.1992 (Permanence of Paper). and can occur deep within the substratum or at its surface as they rasp. In: Freiwald A and Roberts JM. ments.. land. and inozoans (redrawn from Wood 1990. However.g.related differences in the excavating micropatterns of boring sponges. Indeed. Taylor MW. for their lives or livelihood.like scleractinian corals is a good place to find them. it has been suggested that the open needle architecture may limit the extensive microboring typical of more dense carbonates. Nava H and Carballo JL. and so it is with burrowing and tube. for example. Brownlee C and Taylor A. nes is pedal laceration in which the basal region constricts into small pieces that separate and then develop into new polyps. gesting that the matrix may control the process.e. and reproduce. vices. mation and produce amorphous deposits. ment with bleach used to prepare this material. 2005). Jindrich V. parrotfishes referred to above) may learn to prefer the older. calcification may continue independently of photosynthesis as long as oxygen and energy levels remain high (Al. University of Otago. 2006. ments or tabulae (Figures 6–12. ward) sides. and several days after the full moon! Similar synchronized gamete releases occur in the western Atlantic. and that calcification itself may enhance the process of phosphate acquisition (e. Case study for black coral in Hawaii. stout chaetae that can fill the opening of the tube when the worm contracts. are easily accessible to such organisms. South Pacific: geology and petrology of the emerged part of the Society hot spot. ner et al. ing 2002). Sammarco PW.Victoria M and Zea S. Particulate organic matter and plankton as small as 0. or even within the thoracic segments. yellow or tan tube constructed from sand grains embedded in a matrix of mucus and protein. etons. While bota. Reefs and Related Carbonates: Ecology and Sedimentology.bathymetric implications. c2. Two new species of the gorgonacean genus Ctenocella (Coelenterata: An. chaete worms (Pawlik 1983). However. hemispheres. perhaps enabling such corals to change algal partnerships. even though many boring sipunculans do not have these structures.g. ogy and Ecology 322: 67–81. An Introduction to the Study of Re. low. sea urchin spines.and microgametes. and once again produce the haploid gameto. 1972. bacteria may be able to use virtually the entire PAR spectrum and can thereby adapt to a wide variety of ecological conditions. plants are dioe. vores and are potent feeding deterrents. equatorial island of New Guinea provides a striking example of successive bar. biggs WD. NOAA. It is easy enough for a biological scientist to write a paper or a chapter.water species commonly called “glass sponges. Science 228: 488–490. Succession and growth rates of encrust. the term “rockground” is sometimes used (e. Ferrier. Henckel TP. although a few have only a single opening. 2007). to as the S. gen fixation in refractory organic sediments by Caul. 2000.swimming polychaete larva is called a trocho. its composed primarily of their shells. making them less resistant to wave surge (Lewis 1998). trunkfishes. thors have referred to these as guilds. most of which are assembled into chains and cy. While solitary cor. and may contain valuable clues to paleotemperatures and other environmental information (e. atic account of reef types and their evolution was writ. cally.Mansour E. 42 Stromatolite Formation. trophic (e. Sinularia spp. Hartman WD and Goreau TF. New York: Oxford Uni. rather than a change in the motion of the Pacific Plate (Tar. the field of biomimetics. 33 Calcium Carbonates. Charles Darwin (1842) noted that fringing reefs. Sheppard CRC. bonate production: system level studies. Marshall JF. Capper A. The shallow portion may extend to a depth of 10–15 m and is typically the most diverse.axis. Such structures have been forming since the late Miocene. 2007. especially if they are photo. Dictyota) where the gametophyte and sporophyte stages are iden. Blair SM and Norris JN. and some are essentially or completely closed (under. including deposit and suspension feeding. produces prominent tubes that exude toxic mucus.barrier system. onies of Montastraea cavernosa and possibly other hosts that may contain high densities of intracellular symbiotic cyanobacteria (Lesser et al. strata determine postsettlement survival in corals. Depth distribution of calcareous encrusting associations in the northern Red Sea (Safaga. 2005. Puglisi MP. b) Pink sand beach at Warwick Long Bay. 2002. polychaete worms. Porphyra microbor. Two other forms of calcium carbonate occur within the hexagonal system and produce complex crystals. Lang JC. and sexual reproduction. which may be crucial to the main. Launay J. form rather than capitate structures. erect filaments (not shown). Lee JJ and Anderson OR. which are not unlike the intercor. The western side (Les Récifs des Française) displays a prominent double barrier reef.spawning corals of the Montastraea annu. 1989. although specific details remain to be clarified. 2004. Thus as the reef becomes a more complex physical habitat. Photosynthesis Research 88: 259–268. Organisms may induce nucleation and mineral forma. Alkaloids (nitrogen. Discovery of a “living dinosaur”: globally unique modern hexactinellid sponge reefs off British Columbia. 1990. although gold and black corals described above may also be precious. cal marine sponges. ogy. A second group of octocorals that produces an unusual solid skeleton is the blue coral Heliopora coerulea. http://www.builders by damaging or burying them with sediment (McCarthy et al. Carpenter KE. These are areas where ad. mations. Francini. Interactions among Florida sponges.. whereas in low. Gold coral (Savalia savaglia) and gorgonian forests enhance benthic biodiversity and ecosystem functioning in the mesophotic zone. This reinforce. Identi. ing these evolutionary relationships is clear. for example. layered cyanobacterial mats called stromato. men was photographed off central Florida at a depth of about 800 m. Nitrogen fixation products including amino acids may be transferred to the host along with photosyntheti. 1990). a 2.Rivera E.Emperor Chain—revisited. der different conditions. Reproduction in Halimeda and Other Chlorophytes Most green algae exhibit a biphasic life cycle that in. and cool winter temperatures due to its position ~26° north of the equator have been suggested as causes Figure 1–23: a) Reef histories can be characterized by their relationships to sea level changes. tempera. gin. provide a visual screen for both preda. Clownfishes also derive nutritional benefits from their hosts by consuming regurgitated anemone waste. shape. sea urchin spines. called the atoke. recycled into organic form by zooxanthellae. described in the sections below. The latter is the endolithic con. Zale and Merrifield 1989). alcicornis. but are restricted to only a few places. Effects of the benthic cyanobacterium Lyngbya majuscula on larval recruit. Mahoney MM. and brown algae is likely not reflected in the 20. Entire sections of reef can be sculpted. ties produce carbonate by trapping sediment within the sheath. Partial carbon and energy budgets of the bacteriosponge Verongula fistularis (Porifera. other factors that directly stress corals. sylvania State University. Steneck RS and Dethier MN.Water Corals and Ecosystems. they do not form frame reefs by themselves as defined in Chap. and temperatures can have strong effects on larval distribution. genetics and Evolution 47: 433–438. Slow turnover in the soft coral gen. Schembri et al 2007).Furman NE. Webb KL. nate reef substrata that are actively grazed.92537. people. and partly mechanical. 1983.unep. tratentacular budding (Figure 6–29a.g. However. Where do clonal coral lar. diversity. ings of the 8th International Coral Reef Symposium. and propulsion to gametes during reproduction. or separate more com. take during photosynthesis. fibrous connective material called spongin. both of which can create topographically significant structures An Introduction to the Structure and Formation of Modern Reefs 1 2•Chapter One of both massive and large. Likewise. Under favorable conditions of turbulent and turbid water. or cyanobacteria (Lee and Anderson 1991. 2005. Thus. line algae have been recognized since the Permian pe. Helman et al. Ferrier. fied plate. reinforced connective tissue: stiffness. A similar pattern of growth is exhibited on the Ho. geologists are able to distinguish biological stromatolite growth from inor. also known as the soft corals. The relative predominance of western Atlantic shelf atolls is a reflection of conditions and historical differences that characterize the region. Fringing reefs may occur immediately near shore. enabling them to outcompete other sessile invertebrates as well as mac. but especially in response to wave disturbance. US Virgin Islands): applications to the nature of reef systems in the fossil record. Endolithic cyanobacteria are extremely common in tropical marine limestone where populations of more than a half million individuals per square centimeter are common (Schneider and Le Campion. The mechanisms that produce such cements are unclear. clear water and open. Thus. some of which may form true. called the reef’s carbonate budget (see details below). and Montaggioni LF. Some sea anemones are equipped with numerous long. My. thus. or red skeletons (Fig. 2008). Arthur KE. sial. 1999. able substrata (Golbuu and Richmond 2007). colonies produced by fission may dominate over those that are sexually reproduced. and Sanders JB. called the tetrasporophyte. 107 Uncalcified Reef Anthozoans. Becerro MA. cies. although such defenses are typi. Marine cyanobacteria may exist as single cells or as colonies that assume a diverse array of forms.water Octocorallia are placed in two orders: the Gorgonacea. ians. it is sometimes difficult to distinguish biologically induced minerals from laboratory precipitates (Mann 2001). seconds. New York: Chapman and Hall: 68–95. The reef cave dwelling ultraconserva. 2002 a. photosynthesis in marine algae would appear to be CO 2 limited. butions. In: Cato JC and Brown CL. ogy. 2011 In addition.sea sponge groups that are carnivorous (e. Atmospheric nuclear testing is recorded by the ratio of various long. some modern sponges can become frame.laden tentacles suggesting the ability to feed heterotrophically.noelsirivansanti. ries of seamounts (see below) whose carbonate growth is insufficient to keep them at sea level. for her skilled. ments and fragments (Drew 1983). a globally significant marine cyanobacterium. and Norris JN. Impor. Tulsa: American Asso. is an example. incorporated into the reef and how much is stored in sand channels and elsewhere? • Sediment fate b: How much is dissolved or ex. mental Marine Biology and Ecology 336: 211–220. and Andros (Bahamas). one atop the other. but in some genera there can be multiple trichomes within a common sheath (Figure 3–1. and mucus cytochemistry of the scleractinian Myce. there may be geographic variability in toxin production. and provided 36 of the illustrations used in this text as well as the cover photo.building species may occasionally create reef. The sclerites and geographic distribution of the gorgonian Swiftia exserta (Coelenterata: Oc. 381:56–58.edu. Shick JM. 207 Coral. dence of nitrification and denitrification in high and low microbial abundance sponges. Dai C. Archaeocyaths were not known to produce spicules and. and even algae (Lang and Chornesky 1990. Gy. usually as closely spaced pairs. but also modify and store them for their own defense. tives. 2008. 2006. Ecology 78: 146–159. Many of the island bar. where they provide habitat for macroalgae and numerous in.fixing agents. brown (Phaeophyta).edge reef system (St. In rock borers. showing organization of internal chambers and elongated shape. and deep tropical regions at depths of 30 to >90 m. Berkeley: University of Califor. 2007). building from symbiosis with zooxanthellae. Harney JN and Fletcher CH III. and Kühl M. Dullo W. by the time it has settled and metamorphosed to a polyp. 2001). Hawaii. 1996. Sherwood et al. known products of marine and freshwater bacteria. sists year round (Steneck and Dethier 1994. followed by the abdomen and a tapered caudal region (Figure 7–7a). Biological Materials of Marine Origin. which vary in shape from narrow “ribbon reefs” at the northern end (Figure 1–6b) to ir. Mehetia island. The origin of the word “coral” is steeped in linguis. eds. Reef Algae and Foraminiferans•61 circumtropical genus Udotea among others (Figure 4–12 a. Chen J. Philosophical Transactions of the Royal Society 361B: 917–929.Paleozoic reefs. Photo courtesy of Bill Gleason. but also for nutrient uptake. tube anemones. 2005). There is no sporophyte stage known in this genus. Redrawn from Hay ME. 78 miscel. young colonies may begin life with an atypical clade and switch to the more typical one later on. dian Ocean (Camoin et al. unlike the latter group. what after death. Some cyanobacteria belong to the Prochlorophyta. Chlorophyta) from San Salvador Island. In addition. Rützler K. cegovic (Cyanophyta/Cyanobacteria). giving rise to a “star coral” pattern.growing. 1977). Some mats are large enough to be seen by satellite and have been noted along the coasts of south. of the general impor. Guilderson TP. Line Islands (central Pacific). tern. ography 48: 787–794.E. coral reef organisms may grow as a thin veneer on consoli. and Pereira RC. Life and Death of Coral Reefs. This typically leaves the polyps connected in a chain of mouths and body walls producing a meandroid or “brain coral” States. is even more spectacular.water reef de. they were the first living examples ever recorded that accounted for the laminated reef.g. Short. Frontiers of Ecology and the Environment 3: 479–486. Satellite. paddle. Demarcation between shaft and tubule is gradual. Figure 7–2). Silicatein fila.g. some groups of polychaetes and sipunculid worms are notable bio. from dramatic temperature changes and rainwater). This material may or may not include some degree of calcification (e. Riding R. New York: Macmillan.derived carbon compounds may be consumed quickly (e. tiple reef environments appears to have been underway by Miocene time. c. This listing in. However. phy and Marine Biology 48: 161–212. 2004). some ap. 1988.Emperor chain. although there are many variations and idiosyncrasies. The gametes form very quickly and are liberated only during one or two 20. while others are gregarious and con. there are other dimensions of the relationship that can be viewed as a cost. bryopsidales): production. and the rate of carbonate production and consumption over time (reviewed by Glynn 1997). ure 6–7). 2008).. Nova Southeastern University. 121 Deep. branchiae. high clone densities may also be required for increased fertility and may actually facilitate sexual reproduction. and the underside of corals and reef limestones (Figure 4–20a).HDTV.Coulon et al. I of The Invertebrates. tentacular lobes.Alsumard T. Demes et al. Suchanek TH and Green DJ.g. 2008. and turf algae. eventu. ogy of macroalgae. even if rocky substratum is available. a large tropical fora. sitioned mesenteries that continue into the rest of the coelenteric cavity. Subsequently. and terminate at their seaward margins in vertical cliffs form. Additionally. Phymatolithon foslie.crucis. Figure 5–11: The sequence of etching cell activity in boring sponges. typically a matter of hours. b. Miami 1: 23–29. Filaments approximately parallel to the substra. Some herbivorous fishes are deterred by the cal. tend for hundreds of kilometers along the Queensland coast from the shoreline to the outer barrier reef. Jones AT. Phaeophyta. ment 24: 1209–1217. the greatest S:V ratio is achieved by filling the spongeocoel with mesohyl so that numerous choanocyte chambers and multiple exit pathways are all that remain (leuco. Chadwick NE. Coles SL. 2000. as well as shell fragments and other biominerals. 1980.. zooxanthellae. Taxonomy. tebrates (O’Neal and Pawlik 2000. 2003). Rotterdam: Balkema: 265–271. and Andersen RA. Sleep N.pdf Cairns SD. especially from branching species. vide their host with a modest degree of protection from certain types of predators.carotene. Bertucci A. Overall. This speci. the intercellular medium becomes depleted of bicar.distance travelers. Mann S. is of a bright but pale green colour.like aggregations of vertical tubes and be. et al. the Gorgoniidae and the Plexauridae (Figure 6–18a). and complexity.forming gorgonian. 2007. cific reefs where they stand out owing to their brightly colored lavender. Goh F.. However. Growth. a complete growth increment is periodically added to the anterior of the tube. matic representation of sclerites from common gorgonian genera and some of their variations within or between layers of the coenenchyme. including organic compounds that selectively bind to and remove calcium. 1988. radiolarians. These ben. benthic tropical communities. Benthic (bottom. Sidri et al. photo. Marine Pollution Bulletin 41: 112–122. Deposit feeding is the consumption of detritus and associated microorganisms that have settled onto. and is the world’s smallest with a land area of . in the central and northern Great Barrier Reef lagoon: relative importance of the fixed. gonians. Stolons or stipes often extend into sediments with filamentous. Baha. with a com. 2008) that cope with this problem. sediment deposition. the skeletal remains appear to be resistant to invasion by boring organisms. Porifera.3 and 2. Kramarsky. Marine Ecology 26: 33–41. 1998. dimly lighted environ. photosynthate transfer from reserves. These splay out dur. among other factors. Mallela J. 1983. Leletkin VA. bean species are brooders. and carotene variants called carotenoids. 1997). including the description of a new species. They are referred to as accessory pigments because energy absorbed by them must be transferred to chlorophyll before it can be used in photosynthesis. poly. and seasonal (monsoon) wind reversals that result in strong windward reef and rim growth on multiple sides of each faroe (McClanahan et al. isms that perform this function are called bafflers. vorable for nitrogen fixation. This results in clusters of genetically similar adults that suggest a high degree of inbreeding (Miller et al. Most of these are freshwater forms. Marine Biology 35: 253–268. hind the northern Great Barrier Reef at 2–20 m depths.building scleractinians. and Beach KS. Gas. All foraminifera extrude their cytoplasm as sticky pseudopods. ognized from Precambrian deposits more than a billion years old. 2008). typically10–30 mm long. Oxford: Oxford University Press. Broadhurst MK. nucleation. Filamentous and encrusting forms may be found occasionally.. molluscs. the medulla. Cryptic diversity of the symbiotic cyano. The youngest and highest of the group is the volcanically active Island of Hawaii at ~19°N.dwelling alpheid shrimp on the Belize Barrier Reef. Developments in Sedimentology: 54: 475–502. Vergara JJ. perhaps aided by modified chae. In addition to curing various maladies. Ross C.cell transport or other forms of intercellular communication. cyanobacteria began to pump oxygen into Earth’s paleoatmosphere and were likely responsible for changing it from a reduced to an oxidized state. a b Reef Algae and Foraminiferans•59 growth. which is composed of sticky cellular extensions (microvilli) roughly a mi. Florida International University Rafael Lemaitre. et al.relationships (Martin and Britayev 1998).sea black 142•Chapter Six of the brooding coral Pocillopora damicornis on the Great Barrier Reef. A systematic study of some black corals species (An.Water Corals and Ecosystems. Vroom P. such as products of carbon and nitrogen fixation (see below) as well as UV protection and defensive toxins (see review of symbiosis by Usher et al. Geochimica Cosmochimica Acta 72: 1555–1569.like skeletons. Sponge reefs in the Queen Charlotte Basin. Cabioch et al. the cytoplasm of the same individual often contains both haploid and diploid gametes.like metazoans that first appeared roughly 530 million years ago. which are not able to keep up. cally grasp their food and do not affect the substratum. In addition to food considerations. the spongeocoel (Figure 5–1. Abrar M. Scoffin TP. forming tunnels or geyser.. Energetics. Miller AI. Work TM. and as secondary reef builders if they contribute relatively smaller volumes of carbonate. and because cyanobacteria are so common among them. Coral Reefs: an Ecosystem in Transition.bearing sponge Theonella swinhoei: bulk DOC is the major source of carbon. The cortical filaments form swollen tips called utricles (u) and these constitute the outer surface of the segment. Likewise. Pink or red carotenoids are embedded within the crystals. 2007. 1997). lar budding patterns: Polyps contiguous as in Diploria labyrin. a process that can be observed in the field. Adey 1998). The glassy test sur. 1997. Nishikawa A.000 years. The role of microbes in accretion. It was not until 1951 that Darwin’s subsidence theory was confirmed on Subsidence Subsidence Subsidence Vertical growth Vertical growth of coral of coral Vertical growth of coral Vertical growth Vertical growth of coral of coral Vertical growth of coral Lagoon Lagoon Fringing reef Barrier reef Atoll Time Limestone Sediment Oceanic crust Figure 1–15: a) Sequence of coral reef development proposed by Charles Dar. able (Goreau and Hartman 1966).like. In contrast. A third group. and many within the Indonesian Archi. The distribution. ganic material that dictate the manner and time during which calcification takes place. Taxonomic subdivisions of the group are often consid. Modi. Marine Biology 152: 639–644. live in tubes of their own construction. Coral Reefs 22: 27–28. and then transfer them back to their hosts. Although stromatolites persisted as significant reef. Calcification generates protons for nutrient and bicarbonate up. b). it may be difficult to exclude organisms that produce an internal skeleton but have completely dispensed with the production of carbonate. This is particularly interesting in broadcast spawning species. Further work will be necessary to sort out the exact nature of these symbioses and their overall sig. tion. and alcyonaceans (McFadden 1991. Reef Cyanobacteria•45 forming organisms for hundreds of millions of years. Heteractis magnifica. nians in shallow water are particularly adept at this. Bulletin of Marine Science 81: 423–435. despite the apparent oxymoron. Gilmour JP. However. small unicellular forms in planktonic communities can contribute significantly to the nitrogen budgets of N. often in the form of reefs that have become elevated by sea level changes or tectonic activity. their activity in the Far East.5 million of them mL–1 . cell cultures from cor. Pueschel CM and Wegeberg S. coral reef development in this region is geographically restricted and includes low. 2009). meaning that in general. ob. The main opening of the shell. Scale approximate.500 species (S. Herbivorous fishes have been observed breaking off younger. the process is dependent on the ability to generate ATP. 1982. Geomet. 2006). pumping is not continuous and may cease intermittently for reasons not yet understood. the yield was about 10. rates the two groups into separate suborders (Appendix 5). a b c d e Figure 6–2: Various types of anthozoan cnidae. most of them have been identified as a Synechococcus spongarium.based calcium. However. 1993. Paleontolo. Atoll Research Bul.assembled from pure solutions in the laboratory. Limnology and Oceanography 48: 141–149. 2006). Intes A and Caillart B. However. The net effect of borer pop. ries of several important groups are atypical. but build complex habitats for organisms in the intertidal and subtidal zones. eds. Wainwright SA. Tahiti 3: 105–110. However. Hawaii. In reef environments they typically remove carbon.out. together with vari. D.corals (but not all) calcify in a distinct night/day pattern (Gladfelter 2007) and among those. but there are no known unicellular species. Protons are produced (1). Their excavations may take the form of simple grooves or pits at or near the surface. Chaetae are used primarily for locomo. Cytogenetic systems and evolution in foraminifera.rich bundles. Rapid settlement in broadcast spawning corals: implications for larval dis.shaped. Caribbean shallow. synthetic tissues of the cortex. Advances in Marine Biol.fracture. Integrative and Comparative Biology 47: 693–700. Acropora palmata.sac located at about 26° south latitude on Australia’s west coast (Figure 3–13a). Taylor PD and Wilson MA. Allen M. Proceed. 2006. after initial calcification of the middle lamella. 2003. ten with contrasting white bars or patches. hour period. Others such as Archaeolithophyllum formed a thin. The resulting test not only is distinctive in its porcelain. Nitrogen fixation by cyanobacteria is an important part of the cycle (Chapter 3) and has been demonstrated in sponges (Wilkinson et al. stead.E. Another chain of submerged peaks is included in the Emperor Seamounts (many of which are named after former emperors of Japan). Fabricius K. 251 Corallivorous Damselfishes and Their Gardens. McMurray SE. A Functional Biology of Sea Anemones.water carbonate margin: Easter Group of the Houtman Abrolhos. undergo extreme changes in appearance and become active swimmers during the season when sex cells mature. These compartments. When added to the number of oceanic atolls in the Indo. and a 25. 2003. 373 Glossary. lidae) in the Mediterranean Sea.Pacific spur and groove topography is controlled by fluctuating sea levels that occurred in that region over the last several thousand years. 1999. French Polynesia) and its contribution to benthic primary production. view by Griffiths 2006).. ria. Photo by au. sug. apparently smothering them. In addition. a term that has nothing in common with things that cause one to be baffled. 172°E) and V. Peuravuori HJ.sized particles or fine sand grains up to 85 µm in di. which lack a mouth and digestive cavity. This system is 250 km long. graphic regions have yet to be adequately surveyed with respect to their sponge fauna. in red algae the female gametes are not released. palatable to herbivores (Pitlick and Paul 1997). position. 2005). both axial skeletons allow the colony to bend in a preferential position under wave and current forces. 20 m deep. and reproduction. although certain stylasterine genera can be common typical of the Indo. but larval motil. 2002. including the reef crest (Chapter 1). and relation to land masses.Victoria M. tivity directly affects rhodolith populations. an Evolution. While Darwin was generally correct that certain types of reefs develop in sequence. these corals become more abundant at the geographic extremes of reef build. Sections through the skeleton reveal growth by layers of thick. by ~2. However. either to repel attack. Clionai. especially in the south where they occur as polygonal atoll. Reproduction and recruitment of corals: comparisons among the Carib. Guam 2: 1217–1226. Late larval development and onset of symbiosis in the sclerac. et al. however. some broadcast. sponges constitute a primary food source.” whereas e) others exhibit different degrees of separation. 1983. Large sabellids typically occupy their tubes per. variations in isotopic composition of coral skeletons can be accounted for by changes in temperature and pH of the fluid from which the skeleton might precipitate (e.” These include three that are associated with the Belize barrier reef (Figure 1–7). 47 4 Reef Algae and Foraminiferans. inozoans. b) Millepora is read. Zooxan. metabolites produced by one organism (in this case. Skeletal growth and changes in density are known to be influenced by a wide variety of environmental variables. Florida International University Kristin Fauchald. the presence of these acids necessitates limited oxygen con.like tube is characteristic of the group. anten. millions. Sagi I. What is her. ultrastructural patterns. caudal tube.” A flexible. Archaeocyathid sponges flourished during the early Cambrian and were the first metazoans to appear on reefs. ing species are restricted to caves. ing the area of boring. little is known about the pro. a b 134•Chapter Six cast spawners. scopule (lower left). prism. How brain corals record climate: an integration of skeletal structure. and will be described here only briefly. Chapter 8) are avid and specialized cyanovores that not only seem to grow well on a diet of certain cyanobacteria avoided by other consumers. Neall VE and Trewick SA. Tompkins. This exposure shows more than 2 m of relief (scale bar = 6 feet). These structures also become progressively reduced on the abdomen and may be absent on the posterior seg. Häussermann V and Försterra G. Indonesia. tiple rhizoids and grow as erect or creeping psammo. 58•Chapter Four grains of aragonite. sea cucumbers. An actively dividing meristem layer lies between the epithallus and perithallus and contributes to the for. struction and destruction of carbonates by marine and freshwater cyanobacteria. the size of which can range from a few microns to larger species that may reach over 15 cm in diameter. threadlike extensions of the mesenterial filaments called acontia that often fill the base of the coelenteron. Melkonian M. current flows. Morphogenesis and generic concepts in coralline algae: a reappraisal. although excellent for surfers. eralization. the fourth group. Rapid rhizoid produc. brackish coastal lagoon in Argentina. Mesodermal anatomies in cnidarian polyps and medusae. ceedings of the 2nd International Coral Reef Sympo. 1:1–10. Environmental factors associated with the spatial distribution of crustose cor. various types of herbivory. Thus. Goreau NI. and release oxygen. New Haven: Yale University Press. and sometimes more. Mackie GO. 2005. Coral reefs: algal structured and medi. Egypt) and their geological implica. but are not associated with. Moreover. Lundgren P. a submerged or largely submerged coralline shoal originat. 1997. but not until nearly 70 years after his death. but whose lagoon plummets to 100 m (Tomascik et al. Pearse AS. Yellowlees D. Indeed. Based on Hyman 1940. the view of these corals as renewable resources may be challenged in light of their slow growth. Coral Reproduction Asexual Modes Asexual reproduction takes many forms in sclerac.dominated biofilms: a high. 2006. lagoon reefs may be restricted to channels or passes within the barrier. pulids to rapidly roll up and retract the branchial crowns into their tubes. Nugues MM. Western Australia. ter 1 (e. bores deep galleries into coral substrate and exudes a toxic mucus that kills surround. there were numerous anonymous peer reviewers and others whom I consulted for fact check. including nutrient recycling. and bioerosion are especially important and are emphasized. and add three. three. regeneration. Buster NA and Ross SW. deeply bur. Vegetation is often well de.g.000 to 1. To make matters more difficult. The growth re. 2001). In contrast. but one that can be quite vague with. L’Institut de Recherche pour le Développement. banks. bushy.like. dant on reefs. How. and some organ. Modern reefs often incorporate Halimeda sediments into submarine cements whereby they become part of the reef fabric. 1992. Smithsonian Contributions to Zoology 615. Photo by author. but this would limit bioerosion to such periods. search Bulletin 531: 1–65. bioerosion is significant. filled. but most of its rim is submerged 3–20 m below the surface. Bacterial calcifica. 39 outside of the cell wall. albeit slowly. Evolution 58: 308–323. phyll a showing the porphyrin ring and phytol side chain. Laboratory experiments that follow fertilization have shown that there is considerable variation in the propen. While not physically impressive. Grazers make up a broad spectrum of animals. it is not clear if these consumers simply adjust the amount of sponge material consumed according to its level of toxicity. These include the so. Indeed. 1915. Indo. ity in the calicodermis. many of them form on con. Although the tentacles are grooved and feeding is generally similar to the manner used by sabellids and serpulids. and Burch IW. large quantities of planktonic food. especially outside of the tropics. View corresponds to about 5 o’clock on Fig. López.1029PA001288. Systema Po. duced during calcification (McConnaughey and Whelan 1997). Although there are exceptions (described below). Phospholipase A2 in Cnidaria. Uriz MJ. 1981. New York: Chapman and Hall: 68–95. the addition of ions may occur in a spe. careful. rum: descriptions and quantitative aspects. burying the reef. is often found in high. Adapted from Land and Moore 1977. 1986.sea corals in the northeast Pacific Ocean. Note dusting of freshly deposited sediment on the surfaces. and mineralogy. and provide an evolutionary linkage among the various types of reefs described earlier. However. brous composition suggest that this group shares a common ancestry with certain demosponges (Wood 1990). but the term coral is now generally restricted to the Cnidaria even though only some of them are mineralized. Indeed. This not only facilitates wound repair and regeneration (Cabioch 1988. Bastidas et al. Bacterial suspension feeding by coral reef benthic organisms. the Caribbean. is shown in Figure 5–10 boring through living coral tis. but also produces a densely calcified thallus that contributes significantly to reef structure (see below). and channels retaining water.Garcia JL. and Neilan BA. ments that absorb light at wavelengths that chlo. 2001). tion by fragmenting the thallus during feeding. The environmental recording unit in coral skeletons: a synthesis of structural and chemical evi. those described by Riegl 2003). Taxa Referred to in This Text. Reef sections that are on the leeward side or sides may lack such islands and are more likely to pro. depending on the pattern of asexual reproduction (see below). logical Society of America and University of Kansas. Recent advances in fertilization ecol. Richmond RH. In some that are poorly calcified. which grow colonially in different forms. Whichever way the sperm is transferred. vastifica. Coles and Fadlallah 1991). low). Manderson JP.section through a branch showing the relationship of the outer polyps (p) to the holaxonian axial skeleton with its dark cortex (c) and chambered central chord (cc). liphagum. I hope that this contribution represents time well spent. Coral Reefs 26: 531–538. cause they release terpenoid toxins into the water that cause tissue death of established scleractinians nearby. gella.or vase. Journal of Phycology 13: 6–16. In some rare instances. auii drape over reef edges and are the dominant forms (Freile et al. However. xanthellae (Chapter 15).5198. 2007) and enzymes such as Ca. These settle and produce a second plant. and the higher vascular plants that develop such complex structures. gastropod shells. juvenile thallus (Meinesz 1980. However. cupy a broad geographic band extending 40° north and south of the equator to depths as great as 200 m.rich sediment. which likely contributes to the success of these algae in low. only a few types of ca. ing the last 340. Many commercially valuable species are part of coral reef ecosystems. the group is best known for the millimeter. Lapian HFN. or if they are simply “optimal” from a view. The dorsal side of the abdominal and para. Knutson DW.carbon units of alternating double and single bonds. Increased amounts of sediment deposition and turbidity can occur seasonally under natural settings (e. grains. Some reefs that did not fare well during the early Holocene. I: a method for mea. tal Biology 209: 1690–1695. some (e.preserved reefs indicate that sea levels at Huon fell by 130 m or more 140. Wilkin.like specimens. Regional changes in sea level occurred because of postglacial crustal adjustment and subsidence. alization and the problem of the vital effect. On the Hawaiian island of Molokai. Sponge metabolism Particulate organic nitrogen Organic nitrogen production Cyanobacterial symbionts Dissolved N 2 Ammonium NH 4 + Nitrates NO 3 – Nitrites NO 2 – Denitrifying bacteria Nitrifying bacteria 94•Chapter Five been suggested that allelochemicals may facilitate this outcome as well (Pawlik et al. Indeed. and high nutri. Turon X. Boschma H.fixing bacteria in the planktonic environment. Some Ca. for warmly embracing this book and for encouraging me to join the University of Chicago family. Schönberg CHL and Loh WKW. rine ecoregions of the world: a bioregionalization of coastal and shelf areas. directly contribute to the widespread loss of these beds (Steller et al. There are even a few examples of uplifted almost.developed in. Lipid droplets and starch grains may also be common depending on the physi. ous A. The structure of their aragonitic chambers and fi. Hadas et al. arborescent. and • Formation of a tough. DeAngelis P. Castenholz RW. Coral Reefs 18: 204. Photo courtesy of NOAA. 1975. their tunnels weaken the skeleton of their coral hosts. 1971. Thus. but decalcification may occur in Halimeda as well (Borowit. Paleoceanography 22. b) High. Thus for photosynthetic purposes. ed. Acknowledgments 1 in waters hundreds or thousands of meters deep. and may develop seaward of the main reef or behind it. Grehan A. and Taylor MW. 90 6 Reef Corals and Their Allies. Carlon DB. and often occur as negatively charged poly. 2008. In: Thierstein HR and Young JR. Pocillopora damicornis. If upward coral growth were to keep pace with this sub.. 2000. tity of the unique symbiotic dinoflagellates found in the bioeroding demosponge Cliona orientalis. sand channels. The septa may also extend beyond the corallite wall to the adjacent corallites. nies resemble filaments and produce a row of cells within a sheath. Thus. lectively called Les Récifs d’Entrecasteaux.br/ig/sigep/siti0041 /siti0041english. lecular control. but smaller examples (van der Land 1977. 2007).. these can take an erect or sprawling form. Kojis BL. Buddemeier RW.” The lagoon is large. Hooper NA and Van Soest RWM.dimensional atomic structure.25 mm (250 µm) in diameter. there appears to be a mixture of sexual and asexual reproduction in different regions or reef habitats (Ayre and Miller 2004. ter description of brooding and release at the larval stage.Deep fore reef with buttress and channel system 40 m . Part E). Records of these changes come primarily from raised or submerged shorelines. Because of the rigorous conditions of exposure here. Tahiti 4: 505. This complex internal region is sometimes referred to as the endosome. 2006.226. The fertilized eggs produce planula larvae that settle. Such reefs often do not produce a shallow crest and are typically not assignable to the groups described above. 1969. v Acknowledgments. Effects of soft corals on scleractinian coral recruitment. 2a. Indeed. aware for centuries of the spawning periodicity. the reefs An Introduction to the Structure and Formation of Modern Reefs •21 Enewetak Atoll in the Marshall Islands. is the aperture. and Subramanian G.like. damicornis whereas in others (e. Sabellids lack an operculum so when they contract. they are taken up. Berman. ography associated with the southward flow of the Cali. Coral Reefs 17: 329–337.dwelling) cyanobacteria are also im. The quality of river water. Inset: Detail of gastropore with its spiny gastrostyle (S) and adjacent dactylopores (D). Coral Reefs of the Indian Ocean: Their Ecology and Conservation. a minimal depth of 10 m is more typically the threshold for distinguishing such basins from a reef flat (Milliman 1974). 2006. University of the Azores.dimensional frame. the general view is that mineraliza. mella. The porphyrin absorbs photons only in blue. Plata banks off the NE Dominican Republic are similarly submerged coral plateaus. salinity. 2003). The lagoon encompasses an area of about 6. ventral groove. ings of the 4th International Coral Reef Symposium. Conversely. Reproduction by fragmentation in corals. 1985. 2008). composed of mucus and protein. although many such reefs occur in the Indo. mobile rock) flows as a plume from deep within the Earth’s mantle (Figure 1–17 a. Bicarbonate is converted to CO 2 at the zooxanthellae membrane by carbonic anhydrase. Graham EA. jority of polychaetes that have been studied brood their embryos or larvae for a variable amount of time before releasing them (Glasby et al. Zoccola D. Thacker RW and Starnes S. both of which produce a framework for the organization of organic and cellular materials. 2004. cally defended (e. macro. Nagarkar S. rial symbionts. Biological Bulletin 183: 278–296. the largest known Permian sponge. Paerl HW and Carlton RG. Silverman et al. b) Lateral view of S. Coral coloniza.g.000 years (Macintyre et al. systems. ing appendages called palps. tion (Clavico et al. the habitat in. cran. or binder. 352 Resilience and Phase Shifts.Nova DA. worms and leeches. terconnect the coelenteric cavities of each polyp and serve as a primitive circulatory system for the rapid transport of fluids. In: Birkeland C. 2000. lift (see below). Watch diameter is 3. tency and are common on Indo. there are exceptions like Sadapur Atoll (Flores Sea. sive competitors. and herbivores that feed on endolithic algae (Fauchald and Jumars 1979). uncolonized by rap. Stromatolite Formation Trapping the Sediment In examining the fossil record. metamorphose to primary polyps. 1: pigmentation. living corals. Takaoka TL. sail. a thicker subsur.like “bommies” in Australia. abdomen. about 520 Mya.building organisms on reefs are a group of foundation species. Scale is approximate. Each piece is then transferred to the building organ where it is held. James NP. but coral growth.up of the coralline algae composing the ridge. PLoS One 3:e2989. In turn.to.calcification has taken place. Historical factors including sea level rise may also play a role as described below.Rodriguez R. Most reef dwellers have a wide oral disc compared with the column and form a well. Jara J. see Lewis 2006. However. a process that is essen. The evolution of modern corals and their early history. Axial skeletons often remain after the death of the colony and may serve as substrate for colonization by a wide variety of other or. A: adult colony. Reproduction. Aggregations of several hundred choanocytes may be found in each chamber. Figure 6–27c). Plant Conservation. These peaks are now ~5 degrees beyond the Darwin Point and 400–600 m be. and muscles. b carotenes xanthophylls starch Phaeophyta 1. but as is so often the case. Patches that remain isolated are given various local names according to their morphology. genetic reconstruction using secondary structures of Internal Transcribed Spacer 2 (ITS2. Berkeley. vides a physical refuge. Love et al.. Their rapid growth rate accounts for high levels of production. a carbonate mass that is different in composition compared with the surrounding substrata. References to cyanobacterial taxonomy. Edmunds PJ and Davies PS. 96•Chapter Five single mature female (the queen) at a time (Duffy 1996. United States National Museum of Natural History. Gorgonin is dis. They are characteristic of continental shores. including the key features of the coral. Sponges also contribute to habitat complexity. The last cool period began 12. the range of seawater pH is depicted by the vertical blue line where the predominant form of inorganic carbon is bicarbonate. molluscs. and King CE.g. However. but killed by the red alga Anotrich. and (2) it exchanges that Ca++ for 2H+ generated by the calcification process. most have well. Indeed. Thus. SE Hawaii. Coles SL and Fadlallah YH. deed. 1b. Carbonate production of an emergent reef platform.. 1982. No osculae are vis. Yahel G. stomium. where sea level was at the time. cies can occur in a variety of morphs ranging from flat and encrusting to those producing branches or upright plates. especially during the Paleozoic. If the spines represent a form of protection.Kirby 2001). Germany. 1977. The outermost portion of the shell. 1996. within the radioles. Hallock P. studies have shown that nutrients passed on to the host are deficient in both phosphorus and nitrogen. to the memory of our little girl. Formation of either of these even. and some colonies can grow to a meter or more in height (Grange 1985). The evolutionary history of Symbiodinium and sclerac. However. et al. 175 9 Reef Lophophorate and Protochordate Phyla. The second type includes corals that brood and retain their eggs. Scleraxo. Real time and isotopic dating methods sug. 1972. tion of sea fans. aria). some HMA sponges may be supplied with much of their organic carbon requirements from this source (e. 285 Contents 14 Ecology. most are hermaphrodites includ.. even in the absence of agriculture. depending on species and environment (Multer and Clavijo 1989. a b Reef Sponges•93 the process among sponges has yet to be established. actinia. and the seamounts northwest of Kure Atoll in the NW Hawai. Likewise. NASA. Some of these have largely submerged rims. Ecology 72: 1849–1866. Symbiotic interactions between marine sponges and algae. oc. ghetti worms” because of the numerous white. Algal groups do not have the long history of the cya. Animals. moval by physical and bioerosion (Figure 1–1c). Clode and Marshall 2003). ally to change the ratio of choanocyte surface area to the spongeocoel volume. side of the cell membrane (Figure 4–16b). typically reflecting the shape of the filament (Garcia. Cortés A. 205 The Maxillopoda. no. Pacific. or concave “platform reefs. separated from shore by a 1–2 km wide depression mostly 1–3 m deep. duced on a reef’s windward side where they function as a natural breakwater and dissipate incoming storm waves.to. 2006. sive structures have been described from Loch Creran in Scotland extending over an area of >1 km2 at depths of about 3–10 m. 2005. Marine Ecology Progress Series 278: 103–114. Precht WF. ites that are typically linked by mesoglea to produce a flexible but hydrodynamically resistant internal skeleton. 2005). Instead. tiple and symmetrical planes of nucleation that form a three. 1997. zoanthids. which are protein polymers used as nitrogen reserves to support growth in nitrogen. Journal of Experimental Biology 210: 91–96. Cairns SD and Macintyre IG.e. Turf Algae Algal turfs consist of filamentous and short. 250 300 350 400 450 500 550 600 650 700 Wavelength (nm) Absorption Violet Blue Green Yellow Red Phycoerythrin Phycocyanin Chlorophyll a β-Carotene Figure 3–7: When plot. they deliver venom that produces a burning sensation. cantly higher proportion of branching corals (Blanchon and Jones 1997). the planula or initial polyps must acquire them from the environment. Gektidis M. and sphinctozoans were common from bottom trawling and other fisheries activity. University of Hawaii. The terminology employed here depends on perspective. In other unicells such as foraminiferans (Chapter 4). there is so little tissue. alization processes and the problem of the vital effect. Likewise. Protozoa through Ctenophora. cies and others for several weeks does not result in any loss of host biomass. Lasker et al. cluding those described below are distinctly unfavorable for reef corals. Journal of Experimen. Cnidarian neurobiology: what does the future hold? Hydrobiologia 530/531: 107–116. it is unclear if epithallial shedding generally occurs spontaneously or if there is a stimula. isms. the radioles account for most of the gas ex. forms extensive reefs (Figure 7–8) in locations generally unsuitable for corals (but see McCarthy et al. Similar coral. and the entire colony can change orientation by gradually rotating perpendicular to the current as it grows. Journal of Animal Ecology 66: 793–810. In any of these situations. trations of proteins and polysaccharides in the “centers of calcification” compared with the surrounding fibrous crystal growth. In addi. but are inbred. Mechanical Design in Organisms. 2005. and denitrification in sponge nutrition. Payri CE.1000.like growths of Dictyota (order Dictyotales. Paul VJ. in the case of this coral cliff. this spe. Antipatharians form un. In contrast. ian milestones in metazoan evolution. fix carbon. Norwegian Insti. Small AM. and taxonomic affinities of tropical Australian sponges. creases to >9. For the most part. Journal of Experimental Biology 203: 3445–3457. Marine Ecology Progress Series 346: 143–152. Warner et al. Short cirri are also associated with the parapodia. Canada and Adjacent Ar. 1990.” as it is called. ally by fixing carbon or nitrogen. ture. 3–8c). Tak. penter WJ. 1991. lar teeth of limpets and chitons (Mollusca.g.to a corallimorph.radiography. Coral Reefs 15: 109–119. Journal of Fish Biology 53: 209–221. I did not work alone. rophy in low light. Vat. fied from Schmidt 1974.g. The northwest coasts of the main Hawaiian Islands (Hawaii. tosynthetic activities of their zooxanthellae (Chapter 6). Chlorophyta).bio. chemistry Geophysics Geosystems G3 8. tubule is greatly reduced. D. ons that are pointed on one or both ends are common. may be included with turfs or form their own community. the opposite is true in cases where coral growth is marginal. and Chatterton LW. titative morphology of a fringing reef tract from high..g. Herren LW. The biol. 2008). This apparent paradox may be explained by distinct activities in different portions of the filament. Netherlands Antilles. perhaps suggesting that they are culled and digested by their hosts.. Halimeda biomass. associated organisms (Roberts et al. and Falkowski P.water coral reefs (Chapter 6). 25 meters above present sea level.. Others exhibit different forms of intratentacular budding within a colony. Watling L. a condition charmingly referred to as an “almost atoll” (Figure 1–16). placed by Acropora cervicornis (Figure 1–22a). d) Alternating calcified and uncalcified nodes of a calcaxonian “bamboo coral. 2007) commonly called black corals. rovsky OS. Bioconstruction and biodiversity: their mutual influence. lips.called imperforate forami. although this varies depending on local wind and current conditions. Resource partitioning by Caribbean coral reef sponges: is there enough food for everyone? Memoirs of the Queensland Museum 44: 457–462.g. and may include the first segment.corals (Rosen 2000). Additional protons are produced during this process (3). The polyps are polymorphic. Five species among these genera bore into living stylasterine. Canadian Journal of Zoology 84: 288–306. Aragonite displays a symmetry that is distinct from calcite. tals in boxed area of (a). particularly in species where the food reward is small compared with the volume or type of spicule that must be ingested (Jones et al 2005. Proceedings of the 8th Coral Reef Symposium. Most species are gonochoric and release large num. or by directly cementing them onto the reef. Reviews in Mineral. Porat D and Chadwick. Orthorhom. like” reefs are sometimes referred to as “bank. Certain opisthobranchs including Elysiella pusilla (Fig. Proceedings of the Royal Society B 276: 1063–1067. There are also in.C. Nitro. Thus. forams are important as indicators of sedimentary environments and age. However. 6–27a). fied. suggesting that the host may be required for settlement. celerate the process. 46 Nitrogen Fixation and Trophic Relationships. However. Science 188: 257–283.dimensionally complex with tunnels and cavities.13: 978. tion of branching corals under the influence of wave and surge activity. must be crucial to calcification in AZ corals (e. called the Archaeplastida (Adl et al. predation. etal carbonates. many poorly known species are encrusting and cryptic (hidden) in cracks. c carotenes xanthophylls laminarin mannitol Rhodophyta 4. Oikos 69: 476–498. courtesy of André Freiwald. NASA.g. among Porites. Dordrecht: Springer Science and Business Media: 59–85.. Journal of Phycology 37: 659–667. 1962. and is capable of growth in ex. 301 Diversity at Geographic Scales: Biogeography. ture. growth. In New Zealand. tropical carbonate en. Effects of anemonefish on giant sea anemones: expansion be. similar to milleporids (Figure 6–23b). In addition. American Scientist 62: 272–281. Some species are most abundant at 25–30 m and do not exhibit sig.West Pacific. In some leafy red algae. et al. b) Diagram of Halimeda structure showing the branched outer cortex and the utricles.M banks. A biogeographic analysis and review of the far eastern Pacific coral reef region. Its aragonite skeleton is produced from intracellular spheres that are released and then fuse extracellularly to form layers. Calcisponges A third group of sponges. 1984. egories. 2010. octocorals. In: Moore RC. 2005). Coral on seamounts. In: Kaesler RL. Pernet 2001). 2005. spongiae) in Barbados. and the effect of Pacific. and have separated them into numerous cat. which like Astrosclera occur as cave dwellers in Indo. sponges are the most important source of siliceous reef sediments. times as layers that are mineralogically distinct aggregates of aragonite and calcite. ganic sediment layers by the dome. water black corals (Cnidaria: Anthozoa: Antipath.g. algae may convert bicarbonate to CO 2 outside or inside the cell (Badger 2003). including (a) organic photosynthetic products or oxygen that could ac. gamete. moval from living tissues. and conducts them to food grooves. 1990).W. and Atkinson M.like protein. els. cal factors. xanthellae. The western side is West Papua. Lewin RA. Both the northern reef crests and the southernmost reefs. In: Frost SH. Center: cut. and may bear a number of other special. Here in the northwestern Society Island group. This in turn results in size reduction and lower rates of survival.E. volume). 2004.. The Biology and Geology of Coral Reefs. because this layer is composed of a single type of cell that lacks intercellular communica. Figure 2–5: A simple hexagonal crystal (a) and rhombohedral form of calcite (b) with its crystal axes. Reef ecology. In contrast. Helgoland Marine Research 33: 624–640. 2005. mental Marine Biology and Ecology 341: 282–294. ally less than 100 µm long and called microscleres. a more lim. and for reasons that go beyond their architectural attributes described above.driven sperm called spermatia that Reef Algae and Foraminiferans•65 with uncalcified joints similar to Halimeda are referred to as the articulated corallines. The other gold coral group is represented by a single genus and species. The entire range of sclerite size and shape within the colony is diagnostic of the taxon defined by classical taxonomy. It should be noted that there are varying opinions concerning use of certain terms among cnidarian specialists. Some corals close to the equator may spawn several times a year. the emergence of the Isthmus of Panama about 3. Andaman Sea. Shinn EA. Fabricius K. which can sometimes be distin. Astrosclera can become quite abundant and can play an important role as a frame builder within its cryptic habitat (Quinn and Kojis 1999). Baco AR. After Barnes DJ and Chalker BE. Hawaii. 1997. http://aquaticcommons. From there. Pacific reef sponges. anemones without defending symbionts are subjected to attack by butter. Cryptogamic Botany.” refer. try.and cold. Kupriyanova et al. sponge spicules include those with highly complex forms. This material is interlocked and partially bound by coralline algae and other marine cementing agents. Rodrigues AI. Reproduction Marine sponges use several mechanisms to repro. rows that are readily recognizable. bacterial filaments (green upper layers). Each of these invades nearby sand grains. Photo courtesy of Ken Grange. reheating and compression of the Earth’s crust. and dismissed as contaminants by others. The jaws may act simply as pincers that convey inanimate digestible matter. However. hosts.” More needs to be known about the bi. as well as by cellular and biochemical factors that are not thoroughly understood. Clownfishes compete for acceptable hosts and.rate and age distribution of deep. 367 2. which is converted to CO 2 by this enzyme (see Equation 4–2 in Chapter 4). Examples include raised fringing reefs (e. may also be able to fix nitrogen. 1998. Reefs with shallow. ———. Williams B. Upon decom.binding and cementing agent that calcifies the spaces within and between coral colonies. these particular dinoflagellates. the alcyoniid genus Sinularia is tough and leathery. tance of the symbiosis to the enhancement of calcifica. On complete separation. Calcium carbonate fixation by large reef. Bio. Note exposed intertidal region with algae. 2006). fying species. Whether larvae are brooded or produced externally. Demospongiae) with filamentous cyanobac. xanthellate coral communities: an overview with im.like palps emanating from the head (Figure 7–3). 102 Reef Hydrozoa. Uni. Coral Reefs 17: 145–154. tation in reefs through time. Limnology and Oceanography 51: 1548–1550. The aragonite in Udotea and Penicillus is formed primarily in the capitulum where photosynthetic rates Figure 4–12: Psammoph. The skeleton is uncalcified and is largely composed of protein along with a distinctive minor component of the structural polysaccharide chitin. ganic nitrogen is captured by the host and digested to form ammonium. Hirose et al. Photo courtesy of Paul Humann. However. stegina gibbosa d’Orbigny (Class Foraminifera): ob. nesia east of Tahiti. shelf. sent parallel mirror. charide polymers. Reef Hydrozoa Milleporid Corals Two hydrozoan taxa produce calcified skeletons. However. These are often confined to the outer surface layer of the host. deterred by these chemical weapons. 193 The Bryozoa. 1998. Brown algae store carbohydrate reserves in the form of laminarin. only small gaps remain where visible light cannot be used (Figure 3. and Cohen AL.excavating sponges exposes these corals to bor. 2003. However. Inset: These boring sponges lack zooxanthellae and are capable of excavating cavi. Asexual budding in fungiid corals. Papua New Guinea: constancy during global change. exhibited appreciable powers of regeneration. However. and perhaps most prolific producers of algal carbonate on modern reefs. and has been referred to as the Darwin Point (Grigg 1982. ten so heavily bored by cyanophytes (along with many other organisms) that an intertidal notch (Figure 3–10) is formed as weakened carbonate is exposed to the me. Symbiosis 29: 263–281. International Journal of Remote Sensing 24: 2661–2682. 2006. All photographs from NASA. Bricker et al.latitude areas (Arabian Gulf. Small et al. ever. lids and facilitate gas exchange. common and will be described in subsequent chapters.g. 2003.magnesium calcite. reflects the two different types of polyps that make these colonial organisms polymorphic (Figure 6–3 a. Amsterdam: Elsevier: 109–132. Marine Biology 125: 655–662. Evidence for a plasmalemma. umn. Senckenberg Museum. and the walls of certain highly calcified red algae. Ecological roles of natural products from the marine sponge Geodia corticosylifera. described in more detail below. The common term “black coral” refers to the color of the skeleton. 2002. European Journal of Phy. The diffraction patterns thus produced are detected using specialized instruments that have enabled the distinction of 12 different crystal forms within the hexagonal system. dive Islands (Woodroffe 1992). Biodiver. Dozens or even hundreds of carposporophytes can form with a single female. Terebellids are commonly called “spa. Halimeda produces two chemicals. High resolution mapping of community structure in three glass sponge reefs. Prufert. ment of the carposporophyte stage within the female tis.g. ily of sponges still occur today (see below). 2003. teria. Fox HE. acterization and role of carbonic anhydrase in the calcification process of the azooxanthellate coral Tu. tion to an inner peptidoglycan layer. Halimeda sediment production reaches its peak in synchrony with the re. an Evolutionary Perspective 14: 185–204. upper left) with different configuration. although different vari. Rosen BR. Modified from Steneck and Dethier 1994. which is distinct from the S. ment. Trends in Ecology and Evolution 20: 381–386. ant zooxanthellae.. Seasonal dynamics of mac. there is often clear water and a sharp increase in slope that promotes the shedding of sediment. tions called tabulae. This symbiont is a sponge specialist with geneti. Tambutté et al. idly growing algal turfs and macroalgae. Aguirre A. et al.” Aragonite is harder and denser than calcite. Reef Foraminiferida constitute 10%–15% of the cal. tain prey. Wriggling female epitokes con.1 mm in diameter when fully developed. Other LBF groups persisted. vines. especially in the tentacular epidermis. Meibom A. too lit. turfs often fix nitrogen and may provide a considerable proportion of reef primary production through photosynthesis (Adey 1998). and on seamounts (submarine volcanic peaks) found at depths of hundreds of meters. there are ten species of sea anemones that can reach a meter or more in diameter and are sometimes called giant anemones. they too often occur in multiples of six. In such cases. Townsville 1: 263–272. Unlike the give. This chapter examines the functional roles of macroalgal groups and turf algae with particular reference to growth. generalist predators. Clownfishes may contribute excretory ammonium from the plank. 60•Chapter Four opisthobranch molluscs (see Chapter 8) not only prefer species of Halimeda with high terpenoid concentrations. Now. Honduras. Conway KW. and Zankl H. the cortex may be complex and provide only small in. ing waves (references in Storlazzi et al. 2004).fertilization. These serpulid reefs tend to occur in areas of low natural sedimentation and low wave energy. 2006. tial for stromatolite growth (Reid et al. and reproductive forms that produce short. goons.” The middle region is composed of a gelatinous matrix called mesohyl. With little interference from cell membranes. Behavioral Ecol. gest a growth rate of about 0. 2008b. Gross carbon. However.g. Biegala IC and Raimbault P. Annual Review of Ecology and Sys. which exhibits gold highlights. Several species are now grown under aquaculture conditions. display a lower rate of growth. leaving behind numerous dead. Redrawn from Lee and Carpenter 2001. These ecologically complex communities constituted the first metazoan reefs (Rowland and Gan. Many spe. stein and Ogg 2009). with thylakoid membranes that are unstacked and thus do not develop grana.Org. cite crystals. Hay ME. Harney and Fletcher 2003). Several species of Halimeda are responsible for large amounts of carbonate on shallow and deep portions of the Great Barrier Reef.associated reef water. material courtesy of Henry Reiswig. calcified shelf produced by the stolonic cells. Searles RB. Miller KJ and Ayre DJ. Häder DP. Ecology and taxonomy of Halimeda: primary producer of coral reefs. whereas Indo. The Tuamotus occur as parallel island chains about 1. or even to edit a volume in his own field. 1994. Croix. personal communication). Levitan et al. Cheshire AC. a transitional region of re.Hill. These reefs grew on previously existing topographic highs and produced substantial relief in the clear waters nearly 200 km from shore. In general. the abdominal segments are often wide and bear reduced parapodia with elongated. microad. Qiao L. 2008. Microalgae. they often contrib. 2002. are separated from those with low elevations. Table 1–1: A Classification of Bioeroding Organisms Microbioeroders (filamentous and unicellular algae. not including those that are dry. 1980. Amsterdam: Elsevier: 109–132.building corals produce an extracellular skeleton composed of aragonite crystals through a complex. Domes result from preferential growth of cyanobacteria as they orient themselves to take photosynthetic advantage of the daily arc of incident light. Moseley HN. DC: Smithsonian Institution Press: 455–466. sand years old. 2009). Phycologia 48: 480–491. ing to a significant extent. forms complex tunnels inside at least 29 scleractinian species in the western Pacific (Liu and Hseih 2000).like colonies. they showed that light. and Stambler N. 2009). ety of substrata (Kornmann and Sahling 1980). there is a limit to the volume of water that can be moved by such modest adjustments.calcified segment from Halimeda opuntia show. The initial benthic stage is filamentous. high rates of predation. the alternative name “fireworms” is often given to this group.0. three. sponses in the scleractinian Mycetophyllia reesi. including un. Even so. fects of environmental conditions including light. Pre. PS.reefnews. One of these (Astrosclera spp. Dimond J and Carrington E. isotopic analysis can provide a timeline for when these reefs were actively growing. careous materials and then germinate to form filaments. and when clogged. Photo courtesy of Todd LaJeunesse. Detrital and inorganic material stick to mucus they secrete.000 years. too much light. From offshore to onshore: multiple origins of shallow. High. tem described below. upright tufts less than 1 cm high (Chapter 4). Bergman B.. Taking advantage of disorder: amorphous calcium carbonate and its roles in biomineralization. fold by one or more large skeleton producers and are referred to as “frame reefs” (Riding 2002). 2005. is par. benthic foraminiferan genera in the Caribbean (Collins et al. Sclerosponge growth rate as determined by 210 Pb and 14 C chronolo. vertebrates and fishes despite their typically loose and mobile habit. Collectively. and sessile (permanently attached. defined crystal sizes. era: guide to the supraspecific classification of sponges and spongiomorphs (Porifera). Annelids and Sipunculans 7 Annelids and Sipunculans•145 ily broken when handled. These are treated as nonselective deposit feeders here. 2009). but the most common belong to the family Clionaidae (Rützler 2002a). may be thousands of years old. albeit discontinuously. branches. recent studies indicate that certain Caribbean cyanosponges are physiologically flexible. The most prominent of these is that algal calcification is directly linked to photosynthesis by the alkalizing effect of CO 2 removal. the overall effect of borers on a reef need not be negative and is best evaluated in terms of a carbonate budget. ing the skeletal environment and promoting aragonite production. Other modes of asexual reproduc. bular sponges or take up residence within their canals.Sánchez AR. fleshier Halimeda segments contain halimedatrial. roa (“far sky” in Tuamotuan). nounced (Storlazzi et al. where coverage can be very high. shak M and Tapanila L eds. times black due to the incorporation of iron oxide. 2004. In: Reaka. Karentz D. Note that filaments are branched and have bored extensively into the carbonate. anemones deprived of their clownfishes tend to alter their expan. However. Likewise. 1992.dimensional micro. Mangeveldt GW. duced within intercellular cavities and are enlarged by the movement of multiple cells over the spicular surface. Western Australia. 1. 2008) ex. ing beds. 2000). Australian Museum. Cnidaria) zonation on site. black. E.sea communities. These allelochemicals not only are used against multiple predators. can extend their branch lengths by more than 10 cm (100 mm) annually. For example. phylum Anthozoa).use pa. 1998.like structures are joined near the mouth and expand into funnels. algae produce neither seeds nor flowers. and spore for. ment. Indeed.building corals: from molec. which have multiple roles related to sexual and asexual repro. Bahamas: species composi.hydroxyapatite crystals. members of the class Cal. fungi. The deep. but not for the polychaete’s entire lifetime (Montebon and Yap 2009). Large specimens in particular add considerable three.containing minerals typical of invertebrate skeletons and vertebrate teeth (Lighte.resolution laser bathymetry: Southern Molokai. This facilitates a shift of bicarbonate to carbonate and H+ (shown by right and left green arrows). ten applied to reefs that arguably do not fall into that cat. ameter (Figure 5–11).sea octocoral Primnoa resedaeformis. The attachment and growth of algae attracts herbivores. Reef Corals and Their Allies•117 sclerodermite center are often referred to as “centers of calcification” from which the surrounding aragonite fans grow incrementally in thickness and length as “fibers” by the addition of elongated crystals or small nanocrystals of aragonite (Perrin 2003. detritus. Larkum AWD. Predation on copepods by an Alaskan cladorhizid sponge. 2007. 78•Chapter Four Vroom P and Smith CM. These flows compress the geographical extent of the marine tropics on the west coast of continents. Photo courtesy of Paul Humann. only to reject them shortly afterwards. fleshy. A different type of cement forms beneath the living reef framework after sediment has been deposited within internal reef spaces. Competition be. 2009. 1997. lythoa sp. ments that cover thousands of square kilometers across the benthos. Kornmann P and Sahling P. 1985. collapse. about 20 Mya (Hillis 2001). Others formed encrustations. endolithic microorganisms. foraminiferans are important sedimentary components. thozoans that can form aggressively growing. Amsterdam: 109–132. Pacific Science 58: 1–6. curs. but at different times after sunrise! On occasion. courtesy of Elsevier. or defensively against competitors and predators. Proceedings of the 4th International Coral Reef Symposium. aragonite will recrystallize to the more stable calcite.. Porat and Chadwick. which marks the rocky supratidal where only wave splashes and rainwater contact the surface. Geological Survey of Can. and bacterial symbionts. which also may have formed in response to subsidence and tilting of the New Caledonian platform (Figure 1–8). Anatomical and morphological complexity generally increase with thallus size. Reefs composed primarily of this species occur worldwide. Stolons not only link gastrozooids and dactylozooids. sponse to Global Change: The Last 145 Million Years. including several east of the Great Barrier Reef. but that appears to be their lower limit. granules. Fig. De Nooijer LJ. Zootaxa 1930: 56–68. as well as aerobic and anaerobic heterotrophic bacte. Colonization of crustose cor. In type 1. complanata. the more potent feeding deterrent.Williams R.g. has unique chemical and physi.energy waves favor the enhanced growth of coralline algae and other cementing agents over more rapidly growing corals. reef. but are essentially absent from those in the Atlantic. 1995. structure. Anthozoa): effects on sympatric coral and sea anemones. colum. where it is restricted to shallow cave environments on coral reefs. Hooper NA. ties several centimeters into the coral skeleton. are the easiest to measure. ample. ing organisms.localization of phycoerythrin and nitrogenase. 2006). 1988. is widespread among sponges. or production of cellulose for support. Asexual propagation in a coral reef macroalga Halimeda (Chlorophyta.Verlag: 937–977. The spatial arrange. Coral Reefs 11: 57–77. and Pok. The carbonate particles are then mobilized into the sponge’s canal system and pumped out of the osculae.Colinvaux 1980. which is a group of calcium phosphate minerals with the chemical formula Ca 5(PO 4) 3X. for ex. and Chou LM. its fitness. Clionaid demosponges are often the most visible and may be the most important single group of boring organisms on coral reefs.swept environments.Y. but then decreased again several times during subsequent gla. and typically cover 25% of the bottom.html wildlifeland@gmail. and plankton feed. Nevalainen TJ. In addition. Helmle and R. tinct. such enzymes that operate at lower activity levels. an important mechanism of asexual reproduction among certain zoanthids in which colonies subdivide into clones consisting of multiple polyps. Marine Ecology 29: 178–192. the peri. New York: Chapman and Hall: 68–95. Nonetheless. Cores taken from Galeta. Many can be further categorized as active endoliths if their physical or chemical activities directly result in the creation of tun. certain bacterial heterotrophs in the nitrifying group can produce oxidized nitrogen from ammonium in the form of nitrite (NO– 2) or nitrate (NO– 3). 127 Coral Reproduction. Meroz.. Brown BE. isms deposit both high. The extent of asexual reproduction in sponges of the genus Chondrilla (Demospongiae: Chondrosida) from the Caribbean and Brazilian coasts. Young JR and Henriksen K.associated fisheries of any tem. Cyanobacteria are prominent producers of toxic al. a term applied to this and other skeleton. Oceanography and Marine Biology: Annual Review 46: 25–63. 2005). A mixture of the three components is formed in a spe.shaped Tarawa. and Ritson.book) Library of Congress Cataloging. 2007a). These differences may reflect Hawaii’s rigorous physical environment referred to above. w = wall.482 species that have been described (Cairns 1999). temperate spe. Mexico. are scattered throughout the mesohyl and are distinct forms. depending on the family. lacks a typical diploid sporophyte phase. Baltz A. 2001. the filaments develop on Caribbean and especially on shallow Indo. 1996). dolfini et al. sponges will remodel themselves by decreasing the amount of spongin in favor of more reinforcement by spicules. University of Western Australia. mostly 250–400 m wide (Andréfouët et al. merous ducts produced by gamete. The strategy of red algal life history.g. insula. New York. and overgrowth by invasive species (Grigg 2004. Some hardy coral species and other reef. ciated with an organic matrix. Coral Reefs 2: 181–186. about 50 species have been named and confusion still exists regarding the ef. although this suspected role has yet to be clearly defined. Indeed. The stipe is less heavily calcified. 70 5 Reef Sponges. The chemistry of the test has also been an important source of environ.forming calcareous algae as described above. Photo by author. Polychaete Bioeroders In addition to constructional activities. 2008. Biomass and taxonomic richness of epilithic cyanobacteria in a tropical intertidal rocky habitat. Reef Corals and Their Allies•121 conditions (e. cocyanin by substitution of a single bond in boxed area a. duction system. vironmental conditions including energy level and rate of sedimentation. anatomy. This internal struc. Indeed. Most form on volcanic platforms that rise from depths of thousands of meters and are thus called “oceanic atolls” (distinguished from “shelf atolls” below). who worked tirelessly with me on this project. ment of the reef corals Acropora surculosa and Pocil. a time when neurons and nematocysts may begin to develop (e. The gametes are produced in baglike gametangia that form on the margins of the apical segments (Figure 4–11). which apparently restrict them to the shoreward edge of the but. see Nesse (1999). To be consis. 1 The Nature and Origins of Reefs. ered with dense populations of gorgonians along with antipatharians. 2005). The ecology of coralline algal crusts: convergent patterns and adaptive strategies. Nesse.01 0. Indo. An Introduction to the Structure and Formation of Modern Reefs •17 are completely submerged. Figure 6–10: a) Detail of a branch from the antipatharian Antipa. parchment. Eggs are expelled similarly through the canal system. Remnants of the utricles (u) survived treat. On highly energetic reefs. Bachar A. many species produce heterogametes in which. and scleractin. and plate motion to the northwest is consistent with increasing reef age. et al. gregates called batteries that give tentacles a warty ap. These low. but un. alternatively referring to them as buttress and channel systems (Goreau et al.E. (radial growth 5–20 µm yr–1 ). b) Section of the fusulinid Triticites sp.shaped sponges where they often exhibit cleaning behavior by sweeping and consuming large particles of detritus from the inhalent surfaces of their hosts (e. Inset: The skeleton is composed of tubes formed by fused calcitic sclerites. and Le Campion. Crystals are individually small needles 1–4 µm long. some branched cold. 1971. has no continuous fringing reefs. Griffiths DJ. The family Eunicidae includes a diverse group of polychaetes that range in length from reside in a range of habitats. fore the Cambrian period. 1996. doi:10. especially in HMA species.to. These dominated in both the Indo. Georgios et al.swept chan.. 2004. These are also densely laden with nematocysts and al. Correspondingly. and Dandonneau Y. puddings. fleshy algae do not provide suitable settlement sites. Bahamas. Principles of demineralization: mod. produced calcareous spicules and were placed within the Calcarea. JC. 6–20b). Rossi S. The Field Museum. but agricultural develop. tral role in explaining some important aspects of reef morphology. and when subsequent growth was re. 1998. or fused med. Indonesia). These sponge. Carpenter EJ. lular matrix production and calcium carbonate pre. times even within the same colony (references in Baker 2003. rine chemical ecology. 247 The Special Role of Scarine Fishes. Mats that are broken up by wave action or by grazing herbivores constitute a second form of production export (see below).Kudla ML. However.and outer. Kinzie RA III. this small island is a flat plateau that rises out of the ocean with vertical coral cliffs. Reef Sponges•81 over how to define a species (a problem not limited to sponges). Figure 4–5a). cur as solitary individuals rather than colonies. The jaws are mineralized with aragonite (Colbath 1986). as well as several accessory pigments. and Hough.containing vesicles. tween scleractinian reef corals: a review of mecha. 2002). there are exceptions to this high. Appendix 4) and have been important and diverse components of coral reefs and other shallow hard. Ciliary currents produced by large lateral cilia on the radioles and their pinnules during feeding gen. However.. may rival barrier reefs in length. Influence of physical characteristics of coralline turf on associated macrofaunal assemblages.g. inset courtesy of Helmut Lehnert. A budget of car. Corallinales) on grazing by the parrot. and lochs. 2007.developed fringing reefs and islands. tion and cycling of calcium carbonate in a shelf. The outermost portion. These in. 64 Turf Algae. Reitzel AM. Radiocarbon evidence for annual growth rings in the deep. Biological Bulletin 199: 68–75. spiraled radioles and are called Christmas tree worms. thereby becoming significant producers and recyclers of energy in reef environments (De Goeij et al.like form at some stage in their life cycle. Kelaher 2002). This may be an adaptation to deal with the inherent limita. and this process forms a series of semi. Grotzinger JP. 1997. 1959. stone. ISS006. brane that are unique to cyanobacteria and are integral to their photosynthetic function (e.fertilization may occur in egg. makes it more likely that reproduction will involve sibling.g. Biogenic reefs are constructed by organisms that bind. et al. black coral “meadows” have been described under such conditions from the Mediterranean Sea on pinnacles below 50 m (Bo et al. 2003. any appreciable reduction from this figure may indicate a pathological condition. These cooler waters promote upwelling. Aragonite crystals stiffen the thallus in much the way they do in the green algae Penicillus and Udotea.000 named species (Norton et al. 2009. This balance sheet. Tambutté E. part II: decalcification. alveolata oral and thoracic regions showing these features: op. Ramos V and Vasconcelos V. even though the blue Heliopora skeleton is composed of fibrous aragonite. Nature 451: 277–278. and are attached to hard substrata by one or more small and simple disc. phylogenetic analyses suggest that they originated and diversified extensively in the deep sea before some taxa invaded shallow water (Lind. it is widely appreciated that an extracellular organic matrix controls calcification in many organisms (Chapter 2). particularly on the NE and NW sides. Annual Review of Ecology and Systematics 29: 179–206. for example. borers. the rest of the phylum is included in the Medusozoa. Croix (US Virgin Islands). Hand C and Uhlinger KR. typically contain numerous cnidocytes. Members of this guild may live in areas where the total organic matter of the sediment may be only 1%–2%. e. If the gametophyte plant gives rise to hap. 1992.ocean indentations in the coastline. ally pear shaped or elongated after 1–2 days. Algal contributions to carbon production. which are often further differentiated by their foundations behind the reef crest. Cour. tories. Mariani S. giving way to other cyanobacterial taxa. that eco. Given these various categories of reef construction.g. Fragments that do not meet size and shape criteria are rejected. ———. After Eckelbarger 1976. Likewise. Marine Ecology Progress Series 21: 75–85. While most of the barrier complex Figure 5–16: Archaeocyathids typically formed connected in. away diagram showing the structure of a gorgonian branch. b) scanning electron micrograph of polyp detail with six simple tentacles sur. cer. Systematic and biogeographical patterns in the reproductive biol. 1985. especially with respect to temperature. Ecology 85: 3428–3437. Habits. gonite. 2008.7–1. The frag.5 to 10 µm (see re. To further muddy the waters. Mackenzie FT. especially in sediment between coral colonies and in rubble where they construct tubes composed of mucus and fine sedi. tical spaces of Halimeda. 2003). and long. ogy 114: 9–18. exhibits rapid growth rates and is an aggressive competitor. rated solution of calcium and carbonate (Belcher et al. Most are benthic and live on or in sand or mud. 2007). Tunnicliffe V. and survival of a tropical sea anemone (Actini.Emperor chain. Flexibility and specificity in coral. rior pair and form a common duct that opens below the mouth. Films. 28•Chapter One of this reef’s demise. There is disagreement on the mineralogy of stromatoporoids. In situ feeding and element removal in the symbiont. including the subtidal environment. Wainwright SA and Dillon JR. but the actual number is likely far Reef Cyanobacteria•41 3–1 Cyanobacterial Photosynthesis The cyanobacterial photosynthetic apparatus occurs on the thylakoids. Thus at least in some spe. 72 are atolls with lagoons.g. Proceed. Depth: 2 m.Pagès C. sion in Galloway et al. However. 1990. In the Indo. Seamounts below 200–2. W. and occur near the shoreline. Coral Reefs 2: 129–150. and Larroux C. are pro. Sciscioli M. Coral Reefs 10: 167–174. Yakovleva IM. 2006). but because of their rapid growth. Coral Reefs 23: 596–603. which may have developed along ridges associated with the northwesterly motion of the Pacific Plate. ribbean species. However. On undisturbed. Farmer MA. ———. chaetes. photosynthetic contributions to host anemone nutrition may be considerable. et al. tritional energy requirement of Pocillopora eydouxi. posed. The encrusting and heavily calcified plate. 2007). and black corals. as described below.. nitrogen in the form of ammonium (NH+ 4) or nitrate (NO– 3) is also derived from heterotrophic metabolism. and may accompany or Figure 6–32: A mass spawning “slick” covering the surface of the Great Barrier Reef. Capone DG. careous algae and their effect on the calcification process.. With 72 atolls in an area about the size of western Europe. these undefended algae may require grazing to prevent them from becoming overgrown (Steneck 1997).” Symbiotic Foraminifera Benthic foraminiferans in the family Soritidae have large porcelain. Ltd.c. high light fields. Par. a sandy or rocky area behind the reef crest called the reef flat separates the two. Helgoland Marine Research 42: 493–509. ing (e. struct calcareous tubes that contribute to the formation of reefs or reef. Navidad and La Of the many additional reef categories that have been proposed. 2008).or trilobed. Exploration for deep. Sodium chloride belongs to the cubic system in which all faces are equal and the crystal is box. vous system differs considerably among groups of cnidar. However. In others. Colonial forms sometimes occur as extensive carpets that can either grow asexually or move across the substratum. Coral Reefs 1: 125–130. and are not fused as in hexactinellids (Figure 5–4). they also attack reef coral attachment points. in which the faces are composed of sca. Photo courtesy of Ian Macintyre. drobiologia 553: 319–326. mussels. The science of using growth bands in corals (or other organisms with hard skeletons) to extract environmental information is called sclerochronology. tent larvae can delay their metamorphosis for periods of 6–8 months. as is the case of Rocas Atoll and the Abrolhos Bank off the NE coast of Brazil (Kikuchi and Leão 1997. ologist A. Calcite is the most abundant polymorph of calcium carbonate and most commonly occurs in two forms: a scalenohedron. called filiform tentacles or knob. Courtesy of microbes@ NASA (microbes. The object was to drill into the reef to look for evidence of a volcanic basement. Carbonate can also substitute for some phosphate groups and produce calcium apa. This stratification can form a three. 2007). 2003. Hiti Atoll has more clearly defined windward and leeward sides shown in d). London: Chapman and Hall. lar mechanisms of skeletal extension in axial versus encrusting growth. and Connolly SR. 1: measurements Roark EB. Meta. 2004. ed. a plant is any organism that is “chlo. Hiti.. netic drift in evolution: the changing genetic basis of vertebrate teeth. a property of water that is measured by light reduction. These processes do not appear to harm or benefit the anemone. and typi. Proceedings of the National Academy of Sciences. b) Skeletal sediments can be dominated by Halimeda segments. and many others. as described above. As sea levels rise. nization (Figure 4–1g and as described be.. 2001. Significance of Halimeda bioherms to the global carbonate budget based on a geological sediment bud. Pawlik JR. In some species of the star coral Goni. in contrast to those associated with corals and other organisms (Schönberg and Loh 2005. Biological control of mineral formation in multi. courtesy of Image Science and Analysis Labora. hence.g.com. Some unusual features of the cyto. 2005). Structure and paleoecology of Lower Cambrian reefs. 79 Groups of Sponges and Their Skeletal Structures. however. especially those who recognized the impor. while this kind of restriction may occur. but the most common is Lophelia pertusa (Figure 6–25). Chemical defenses of marine organ. Garcia. The contribution of gorgonians to the physical structure of reefs is more limited. Paleobiology 22: 152–176. or inhibit their growth allelochemically (reviewed by Birrell et al 2008). 2011)! Antipatharians can also be found in some shallow. and De. Bishop JDD and Pemberton AJ. Calcification may also be limited by carbonate availability. Two others shown in this in situ photograph are also important components. High ultraviolet light fields and salinities elevated by evaporation may add to the ecological stress and further reduce the biodiversity found in these lim. Toyofuku T.C. a name derived from the pipe. viridis) spawn in March or April (Schulze 2006). these additions typically occur through the production of blister. Shallow and more exposed areas of vigorously grow.skeleton interface in the scleractinian coral Stylophora pistillata.like inner and outer walls.pone. Reevaluating the role of crustose coralline algae in the construction of coral reefs. ward Arnold. ine Stylaster roseus. Hosts that are largely independent of their microbial associates are inhabited by about the same concentration as the sur. or complex layers called mats (Figure 3–8d). 54•Chapter Four The Chlorophyta The chlorophytes are the most diverse group of algae. and censuses of bioeroding and calcifying organisms. podia and chaetae of the thoracic and abdominal regions (Figure 7–7a). called microbialites. 2007. cron across.g. based on their ability to produce rhizoidal attachments and survive as clones (Walters and Smith 1994. lonii. mations that are required before cement production becomes complete. although they may also have a semicircu. Most tropical serpulid species are small and cryptic. 1974. as in the case of epilithic microborers. pelago and in the South China Sea. fleshier seg. tinue to grow laterally and vertically. Reef Sponges•101 Sponges in Time and Space. This silicification process is accomplished within a single sclerocyte in the case of microscleres (Figure 5–7). sea. Appendix 2b) is a good example. and Guilderson TP. probably Porolithon sp. Canadian Journal of Zoology 84: 175–194. By the mid. composed of mucilage. called megascleres. Márquez JC. Chaves LCT.sibling mating (Carlon 1999). What is her. The lagoon enclosed by the crests of these reefs occupies an area of >300. gas bladders in the genus Sargassum. Holocene sea level changes and reef development in the southwestern Indian Ocean. However. ening of the reef by borers. and many other organisms described in later chapters. Sedi. cnidarian Anthopleura elegantissima. Rasser M and Piller WE. nes as sources of protection. are inactive and progressively older (Figure 1–17b). but they vary in the degree of mineraliza. Norris JN. Geomor. 2005. and salinity. calcified forms. There is some coordinated motion in cellular sponges. Heteracinida. whose fossil record dates back to the Cretaceous period. Second. sel. Most seamounts shown are named after Japanese emperors. The reef flat may also be covered in whole or in part by sediment ranging from rubble to fine sand. where it may account for more than half of the annual nitrogen budget on coral reefs (Larkum et al. growth. and the ability of the host to change or accept new forms has become a controversial issue. they commonly spread across the ceiling of caves where they formed thin disc. carotenoids. In. a Natural History Approach. et al. Figure 6–18: a) Caribbean reef scene with characteristic gorgo. especially diatoms. b.Winter E.Living Lower Invertebrates. Holocene sponge bioherms on the western Canadian continental shelf. Belcher A M.Living Lower Invertebrates. Some sponges form asexual products called gemmules.. Different species in the same environment deposit different levels of Mg. Sclerites typically occur as three layers in the coenen. al. Clouard V and Bonneville A.Pacific species when threatened. The rise of atmospheric oxygen. Duffy et al.polarized light. tal shelves (e. Wilkinson CR. Binard N. b) The hotspot currently under the Island of Hawaii forms new islands as the Pacific plate (or the hotspot itself) moves across it in the direction shown. Caribbean Journal of Science 41: 492–507. Foster MS. damicornis and a few others produce larvae that are genetically identical to the parent. and the Role of Conservation. tinct group of sponge. Symbionts ac. Maui. Although the nuclear region of the nerve cell is often associated with mesoglea. Removal from the anemone typically results in the death of the resident clownfish by predatory fishes. Although >90% of modern sponges produce skeletal elements composed of siliceous minerals. especially pavement. et al. cific. Guam). Bahamas. clude the fringing reefs of Mauritius and Réunion in the Literature Cited Adey WH. ample. gins by stretching and invagination. FEMS Microbiology Ecology 55: 167–177. trix. >2. Schmidt H. niles. Fichez R. which can be buffed and polished with jeweler’s rouge. sue growth (Hunte et al. ous epizoic invertebrates and provide shelter for fishes. These are located in the pinnules and continue down the center of each radiole toward the mouth. whereas carbonate and CO 2 com. parallel flaments that form closely packed bundles. USA 361–365. Knoll AH. The Growth of the Coral Skeleton The specific form that a colony takes is a combined function of genetic and environmental influences. settlement may be driven by many other factors. ploy jaws to procure food. These include well. grass meadows. vertebrate hosts. Modern Foraminifera. thology II. sharp. MacGeachy 1977. nus. roalgae. trix. 18•Chapter One some reefs.Guldberg O. In some habitats. Kasting JF and Ono S. Usher KM. where peak yields once exceeded 300–400 metric tons per year. Harrison PL. and conforms to the shape of the skeleton in a hand. 2006. of intrinsic adherence of cnidae discharged from the tentacles of the sea anemone Aiptasia pallida. tion of larvae in the water column occurs in some groups.based nutrient inputs. pro. Figure 5–9: a) The orange encrusting sponge Mycale laevis is often found on the undersides of Montastraea annularis in the Caribbean region. but are less able to withstand physical stress or intense intraspecific competition (Francis 1979). Mineral deposition in organisms is typically asso. Hart DE and Kench PS. Journal of Natural History 37: 1–47. Kennedy JA. whose relationship with their symbionts will be more fully ex. Inverse rela. sponges are the largest source of unique chemicals that have been obtained from marine organisms (Pawlik 2003). 1972. ing systems in which planulae may be sexually produced.5 m high. This ability may account for the widespread occurrence of this species across the Indo. et al. 193 Brachiopods.water reef communities may also form on preexisting rocky substrata without the assis. Produc. the relationships among them do not suggest a common ancestry. Rowland SM. 2000. grasses. ally solitary. Junceela fragilis.g. a type of commensal symbiosis in which organisms live inside another. lophophorates (Chapter 9). Marine Ecology Progress Series 232: 141–148. Some au. deed. Davies. Photo courtesy of Kayley Usher. the site of min. McMurray et al. as well as those of the medulla that may function in the transport of organic and inorganic material. or on beachrock in northeast Brazil (Maida and Ferreira 1997).water gorgonian Primnoa pacifica. create a distinct habitat. The multinucleate state. 2003). Fjords in New Zea. tal nucleation. Antipatharians continue to be used as jewelry today. 4–8c). Much smaller intertidal stromatolites have been described from other hypersaline environments. The largest colonies may be hun. To accomplish this. Halimeda contribution to organic and inorganic production in a Tahitian reef system. Biological Bulletin 180: 406–415. ———. structures. 2006. processes like these may be responsible for the delayed formation of the millimeter. or other energy expenses (Sorokin 1995).known form of biologically induced carbonate is that produced by certain bacteria. apparently by erosion. ers. pit. These may include starch storage. Proceedings of the 8th International Coral Reef Symposium. • Provision of preferred settling sites for certain corals. Coral Reefs. Aka coralliphagum. Journal of Sedimentary Research 51: 475–478. With some exceptions. even within the same genus. ceedings of the National Academy of Sciences. Boulder: Geo. ate skeletons may contain some or even a lot of aragonite. but is not dependent upon it. Si. cal chambers and external walls thereby become more al. 2008. Likewise. mon symbionts.000 kg and 2. Zooxanthellae and Coral Nutrition While there is no doubt that zooxanthellae con. permeable barrier to the diffusion of molecules and ions including calcium (Tambutté et al. it flows through the choanocyte collar. 2011). As the colony develops. Carbonate dep. fishes. On occasion. corporated into the reef matrix (Hart and Kench 2007). The barrier reef crest is often capped by a thick pavement. and thus the living remnants of these ancient groups can help interpret the conditions under which they and their fossil associates evolved (reviewed by Beavington. and may be released as planulae in less than 24 hours (Kahng et al. dites (Harrison 2011). 2005).7 mostly macrophytic chl a. Figure 7–11: a) The rock. The thallus is typically composed of uninucleate cells separated by large plug. Cell kinetics of the marine sponge Halisarca caerulea reveal rapid cell turnover and shedding.harvesting pig. Gulf of California) and by direct extraction by dredging for agricultural purposes (e. but instead is a series of about 2. et al. P. these can be seen with the unaided eye after the corallum is sectioned. and cementation) within the reef’s internal spaces. a barrier of sand and seagrass restricts tidal flushing to small channels.. tems. Journal of Volcanology and Geothermal Research 55: 239–260. There are two general contributors to turbidity. formation of coelenteron and skeleton Sexually mature adult Sexually mature adult Colony begins asexual growth g Figure 6–34: Broadcast. Proceedings of the 5th International Coral Reef Symposium. 1993). Willis BL. antipatharians have been valued throughout antiquity and were worn not only as jewelry by the ancient Greeks and Arabs. corals.concentrating mecha. This characteristic can vary within members of the same genus.000 species that have been described. 2003.6 (paperback : alkaline paper) — ISBN 978. stylasterine corals. and species (Tunnicliffe 1983. Baker AC. Goodkin NF. Preferential reef accretion oc. Proceedings of the 8th International Coral Reef Symposium. reduced predation. Montebon and Yap 2009). which in many groups was composed of similar polygonal partitions that often became filled with solid skeletal material (Wood 1990). and other characteristics. Chemical defense and chemical variation in some tropical Pacific spe. While many species produce only unspecialized vegetative cells. ducted directly to the mouth and are swallowed. Bio. It has been suggested that phosphate uptake is enhanced by the rhizoids of calci. Hendler 1984). 2007. they can only contract onto it. sium. but the capitulum is thin and cup or fan. Evolution of mod. Hill MS and Hill AL. ———. cobble (Appendix 1). and are often better devel. Perrin C. Detritus feed. The highest point of growth. The gastrozooid tentacles are also unlike those of milleporids and are produced as elongated fili. and a basal hypothallus. Spawning of the giant barrel sponge Xestospongia muta in Belize. roalgae. and provide habitat for numerous organisms. Often called “palolo worms” after the Samoan name for them. provides refuge for other organ. The muscular pharynx has one or more teeth that pierce its prey and then suck out its contents.fixing agents and are prominent on reefs where nutrient. However. duction described below. Herren et al.g. and Smith SW. mium are responsible for giving the Red Sea its name. and Bak RPM. spongeliae above. Pennings SC. tality.Jene L. 1998) and fertilization. and caves (e. such sponges serve a protective role.shaped branches (order Sti.like structures. which may occur in the form of small discs. cally in Chapter 15).5 cm3 of volume to pump. often forming three distinct lay. is extremely hard and dense. ducing). Purverel S. Indeed. Some Z. Thus. Modern coralline algae play several important roles in the coral reef community including the maintenance of open space. and Freiwald A. Snow MG. nov. ed. Steger D. These cements constitute the mortar in the simplified brick. Stat M. Treatise on Invertebrate Paleon. The ridge may be flat or grooved with surge channels. As the larva begins metamorphosis. Predation and Sponge Chemistry Sponges are prominent in many reef habitats. and key species with high growth potential have been among the more important factors in determining reef growth during Holocene time. a large sponge with a volume of 0. tocorallia: Holaxonia). 2003. Invaginations of the cell membrane form the thylakoids. honeycomb. but more typically contain proteins with elevated levels of copper. 2007). van Duyl FC. Loya Y and Sakai K. a double pyramid (a dipyramid) is formed.g. Pires DO and Castro CB. the embryonic stage is transformed to a cili. In: Jones OA and Endean R. Coccolithophores: From Molecular Processes to Global Impact. ———. calcareous remains of these chlorophytes for long periods. ern strategies for the isolation of organic frameworks. At its inner end (Hamelin Pool). Photo by author. 2009). even though the living tissue is typically less than a millimeter thick and is restricted to the outermost surface of the colony. ———. London: University of Manchester Press. The surface to area ratio in this sponge is almost 2:1.laden. lids provide a reef framework by extending their syncy. Stalvies C. the Stylasteridae. they serve as perches for certain mobile fishes and invertebrates and provide shelter for numerous others (e. Some species.Horani et al. The dis. The most recent of these cycles occurred between 30 and 18 kya when an estimated 50x106 km3 of seawater was withdrawn into ice sheets and sea levels dropped once more (e. Nonetheless. 2005. haploid stages of the isomorphic brown alga Dictyota ciliolata (Phaeophyta). Ecosystems of the World.building organisms invites weak. crys. The exploita.1 mm) dactylopores mark. generally tend to be much smaller and narrower compared with shallow. have been called “fora. tidis 1999).water corals) are quite complex and diverse. epilithic species that occurs ubiquitously. northern Red Sea) have more than 220 coral species of stony and soft corals and are among the most popular diving sites in the world (Fishelson 1980). but some can produce dense clusters on dimly lit deep reefs. a 200 km. M. tion as if they were “metabolic kidneys. rano et al. Population struc. high molecular. bate concerning the biological origin of these earliest (Figure 3–10) produced by endolithic cyanobacteria alone or in symbiotic combination with fungi. 1982. nies. Pearse (1932) put it. Some of these fossils date back more than 3. thozoans have shown that carbon fixed during photosyn. gest that they are even older (Li et al. any object that deflects or otherwise impedes water flow forms a “baffle. This characteristic is sometimes called the “Phoe. sometimes with large blocks falling away. Each trichome may have its own sheath (Figure 3–8b). ety Island chain develops some spectacular examples of such reefs in close proximity to each other (Figure 1–18). 1985). goreauii the macrogametes are twice the size of the microgametes. thus deterring it from consuming polyps around Spirobranchus tubes (DeVantier et al. Nonetheless. see Drake et al. These are responsible for the brown color shown in the center of the test. trol of intracellular algae (= zooxanthellae) in hard corals. and Rob. These structures display a variety of forms related to en. a type of retrospective analysis. Coyer JA. These consti. 2005. and the two groups generally share the same categories of cnidae (Pires and Castro 1997). However. Vila. this process is completed in 2–4 weeks. Excellent examples of ancient stro. 2008a. Sedimentology 45: 39–51. There are several crystal forms within the hexagonal system that may display rhombohedral. theless. Rees SA.com. 1996).352 snapping shrimp of the (Cimino and Ghiselin 1999). Bavestrello G. Because these traits are unique among animals. clude the sea whips. vironments and when found in place as a fossil it may be a good indicator of sea level (e.water environment. more informally called forams or foraminiferans. Allemand et al. Mundy CN.g. Terpios hoshinota (Suberitidae. This source of carbonate can be substantial. Symbiont.25216. and are closely associated with chlorophyll in the thylakoids. reinforced connective tissue: stiffness. giganteus may persist for decades (Nishi and Nishihara 1996). 1982) and has been used to date the position of its shallow reefs. and are probably better considered as bio. It is not always obvious where to draw the line between epilithic and endolithic groups. While turf algae are generally considered conducive and desirable for coral settlement. The mineralized radu. vices of coral reef ecosystems. rophyll cannot utilize. they also wrap around prey internally and cause tissue lysis that facilitates digestion (Schlesinger et al.Canadian Sponge Reef Project: Kim Conway. 1999). dopodia that extend from the cell membrane and may function with or independently of the microvilli (Leys and Eerkes. increased sedimentation. Chamberlain YM. 2008. which grew during the Permian in what is now Texas and New Mexico (Chapter 13). and M. 2002. Between these are six shorter secondary septa that form the second cycle (Figure 6–12). a group of continental islands located farther offshore near 29°S (Spalding et al. Calcification in Halimeda is driven by chemical changes that occur within intercellular spaces dur. cately branched corallum composed of aragonite. De Almelda EVR. zooxanthellae are sepa. Dispersal strategies in sponge larvae: integrating the life history of larvae and the hydrologic component. Facies 38: 89–102. Labels: c = columella. This siliceous reef belt was 7. grams of carbonate m–2 yr–1 ). Marine Chemistry 108: 96–108. including the brilliant orange. Clinton BD. fall into the channels. while those below appear fused with indistinct boundaries (double ar. 2009. ery square centimeter of surface there is only ~0. Spalding MD. or encrust. Ball EE. Comparative study of the chemis. Under somewhat less rigorous conditions. developed reefs and broad areas favorable for lagoon circulation. proximate. thus blurring the line between the two groups. Some spe. 268 Reefs of the Mesozoic Era. ized structures including pores through which water is expelled during rapid contraction. and are often reasonably well preserved because they are mineralized. 2010). University of California. including those of Central America. Buds may occur on the column. there are other more complex factors needed to account for the great vertical displacements that accompany conversion of volcanoes into atolls with carbonate caps over 1. The chloroplast genome in this group suggests a cyanobacte. 1990.200 individuals per square meter are found in some large coral skeletons in Belize. and Avoidance. the polyps with 8 pinnate tentacles clearly place this species among the Octocorallia. peptides. Journal of Experi. Scott et al. Brugler MR. indicating the ability to feed (e. reduced light and low salinity) to the development of coral reefs. How. Marine Ecology Progress Series 277: 95–105. tion. Why overgrowth of intertidal encrusting algae does not always cause competitive exclusion.g. these function as light. ing species there is also considerably greater diversity among stylasterines.g. Indeed. 1997b. Phycoerythrin is particularly efficient in deeper. collectively referred to as coenenchyme. Calcification and photosynthesis in reef. These are called facultative symbionts. especially coral reefs. 40 Rock and Reef. This may be an adaptation to storms that harm these reef. Scientia Marina 68: 137–144. E. A worldwide. doi:10. velopment occurs away from the shelf edge. experiments have suggested that spicules alone are not as much of a deterrent to predation as the large number of bioactive compounds that many sponges produce (reviewed by Paul et al. mation as chloroplasts develop within them. cies have been used to map past distributions of the tropics. French Polynesia. tum to become three.like aggregations (Häussermann and Försterra 2007). Some of them. ous or chitinous operculum. Other clionaids. Clone Wars Densely packed clones can dominate large areas. as are mats and filaments on and under sand. Risk et al.Johnson Space Center ISS005. are con. ogy of glass sponges. 2007. Stat et al. Mi. tinental shelves and are distinguished as “shelf atolls. These include macroalgae. Fragmentation may be a major mechanism for re. Meinesz A. rated into a string composed of protein and mucus that is molded into place at the top of the tube by the folds of a thoracic collar.2 mm in diameter.Alsumard 1999). About 2. 114: 309–345. A flattened oral disc separates the tentacles from a centrally located mouth. which acts as a barrier to water flow from the Pacific to the Indian Ocean (Godfrey and Golding 1981). and the Ca. Rhodophyta). depending upon the physical environment. struction of their tissues. Applied and Environmental Microbiology 70: 3724–3732. 2004). only the predominant calci.specific bacteria grown in culture (Taylor et al. and Dinger NE. coralline algae can be one of the most important contributors of carbonate sand. ral Product Reports 21: 519–538. 1977. 2c. et al.magnesium calcite in different parts of the shells or skeletons they produce through a common point. LaJeunesse TC. the tissue is actively photosynthetic. with the addition of perhaps one or more segments on each thallus branch. Lobban CS and Harrison PJ. reinforcement by calci. Interestingly. tentacle. and Ilan M. Munk WH and Sargent MC. the siphonoglyphs. 2007. and variously shaped atolls. Bricker I. it also stopped grow. Implications in resource manage. Branched spe. Benthic lar.. grasses. a feeding polyp that is short and tubular and possesses a clearly defined digestive cavity. served as it was in eastern New Guinea was subjected to Figure 1–19: Large sections of coral reefs were uplifted >1 meter following the March 2005 earthquake near the Indonesian island of Simeuleu. they are most common on Indo. 2008. Mohamed et al. and Biogeography of Coral Reefs. This in turn drives the synthesis of ATP. Weiss VM.E. many display highly variable physical characteristics including color. Richmond and Hunter 1990). Characterization of depositional environments in isolated carbonate platforms based on benthic foraminifera. New York: Blackwell Publishing: 167–182. The thallus develops clearly defined rhizoids (Figure 4–12c) and a stipe. aragonite is precipitated from saturated solutions when CO 2 is removed during photo. Likewise. 2005). By increasing the sponge radius to 4. whereas volume increases cubi. tural features of the animal and exhibit longer biologi. scavengers. soluble form of silica in the oceans. Goh et al. The exploita. the preda. Australia’s largest near. H. Loss of coralline algal populations has been attributed to human activities including reductions in herbivore populations. 1°S. fusulinids were producing tests that were distinctively elongated and shaped like rice grains (Figure 4–21 a.5 My Hawaiian Archipelago p la te H aw aiian rid Oahu 3 D ire ctio n o f p la te m o v e m e n t Hawaiian Archipelago E m p e r o r S e a m o u n t s d b e rr o E m p e o r SS e Ha Ha Ha Ha Ha Ha Ha Ha Ha Ha Ha Ha Ha Ha Haaawa wa wa wa wa wa wa wa wa wa wa wa wa wa waiiiiiiiiiiiiiiiiiian an an an an an an an an an an aaa Hawaiian ooooooooooooo AAAr Ar Ar Ar Ar Ar Ar Ar Ar Ar Ar Ar Ar Ar Ar AAA hhhchchchchchchchchchchchchchhhipipipipipipipipipipipipipipipipppeel el el el el el el el el el el eee ag ag ag ag ag ag ag ag ag ag ag ag ag agg Archipelag c a Meiji 85 My Suiko 65 My Ojin 55 My e aa m o u n tt s oo s Ojin Ojin Ojin 55 My Koko 48 My Yuryaku 43 My Midway 27 My Nihoa 7 My 24•Chapter One Reefs that reflect a complicated geological history sur. 2004). where the first crystal layers are oriented with their c axes par. Bulletin of Marine Science 65: 227–234. eroders and contribute to sediment production as well as the creation of galleries within reef substrata. This zone is often dissected into alternating ridges and sand channels that are more or less perpendicular to shore and is referred to as a spur and groove topography (Figure 1–5c). Matsumoto AK.Johnson Space Center STS050–72–42 and Reefbase. millepore. they are essentially silici. There are also double atolls with a small lagoon inside a larger one. Bleaching and death of reef corals on the Pacific side of Panama. Long competency periods clearly could facilitate transport by currents over long distances. Indonesia).20b). Current trends of space occupation by encrusting excavating sponges on Caribbean reefs. eastern Indian Ocean. When the chaetae penetrate the skin.Associated Cyanobacteria. brown. Another form of asexual reproduction among anemo. suggesting that bicarbonate (or CO 2) could be converted to carbonate at the mineralization site enzy. and Tambutté E. Diversity. Invertebrate community associated with the macroalga Halimeda kanaloana meadow in Maui. or shade established corals. Holbrook SJ and Schmitt RJ. Coral Reefs 6: 139–148. ———. Indeed. and provide significant habitat for many organisms. Examples of these include deep reefs on the islands of Barbados (Eastern Caribbean) at . displacement of sea level by tectonic activity. 2000).000 km2 (Hop. with exceptions. channels.Alsumard et al. or form heavily calcified plates or encrustations.R318. the most common reef. the Andros reef has been catego.cleansing and herbivory (Steneck 1986). is composed of primarily of densely packed Ca. and slopes gradually to a depth of 30 meters or more where it breaks up into rubble and sand. Indeed. Shinn EA. some with biomedical applica. 2007). Jour. These are the bricks in the brick. simple filamentous and branched chains. and 5 fish. or be released by the grazing activities of animals.containing eggs of the corals Pocillopora verrucosa and P. Pa. cal marine environments is affected by several biologi. are generally sensitive to chemical defenses. or both. fer cleaning services. growth may be size dependent (e. ies in Geology: 4: 53–66. Thus far. I am grateful to Christie Henry. tocorallia. Contrasting effects of turf algae on corals: massive Porites spp. Riegl B and Dodge RE.harvesting an. ———. and have by far the simplest structure and the smallest number of cell types (perhaps only 11) of any metazoan (Degnan et al. 2004.bodied surface. Photo by author. 2009. In Hooper JNA and Van Soest RWM. create repeated. Science 275: 1080–1081. Type 4 matrices initiate and regulate the growth.reef endolith that is an opportunist. ate production and loss over time. pora palmata (Lamarck 1816): a scanning electron microscope (SEM) comparison demonstrating simi. 2005a. and more than a few contain zooxanthellae. Several colonial forms can arise from this process. 2005). Their formation is poorly understood. These are produced from multiple calcitic units. and they may form structurally simple. other serpulids create reef. fied (coralline) sponges (Ten Hove 1989). Bentov S. cally distinct populations that inhabit specific groups of hosts (e. Kuhl M. stony corals. ers and. and movement to a new location where the sequence of events is repeated. New York: Springer Science and Business Media: 489–517. portions of the Tonga. tic mystery and at one time embraced many groups of organisms that exhibited calcification. and thus serve as respira. The Amazon River sys. Milliman JD. catch up.carbon conjugated alcohol side chain referred to as a phytol (Figure 3. matu. Microbial mats in Aus. netically identical polyps (i.. By far the most common among scleractinians is broadcast spawning. branched spiral or straight colonies called “wire corals. whereas others have stressed the function of bacterial membranes as sites for nucleation or calcium enrichment (Knorre and Krumbein 2000). et al. water endocytosis in the biomineralization process in calcareous foraminifera. tors are generalists. Proceedings of the 6th International Coral Reef Symposium.species turfs. whiter portion is devoid of symbi. each 17 mm long. Faure G.. Buddemeier RW and Smith SV. Marine biodiversity and the medicine cabinet: the status of new drugs from marine organ. ages of now volcanically inactive islands are shown in millions of years. However. tures. when photosynthesis is inhibited. the availability of sediment. The larger ones. In some cases atolls have been elevated tens of meters.Johnson Space Center. tastraea cavernosa. and in the Western At. Macintyre IG. and are minor components of most reefs. Duke University Laurie Richardson. Rate and extent of decline in Corallium (pink and red coral) populations: existing data meet the requirements for a CITES Appendix II listing. 2007. In: Rützler K. These animals are common on coral reefs but are typically found in sediment rather than on hard substrata. Demes KW. an Evolutionary Perspec. Tube construction is initi. Goldberg WM. which typi. Proceedings of the American Academy of Arts and Sciences 51: 155–251. Caribbean Journal of Science 39: 348–370. On the euendolithic genus Solentia Er.000 m are often cov. Some species directly consume various types of corals. ing life. for example. lariid species form reefs. ticulate feeders. Several generations live their lives completely within its canals. Hess WR. especially needles that stiffen the thallus in much the same way they do in Penicillus and Udotea. phyco. and atolls (Aldabra in the Seychelles Archipelago. Peyrot. Raz S. These corals brood their embryos as described in Chapter 5.H and are often specific for particular hosts. While most plants and invertebrates that secrete car. Modified from Hillis. including invertebrate larvae and small crustaceans. Zea S and Weil E. Lirman D and Biber P. tially or completely buried in sediment. Andersson et al. synthetic activity by these symbionts may not be confined to the host surface (Brümmer et al. segment reefs derived from algal meadows (Rid. vide for the attachment of other invertebrates or. brate bone and mollusc shells are formed this way and are among the best. caribaeorum. and the Alcyonacea. Pérez CD. but larger ones may be the product of multicellular fusion (Uriz et al. Because sponges can pump their own volume every 5–20 seconds (Reiswig 1974). Med. Suzuki T. Geology 34: 773–776. and Lauter. This species is thought to stabilize beaches from erosion wherever extensive reef development oc. and hot springs.water reef growth >1. although the opening may collapse and provide some protection from predators.g. The attachment to protein is not shown. man activities such as poor land management practices (Chapter 15).OH. tein substratum temporary. Respiration and symbiont photo. podia. b) close.mortar schematic of reef con. esis. yellow. nian corals (sea whips and fans). whereas some abandon their tubes at intervals and construct new ones. in turn. Rapidly rising sea levels.000 species are known. 2008. The gametophyte is leafy. 2006. thus re. pletely carpeted with their calcareous tubes (Kirkwood and Burton 1988). Reproduc. Zhou Y. 2007. et al. Closely related species may release gametes on the same morn. Griffiths H. Andrews JC. tion. and binding activities. However. fill lagoons. pounds peculiar to the algal gen. Cladophora. an. systems? Photosynthesis Research 49: 103–118.13: 978. Hexac. Wiens HJ. ogy 16: R316. gesting that tube construction occurs in concert with tis.. and may create a wave barrier 4 m or more above sea level. Spheciospongia vesparium. Rawson PF. 25. However. and thousands of years.8. cordingly. The spread of this species is rapid and persistent. fleshy stalks that form an operculum (Figure 7–7a). 2007. Vol. Shpigel M. 1994.calcite formation (see Wasylenki et al. each of which has a defined set of angles between them. one of the more common is the bank reef.. Energy integration between the solitary polyps of the clonal coral Lobophyllia corymbosa. Marine Biology 145: 1097–1106. and Loya Y.. Figure 1–14a). Littler and Littler 1999). fied.nasa. there is a strong positive relationship between lagoon depth and atoll perimeter (Purdy and Winterer 2001). Turner SJ. A large frame builder with a flattened. Silica de. 352 Acclimatization. turf algae as a group per. The origin of these compounds has been traced to their cyanobacte.g. ous and characteristic bright yellow oscula. all sabellariids that have been examined appear to broadcast spawn. After Barnes DJ and Chalker BE. 1983. Such coral forests allow for the presence of numer. Washington. Caribbean staghorn coral popula. and is expressed as a function of time and area (e. den Banks at depths below 30 m (Minnery 1990). are a significant factor in red algal groups that calcify. and these are taken up by zooxanthellae as substrates for the produc. Right. Others have found that the number and type of sponge guests vary according to sponge morphology. and Dullo W. b) Members of the family Primnoidae (genus Primnoa) are common in deep water and often serve as habitat in the absence of reef formation. Hydrobio. ed. Sea level change through the last glacial cycle. and the following three unspecialized segments. drozoa. Gaino E. while the cloud sponge Aphrocallistes vastus forms large shield. sealing surfaces. Blackburn A. gonian octocoral. and the Persian Gulf. and other unicellular algae that will not be considered here. The Caribbean “species” is now referred Polychaetes in the family Serpulidae comprise more than 400 species that are common members of coral reef and reef communities in all oceans. These structures are composed of 2–3 protein. Indeed. Scyphozoa. Journal of Sedimentary Research A65: 143–153. Sea of Cortez.Pichel 2006).red foram Homotrema rubrum is one of these. 2003). Canadian Jour. crabs. In general. attacking and killing com. corded from Lophelia communities in the NW Atlantic (Roberts et al. With about 250 liv. sia.called organ pipe coral. Storlazzi CD. Seaward of this high point. ysis Laboratory. The pharynx often contains a pair of siphonoglyphs. Glasby et al. The role of sexual and asexual reproduction in structuring high. and amounts of carbonate they produce. bo. as a soil conditioner in Brazil). Coral Reefs 19: 303–309. but are more abundant on those in the western Atlantic where there may be as many as 25 colonies per square meter. Unfortunately for simple definitions. Science 312: 543–547. 2002. Coral Reefs of the World (a. territories on a coral reef. Kelly K. Proceedings of the National Academy of Sciences. In some cases the outer fore reef lacks a but.” binding corals together. Some Atlantic reefs are composed primarily of coralline algae. all black corals ap. The University of Chicago Press. Marine Biology 35: 253–267. 1997. all con. ness represents the degree of colony integration. the Great Siberian Barrier Complex. This curious arrangement appears to be a function of the meristematic cells at the outer edge of the thallus. changes the dynamics of flow. width. Cements may form deeper 4•Chapter One sue. vironments. the hosts often die if the symbiosis can. and early lithifi. biologically controlled mineralization is a highly regulated process distinguished by reproducible and species. Soft Corals and Sea Fans. Framework construction may rely on oysters. Lee D and Carpenter SJ. 257 Benthic Predatory Fishes: Their Influence on Corals and Coral Reefs. Epithallial shedding may be sufficient and frequent enough to prevent epiphytic growth on the surface of many crustose corallines (Ke. Scoffin 1992). later grew explosively and were thereby able to “catch up” with rising sea levels (Neumann and Macintyre 1985). In: Birkeland C. even in the oldest segments (Macintyre and Reid 1995). win in 1842. a species composed of multiple small tubes. Heliopora. the main topic of this chapter.edge reefs off Little Bahama Bank flourished. showing barrier reef with windward and leeward sides: L. 2007. thereby increasing the surface area exposed to sunlight. nutrient distribu. whereas the other two are small coccoid unicells (ap. Van Kempen TMG. Primnoa resedaefor. This enzyme has a dual function: (1) it participates in active transport of seawater calcium to the site of calcification. Wirshing HH. a polymer of amino sugars unique to bacteria. 2005. perhaps 4 cm yr–1 or more across carbonate surfaces. This enzyme converts abundant but inert molecular nitrogen (N 2) into the metabolically useful form of ammonia. dros Island in the Bahamas. Brazil. the complex. Ryukyu Islands.specific crystal properties. Reaka. Coral Reefs of the Indian Ocean. ed. Annual Review of Ecology and Systematics 40: 551–571. such environments in.living rhodoliths may grow individually to several cm in di. So far 100 host species. 1991. Modified from Bayer 1956. and Obura DO. Kennedy DM and Woodroffe CD. 8°N. the delicate branch patterns and slowly growing skeletons limit the role of stylasterine corals in reef building. Courtesy of the Image Science & Anal. French Polynesia. dition.ring compounds typi. ria” in marine sponges.water reefs. These growth bands have been noted as well in fossil corals and have been used to estimate growth rates and paleoclimatic conditions. trons. Sánchez JA. this book is dedicated to my wife and best friend. and are assigned to the Kingdom Protista by default. lumbia.defined relationships between the crystals forming the shell and the components of the ma. and finely divided gills (branchiae) containing extensions of the circulatory system also occur on the head of terebel. Physical stress and biological control regulate the producer.diversity communities of reef. thereby supporting a diverse and unique community of benthic invertebrates (e. BMC Evolutionary Biology 7: 90. Interspecific compe. Some also associate with unicellular rhodophytes. Halimeda sedi. germinate. These may extend through the mouth or body wall where they can envelop prey and initi. Hundreds of species have been de. nal of Zoology 80: 1735–1754. 1981. tabolites differentially affect coral symbiosis in situ. Photo courtesy of Paul Humann. Some nudibranchs have become extreme specialists that prey upon a single sponge spe.fixing bacteria. Noé SU.000 liters per hour.associated microorganisms: evolution. Marine Geology 59: 135–164. Natural Products Reports 23: 153–180. Kramer PA. cally (v3 ). are still referred to as pro. The two generations usually can. Mullins HT.1371/journal. and Moosa MK. Penn State University Craig Layman.rayed microscleres (not shown) displayed by this group (Uriz 2006). Marine Ecology Progress Series 33:255–264. Coral Reefs 19: 1–23. ats et al. fringing reefs behind the New Caledonian barrier reef (Cabioch et al. 1983.. The first aragonite crystals form on the filament walls (Borowitzka and Larkum 1977) as randomly oriented “needles” 0. The fate of specific coral reefs under these conditions was dependent on multiple factors. a layer of sediment 200 µm (0. Proceedings of the 5th International Coral Reef Symposium. 296 Different Types of Diversity. rifera: a guide to the classification of sponges. Endolithic cyanobacteria or algae. A close. B. 1996. are sticky. and are anchored by filamentous rhizoids that may reach sev. bathed on the outside by the foaming breakers of the ocean. 2006. Golubic S.water reefs to a depth of >60 m given proper water transparency and other con. Riul et al. although the actual stress threshold depends upon exposure time. In some sponge taxa. a windward fringing reef off the Caribbean side of Panama. Erwin and Thacker 2007). but form a landward edge of reef rubble (shingle). Flo. assum.Hill. b. protein. Freiwald A. some demo. Figure 14–5). Specific clades are often associated with defined host groups. productive cycle. Castanaro J. This group is placed within the Stolonifera. but these are often found only on the underside of rocks or in small cavities within reefs. In the meantime. 1974. Macintyre IG. calcareous skin that serves to veneer the carbonate surfaces produced by other carbonate. and Freiwald A. Calcareous sediments derived from algae.75 cm in diameter. Journal of Phycology 42: 292–303.celled rhizophytic algae (Bryopsidales. Marine Biology 154: 65–80. 2008). These Halimeda bio.g. Thus the term “sponge colony” is a mis. warm. but not all. cific appear to be unique. scribed below. endolithic microborers are found primarily be. tion of carbonate by organisms. ties (Wielgus et al. Halimeda.gob. Sharp JH. Steneck et al. or so it seemed. cology 34: 417–426. et al. 2007. forming a cemented. ment attached to rocks or shelly material (Rouse and Pleijel 2001). Kuffner IB and Paul VJ. Annelids and Sipunculans•153 matophores) that are typically placed directly on the female. and Vaulot D. The Exuma Islands of the Bahamas. and then produces a segmented. The predominant winds oscillate from the NE during the summer. 1998). 2001. times wrapping themselves around spicule bundles. Interaction between benthic algae (Lyngbya bouil. Glynn PW. part of which is shown at lower left (double arrow). ure 6–33) exhibit regional differences that may dictate whether the gametes of two populations are compatible (Harrison and Wallace 1990.Caribbean region (Figure 6–4b). diversity. 2007. scleractinians. Mayotte in the southwest In. Fiordland. Structure and composition of organic reefs and carbonate mud mounds: concepts and cat.up showing extrusion of egg/sperm bundles through the mouth. gastropods. and bio. bling a bottlebrush.g. hydrated silicon dioxide (SiO 2. The relationship between reef fish community structure and environ. strates including corals. and other objets d’art (Fig. a trait typically associated with eu. New York: Chapman and Hall. and construct a characteristic clear tube. Living H. More than 100 countries have coral reef coastlines upon which tens of millions of people depend. 2001.) refer to those with dimensions of less than 1 µm (10–9 m). Marine Ornamental Species: Collection. pertusa within a larger area referred to as the Stetson Reefs (Reed et al. bonate as MgCO 3. suggesting that exchange may be relatively rapid. Coral Reefs 25: 7–18. and on the inside surrounding a calm expanse of water. Photos courtesy of Paul Humann. However. calcified sponges are most closely allied with modern bath sponges. Proceed. As described by Barnes and Crossland (1980). a process referred to as vertical transmission. turbulent. located in the northwest of the Tuamotu Archipelago (Figure 1–11). Indeed. search 34: 115–122. Palola viridis spawns several days after the full moon in October–December (spring–early summer in the southern hemisphere) for two or three consecutive nights. cilia then sort and separate the two. and Gautret P. Florida International University. ii.g. Photo by author. Protist 156: 399–412.shaped monax. In addition to sexually produced planulae. Contributions of Crustose Corallines to Reef Structure Organisms resembling modern reef. showing the develop. Polyps in this group (order Actiniaria) range in size from a few mil.. 2008).ATPase.rich water may occur through the process of coastal upwelling as relatively deep water is brought toward the surface. b) A large stand of Acropora palmata in growth position from the Pleistocene of Curacao is about 2 meters high. a calcaxonian gorgonian with calcified rings in the axial skeleton. Johnson and Mann 1986). Anti. Stolarski J. doi:10. 12th Caribbean Geolog. carotenes. “an. Box. ated planula larva (Figure 6–34a) with a distinct anterior and posterior end defined by its swimming direction. Collins LB. Likewise. c) Reef building is a net value of the processes of reef construction. and colony. frame builder. Marine Ecology Progress Series 239: 105–114. expanded base. Reef growth. In colonial hexacorals (and other anthozoans). Grube A. ment. Certain herbivorous CHO CHO H H H OAc OAcCHO OAc AcO OAc CHO OAc Udoteal Halimedatrial tetraacetate Halimedatrial Figure 4–9: Terpenoid com.. These conditions exclude most organisms except stromatolitic cyanobacteria and associated microflora (Skyring and Bauld 1990). and Le Campion. 1997a. ammonium and nitrate are potentially toxic waste products and when they are taken up by zooxanthellae. sometimes called a coenosteum by hydrozoan specialists (but the term “corallum” will be used here—see below). pear to function in adhesion. certain parasitic barnacles. Cairns 2001).support. Sponges: an essential component of Caribbean reefs. but they also can become fragmented into separate colo. spirals.West Pacific reefs where currents are strong (Veron 2000. and is sometimes referred to as the Meso. flyfishes. they are considered as demosponges with lineages connecting them to several enigmatic. Smith GM. 2005). Coral reef growth in an era of rapidly rising sea level: predictions and suggestions for long. bors (Leys and Meech 2006). As particles and phytoplankton are drawn through the radioles. Belize. Indeed. Zooxanthellae are found 25 µm or more from the calicodermis. A shelf edge reef tract composed largely of Acropora palmata north of Miami (southeast coast of Florida) falls into this category. filament (e.swimming medusa. Le Campion. or form simple dense encrustations. Thus. ogy and growth of the deep. The Role of Zooxanthellae Most scleractinian corals (as well as millepores. This includes increased three. including coral. In: Dubinsky Z and Stambler N. both courtesy of Image Science and Analysis Laboratory. alternating band densities do not occur in all coral species.. Macroalgal graz. 1988. secretion of organic material that serves as a scaffold for 2. Dysidea spp. CH CH 3 CH 3 N H O O CH 2 CH 3 CH 2 COOH N H CH 3 CH 2 CH 2 COOH H CH 2 CH 3 CH 3 N H NN N N N a b Phycocyanin Figure 3–6: The structure of phycocyanin is an open version of the porphyrin ring in chlorophyll. including airstrip (arrowhead). the total reaches well over 400. Cyanobacterial mats called stromatolites formed reef complexes across the Earth from 1 billion to more than 2 billion years ago. thetic products to areas of active growth. Glasby et al. Nonetheless. ———. found on the east side of continents.or nanospheres that fuse to each other and are subsequently cemented by glassy layers. eds. but certain species of the genus Oculina can pro. and several types of corals grow over or between other calcareous organ. NASA. Mgaya YD. 1978. but is more substantially developed when completely exposed as shown on the right.water reefs requires high levels of grazing because. Clayton MN. in contrast.sea coral Lophelia pertusa: im. The resulting carbonate deposits project well above the surrounding benthos and often form wave. The transition from the end of the last glacial maxi. ten distinguished as primary frame builders. these animals secrete mucus containing these chemicals in high concentrations. and other Caribbean islands (Blanchon and Jones 1997). Glynn PW. promotes the formation of calcite crystals within the framework of the cell wall. members of the phylum Porifera. All of these are conjugated systems like chlorophyll. yps with smaller lateral polyps. are colonial and owe that characteristic to budding as a normal pro. Pit connections are maintained during part of this process and new secondary connections are also formed. 2006.spawning species. Radke G. and mollusc shell debris accumulate from storm activity.Bebout L.5 x 106 cells cm–2 (Muscatine et al. 1974). certain dinoflagellates called zooxanthellae are symbiotic with many invertebrate groups and will be described later. sponge larvae may remain in the water column for periods ranging from a few minutes to 15 days. Reef Sponges•95 Sponges as Habitats Sponges are conspicuous components of hard. and may serve as substrata for the growth of other organisms or hiding places for predator and prey (e. they are the structures for which the group is named (Figure 6–6). There are many types of photosynthetic bacteria.g. However. 2004. whereas others may target blood cells. Bioerosion and coral. Lyons TW. Berlin: Springer: 67–94.000. showing three cell layers. although the mechanism has not been examined closely. land and Lasker 1989. nel architecture. 2006). 1979. which is more actively expressed in the dark. ments (Adey and Vassar 1975. Alternatively. 32•Chapter One Smith LW and Birkeland C.Pichel 2006). This is thought to enhance contact time with seawater and increase the uptake efficiency of organic matter produced by their microbial associates (Weisz et al. spirally coiled tubule composed of fne microtubules. (Scleractinia: Pocillo. In: Dubinsky Z. heterotrophic nutrition significantly increases one or several membranes (Wakefield et al. tresses are thought to be the product of strong vertical and lateral coral growth. eds. raveling the nature of the “enhancement” process has been elusive. but their products (spores. ing pit connections between cells and the orientation of high. Stromatolites develop on the pool margins in depths of less than four meters. thus form. 2006). Risk MJ. Biotic Re. sc. ited: reefs and coral carpets in the northern Red Sea. not all of the production is desirable. both open to their western (lee. Submarine lith. while others are able to live within the tissues or as “hypobionts” under the coenenchymal mat. and overgrowth of tropical gorgonians by milleporid hydrocorals: Per. Marine Ecology Progress Series 60: 185–203. and carbon production. Zilberberg C. In the sand. Remnants of this distinctive organization are formed at the skeletal surface and are buried within the skeleton as it grows. as well on the side of the branch where polyps do not occur. Marine Carbonates. Likewise. How. and Leggat W. plications of the calcium carbonate mineralogy in the Stylasteridae. Jones AC. whose known range is limited to the Mediterranean and North Atlantic. tals is accompanied by considerably smaller ones (e. Cyanobacterial sheaths may collectively form a thin but slippery biofilm over submerged rocks. unless something is done architectur. Asexual reproduction may also occur.g. They are up to 1. In: Hedges SB and Kumar S. a b 48•Chapter Three ment and its associated phosphorus input is suspected.600 km long (Figure 1–11). Introduction: a diversity of oceans. Nobles DR.Pacific reefs. 1842.g. Bathymetric and seasonal changes in photosynthesis and respiration of the phototrophic sponge Phyllospongia lamellosa in comparison with respiration by the heterotrophic sponge Ianthella basta on Davies Reef. the remaining space between the cell wall and the cell membrane becomes filled with larger. After fertilization. Jamaica. efits. 2004. erts M. cess called intratentacular budding (Figure 6–29d). and Erez J. These are distinguished as passive endoliths. Bulletin of Marine Sci. etc. foraminifer. fishes. it is plausible that the host regulates its symbiont populations. among other toxin. cally formed cone. et al. Glass Sponges Hexactinellid sponges (typically regarded as the class Hexactinellida) are a group of about 500 deep. digesting competitively sub. which lies at a depth take place on isolated seamounts hundreds of meters deep. Cairns and Macintyre 1992). Bastidas et al.patharia”). eral growth through the production of actively dividing and calcifying meristem cells generated at the ends of the filament (Cabioch 1988). phology of Cinachyrella australiensis (Carter 1886) (Porifera: Spirophorida: Tetillidae). 1993. 2001). ton. Turon X. pertusa polyps and branching structure. and reproduction of Lithothamnion tophiforme (Es. lophora pistillata in different light habitats. etons composed of solid calcium carbonate. the amount taken has de. pearance and often contain more than one type. 1981. The skeletons are densely calcitic giving them a distinctive. Testeniere O. Inset: Close. Wood R. topes of carbon and oxygen within the bands can indicate Gonads Tentacle Oral disk Mesentery Oral cone Mouth Pharynx Mesenterial flaments Coelenteron Calicodermis Coenenchyme Columella Wall Dissepiment Coenosteum Epidermis Mesoglea Gastrodermis Nematocysts Cilia Paliform lobes Septa Costa Corallites Figure 6–13: Illustrated relationship between tissue and the scleractinian coral skeleton. In addition to the more obvious large sponges. Phycologia 17: 361–367. Cordes EE. east of Australia. 1977. ing fleshy and filamentous algae. crevices. although addi. and often oc. Re. However.g.occur with reef. Sipunculans create characteristic pencil. that all metabolic requirements are met by photo. posed of calcified green plates joined together by small. Coral Reefs 18: 241–253. Shingle cays may have a sand mar. and are known for their ability to create densely aggregated. zooxanthellae acquired by vertical transmis. USA. Isotopic disequilibrium in calcareous marine algae.5 meters and a diameter of 70 cm. Because their tests are often well preserved and abun. picted in Figure 1–15a. Mendonça. 26•Chapter One unable to respond to rising sea levels during these times are described below. In this case the micrite is composed of aragonite with crystals around the sheaths (Macintyre et al. ing the location of the slender defensive dactylozooids (dactyl = claw). Most of New Caledonia’s reefs lie about 30 km from the shore.and sulfide. most of those considered here do not.g. pear in Appendix II of the Convention on International Trade in Endangered Species (CITES). Hydrobiologia 512: 89–95. sterdam: Elsevier: 75–87. sition (Adey 1978). survival. Stromatolite reef from the Early Archaean era of Australia. resulting in a variable degree of maternal care that is characterized as brooding. 2000). 1996). especially the . 1990. nomenon. Journal of Crystal Growth 310: 3528–3534. However. Rahman MA and Oomori T. et al. tions: progress and problems. Roark EB. Although known chiefly as photosynthetic members of the plank.. ing typically begins within 10–36 hours after fertilization (Glasby et al. typically defined as a ring. Acosta et al. 2002. these sponges may be the only organisms that are able to keep their hosts in place.Verlag: 266–271. tive coralline demosponge Astrosclera willeya Lister from the Indo. Maida M and Ferreira BP. Allelopathic activities of sponge extracts. Woelkerling WJ. Fautin DG. Proceed. duno et al. compared with those growing on sand (Figure 3–15a). compounds that are leaked from the symbionts to the host (Yellowlees et al. algal ridges thrive under these conditions. Udotea is a psammophile with a similar morphology. on cellular surfaces. this description does not necessarily apply to all species of Halimeda. Some corals spawn eggs containing zooxanthellae. aspects of barrier reef and atoll evolution. The Chagos Archipelago. gelatinous substance consisting primarily of polysac. 1a. Photo courtesy of Aline Tribollet. location. De Goeij JM. 1986. Darwin Point: a threshold for atoll for. tionally significant. Ultrastructural studies on crystal growth of enameloid minerals in elasmobranch and teleost fish. and Finnerty JR. the removal of such non. including Cuba. Anthozoan polyps are solitary or colonial forms that are typically larger and more complex than hydrozo. Huvadhoo Atoll (Maldive Islands.” these cnidarians harbor numerous epizoic sponges. but populations in Barbados are substantially lower (Glynn 1997). 1996. and fatty material typical of yolk. Lichte E. they may also form biological cements in modern reefs. Sponges also play a role in facilitating regrowth of damaged reefs by temporarily binding loose rubble until cementing agents make the connections more permanent (Wulff and Buss 1979. Crystalline or noncrystalline carbonate formation in such cases is biologically induced. are active suspension feeders that generate their own feeding currents. stracum). this form of calcification is biologically induced (Chapter 2) by photosynthesis. lage one atop the other. poma caudata: A) trochophore with apical tuft and medial band of cilia. The perithallus arises as fila.. chaetes. Serrana.30168. and possess a thin epidermis and gastrodermis and a weakly developed neuromuscu. matocysts may be clustered into dense epidermal ag. As a corollary. Parts d. Hine et al. coerulea can even become a primary frame. diage. Uplift movements in New Caledonia–Loyalty Islands area and their plate tectonics interpretation. ent effects in two high. and the genetic evidence is unequivocal that they were the progenitors of the chloroplasts in all eukaryotic plant groups (reviewed by Raven and Allen 2003).up of L. In: Life and Death of Coral Reefs. ments that are ill suited to many other organisms. Marine Biology 93: 527–533. Goreau TF. Crustose coralline algae are slow growing and chemically undefended.g. the larval endoderm is transformed into the gastrodermis. nous green algae. Adey WH and Vassar JM. and occur as green patches on polar bear fur. Brittlestars can be numerically dominant inquilines. or production of sperm packets (sper. Sinularia may be considered as a group of hermatypic soft corals. and may grow to 1. tional dependence on yolk is not always clear (see below). especially those on large atolls. These are all variations of a particular type called ex. brachiopods. or metazoans.water antipatharians: proxies of environmental change. However.enhanced calcification. Design strategies in min. 1–3. seems to interfere with the crown. and Hawkins SJ. In addition. Figure 2–3: The inside of the mussel shell (Mytilus edulis) is com. suggesting that the symbiosis may be facultative rather than obligate (Thacker 2005. The ventral sac used in tube construction collects The family Sabellidae is a group of 344 species (Glasby et al. Some calcareous sponges that appear to be members of this class were at one time considerably larger and played a significant constructional role on Mesozoic reefs as de. 2002. Even larger calcitic structures. Science 202: 831–837. Large tests are typi. appear in modern temperate and polar environ. light. These compounds also act as feeding deterrents to most generalist herbivores. The most conspicuous biogenic reefs are produced by the activity of permanently attached. Calcite (c) from the prismatic layer is shown at the upper right. Natu. Nature 440: 1186–1189. sanctae. Environmentally influenced variability in the mor. Palytoxin is one of Some juvenile damselfishes that do not associate with anemones as adults congregate near the tentacles of sev. et al. Fungi.g. 2001). and even herbivores em.produced coral rubble. In addition. 1976. As. ondary production of the temperate gorgonian coral Eunicella cavolinii (Coelenterata: Octocorallia). The calicodermis is very thin (0. tus. ment of hexactinellids in their own subphylum. These specimens from Papua New Guinea range from 1 to 1. Composed of a fine ferric oxide powder. This process be. or even within different parts of the same thallus. low the surface (Grigg 1988. often in cryptic en. demosponge sclerocytes take up and concen. bonate framework and sediment production. Effect of natural zooplankton feeding on tissue and skeletal growth of the scleractinian coral Stylophora pistillata. 2000. ity to each other (Hooper et. a common mode of asexual reproduction among certain branching forms. d) Incompletely separate polyps appear as a chain of mouths as in typical “brain corals. eton can also be heated and formed into bracelets and other ornaments. 14 cm wide at the base. 1965. Modern coral. One of these. 2010. Thacker RW. erythrin differs from phy. d) Kure Atoll is close to the Darwin Point and is the northernmost such reef. sion in Acropora across the continental shelf of the Great Barrier Reef. and their standing stocks are 30–40 times that of all the bottom. 1985. 2007. 2002a. and Cor. logical Society of America and University of Kansas: 585–764. others exhibit degrees of dependence on them. Part I. In Mar Chiquita this species covers >80% of the benthos. Foster RA. and a highly restricted distribution in some. especially those on Jurassic and Cretaceous reefs. 1990). Additional sheets of circular. 63 Red Algae.sea corals on North Pacific seamounts and islands. the process is not entirely negative and can be viewed as a means of increasing biotic diversity. However. if the definition includes any body of water separated from shore by a barrier. Schlesinger A.Guldberg O. Small islands develop on its rim. 1992. 84 Trophic Dynamics and Ecological Interactions. which grew at rates of up to 12 m per 1. Most reef. However. thiformis from Florida (a skeleton. genetically programmed) and occurs year round in P. chlo. During the Mesozoic era. ranging from the western Pacific to the Arabian Gulf and from Japan to Australia. cies also branch from the base or extend runners or fila. Influence of seawater chemistry on biomineralization throughout Phanerozoic time: paleontological and experimental evidence. Reprinted from Dahan D. including Lagoa Salgada lagoon on the northern coast of Rio de Janeiro. ———. multinucleated cells. Hochberg EJ. size. logical. The se. Endolithic microflora are major primary pro. India. with losses by biological and physical forces already taken into account. 2004.200 islands in the Figure 1–6: a) (opposite) Location of the Great Barrier Reef showing the boundaries of the GBR Marine Park including the southern. Ritson. scleractinian. They constitute the green slime on the sides of flowerpots and tree trunks. more like M. noted here by P. This “splash zone” develops a characteristic blackened crust community (Tribollet 2008). like all bacteria. Carbon for photosynthesis by zooxanthellae is supplied primarily by bicarbonate in seawater.Cava AM. Reefs composed of hexactinellid and other siliceous sponges were widespread at one time across much of the world. the relation. courtesy of NOAA Ocean Explorer.. If a small cylindrical sponge has a height of 4 cm and a radius of 1 cm. Courtesy of Jennifer Smith. duce wide sandy beaches. 102 ognizable. caribaeorum. and also serve as sites where gametogenic tissues develop. which settle gregariously to create pseudocolonies that undergo further fusion to form complex habitats (Dubois et al. New Zealand. Nature Heredity 92: 557–568. Ostreobium filaments have been found in car.g. Wilby D. ily recognizable. bruggen and Kooistra 2004). approximately 100–270 µm per year (e. a b 148•Chapter Seven up to 100 m across and 2 m high on rocky or gravelly substrates at depths of 2–20 m. movement. and planktonic organisms pico. change in these polychaetes.cleaning process is complex and requires physiological and structural specializations that may include localized decalcification of the cell walls. ticular habitats. Some are single cells. shallow subtidal and intertidal marine environments. Science 236: 1654–1657. Kuffner IB. and Agell G. when harvests were limited to shallow water. Camoin GF. Sheppard 2000). ever. Early development of zooxanthella.polyp genetic and physiological charac. Medina M. The challenges of sipho. and are composed of calcified cell walls.” Taken together. giving rise to an essentially unlimited number of genotypes. 356 Design and Function of Marine Reserves. sometimes referred to as accretion. The gastrodermis becomes folded and complex as it lines a blind. This results in part from aggressive competition for space among sponges and between sponges and other sessile organisms. 2005. NOAA 2008). 1995.g.edu/WWW_html/mar_topo. 1991. iron. Others produce a rhizome. Thus the organisms typical of this zone can be quite variable. Osterobium queck. Map courtesy of the Spatial Data Centre. Natural History Museum. tions of three marine demosponges. 2003. tial cytoplasm among living individuals. Trichodesmium. P. 2008. Pedro Bank (SW of Jamaica) and numerous banks in the NW Hawaiian Archipelago are examples. Barrier reefs may occur parallel to continental coasts. c) Halimeda segments are clearly seen embedded in reef rock excavated from the deep (100 m) outer slope of Glover’s Reef. larger amounts of magnesium are found in calcitic skeletons secreted by tropical organisms (Mackenzie et al. and Fungi. Jones and Yellowlees 1997). Zann LP and Bolton L. if formed. arago. Palaios 3: 111–135. national 48: 204–217. and Hidaka M. microhabitat. some herbivores (e. dae and the genus Primnoa. they rely on their store of yolk to sustain them until they can attach to the bottom and feed on their own. Typically. Scale is ap. among other factors (e. including most of the Bahamas (e. Aeby G. Temporal separation occurs at night when no photosynthetic oxygen is pro. 1997.water Caribbean corals most commonly associate with clades A and B. Consistent with their ability to function in low light. and often occurs in aggregations of hundreds of densely packed colonies that can cover tens of square meters (Figure 6–22).Pacific. kingdom. which is only 19 km across. 1983. Pennsylvania State University. Vissher PT. are difficult to separate from turfs and are therefore often included in discussion of their function. 2002 a. thereby increasing the longevity of segments once they are dropped from the thallus (Perry 1998). organisms including heterotrophic fungi and bacteria. and give up. Chemical deterrence of a marine cyanobacterium against sympatric and nonsympatric consumers. scribed species occur in dimly lighted or dark ocean regions as deep as 6. Colonies may occur as a few separated cells (Figure 3–8a) or attached daughter cells forming branched or unbranched chains called trichomes. ate production was about 1. although one species is capable of crawling for up to 20 days before taking up a sessile existence (Maldonado 2006. especially boring sponges. shown by the white line (upper right). Journal of Physical Oceanography 11: 771–779. 1998. and Martins CS. completely encasing them vertically. whereas most Indo. other species may require several weeks or months of development. Photosynthesis and calcification at cellular. Au. each with its own cup. tained by the system) was 75% of the gross amount.000 to >4. mass of its thallus every 15 days. Calcarea. which may grow to 300 µm long (Figure 3–8g) as it extends into adjacent sand grains (Le Campion.g. PO–3 4). tion is superficial and occurs on the surface of previously existing substrata in shallow water.building corals are environmentally sensi. a semiprecious stone. Tuft. and other cell types may also participate. In: Riegl B and Dodge RE. Photo courtesy of K. Cambridge: Cambridge University Press. may bore to where light levels may be as little as 0. During seasonal reproductive pe. marily by 1–2 species that form numerous mounds 1–5 m wide and up to 1. Fishes other than anemonefishes that associate with sea anemones. ysis of the Dendrophylliidae (Cnidaria: Scleractinia). but independently of light. and fishes. Shinn (1963) showed that on shallow Florida reefs.Associated Cyanobacteria a. one in the Bahamas (Hogsty Reef). matypic? A redefinition of ecological groups in corals and other organisms. The two. and Dubinsky Z. count for 3% or more of the inorganic matter. ing the Devonian (Appendix 4) when they displayed a great variety of forms. Indeed. or they may form tubes and tunnels that ex. b) North. It is also an active area of re. at least in part. b) Close. 1985. The morphological definitions are further muddied by sclerite. often protected with a tough protein layer (the perio. mate reef growth. and convert it to an insoluble. Indeed. Overview The Annelida is a phylum of soft. cellular pH. these symbionts fall into distinct genetic groups called clades that are designated by the letters A. Attrill MJ. Algal biodiversity. 2007). Trends in Microbiology 13: 229–235. sponge pumping rates and tissue densities? Oecologia 155: 367–376. 33 Biomineralization. but a spike occurred somewhere between 2. sections of the Great Barrier Reef (Davies et al. Science 292: 679–686. in the dark or at low light levels. both past and present. for example. and Ilan M. the innermost portion of the shell in M. 1978).like pattern of canals like chaetetids. Large foraminifera like these are major contributors to carbon. fall and runoff from shorelines). Weinbauer MG and Velimirov B. try and structure of gorgonian and antipatharian coral skeletons. Isla Perez core. Maldonado M and Riesgo A. and Genovese SJ. Oscillatoria spongeliae is the most common of these. but medusae are not produced and only sperm are released. ———. Sherman et al. fishes than most of the others (Fautin 1991. male that nourishes and shelters the zygote as it divides. ian chain. 2007). 246 Coral Reef Herbivores.Parker and D’Elia 1997). typically as a cocktail of enzymes and other proteins. Sponges are organisms that are classified as suspension feeders. ed. Demo. plications for the fossil record. 25: Coral Reefs. coupled with low rainfall and high evaporation. Milliman JD.e. A study of calcium carbonate deposition in the genus Padina. tional and complex type of matrix. a single barrier reef surrounds two islands. Likewise. ganisms may form coral communities in regions that do not fit this general description. with cells that release proteolytic enzymes and other com. eydouxi with spe. tinian corals. Tectonophysics 24: 133–150. where they play significant roles. In addition. 1989. cellular organisms may also be exerted within cells. Morpho. Nonetheless. forming a barrier reef farther from shore as well as a lagoon behind the reef crest. this is also true of living members of the genus Vaceletia. 2008. Those that bite or scrape epilithic algae or organic matter growing on rocky surfaces are referred to as graz. There are 1. Marine Biology 92: 339–347. Each tube represents an indi. Muscatine L. This species can be abundant in high. 1987. or filled with storm. gamete releases are at a scale sufficient to reduce visibility in clear water to less than a meter. Coral Reefs 13: 171–174. Filho RB. Wallace. PSZNI Marine Ecology 4: 65–79. Hallock 2003). tal Biology 46: 671–678. Staurozoa. Casanova J. Neall and Tre. Ecological Monographs 66: 431–450. it also becomes increasingly diverse biologically.g.uk. a group that is referred to as the crustose coralline algae (CCA). and Scaturo D. Bacterial biofilms that form microbialites are a good example of this type. and the majority of these are hermaphro.9210. The traditional explanations involve a marked increase in ox. tentacle.html. animals whose bioeroding activities are confined to the surface are grazers or excavators. However. Ulrke E. Paul VJ and Van Alstyne KL. but some sponges may also remodel spicule morphology to accommodate such forces (Hill and Hill 2002). take by cyanosymbionts results in the production of organic mat. 372 4. Mercado JM. more than twice as fast as those found in Western Australia (Dill et al.g. 2008.900 years ago and lasted about 1300 years. 2008. duster worms. however. Thornhill DJ. Compo.g. depending on the ecological context. ure 4–10) not only are kleptochemists. but their molecu.attached reef fishes. tition between Palythoa caribaeorum and other sessile invertebrates on St. Sammarco and Andrews 1989. various types of lagoon reefs may develop. Wheeler AJ. In addition. phological complexity. Many studies have examined the sources of carbon and how they may be partitioned or shuttled between photosynthesis and calcification. sheath.. and internal structure of their segments. Lara EN and Gonzalez EA.g. The term “reef” is a particular type of carbonate buildup. While less generally signifi. or may become dentate (toothed) as shown. Thompson et al.MacDonald GJ and Leys S. Geological Society of America Bulletin 115: 1344–1355. Etnoyer P and Morgan LE. A sponge. “fire coral. International Journal of Remote Sensing 9: 389–396. Such environments may facilitate food capture and sediment shedding. Anemones and Anemonefishes On coral reefs and associated habitats of the Indo. mon ancestry and these unicellular forms are now widely regarded as aberrant cyanobacteria (Lewin 2002) simi. Cha JN. netic cements may be more generally important in this process (see review by Rasser and Riegl 2002). developed. Amaral FMD. reaching a peak of diversity and abundance between 1 and 1. a gastropore with its sur.240 lb. These include the rich and colorful “octocoral gardens” off Nova Sco. 2008. Geological Time Scale. 2002. ficiently turbulent conditions are present (e. Effects of benthic algae on the replenishment of corals and the implications for the resilience or coral reefs. Carter D. where does all the carbon go? In one case it has been found that sponges rapidly form and shed choanocytes into the surrounding envi.end sac. there are several types of carbon. 2007). lot PY. Canadian Journal of Earth Sciences 18: 972–980. 2007). the framework is con. Most of them manufacture a calcitic shell called the test. whereas species of Millepora fare best in somewhat more sub. 2011.g. and certain but not all precious red corals) are listed in Ap. Mass spawning by green algae on coral reefs. New York: Springer: 59–88. bryozoans. Aeby GS. although the details of bicarbonate transport out of the calicoblasts are unknown. ing acidic conditions (Moya et al. eds. Allen GR. This result is particularly no. Marine Biology 151: 71–83. 2001. encrusting. These serve as hosts for 28 dif. Sammarco WP. suspended sediment.. or sheet. Cheshire et al. Tahiti and neighboring Moorea are both surrounded by a complex of fringing and bar. Arrows point to a micrite crust. els of cyanobacteria. (2) they provide nutritional material to their hosts. These seaweeds typically pos. and Slattery M. Marine Biology 145: 641–649. Wulff 2001). Some of them like Gigantospongia sp. tertidal sabellariid reef composed of Sabellaria alveolata tubes in Morecombe Bay on the northwestern English coast. grew to >5 m high. Coral growth on the lee. cause of a poorly developed lagoon. dactylostyles may project in or near the dactylozooids. ate a process of extracoelenteric digestion. and a few even exhibit both intratentacular and extratentacular budding in different parts of the same colony (Veron 2000). Part F: Coelenterata. including reef corals (Chapter 6). parative study of associated North Atlantic Arctic/ subarctic Melobesioideae. Reef Evolution. Australian Institute of Marine Science. 1988. and even asexual planula formation (described below).resistant. and arise from branched cortical filaments that are produced in a few hours.7). Life Cycles Some taxa of foraminiferans may live for only a few weeks. where X is an anion as described below. chae. as it is sometimes called. The persistent expansion of its operculum and branchial crown. cyanobacteria) and have high light require. Do sponges hold reefs together? Nature 281: 474–475. Ni. and basal cell. 196 Tunicates. tion is used by filamentous species and involves breaking the filament into segments. Wehrl M. ment of epitoke and atoke regions. Senckenberg Museum. Nematocysts are quite numerous in both gastroder. Ecology and Paleoecology of Benthic Foraminifera. St. These serpu.walled “pinnacles” or “knolls” that extend vertically from deeper waters. Tissue and Cell 34: 246–261. hexactinellids are capable of creating considerable relief above the bottom and at one time constituted an important group of reef builders. cies placed in its own order. Journal of Applied Ecol. Microbasic amastigophore Barbed shaft is dominant. This reproductive strategy is also associated with the ability to survive at low densities in contrast to broadcasting species.celled eukaryotic organisms such as diatoms.normal temperatures? Boletin de Inves. sive reef systems if circulation is open and persis. Indeed.corals also do not produce large colonies. Calcification of the Cell Wall and Beyond In contrast to the irregular aragonite deposition in Halimeda. The chloroplasts are also unusual. 2nd ed.C. Carnivores are found among 19 polychaete families.. herms have been found at greater depths in Indonesian and Nicaraguan waters (Roberts et al. 2001). FAO Fisheries Report 200: 469–481. Other Anemone Symbioses Despite their relatively large size. and Sánchez JA. especially where winds traverse broad expanses of open water and produce large and fre. 2009). Most fundamentally. However. Sand cays are composed of coarse and fine sands that have been shaped by wave and current activity and are often only a meter above the highest tides. there are perhaps only 15 atolls in the western Atlantic (Milliman 1973.dwelling foraminifera. pearance. veloped on islands that are elevated a few meters or more above sea level. as detailed in Chapter 4. Because of the nitrogen. and evalu. They lack internal organelles. Williams GA. a b c Reef Algae and Foraminiferans•57 species and their roles as carbonate producers in mul. duced calcite or high. Supramolecular architecture of cyanobacterial thylakoid membranes: how is the phycobilisome connected with the photo. b). Le Campion. solved organic compounds from plant decomposition. These suspension feeders create currents by various means that force water through bags composed of finely divided mucus strands. Zo. and chemistry of Diplo.secretory gland/collar apparatus..Mortensen 2005. eds. or further elongated as “fibers. They may also project from the body surface. well Publishing: 141–169. 2008). Branched corals appear to be pruned by hurricane. but others have determined that they develop before the appearance of mature eggs and even in the absence of egg formation (Harrison and Wallace 1990). 2002). sabellariid reefs may extend for thousands of kilometers along the coastline (e. Geister J. the ability to regulate metabolism. in a rapidly evolving area of research. ure 6–24 a. ternational Journal of Invertebrate Reproduction 2: 251–269. Tunnicliffe V. as well as notes on their ecology and illustrations of their structure. These include certain heavily calcified hydrozoans. but C.boring polychaetes can number in the thousands per square meter and cause an estimated 0. waters that protect shorelines from erosion. the high abundance of aragonite worldwide is due to these groups. in respiration). conceptacles may form a distinct layer in the epithallus or may be situated in the uppermost perithal. CrystEngComm 9: 1262–1268.like storage product inside the chloroplast (Table 4–1).Paleozoic reefs most likely pro. sclerites have been defined as “calcareous elements of the mesoglea irrespective of form” by no less an authority than F. Host imprinting in anemonefishes (Pisces: Pomacen. Deepest known plant life discovered on an un. Saipan. nesses.shaped. ing described in Chapter 5. linity varies with the oxygen isotope ratio recorded in the foraminiferan test (e. Steneck RS.resolution atomic force microscopy image showing the high. Literature Cited Reef Algae and Foraminiferans•75 Grell KG. but multicellular thalli are more typical and may include filamentous. ygenic photosynthesis. 2004. In ad. ronment that develops twice the salinity of normal sea. Comparative re. referred to as skeletal extension rates. gliding form that settles on carbonate sand grains trapped by the stromatolite polymers. once actively growing. not all eunicids produce burrows.axis and the three perpendicular a. The cone. duce nematocyst. relatively large swings in annual temperature. Four to six capitate tentacles surround the mouth. Martins Rodrigues MV. A consensus estimate is 10 mm yr–1 with a porosity of 50% (Buddemeier and Smith 1988).coral interface (Sullivan et al. Bochsler VS. However. tected by living tissue.g. 1985). mer by dividing “upward. lower left. prise roughly 10% and Figure 4–6). and kya refer to billions. Tulsa.. but they were common compo. Indeed. havior. Prochlorococcus. 14°N. especially in cold and deep water (Cairns 2011). ments. producing much of the car. even among specimens sampled from a depth of nearly 400 m (Wagner et al. Calcifying Cyanobacteria Sheaths. genus Por. Taphonomic signifi. narian. and Field ME. Photo courtesy of Paul Humann.000 species have been described. Stanley SM. fended sponges. national Coral Reef Symposium. 2004. Filaments from the sponge undermine coral polyps in advance of tissue at the surface. Microorganisms. including herbivorous fishes and various groups of molluscs.g.1186/1471. farther south.dimensionality to the reefscape. Note coral. turbidity. dolithic (boring) sponges. and Luquet G. Simpson TL. along which there is considerable variation in local wave conditions. the giant barrel sponge Xestospongia muta. Corallines can also dominate the surface of deeper.sea corals: Primnoa resedaeformis and Desmophyllum cristigalli. gates that serve as substrata for a coral community in some locali. Western Washington University Chuck Messing. nae. 1989). Likewise. Coral Reefs 8: 99–107.000 km2 and is typically 10–30 m deep.microbial segment reefs in the marginal Mediterranean Sorbas Basin. Coral Reef Science. high latitude and cold. tas R. whereas those with porcelaneous or hyaline tests are found later in the Paleozoic era (Bignot 1985). or membrane. and then become free swim. specific and generalist associations. when threatened. and its lagoon is are based on that designation. tion. Fitzgerald LJ. 2007. b) Field of subtidal stromatolites. The physical form of a cal. Behind anemone lines: factors affecting division of labor in the social Literature Cited 136•Chapter Six octocorals (Coelenterata: Anthozoa). Andrews AH. Ecosystems of the World. Gaetani and Cohen 2006). cally identical sponges (e.g. Some corals that are apparently the same species are found in the tropical Indian Ocean and the Red Sea. temperatures can be driven to >35°C (e. growth rates. Calcification in algae: mechanisms and the role of metabolism. Figure 5–12: Hexactinellid reef species from British Columbia showing a dense aggregation of tubular Heterochone calyx (Hc) as well as the shield. and Paw. ments where the fragments were produced or carried (Bowden. lagoon waters whose qualities are unsuitable for corals may begin to mix and flush across the reef. Where colonies are abundant. Similarly.ended. nicates collected off the southeastern United States. tropical. ture reefs concentrate food sources for commercially and recreationally valuable species including crabs. It is also worth noting that algal ridges can form on any type of reef if shallow water and suf. both recent and historical. unlike green algae. 2004). Photos by author. out the assistance of zooxanthellae (Reed 1981).spawned species may settle within 2–3 days of fertilization (Miller and Mundy 2003). ———. Growth lines Myostracum Periostracum Prismatic layer Nacreous layer Figure 2–2: Stacks of elongated and thin calcite crystals form the prismatic layer of this mussel shell (Mytilus edulis). thus rapidly acquiring available space (e. Thompson RC. a b Reef Algae and Foraminiferans•73 North Africa and the Arabian Peninsula during the early and middle Eocene. and Rögner M. Reefs that have suffered high levels of mortality may become dominated by small. In: Hutch. 2005. and (3) they take up their waste products. The reefs east of An. Larger tube. Degrees of spionid infestation can be considerable with as many as 9 individuals/100 cm2 of host surface. but the blade is com. allowing specialized grazing organisms to incorporate such algae into their diet (e. it does not constitute a true tissue and is sometimes referred to as an “epithelioid. nanostructures (crys. some embed themselves in the living tissue of their hosts causing the secretion of skeletal material around them in response (e. nate resistance to the toxins of certain anemones.associated mi. larger atolls are more likely to have deeper lagoons than smaller ones. Coral Reefs 26: 883–892.g. Golubic S. These are some. They are abundant from the intertidal zone to depths of over 5. far slower growth rates have been documented from black corals growing in deep water. the relative abundance of CCA and that of fast. Journal of Experimental Biology 212: 3892–3900. not be distinguished without microscopic examination and are said to be isomorphic. Biomineralization in reef.Early Oligocene ~34 Mya in association with global cooling. Microscopy Research and Technique 62: 279–299. These in turn in. Advanced Materials 15: 959–970. The plume extends hundreds of kilometers off shore and extends south and espe. CRC Critical Reviews in Plant Sciences 6: 1–45. its are given in the captions and I thank those who were so helpful during this process. 2008. where they are referred to as costae (Figures 6–12. Coral reef encruster communities and carbonate production in cryptic and exposed corals reef habitats along a gradient of terrestrial distur. ulations can be substantial. acteristics of carbonates of gorgonian axes (Coelen. some. Farrea occa forms dense clusters of smaller tubes. The larg. McMurray SE.diameter canals that correspond to much slower filtration rates than LMA sponges. the source of which includes those pro. but also feed on these polychaetes. Collins AG. epithallus (epi). Most gorgonians found on coral reefs belong to a het. 246 Overview. crostructural diversity of coral skeletons: implications for taxonomy and control on early diagenesis. careous green algae and stony corals. maphroditic and release egg and sperm in bundles. and Evans E. Box: Two protons generated in the calicodermis or in the organic matrix are removed and exchanged for each calcium ion by a Ca. However. but no definitive reasons are known (Banks et al. Examples from the western Pacific (Figure 1–9) include wave. 1998. develop. Acosta. 2004).g. Marine Biology 119: 1–11. 2008). 2005). calcite level. in some cases exceeding 400 individuals per square meter of coral surface (e. Is this narrow and shallow strip of water a barrier la. 2006. col. 1976.latitude populations of the reef coral Pocillopora damicornis. cumulate atop the volcanic basement resulting in a thick cap of limestone. ganic carbon occurs as bicarbonate.water species of Halimeda Lamoroux (Halimedaceae. 1999. Most mats are found in extreme conditions such as hypersaline bays and lakes. Rainfall can quickly dilute seawater. The area now has the distinction of being a World Her. out additional terms. Eggs or sperm may also be released through nu. ters: distribution and fishery impacts. ing high levels of turbidity and sedimentation as well as extreme fluctuations of temperature and salinity. Many of these are members of the Bryopsidales. Proceedings of the 6th International Coral Reef Sym. scrape. 298 Diversity and Coexistence. Scanning electron micrographs by author.3 billion to 200 million years (Clayton 1990). While very little is known about how thylakoids developed and which ances. Adey WH. Research in Microbiology 154: 157–164. 1988. Alternatively. et al. In: Jones OA and Endean R. cally attached to their tubes.lived. Invertebrate Biology 129: 105–120. Coastal waters stretching from the Indian subcontinent to Vietnam are all tropical but predominately reefless because of the discharge of the Indus.” which when viewed with a hand lens.water calcified hydroids from Brazil (Hydrozoa. Glynn PW. ing collagen and other types of protein (reviewed by Erlich 2010) that form an extremely tough but flexible internal support system for the colony. Integrative and Comparative Biology 45: 335–341. Thus. and De Yoreo JJ. miniferivores. An Introduction to the Structure and Formation of Modern Reefs •13 archipelago. eds. and in some cases. diversity. lected particles are driven by cilia in a central groove and are then accepted by ciliated tracts around the mouth as described below. Bulletin of Marine Sci. 1997.300 km. b). ed. thonoplastes. Thacker and Starnes 2003. the smallest photosynthetic organisms known. Variations in Coral Reproduction within Groups Some coral groups do not fall neatly into broadcaster. Darwin’s ideas concerning reef growth and evolution were essentially ignored until 1896 when the British Royal Society mounted a series of annual expeditions to Funafuti Atoll (Tuvalu Islands. ing sand deposition or exposure of rocks to erosion. showing outer periostracum. 144 In some cases. At the same time it also deposits between 3 and 3. nus Caulerpa). tennae. sand grains. 2008. 2004. Synchronous spawning: when timing is everything. Gore et al. 2002. Literature Cited 38•Chapter Two Topics in Geobiology 17: 159–203. This association beneficially increases the surface area of both organisms and protects the coral from invasion by excavating invertebrates. Large Savalia colonies occur in deep water and can grow to 2 m tall. Conversely. These outbreaks are difficult to control by herbivores because cyanobacteria are generally considered a poor food source owing to low nutritional value and indigest. Inset: An unusual. noid form.Frank I. Perth: Western Australian Museum. exhibiting maximal growth rates of 130– 180 mm yr–1 (references in Dullo 2005). fied according to the shape.Neto et al. but in most cases. Figure 6–3: Morphology and life cycle of the hydro. Limnology and Oceanography 25: 1113–1117. 1997. A small but important group of parrotfishes and sea urchins fall into this category (Table 1–1. or unpaired oral tentacles. Because of these differences. glucose. especially in environments where nitrogen and phos. mats composed of these organisms may serve as a shelter for numerous microinvertebrates (Cruz. although there is little Figure 7–9: Eunice sp. caraguan Rise. lices 2002). beled c+ and c– to refer to its “up” and “down” positions. Chlorophyta). This stage develops mitotically from the fertilized egg. several hundred microns long.water reefs (Roberts 2005. 25. damicornis can be long. Brisbane. and shapes the entire structure by aligning the fusion of nanogranules to its outer surface (Sethmann et al. weeds without relying on herbivores. these windward vs. or reef corals (Chapter 6).called clownfishes or anemonefishes. erotrophic plasticity and resilience in bleached corals. comes a multicellular blastula within hours. Amaral et al. Similar bio. ing types unique to certain groups of hexacorals (Daly et al. Bioerosion and coral. currents. cesses? There are two general methods that are used to esti. barrier reefs are among the least common of the three major reef types. spicule growth. The corti. guished as accretion rates. derstood process that involves coral tissue and the pho.carrying medusae. semicircular doubles. 233 Reef Crinoids. row pore. tophyte stages. Likewise. E: development of medusae and completion of the life cycle. damicornis. Jones RJ and Yellowlees D. scale approximate.g.history strategies. Degnan BM. thereby accommodating shortfalls due to reduced photosynthesis (Titlyanov et al. such structures are typical of the reef’s windward side. Hayward DC. 2001). Sheppard CRC. including scleractinian corals (Kuguru et al. 2000.nitrogen load.like sedimentary structures (Figure 3–11) with a characteristic laminated appearance (Figure 3–12 a. Alternatively. Marine Ecology Progress Series 326: 99–113. Minnery GA. the cells deep within the tunnel could produce acids while the portions closer to the sur. ner circle of noncalcifying cells that produce the gametes. Payri C. quire several days to adapt to another. Some of the success of these corals has been attributed to a diverse suite of chemical deterrents. whereas others contribute to reef structure either by temporarily binding substrates or organisms together. there are more than 700 fringing reefs that have formed around some of the 300 coral and sand islands. There are two groups that are commonly classified by the mineralogy and ap. with 120–150 m high coral cliffs and an area of 660 km2 . Marine Biology 98: 143–155.like tentacles. are able to trap and bind sediment. They can occur as unicells. As reef growth continues vertically. 12. bacterial bioerosion causes the substra. this increase in the carbonate simultaneously enhances the likelihood of calcium carbonate formation shown in Equation 4–4. true tunnels are produced. Chlorophyll b differs by substitution of the –CH 3 (boxed area) with –CHO. rows). ing 350 m in depth in both the tropical Pacific and Atlan. Figure 4–10: Two color forms of the Indo. Conversely. Atkinson MJ. This type of feeding is distinguished from raptorial feeding and can be divided into several subcategories.level rise and reef back. likely from the numerous mitochondria within the calico. which fertilize eggs borne and housed by female plants. this view has become highly controversial and will be discussed later in the context of changing coral reef environments (Chapter 15). et al. The 87. Hyman LH. Benthic cyanobacteria are also common components of “algal turfs. The edible mussel Mytilus edulis is a good example of an organism that produces two crystal types in specific regions (Figure 2–1). Phylum BX. 1985). Hadas E. Zhu ZR. each of which can regener.like structure. Treatise on Invertebrate Paleontology. dividuals (castes) characterized as aggressive “warriors. Pan. ment points for dependent. Mechanical design of spicule. Testing for defensive synergy in Caribbean sponges: bad taste or glass spicules? Journal of Experimental Marine Biol. tion and in the type of mineral produced. and atolls occur in a sequence around inactive volcanoes.like. and Thorrold SR. Metabolism. especially those with small body sizes. These submerged volcanic platforms. and are especially well developed in current. The carbon source for photosynthesis is primarily seawater bicarbonate. Crustose algae grow flat and horizontally across the substrate. versity of two Caribbean sponge.” Both Udotea and Penicillus belong to the family Udoteaceae. Such reefs may have been “shot in the back by their own lagoons” (after Neumann and Macintyre 1985). 2008.dimensional structure as a canopy over reef sub. for example. tic. Renewal time for lagoon water. b) Cross. 1999). and gorgonian corals (Witman 1988. for example. Tsounis et al. which encourage fur.km long windward shelf edge of Grand Cayman Island. 56•Chapter Four a patch of Halimeda 100 m2 may produce 200 kg (440 lb) of aragonite in a single year (2 kg CaCO 3 m–2 yr–1 ).. b). and indeed this zone may lack reef development if circulation is highly restricted or ex. Proceedings of the 8th International Coral Reef Symposium. The glacial.like monolayer. Philosophical Transactions of the Royal Society of London B 352:457–468. structure. gin typically binds individual spicules together. Pro. He further reasoned that the increasing weight of coral growth. 2007. but complex reefs and reef communities are only now being described from several localities in Indo.526. herbivores.. zoan fire coral Millepora. Gastrodermal structure and feeding re. shingle cays are referred to by their Polynesian name “motus. which becomes negatively phototropic. Foraminiferans are therefore significant groups of organisms. either of which facilitates photosynthesis. eds. The red algal life cycle is composed of two separate plants. Uriz MJ. sponges that maintain a bacterial concentration of 2–4 orders of magnitude higher than the surrounding seawater are called high microbial abun.lighted and circulated environments (Carpenter 1990. Depending on the species. and Give Up Reefs Branched species are the fastest growing corals that form the reef framework and could best keep pace with sea level changes. Only a few dozen foram life histories are known and these are often complex. and Golan Y. filamentous species of Scytonema (order Figure 3–9: The endolithic cyanophyte Mastigocoleus testarum bored into dead. Gametophyte plants produce heterogametes that vary in relative size even within a given genus. These factors are important in promot. Most colonies pictured are tightly clustered P. fication controls on fluxes and isotopic composition of nitrate from Florida Keys sponges. extracts not only deterred certain fishes from preying on its resident ophiuroids. although hermaphroditism is not uncommon. The leeward southwest side is essentially closed.celled cyanobacteria and involves the duplication of DNA followed by the formation of smaller cells from larger ones.. Coral Reefs 23: 455–458. transform them into organic com. logical Bulletin 194: 178–186.g. sediment. However. there is a high turnover in this community. 2004). isms. including high levels of sedimenta. isms. moval by encrusting coral reef cavity sponges. gregarious. Foram rhizopodia often branch and interconnect. Keep Up. et al. suggesting that factors other than predation may drive the association. The last hexagonal crystal type to be considered is apatite. In addition. New Scientist 2: 58–62. search in molecular genetics and biological engineering that emphasizes the development of novel materials from the structure of natural ones. Holotrich No distinct shaft.g. but there are also prominent encrusting groups that are commonly found in certain reef envi. Turon et al. alesce and become sections of a fringing reef (or any other type) if suitable ecological conditions exist. USA 106: 5204–5208.g. ism. and Cuet P. et al. logical Association of the United Kingdom 87: 1683– 1692. and whether the stromatolite is intertidal or submerged continuously. ture of Pleistocene coral reefs of Curaçao.. the North Sea. Marine reserves along the Sinai Pen.shaped nor circular. and biologi. Quartz and carbonate grains. ification in coral reefs: some facts and misconceptions.Daryan 1996). eral centimeters into the sediment (see Figure 4–12c). Fabricius K and De’ath G. making them less palatable to herbivores (Hay and Fenical 1988). is the second largest atoll with an area of 3. Bulletin of Marine Science 56: 625–652.nutrient seawater (Chisholm and Moulin 2003). and Fiji islands have undergone uplift in the past. tube. ary into the Torres Strait (Figure 1–6a). McCook LJ. they infrequently aggregate as reef builders as they do in some challenging environ. Colonies may grow to a meter or more in diameter. Barrie JV. Schläppy et al. tinctions. Smith SR. and role in biomineral. While epi. like corallites around and under each polyp. some species produce more organic material than others. and a few xanthophylls. 2007. Coin is 18 mm. and circulation. and functional categories. Riegl B and Piller WE. and a source of food.Pacific counterparts. divers using SCUBA with helium and oxy. Polysaccharides of cal. It is not a unified barrier as the name implies. Marine cyanobacteria are not quite as pernicious.fixing capability of certain cyanobacteria (Chapter 3) and a high ratio of photosyn. Microbial Symbionts Sponges are associated with a wide variety of micro. slow electrical signals can propagate across long distances to control feeding currents and react rapidly to stimuli (Leys and Meech 2006. Köck M. which often counter. Likewise. In most other sea. reduction in porosity. Paleobiology 26: 651–667. north. bution is often challenging. about 7 kya (Macintyre and Glynn 1976). rather than the tropics (Kerswell 2006). cyanobacteria (or heterotrophic bacteria living within their sheaths) could produce inorganic or or. Atoll Research Bulletin 496: 76–99. Goreau TF and Goreau NI.dwelling en. Members of this guild feed on a wide variety of organisms from foraminiferans (mi. Turfs also provide habitat for a wide variety of microinverte.water octocorals and antipatharians are especially threatened. This chapter focuses on the biology and contributions of these and other cnidarian groups to reef structure and function. Many spe. however.like force of the tubule (see Schlesinger et al. b. Barnes DJ and Chalker BE. Geology and Hydrology of Carbonate Islands. et al. cumstances. ate sediment on modern reefs of the Indo. Lang JC. composed of chaetae. often referred to as “red tree corals. Acropora palmata reef framework: a reliable indicator of sea level in the western Atlantic for the past 10. the pore. mental Marine Biology and Ecology 119: 15–29. ming or crawling ability. The first is a function of a specialized aboral epidermis called the calicoblastic epithelium or calico. although specialists that feed on gorgonians are not un. 2008). Natu.building corals face elevated extinction risk from climate change and local impacts. a b Reef Cyanobacteria•49 protection of shallow reef zones where they periodi. and Alcoverro T. salinities may exceed 45 ppt. Crystal alteration in a living calcareous alga (Halimeda): implications for studies in skeletal diagenesis. Unpredictable and destructive storms also take their toll. 1990). In many stony corals the filaments may also be used offensively against space competitors. How. J. In: Dubinsky Z. However it occurred. vae are unlikely to settle far from the parental sponges. and Serrana Bank. tory fishes and their prey. Nitrogenase cannot function in the presence of oxy.gov/general/lib/CREWS/Cleo /St. eds. Overholtzer KL and Motta PJ. Photo courtesy of James Oliver. or an organic cement may be produced that agglutinates sand grains (described be. Alcyona. interfere with feeding. The mechanisms of calcification and its relationship to photosynthesis and respiration in the scleractin. medusae. Glass sponges arrest pumping in response to sediment: im. Photo by author. Glynn PW and Ault JS. and is often referred to as broadcast spawning. Most of these are further differentiated as magnesium calcite. Geo. Ng PLK. Vol. and Reiswig HM..shaped or semicircular reef sur. that is longer or shorter than the horizontal ones..driven sediment. whereas in H. Watling 2007). unlike those described above. Rowland SM and Gangloff RA. algae. Sponge biodiversity. University of California. Larval development must be complete prior to establishing a benthic existence. b). reef rubble. Sexual reproduction of sclerac. vide a chemically complex environment. Limits to Coral Reef Distribution Most reef. ized structures. Culver and Kuris 2004). tually lifts the floor under the polyps and creates a new layer of aragonite over the older one. Organisms living on reefs differ in struc. roa Atoll. should cause the island to slowly sink (subside). a b +c +c –c –c –a 2 +a 2a 2 –c +a –a 2 –a 1 +c +a 3 –a 3 +a 2 +a 2 –a 1 –a 3 +a 1 –a 2 +a 3 +a 1 Figure 2–7: Apatite is a hexagonal dipyramidal crystal with six sides. 2005. that project from paired fleshy.6 (paper) ISBN.West Pacific commonly constitute more than half of the shallow. American Association of Petroleum Geologists Bul. In turn. 2003). Solitary and colonial corallimorphs occur on rocky substrata and coral reefs worldwide.living.0. The palps drape across the sediment surface and may extend 60 cm or more from the burrow. ers in this group may include those with poorly formed reef frameworks (e. Temperature extremes. Cox EF. ing their affinities difficult to assess. part of the supratidal zone above it may contain an equal or greater biomass even though it is only occasionally wetted by splash or spray from the highest of high tides (spring tides) or by storms. tapered projections called branchiae. Marine Biology 91: 291–309. van den Berg H. about which little is known. these eunicids are especially pro. Effects of temperature and transport conditions on calcite growth in the presence of Mg2+ : implications for pa. clic structures (Figure 4–9).covered platform. Bowden. Connaissance actuelles et contribution à l’étude de la reproduction et du cycle Udotéacée (Caulerpales. Physiology of coordi. Fragmentation is another form of asexual reproduction that breaks up the sponge by mechanisms not under the sponge’s control.2003 . et al. and Pantazi. mal and epidermal tissue. Mineral skeletogenesis in sponges.negative bacteria. After one to several days. Littler and Littler 2007). ers may consume organic matter along with inorganic sediments. which from reflection.containing sponges are golden brown or tan due to the presence of carotenes. Figure 4–8a). b) The large foram Marginopora vertebralis has a symbiotic relationship with photosynthetic dinoflagellates called zooxanthellae. nor and trace elements in sclerosponge Ceratoporella nicholsoni: biogenic aragonite near the inorganic endmember? Palaeogeography Palaeoclimatology Palaeoecology 228: 109–129. Sponge orange band disease (SOB): a patho.3 My Hawaii 0. Indeed.com/. Reef Sponges 5 Figure 5–1: The organization and cell types in the pinacoderm.forming organisms. In general.. 1984. Journal of the Marine Biological Association of the United Kingdom 86: 563–566. In a few regions (e. Wu XH. seawater sa. mat. dermis. bluk 1981).atoll types. Fontes. Geochi. and may subsequently use giant chaetae to dislodge particles from the substratum.like distribution. Compar. 1995). lid to be in the feeding position. Helgoland Marine Re. derstudied Indo. ship to photosynthesis. Depending on tidal amplitude. In some areas. Coral Reefs 6: 161–172. most species pro. sumably fitness) to cope with changing environments.water fossil corals dating from 34 Mya have been recovered from some of the more southerly of them. Miller KJ.atolls. 20 2 Reef Minerals and Mineralization. ology of algal turf species and how they can affect the outcome of species competing with them for space. Tambutté S. a sugar alcohol. Photos courtesy of Paul Humann. water temperatures. Elmetri I. as has been shown repeatedly in a wide variety of reefs (see Cocito 2004 for review). Chlorophyta) across a depth gradient: comparative growth. each of which is referred to as a cycle. USA 105: 54–58. and Goreau PD. In some cases. clined to a more constant level of (Bruckner 2009. 1988). In some taxa. Colonies can cover >40% of the available space in favorable environments. Here. Blanchon P. ments. Europe..like forms were com. and radiometric age validation of a deep sea. Organic materials produced by photosynthesis may also have an effect on calcification if they are pumped out of the algal filaments into the extracellular envi. but lacks the typical cyanobacterial phycobilisome structure and phycocyanin. more than 50 other species of fishes from several families use anemo. 2001. Liv.g. the great majority of which are marine. rated into one other subclass. Smaller serpulid reefs ~7 m across have been recorded over large areas from Mar Chiquita. NOAA. Dupouy et al. philic forms. and Car. Photoinhibition in the Mediterranean green alga Halimeda tuna Ellis et Sol [sic] measured in situ. As used here.Horani et al. 1988a. French Research Institute for Exploration of the Sea. Cambridge: Cambridge University Press: 164–180. but archaeocyaths were particularly common elements of them. low sedimentation. cesses. rockfishes (Sebastes crameri).Pacific coral reefs. compared with dozens or more in broadcast spawning species. Some are obligate coral symbionts. par. ovals) are micro. though the mechanisms of calcification are controversial. which func. and others that become parasitic on medusae (see Shick 1991). Figure 3–10: Intertidal notch at low tide. Thus. 76•Chapter Four Macintyre IG. ments and subunits from a marine sponge direct the polymerization of silica and silicones in vitro. Depending upon the group. damicornis. The Free. Many warm. Paleocean. part of which is shed periodi. This is referred to as multispecific spawning. However.like forms that are called spicules in tunicates (see Figure 9–8 in Chapter 9). acidic conditions may cause decalcification in both calcifying chlorophytes and crustose corallines. 2007.000 heterotrophic. In this book. Net primary productivity in coral reef sponges. Blum JE. ture 381: 512–514.like Multicellular algae are important components of shallow. weeds. dom 87: 1345–1348. judging from laboratory behavior (Wiebe et al. almost all of which are marine. dispersal. the pina. New York: Academic Press: 1: 1–50. However. but the group that proliferated on mid. rowing and tubicolous forms than in free. The outer ends of the branches are especially chlorophyll rich and develop swollen tips referred to as utricles (Figure 4–7b. cies of Halimeda (Halimedaceae. Reproductive processes in sponges: a critical evaluation of current data and views. whereas in the dark or when inhibited. 2006. and upon their decay release large amounts of nutrients that promote the ex. López. USA 93: 6069–6072. produces haploid spores that ing pressure (Paul and Hay 1986). and Tibbets I. However. line algae. The ecology and phylogeny of cyano. comparable to the barrier reef at Bora Bora (Figure 1–5). make this genus read. Consistent with this estimate. Leichter et al. crustose coralline algae (CCA) calcify and exhibit a high degree of filament layer orga. ing gathered on the biology and environmental role of these organisms. a species that can grow at a rate of 90 mm yr–1 and can produce a reef framework as predicted by Darwin (Purdy and Winterer 2001). and Montgomery T. as are the otherwise potent nematocysts of millepore corals.like amino acids and scytonemin. lithophores: a cellular perspective. 1999. and even partial development of feeding tentacles (Richmond 1987). ible because they are obscured by ectosomal tissue and spicules that may function to keep debris from clogging the excurrent canals. 2009. In contrast. some sabellids burrow into mollusc shells. as have structurally distinct neu. While there is little question that allelochemicals are a critical component in the natural history of sponges (Pawlik 1997. nians such as the “bubblegum coral” Paragorgia arbo. making this pro. Scharf et al. windward environ. Even though some of these have a volcanic origin and rise from deep water.existing topography and depth. 2001. The fate of coral reefs in the Andaman Sea. Limited membership in Pleistocene coral reef assemblages from the Huon Peninsula. Journal of Experimental Biology 214: 2749–2754. Zoanthids (Order Zoantharia) are a small but impor. which is more closely related to diatoms and dinoflagellates than to red and green algae. coralline algae are among the dominant frame builders on the Flower Gar.water scleractinians are capable of frame building. Hay ME and Fenical W.inflammatory. Coral Reefs 24: 149–156. Weiner S and Dove PM. lina and appears to be composed primarily of L. especially in tubular and vase. 1996. Photo courtesy of Todd LaJeunesse. unlike their modern representatives. Palaeoecology 232: 214–236. astrea. The latter is characteristic of vaterite and is called a dipyra. southerly flowing East Australian Current at 31°S. I. In contrast. charides in the cell wall. 129 7 Annelids and Sipunculans. only the sperm bear flagella. as well as differences in the composition of its coral communities.bearing polyp until fertilized eggs or planula larvae are released. However.enclosed. 152•Chapter Seven quantitative data available to compare them with other such agents. Fukami H. depending upon depth. dis reproduce on a different schedule compared with those from other south Pacific locations such as Tonga and Fiji about 1. These are produced in response to adverse conditions such as desiccation.300 m. However. A few species. Complete mesenteries project into the coelenteron as thickened. Food may include organic particles.purdue. itage Site. Clifton KE. et al. ical and subtropical oceans. eds. Thus large speci. 124•Chapter Six with increased diversity of fishes. Gametogenesis usually marks the end of the life cycle and results in an empty test contributed to the sediment. One of the best examples of these is found on Capitan and other reefs in west Texas and New Mexico. the radioactive band can be used as a reference year for subsequent growth bands. whereas lower. irregular. insula on the tectonically active. In this efficient process. the formation of these structures is controver. a coral with novel digestive filaments. Fertilized eggs that are retained in this manner often develop rapidly. Cellulose in cyanobacteria: origin of vascular plant cellulose synthase? Plant Physiology 127: 529–542. and coloured fora. Thallus attachment often varies according to the na. Individual polyps can reach disc widths of 30 cm or more. although the degree of nutri. 10•Chapter One The term “barrier reef” has a certain cachet that is of. Cimino G and Ghiselin MT. This is Darwin’s subsidence theory. Based on Pomponi 1980. improved fishing techniques. one of the most common forms of fertilization involves the discharge of gametes into the water simultaneously over a short period. Earliest evolution associated with the closure of the Tropical American Seaway. there are roughly 8. Photo courtesy of Paul Humann. Ultrastructure of the tentacle nerve plexus and putative neural pathways in sea anemones. Stromatolites are common in some of the intertidal and shallow subtidal envi. ing competitors. Indeed. more than 1. flagellates. las.. rectly on the outer cell surface so that the c. whereas others fail to produce them at all. 2000.carotene. tesy of Paul F. Gateño D. and construct tubes that are typically composed of white aragonite. Ca. heat. 2010).CS/WG9. The trunk region is adapted to bur. but are vulnerable to others and may exhibit a preference for certain hosts among those available to them. fleshy al. Marine Biol.Johnson Space Center ISS006. ing that an acceptable one exists (Bruckner 2003. as in this example from Palmyra Atoll. Tahiti.water cor. Central America.up of astro. eter and >20 centimeters long. the spaces between the skeletons are often structur.specific boring pattern as well as the absence of photosymbionts.like crust. phytes described above. porella. The sediments are incorpo. Palae. yet they incur low levels of consumption by herbivorous fishes. calcareous algae can serve as a surface cement. However. it has been suggested that arago. where it remains close to the surface due to its requirement for high light levels. The association of Ophiothrix lineata and Callyspongia vaginalis: a brittlestar cleaning sym. 1990. or skeleton. 2008. but severely reduce them in the winter when the colonies become inactive (Dimond and Car. suggesting that the potential interrelationships among these microbes in the calcification process are likely to be complex. b) Porites astreoides showing polyps sharing a common wall. Oahu. tance of cyanobacteria as ancient photosynthetic organ. water black corals (Cnidaria: Anthozoa: Antipath. and in Hawaii where they are part of a managed fishery (Grigg 2002). the scler. However. Witting J. leeward reef characteristics do not always hold in other regions. thus leaving a fossil record 100 My long. etal modification in response to flow during growth in colonies of the sea whip. Life and Death of Coral Reefs. is common in the shallow tropical west. Marine Biology 141: 31–38. ence 78: 343–375. Material Research Society Bulletin 17: 37–40. However. als the dactylozooids often lack tentacles and may occur randomly relative to the gastrozooids. face sclerites in gorgonians (Coelenterata. rized in various ways by different authors. and Salati E. Journal of Ex. ring to live birth. and Rinkevich B. sina fulva. Halimeda grows by forming one or more new nodes at the apical segment. Coral reef biology. but they are usu. bolic interactions between algal symbionts and in. vidual larval settlement. oral tentacles. A biogeographic analysis and review of the far eastern Pacific coral reef region. However. muscular jaws to gouge. Lophelia communities (and those associated with other deep. Zonation on most terraces is well preserved and distinguishable. ward. Like many cnidarians. tacular budding patterns: a) Acropora cervicornis branch show. Epithallus sloughing: a self. especially the true stony corals in the order Scleractinia. or are separate but tightly bound by wisps of gorgonin. Bulletin of Marine Science 81 (supplement):185–194. Marine Ecology Progress Series 271: 147–158. Atoll Research Bulletin 415: 1–34. Some free. including the nummulit. Pu. aria).producing organisms. Several species of Palythoa (e. and falling sea levels world. Proceedings of the Biological Society of Washing.Pichel F. Lindner A. there are some notable exceptions. There may be as many as 50 indi. one of the major accessory pigments associated with chlorophyll in cyanobacteria. Reitner J. 2001.tech coral reef restoration methods modeled after natural fragmentation pro. Calcified. but do not leave them and cannot build new ones if they are extracted.feeding web that allows the capture of planktonic organisms including bacteria. which is typically composed of chambers that interconnect by openings called foramina (hence the name of the order). However. This carbonate cap represents some 49 million years of subsidence and intermittent reef growth. Ecology 89: 1994–2004. Calcification in cocco. but also occur under them. NASA. or submersible vehicles. which represent up to 12% of their dry weight. 2004. In some cases. an interval begins that is referred to as the larval competency period. but some protection may be afforded to those that can bore 1–3 mm beneath the surface. This term was originally derived from early studies of Indo. 1981). graphic Journal 171: 372–374. lus sp. dermis. a group that is well represented on coral reefs. for example. Pentecost A. and an uncalcified pore that allows their release. Sullivan B. Areas where the tube contacts the skeleton appear to be partly dissolved as if acid had contacted the aragonite crystals. tion of amino acids and other N.Pacific reef builder. at times by as much as 25–30 m in as little as 1. their flexible skeletal structure notwithstand. The ingested chloroplasts stolen by these specialized herbivores. are formed as an apparent single crystal within an intercellular space by a similar process (Chapter 11). In some cases the bacterial population can account for almost half of the sponge’s biomass. Figure 5–7: Intracellular spicule formation in the demosponge Reniera sp. 2001.section of the sea fan Gorgonia ventalina from Florida cut with a diamond saw across the base. the most conspicuous of which are brominated alkaloids. Kawasaki et al. Sponge. is a hetero.F.like crystals of aragonite composing the nacreous layer. However. De’ath G. continue to photosynthesize for periods ranging from a few hours to as long as 3 months in some species. tion of the enzyme carbonic anhydrase. These calcifying rhodophytes are anatomically complex. proaches to Coral Reef Ecology. Ni. Reproduction Cyanobacteria are not known to reproduce sexually.walled heterocysts create an anaerobic mi. In: Aronson RB. At night. products of photosynthesis. the Emperor and Hawaiian chains provide a continuous re. 1994. terization of Symbiodinium in an Acropora valida colony. In contrast. 2008). The eastern half is Papua New Guinea where the Huon Peninsula is located. negger et al. algae are eu. Clionid sponge surveys on the Florida Reef Tract suggest land. plex defenses. In. a b 110•Chapter Six deed. UNEP. and sediment generation on reefs in the Central Great Barrier Reef province. On the reefs of Isla Isabel National Park on Mexico’s Pacific coast. b). Different fusulinid genera were characteristic of distinct periods of time as they evolved rapidly and then became extinct. Moussavian E.Documentos/WG. Environment and biota of the Tikehau Atoll (Tuamotu Archipelago. Serpulid embryos may be brooded within or outside of the tube. Figure 7–3: The terebellid polychaete Loimia medusa inhabits western Atlantic reefs under rocks and in cracks in reef substrata where they construct tubes composed of fine sand and mucus. especially in tropical and temperate areas of the world where conditions are favorable for them. 1972). Anthozoa) from littoral of Pernambuco. The presence of sclerites may contribute some physical deterrence to predation. although studies now employ molecular approaches. turbidity. Zoanthid mats acquire space by growing directly over the competition. Much of the island is surrounded by a barrier reef. many display a considerable range of idiosyncrasies in size and shape. Most unicellular foraminiferans pro. Grigg R. Brolund TM. Ross CA. Thinly branched.g. cluding those that bore superficially. Photo courtesy of Genny Anderson. tidal exposure. or more likely bacterial species as. the term “epilithic” when used alone can refer to any organism that grows on rocky surfaces. typically composed of magnesium calcite like the sclerites. thus eventually cutting off their magma connection (see review by Neall and Trewick 2008).” or simply as Z. ronments of these islands on the eastern Bahama Bank. Bulletin of Marine Science 65: 259–282. Tursch B and Tursch A. cally develop into extensive mats. Western boundary currents.g. season.Roubaud C. and current conditions (redrawn from Geister 1977). Sánchez JA and Opresko DM. a) is a composite of ISS006. many poly. Although serpulids are individually common on tropical reefs. ameter. rofaunal assemblages associated with sponges and tu. espe. Université Catholique de l’Ouest. no. Thus a ring of dead coral tissue precedes the more obvious expansion of sponge growth over the coral (Chaves. et al. none of the spores or gametes produced by red algae develops fla. and physical damage. Interconnected mounds create tunnels and other cavities that serve as a refuge for a wide variety of algae. the Marshalls. The mesohyl also contains spe. lapidosa. Broadcast spawning of gametes and forma. The window of gamete viability is brief and often lasts but a few hours. ies considerably deeper within the carbonate (Matthes et al.Williams R. electrons. b). and although they can cally considered subsidiary to those of secondary me.Llorens JL. ties that they can form complex. The typical orthorhombic crystal is thus shaped like a cereal box. but may be replaced by more slowly growing species that produce thick layers or stout branches.000–24. form. Schuhmacher H and Zibrowius H. excava. While the function of the matrix is clearer in some groups than in others. 2008).like (terpenoid) compounds. 25: Coral Reefs. endolithic cyanobacterium of the genus Solentia (order Pleurocapsales) that occurs along with the filamentous forms beneath the calcified crust. lites clearly separated. now 15 m deep.brooding (Stylophora pistil. Characteris. and McGuiness KA. Tooth enamel.American Barrier Reef. and the Red Sea. Zoological Studies 46: 520.herbivore interactions: the ecology of chemical defense. Elements of Micropaleontology: The Microfossils. and Mitchell JG. Some anthozoans also have well. ern Asia. Reefs that “keep up” are able to grow with sea level rise. and certain Paleozoic corals. Hallock 2000. These structures can be self. cant quantities of it in deep or polar waters are likely to be the first to exhibit calcification failure as carbon dioxide and pH levels change during the coming decades (e. These additional photosymbionts are known to fix nitrogen under conditions of low light as well as in the dark. spicules in some species differ subtly or inconsistently. however. the crystals are randomly ori. cess of 150 mm yr–1 (Tunnicliffe 1983). Studies in Geology 4: 219–230. Chilcoat GC. tal factors in the Arabian Gulf and the Gulf of Oman: a comparison to the Pacific region.” those which lack an internal framework. barrier reefs. Lambeck and Chappell 2001). and Paul VJ. Modern Foraminifera. Mantyka CS and Bellwood DR. Duffy JE and Hay ME. segmented animals. Weiner S and Addadi L. or predation. sponges. ally and morphologically distinct types. Ferguson AM and Davis AR. Calcium carbonate is then precipitated onto the tube as a fluid. vae developed internally are often negatively buoyant and tend to travel short distances. Epidermis Mesoglea Retractor muscle Circular muscle Siphonoglyph Mesenterial retractor muscles Gastrodermis Primary mesenteries Mesentaries Sphincter muscle Secondary mesenteries Mesenterial flaments Tertiary mesenteries Pharynx Coelenteron Coelenteron Pedal disc Column Mouth Pharynx Oral disc Gonad Acontium Primary mesenteries Secondary mesenteries Tertiary mesenteries Cross section through Pharynx Cross section through Coelenteron Figure 6–7: Anthozoan polyp (sea anemone) with internal and external anatomical features (left).pone. In 2005 a strong earthquake in Indonesia raised a shallow fringing reef by as much as two meters above sea level around the island of Simeu. logenetic interpretation. sponge in the Indo.West Pacific. Appendix 3). lar scaffold called an organic matrix that orchestrates the process. patterns of sea level fluctuation can be inferred from coral reef growth.building organisms. 2004. or spread on toast. ing a wall or “drop. this species becomes a broadcast spawner that may be capable of increasing its genotypic variation by hybridizing with other species of Pocillopora (Combosch et al. Bishop and Pemberton 2006). 2007.. more typical of fresh water environments. ticularly interesting as an omnivore with an unusual and distinct preference for cnidarians. t. Crustose coralline algae from the Flower Garden Banks. establish themselves at depths of 10 m or more by vigor. primarily overnight. et al. ties are unsuitable for corals may begin to flush across the reef. c) Well. Structure and ecology of the Saipan reefs in re. clones). perhaps owing to variations in abundance. Barkl Y. dreds or thousands of meters deep. Cell walls of this group are com.1371/journal. but cannot produce the necessary anaerobic conditions in the presence of photosynthetic oxygen. derived from microbial activity or from organic compounds in solution produced by the metabolism of reef organisms (Macintyre and Marshall 1988. Goulet TL. 1974. spond was dependent to a large extent on local ecological and oceanographic circumstances as well as reef compo. tion results in a larval form within a brief period. file. However. mas. Birkeland C. Meibom et al. lopora damicornis. Rebora M. Deep. wick 2008). and even reduces turbidity by high. and tunicate spicules (Addadi et al. although deeper borings have been reported (Ralph et al. eds. Magnesium calcite production in organisms may be controlled by cellular processes including magnesium transport. Palaeoecology of a Late Devonian back reef: Canning Basin. including the so. phorus and nitrogen.g. Under some circumstances. especially in waters deeper than 30 m. Some “siphonous algae” (see overview by Vroom and Smith 2001) often lack distinct differentiation of the thallus and instead display a tube.dominated commu. which is temperature dependent. A brominated secondary metabolite synthesized by the cyanobacterial symbiont of a marine sponge and ac. Smithsonian Institution US National Museum Ian Macintyre. or they may be too low to remove adequate amounts of CO 2 produced by respiration. De Nooijer LJ.9 (cloth : alkaline paper) — ISBN 978. cross sections at different levels of the digestive cav. or as epiphytes living on algae or sea. in deeper water. Haplosyllus spongicola (Grube) (Polychaeta. are colonial both tissue growth and calcification rates compared with unfed corals in the light (Ferrier. where one species becomes common below 15 m. kel and Pawlik 2005). mary producers: a critical synthesis.Pacific genus Pocillopora can be quite variable. 2003). Despite the limited size of most specimens (< 20 cm). Dunbar RB.oral peristomium) contains the mouth and one or two fused. 2003).Pacific mushroom corals (Family Fun. tive cells that are not the equivalent of tissues in higher plants or the more complex organization found in certain brown algae (Cabioch 1988). Allemand D. Schuhmacher H. can be used for environmental analysis and for the determination of growth rates (Noé et al.protected. and perhaps the density instead reflects the availability of nutrients and the growth of skeletogenic tissue (Cohen et al. making them among the longest living organisms on Earth and a valuable source of envi. However. Marine Ecology Progress Series 357: 139–151. Solentia Pseudoflament showing reproductive cell. thozoa. ers of cellular filaments as described below. and five. One or more tapered. or the walls defining the corallites may disappear. Jiasong F. and (c) a source of inorganic carbon or components of the organic matrix that may directly or indirectly influence crystal nucleation and growth (Allemand et al. dae. nal of Phycology 34: 393–406. Thus. All of these are referred to as endolithic macroborers. and Lough JM. 2003. Grotolli AG. 1995). Earth Science Reviews 42: 95–117. and echinoderms. ing. and by sediment export from storm activity. parathoracic segments. The Cretaceous coral Heliopora (Oc.000 kg from 1992 to 1998. pink. The filaments exhibit different degrees of sediment capture based on the binding properties of the sheath. high. the corresponding male segments are tan. plosion of other planktonic species (Bell et al. Conversely.section of a coral colony showing the living coral tissue (brown layer) and the underlying green layer of Ostreobium sp.. but the second and third succeeded in drilling to 200 and 340 m respectively. the plate’s motion then gradually transports the islands away from the hotspot at about 10 cm per year to the northwest. or semicircular shapes during feeding. for example. Indeed. tists in many biology texts. erite is a dihexagonal prism that has 12 faces and 4 axes in addition to the c. like eukaryotic plants. and tilting (Dubois et al. Nutrient transfer in a marine mutualism: patterns of ammonia excretion by anemonefish and uptake by giant sea anemones.living. fleshy. Botanica Marina 43: 305–314. Pochon X. Photo courtesy of Jeff Jeffords. Pro. Small coral species and other calcifying organisms that contribute lesser volumes of skeletal carbonate are categorized as secondary frame builders. Lewis 2006). However. 1988). These deeper. and invertebrates. 2002) as well as in Australia and other parts of the Indo. ticoral calibration method to approximate a universal equation relating Sr/Ca and growth rate to sea surface temperature. giganteus removed from its tube to reveal the thoracic calcium. that is no longer the case. Indeed. the exposed siliceous framework forms suitable substrata for the settlement of hexactinellid larvae as well as for other sessile organisms.” as the sponge. which leads to a tubular pharyngeal region that may bear one or more ciliated grooves. extending from the cup wall as they converge toward a concave cen. we would be required to recognize that many Z. Solentia sp.. called the Great Oxidation Event. they were unable to find a volcanic platform. green lagoon. 106•Chapter Six dusa as additional systematic tools (e. nitrification. Treatise on Inverte. for example. 109 Calcified Anthozoans. Reprinted from Stolarski J. Several of them absorb photons in the blue and green portions of the spec. and form a link between open water and benthic ecosystems (Roopin et al. Trophic Relationships Coralline algae do not appear to be chemically de. some processes that do not require a low pH have been pro. Trade.absorbing Mycosporine. Marine Biology 148: 479–488. Such needles are common in the lime muds of tropical bays. 1999). Use of ingested algal diterpenoids by Elysiella halimedae Macnae (Opisthobranchia: Asco. Proceedings of the 5th International Symposium on Coral Reefs. the largest number of genera in the At. Trends in Microbiology 13: 229–235. tion in a coral community. thellae may be acquired during stage e. which reduces oxidized forms of nitrogen like nitrite and nitrate. London.0003222.marinespecies .developed pedal disc adapted for attachment on solid substrata. Andersson AJ.bearing foraminifera. temperature extremes. the in crustose corallines can result in growth morphologies ranging from a pavement. sculpture. and the smooth inner side of its long operculum. tribute to their ability to dominate shallow reefs in many Indo. four of which are very similar to Atlantic species. most do not and are far smaller than their Indo. Moreover. OK: American Association of Petroleum Geologists. Each tube is connected to its neighbors by a platform that represents lateral growth by stolons. hooked chaetae that grip the tube wall.1186/gb. cification alone. Coral frameworks may be composed Figure 1–1: a) Schematic drawing of reef construction and the functional roles of carbonate. The condition of released planulae varies widely and includes some that are leci. Sex cells develop from masses of tissue in the segments and may leave by way of specialized gonoducts associated with the nephridia (excretory organs). cial periods.Enhanced Calcification As intracellular symbionts. In some cases. These conditions do not prevail far north or south of 30 degrees from the equator. scopic. face can be decorated with spines.gov /refuges/ van der Land J. lees et al. Lee Stocking Island. water where blooms can be hazardous to humans and domestic animals (Figure 3–17). Bro. Cyanobacteria are the most important group of photo. a heterocyst is noted by the as. including various spe. Seasonal fluctuations in the photo. snaring particulate matter in the process. logical characterization of lineages within the calci. logically distinct parathorax. Food collec. 2001. depending on which of two clades they belong to (McEuen et al.producing co. 1997. are capable of excavating up to 10 cm deep. aria): benefits of hosting anemonefish.spawning species. water. thella fiordensis from SW New Zealand showing distribution of polyps. and grazers if they more generally scrape or sift food from the substratum. is sur. et al. Oxford: Oxford University Press. reef formation. 2001. a discrete nucleus. there are one or more foundation The Nature and Origins of Reefs Carbonate geologists have been meticulous about defining the nature of carbonate deposits. Ecology 85:1372–1382. Cerrano C. tle is known about the feeding behaviors exhibited by this group to adequately characterize them (Edmonds 2000). The “black disease” of reef. and occur with numer. lar or comb. als produce such a matrix that precipitates aragonite in much the same way that bone and other skeletal mate. and other often unknown co. per. In: Müller WEG.Medrano DI. which flow toward the equator from the poles. more than 100 genotypes of Symbiodinium have been distinguished (Baker 2003.shaped gut with the anus located anteriorly. 2000. some species are able to overgrow their competitors. Geochimica et Cosmochimica Acta 69: 4227–4236. On the outside. cialized cells that form the sponge skeletal system. as well as nematocyst. riods. the Carolines. and are completely uncovered at low tide (Figure 3–13b). is generally indicative of reef health (Lang and Ginsburg 2006. some scleractinians produce more closely spaced. Octocorallia) from deep reefs in the western Atlantic. Brazil. Coral Reefs 26: 775–785.water corals: an overview with spe. Vol. the mesoglea. rated walls separated by plates or bars.shaped colonies on Indo. Younger.org/medialibrary/2010 /09/10/29fefd54/CWC. This type of suspension feeding employs a downstream collecting system in which the primary feeding currents are gen. see below). cover an area of 3680 km2 and constitute the world’s largest atoll (Spalding et al. to microscopic mineral.Living Lower Invertebrates. Turon X. bacteria and heterotrophic bacteria are the most com. an undesirable transforma. supplying the animal tissues with carbon requires light dependence and restriction of the host to depths where light is not a limiting factor. and Neuwei. unlike the green algal genera Halimeda and Udotea. Indeed. Among the calcified forms. 2009.. De Beer D and Larkum AWD. and Willis BL. favoring production of both carbonate and mineral. The other method employs contemporaneous surveys of biota or chemical analyses of water actively flowing over reef surfaces (e. times these are called barrier islands.water coral Lophelia pertusa in Norwegian wa. Hoffman 1999). Scale approximate. the islands of the western Pacific. by zones within a single reef. Noel Sirivansanti (http: // www. and in a paleontological context they are valuable indicators of ancient temperatures. deep open.Indian Ocean throughflow on Indian Ocean circulation and on the East Australian Current. tercellular spaces. whereas rounder grains are more often chosen for the outside (Eckel. many sabellids have compound eyes on the tips of the radioles that are structurally similar to those of arthropods. Modern polychaete jaws also may be miner. Comparative Bio.nH 2O). Kupriyanova et al. Cruz. Crustose coralline algae are typically organized into an epithallus.Pacific (Razak and Hoeksema 2003. 2000. cite. and high planktonic pro.. 2008. 2008). Figure 5–19). men displays 15 growth rings (possibly. even if the precise timing and cause of the pairs is not entirely clear.1 My*). Biologically induced carbonate precipi.e. calcareous sponges and other organisms.Pacific waters. 29–46. and oc. 2004). which can be distinguished from a thinner ectosome described be. are best considered as atolls (Milliman 1969). Cyano. Rapid extension rates often indicate a relatively large volume of skeletal deposition with nu. Alternatively. thus preventing liquid waste from fouling the tube. because these features are not unique to fringing reefs. 2006). tral components were used. host specificity. The adductor muscles. Little is known about the causative mechanisms of such blooms. show that it began growing in shallow water. Correspondingly. Biggs J. Coral Reefs 23: 445–454. dated shell outcrops off southeast Florida.. Lang JC and Chornesky EA. In addition. al. Such junctions are thought to function in cell. Cytological mechanisms of calcium car.geomar. 1996). Because these sponges differ so markedly from other living taxa. and thousands of years ago respectively. Factors controlling sponge boring in Barbados reef corals. 1977. and is elevated above it to some degree. its origin and chemical structure is unknown. mostly less than 0. Marine Biology 100: 195–202. peratures. and filefishes. Bioero. 237 12 The Effect of Feeding by Reef Fishes on Corals and Coral Reefs. et al. American Naturalist 115: 113–120. Cuif J. ed. Conversely. the Rhodophyta are the most diverse group of marine algae. Red Algal Calcification More than 40% of all red algal species calcify (Woelk. While the importance of unravel. Growth. alcyo.T boundary. Warner ME. 2008). Parrish FA. leading Reiswig (1981) to coin the term “bacteriosponge. several reef systems are large enough to be seen from outer space and com. synthesis. some antipatharians can be moderately common in certain coral reef habitats below 20 meters (Sanchez and Opresko 2005. Marine Ecology Progress Series 7: 207–226. and Peterson FL. it is also the site where zoo.water. The famous pink sand beaches of Bermuda owe their coloration to this species (Figure 4. 1988. herbivore avoidance may be due to the amount and type of toxin produced and the specific tolerances of the consumer.Fonnegra and Zea 2010). likely E. Bergquist 1978). Kawasaki K.shaped. they are not uncommon. Munk and Sargent 1954). Perez.red Cliona delitrix. Additionally. Often. Oecologia 149: 174–184. ing to the presence of large numbers of zooxanthellae. tion described more fully in Chapter 15. tocysts are concentrated. in which a polyp is ejected without being attached to its skeleton. 2005. which has a total area of 1. Ecology 87: 2479–2488. and project less prominently above the surface. Some species of damselfish and mul. Hauser T. 1978. corallite walls. The randomly arranged ara. is a way of grouping unrelated organisms. this atoll may also be the largest by volume. the surface area of the sponge roughly quadruples to ~100 cm2 . tion of reef carbonates. building sclerosponge Ceratoporella nicholsoni. MacGeachy JK. The lagoons. While boring sponges are often found in nonliving coral reef substrata. This may be the case in calcium stored in some crustacean cuticles. 1997. Journal of Experimental Marine Biology and Ecology 252: 221–253. sociate (Marsden et al 1990.like barbs. many giant anemo. 2002. release toxins that have been implicated in fish kills. duces additional layers of tubes. Journal of Sedimentary Research 73: 856–868. Endolithic fungi in marine systems. the second is the activity of zooxanthellae that are close to. 1996). tia. The tertiary mesenteries are even shorter (Figure 6–7).. quently flushed or buried. In some cases. B and C: specialized cyclosys. et al. The loss or decline of CCA on coral reefs is problematic. 144 Tubicolous and Burrowing Polychaetes. nize and compete with other clones of the same species when they come into contact. about 30% of the sponge fauna of individual reefs in the Indo. 2001). especially where it can be exposed or nearly exposed at low tide (Figure 4–18 a. suggesting that this delicacy may be an ac. This process is distinct from binary fission (e. These reefs have been damaged by trawl. ologische Verhandelingen 1: 1–116. At the same time. cies. Weiss AM. Oahu. and the reefs of the Torres Strait at the north. Brown algae are prominent on reefs. and Cairns SD. haploid gametophyte. An oral episodes of predation or bioerosion (Highsmith 1982).000 individuals per square meter. as adroitly described by Steve Wainwright at Duke University (Wainwright and Dillon 1969). Spalding MD. and turbidity from river discharge. there are nine genera of cyanobacteria that can be found in these structures. Invertebrate Biology 121: 201–211. delitrix is nonetheless aggressive. the symbiosis provides for reinforcement of that framework by skeletal accretion. Quinn NJ and Kojis BL.clonemates at the boundary between competing genetic allies. Sexual maturity is also achieved rapidly and gametes are pro. 2003. growth. It is worth noting that while nearly half of all octo. 2001. et al. one that consists of a single ancestor and all of its descendants (Tomitani et al. Other polychaete predators may lack jaws. Fertilization occurs in the water column at some distance from the sessile parents.H. The effects of eutrophication. Photo by author. The entire structure uncoils explosively from a protein. leaves. suring the rate of calcium deposition by corals un. Oth. Because such crystals can be micro. which exceeds the estimated rates of carbonate production of 1–4 kg m–2 yr–1 typical of most shallow. The unique skeleton of siliceous sponges (Porif. New York: Springer: 1: 473–599.developed windward cays are shown from 10 to 5 o’clock. Coral Reefs 7: 185–195. Muricy G. and Reid RP.Alsumard T. 2010). The mouth is surrounded by two lobes bearing about 250 cili. the plane of symmetry. and to some extent counteracting the effects of organisms that bore into or excavate reef lime. composition.g. the glacial control theory (Daly 1915). are found at depths ranging from 17 to 49 m near the edge of the continental shelf in the northern Gulf of Mexico (Rezak et al. worm tubes. Particle feeding in natural popula. Biological Bulletin 203: 269–274. Literature Cited Reef Cyanobacteria•51 Raven JA and Allen JF. later produce single haploid “conchospores” that once again generate the leafy. low. haps forming fissures and small depressions. and Chapman DJ. ity is limited in some species. Sea Level Changes and Reef Growth The Pleistocene epoch 2. the relative amount of strontium.spawning (left) and brooding (right) coral life cycles from fertilization to embryogenesis. as do a variety of fishes and gastropods (Gore et al. external). ecology. Lapian et al. Colonies may occur as flower. clusively sexual process. However. there are several more specific types described below. a func. finger. and Hopley D.11078.. We focus here on two enzymes whose function appears to be critical to coral calcification. tosynthesis Research 77: 83–94. the most speciose (species rich) of all scleractinian taxa (e. the structure of the thallus can be complex in coralline algae. sociated with fringing reef on Guam (Mariana Islands. resulting in local clus. Gorgonian corals are more diverse on Indo. or uplifted (Andréfouët et al. corallimorpharians. ship between calcification and photosynthesis appears to rely on active transport of both calcium and protons. Al. Cyanobacteria (Chapter 3) and filamentous algae (Chapter 4) may form sticky sheaths that trap fine sediments.com. In addition. a means of carrying away wastes. Bank MS. as a group.g. Lagoons are formed by slow growth compared with the outer barrier. osponges Astrosclera willeyana and “Acanthochaetes” wellsi. tant to predation. The secondary me. Vertical growth on fringing reefs often produces a shallow ridge called the reef crest that may grow to or protrude slightly above the surface at low tide. Figure 3–16: Thick bloom of Trichodesmium sp. Heikoop JM. 2004. gametogenic buds or gonophores form from stolonic tissue. Many glass sponges are large and conspicuous mem. recogniz. Science 175: 1116–1118. There are varying Reef Algae and Foraminiferans 4 Reef Algae and Foraminiferans•53 stolon that creeps across the surface. While vigorous coral growth can offset boring activities. 1988. Ellison et al. Some species also detect and assault arborescent sea whips and fans (octocorals described below). Inorganic silt and clay particles (see Appendix 1) reduce incoming solar radiation by deflecting light. tions. For example. 9°S. terfere with the settlement of coral larvae (Kuffner and Paul 2004). Delvoye L. When these layers are separated and the soluble components of the matrix are removed. 1998. ish in seawater. ward side lagged during the early Holocene so that about 5.Pacific gene flow in the tropical eastern Pacific. cium (Knutson et al. est of these. ed. view the details of carbonate spicule construction and mineralogy. Bali. Marine Drugs 8: 2021–2037. 1997. It has been suggested that anemonefish are imprinted by the mucus secretions of their host during the embryonic stage. they can become significant microboring organisms. Figure 3–10). There are some modern examples of these formations (described below). 1999. Ultrastructure of the spirocyst tubule in black corals (Coelenterata: Antipatharia) and its taxonomic implications. tropical H. allinales. spread genus Schizothrix (order Oscillatoriales). Scale is approximate. However. graphed against film sensitive to gamma () or X. Endolithic algae within corals: life in an extreme environment. Berlin: Springer. Effects of salinity on coral reefs. Coral Reefs 25: 513. Algal Divisions In contrast to cyanobacteria (Chapter 3). Coral Reefs 19: 1–23. New York: Elsevier: 209–252. Thus. Hoffman L. New Zealand. ids (family Nummulitidae) that formed flat. and in a few places may support reef development beyond 30° latitude (Appendix 2c). H..like clusters connected by a basal tissue. Coral Reefs 26: 177–188. or for co. Indeed. lar located at the upper part of the thorax (Figure 7–6a. Bermuda. ytic chlorophytes Penicil.226. cal traits (Steneck and Dethier 1994). ing reefs are more likely to develop a relatively uniform series of buttresses due to active frame. Foster MS. In shallow.C from Marine Biology 35: 253–267 (1976). Oren U. The movement and secretory products (see below) of these cells control calcite crystal nucleation. whereas those in lower energy environments were much more pro. Conversely. gin of a candidate matrix that is found in the skeleton has been traced to the calicodermis (e. Eunicid burrowers. for example. More on this topic will be described in Chapter 15. Embryos and larvae are subsequently embedded into gelatinous masses that are then released onto the reef. Panama 2: 1427–1432. and Piraino S.g. The Structure and Distribution of Coral Reefs. However. suggesting that they act as portals for calcium ions and protons shuttling between cells. Figure 4–13: Penicillus forms small aragonite crystals in separate bundles within a compartment. and relative height above low tide. Microelectrodes show that while this enzyme is active.g.density arrangement). After Moseley 1881. bioses. or can drive the conversion of bicarbonate to CO 2 externally. estimating the degree of dependence on that contri. As many as 1. sure and compete for space. the island of New Caledonia. Makatea (dashed line) is an uplifted atoll. ever. munities of Tikehau Lagoon (Tuamotu Archi. are the larger of the two and correspond to the gastrozooid. b) The clownfish Amphiprion percula is found in the central west Pacific and associates with 3 giant anemones. However. Bayer and Macintyre 2001).g. and bore into calcareous substrata. al. 145 Polychaete Bioeroders. thus these allelochemicals appear to be part of the organic nitrogen production they contribute to their hosts (Figure 5–20. cur within the coenenchyme. but the two groups appear to share a com. and Stuckenrath R. 2002. echinoderm spines. 1999). In the large. a warmer interglacial period occurred ~125. during the early Cam.known regions including the vastly un. and in such cases. lantic is found along the eastern coastline and extends from Europe to North Africa. oceanography. though they are thought to function externally and de. trilobites. ule armament and physical defense in temperate 98•Chapter Five wan: outbreak of a cyanobacteriosponge. Serpulids are 2 mm to >100 mm long. 1–5a. and Palardy JE. 1999. 158 Mollusc Shell Structure. 140•Chapter Six McFadden CS. arguably THE most important role in the biological breakdown of carbonate on many coral reefs. Microbial Ecology 55: 569–580. not necessarily specifying those that etch or tunnel. plate. Bioerosion and coral. Nitrogen fixation and photosynthetic oxygen evolu.” whether oceanic or continental. Pernet 2003).01% of the surface. the endolithic com. times in large numbers.coral calcifica. Indonesia. cemented reef. respiration and ATP production are considerably higher in these cells when exposed to light. 2008 2009). Nature 431: 56–59. Ferreira CEL. Morphologically similar but smaller stromatolites 0. The kidneys may be reduced to a single ante. factors that control the forma. Fabricius K and Alderslade P. 2006. or habitat structure may favor corallimorphs. For example. Large colo. Science 324: 50–53. rayed pentactines in some species. a new coral species from Brazil. constitute a barrier. Berlin: Springer. Faulkner DJ. a group of more than 1. The large shells are brachiopods (Chapter 9). often built by a single spe. chemical solution may constitute a significant proportion. 2010. diploid spores are released that germinate. ed. ter representing the area occupied by the coelenteron. 2005. 2006). and critique of management strategies. 1985.. Gigantospon. geography. cies grow by extension of length as well as thickness along multiple axes. An energy budget for Porites porites (Scleractinia). described below. Fringing reef growth and morphology: a review. Multicellular algae. For example. thelial cells with myofilamentous bases. The demosponge skeleton is composed of noncrystal. Isoprene is the building block of terpenes that exhibit ichthyotoxic and antimicrobial properties (after Gaspar et al. Coral Reefs 4: 221–224. but rhodophytes and chlorophytes are rec.Pacific. It has been suggested that such control may be exerted by different concen.ecsd. tapered as needles.dwelling algae and cyanobacteria that etch or pit carbonate substrata are epilithic microborers and are distinguished from endolithic microborers that form tunnels and cavities. the GBR was designated a World Heritage Area by the United Nations in 1981. lagoon water whose quali. cess in cyanosponges. dance. Calcification in the green alga Halimeda. gamete form. University of Queensland. tively form an “inhalent” surface. Historical contributions notwithstand. Lagoa Salgada (Rio de Janeiro) re. Cryptonemiales). when it reached a substantial fraction of what it is today (see Anbar et al. but this group occurs worldwide throughout the tropics and subtropics. foraminifera. the crest is typically colonized by low. A functional group approach to the structure of algal. 2004. Journal of Pale. Layered Microbialites Microbial carbonates called microbialites are well. ization.shaped fertile segments. not the hexagonal system as are the carbonate polymorphs previ.like system over 200 km long. mucus cells. Journal of Ex. tors including local Ca:Mg ratios in seawater. halimedatrial and halimedatrial tetraac. Santa Barbara Community College. building properties. although they also occur in the polyp tissues around the pharynx and even in the tentacles. Cold. Reviews of Mineralogy and Geochemistry 54: 189–215. Few corals can grow and com.section through the axial skeleton of Ctenocella schmitti (see Bayer and Grasshoff 1995). they smother corals in shallow water. more heavily calcified segments that have less nutritional value but are also less repugnant. two polychaete families employ a mucus filtra. lids. among many other invertebrate host groups. reefs that fail to keep pace with sea level rise “give up” and are drowned by increasing water depth (after Neumann and Macintyre 1985). Banks KW. but may re. 1997. in the tropical eastern Pacific. Part E. vironment for their beauty. hypercalcified skeletons. 2002. Ma. This tight cycling of organic and inorganic components is thought to contribute to the success of the coral symbio. Goldberg. The rhizoidal network is significant not only for anchorage. ———. which in calm weather may extend for several kilometers and may persist for 20 hours or more (Figure 6–32). Borowitzka 1982.Cuetos L. 2004. Circular DNA and prokaryote. 2003. coralline algae. the polyps may share a common wall and produce closely packed corallites (Figure 6–30b). These in turn may be interspersed by a third cycle of 12 smaller tertiary septa. Photo courtesy of Pe. Food captured downstream by sabellids is sorted into three sizes by lips that surround the oral region and join the bases of the radioles. 1990. Close views reveal flattened sand grains lining the inside of the tube. Diaz MC and Rützler K. but these struc. Macroalgal feeding characteristics of the sea urchin Diadema antillarum Philippi at Discovery Bay. come primary reef builders. The two most common of these are agar and carrageenan. Piller WE. dermite and constituent crystals are associated with or.NOAA. ———. Alternatively. peristomium. and light reduction. likely the product of symbiotic zooxanthellae. Biogeosciences 4: 219–232. Penicillus is a psammophilic genus found in seagrass beds and sand flats. Zooxanthellate corals typically continue to produce carbonate in the dark. Marine Biology 142: 419–426. In: Riding RE and Awramik SM. Scoffin and Bradshaw 2000). Grigg RW. One. acteristic dead zone around the sponge. Photo cour. These elements may be more or less continuous with those in shallower water. work typically composed of spongin fibers.shelf islands of the barrier lagoon. Herbivorous fishes and invertebrates graze heavily on turfs resulting in low biomass. This is apparently mediated by a preexisting extracellular organic matrix composed of the cellulose wall and its associated acidic polysaccharides (references in Adey et al. erous canals before they escape in the excurrent flow (Maldonado 2006). and bacteria) a. The eggs may be released as a zygote or as planulae. The primary character that unites these diverse forms is the absence of any supporting axial skeleton of any kind.containing form. Zehr JP. Lewis JB. they are found in seawater of normal salinity. Croix reefs began to re. Invertebrate Biology 123: 23–42. 2002b. which may include a range of unusual peptides and alkaloids. some of which have been intensely fished by commercial trawlers. but are becoming understood by examining living forms. whose feeding will be described in more detail elsewhere in the chapter. Most polychaetes are free. Freile D. Well.developed patch reefs in its lagoon. HMA sponges have a denser mesohyl and a more complex aquiferous system composed of longer. ters or more. corals are capable of calcification regardless of light conditions. Ravilious C.km2 area where the bottom was covered by deep pools of its calcareous seg. but all of the interfilament spaces become calcified in mature segments. The photosynthetic environment is dynamic and therefore the dependent process of calcification in Halimeda lacks daily and seasonal uniformity. There is considerable debate about the transition from this stage to the final step in stromatolite cementation. Gladfelter EH. Thus. als Acropora cervicornis (center) and surrounding elkhorn corals Acropora palmata have been the primary reef builders in the Caribbean for at least the past 125. ments across the substratum. where magnesium substitutes for some of the calcium in the crystal. However. cepts in Bioinorganic Materials Chemistry. 2006). Other acidic polysac. and individual mats (Figure 6–9a) can grow to 30 m2 . and Muller WJ. they a b Figure 6–23: Hermatypic octocorals: a) the organ pipe coral Tubipora sp. are found in the intertidal zone on several of the Exuma Cays (Reid et al. bacterial symbionts of marine sponges. Sperm and eggs may be released synchronously from different indi. along with hemispherical forms on the sides and the top. MAR. 1986. New York: Kluwer/ Plenum. b) A dihexagonal dipyramid is formed when the sides of the dihexagon are bent at the plane of symmetry. NASA PHOTO and Reefbase. These collec. Atolls associated with these reefs develop at the northern end of the archipelago. geography.marinespecies . Inset: Close. miniferans.like growth chambers (Figures 6–19. and even when attempts are made to allocate carbon to these diverse compartments. buttress and channel structures become more variable. Phycologia 35: 308–326. cation of modern marine stromatolites. and assuming the form their hosts had when they were alive (Wahle 1980). but less spectacularly. et al. They lack a digestive cavity. note lens cover. 2001. ecology. 2003. copiosa and the Caribbean endemic H. tent. Annelids and Sipunculans•147 an organic matrix that attaches to the substratum. but its growth ceased within about 3. in addition.. along with many alcyonaceans and other organisms. and are found in tubes with the ventral side fused to the substratum. cult to separate the two groups ecologically. Godinot C. water Research 53: 79–84. Alácran reef. Figure 4–19: a) Marginopora vertebralis. From their initial appearance beginning about 3. thid Palythoa caesia. The Status of Coral Reefs in the Western Atlantic and Gulf Rapid Reef Assessment (AGRRA) Program. other algae may dispense with proton cycling as described later in this chapter. micritization is super. while significant as a taxonomic tool. Meylan A. 2005. whose demise could result from overflow of its own cyanobacterially rich.off. these early Holocene reefs were killed ~9 kya.. destroy tissue. verse) described earlier. the nematocysts in many others are more benign and asymptomatic. This is referred to as light. Marine Ecology Progress Series 417: 97–120. Tectonic Forces Large atoll groups like those described above are associated with chains of inactive. lution of skeleton. and shape (e. whereas omnivores feed on live animal and plant material. 2001.Emperor hotspot track: inheriting the mantle wind. Indeed. suggest that despite considerable morphological diversity (described below). Botanica Marina 48: 116–121. intensely pigmented areas where photo. onions. Palaeontology of sponges: a review. mum to the present was marked by climate changes that included warming followed by sudden temporary reversals in temperature. 126•Chapter Six confined to the tropical Indo. Inset: Dead segments (white) can form on living Halimeda plants owing to a variety of processes. but unlike millepore cor. Perhaps because of the delicate and vulnerable con. Coral Reefs 22: 99–106. is enjoyed raw or fried with butter. and although there are few quantitative studies. Marine Biology 152: 395–404. tion among corals (see review by Fautin 2002) include polyp expulsion. Riding 2002). ing the largest extinction event ever recorded (Chapter 13). the medusae are released synchronously in several pulses within a period of a few weeks during spring or summer (Soong and Cho 1998. 2000). following low tide. whereas others are associated with specific hosts or environments.5 kya when sediment that had accumulated in the lagoon was transported seaward by rising sea levels. and depth. Benavides and Druffel 1986). cluding those that function as bafflers. Published 2013. despite their contri. pigmentation. Chlorophyll a. In some cases corals can be overgrown by neigh. a spe. Coral Reefs 1: 139–150. Different species of Halimeda cluster into 5 lineages that are characterized in part by distinct patterns of filament configuration (Ver. als and other anthozoans. Berman. who not only gave his time freely. Curry JD. and (paleo). Similarly. and develop within specialized skeletal chambers called ampullae. less than 10 cm in height. Studies suggest that these skeletons form very slowly. Hughen KA. tute of Marine Research. arbo. 1968. for example. wherein the polyp is halved and regrows as a separate individual. A rocky flat Figure 1–3: Satellite photograph of Mayaguana Island (Bahamas) with its adjacent fringing reef. ada. Submarine lith. Furla P. 163 Bivalves. whereas fungi may produce spiderweb. Wa.axis. siliceous spicules that are mineralogically similar to the hexactinellids and are formed in a similar fashion (Müller et al. Littler DS. pear as if they formed from an extratentacular process (Figures 6–29e. (a) and Udotea flabellum (b). phous hydroxyapatite is deposited (Lewis et al. Grange KR. other bacterial symbionts. are 75 km or less from shore. Israel A. Nature 457: 718–721. 2006. Marine Biology 49: 147–159. a massive network of sto. How. ventral (C) and lateral (D) views of 30. With the provision of a complex framework. ———. a b 112•Chapter Six on this hypothesis (see Medina et al. The zygote develops into a second stage called the carpos. or are the result of asexual reproduction. Seaweed adaptations to herbivory. Those producing calcified segments Figure 4–15: The triphasic coral. a series of even older.Pacific and Atlantic spurs and grooves revisited: the possible effects of different Holocene sea. a solid is formed through a process called nucleation. and Conservation. 1956. Thus.g. retention of the eggs is common and may be accompanied by shedding of free. Interocean differences in size and nutri. Maury RC. ary metabolites of the cyanobacterium Microcoleus lyngbyaceus and the sea hare Styocheilus longicauda: palatability and toxicity. 1983. Great Barrier Reef. and larval biology in scleractinian corals. Williams D and Hallock P. or other colonial morphologies. and Hua T. 2001). nal of Experimental Marine Biology and Ecology 163: 277–286. Journal of Phycology 38: 4–10. Sea level 10 m . Precambrian sponges with cellular structures. tion SEM study. and Becerro MA.org/porifera. New York: Oxford University Press. sperm and eggs within a bundle) compared with the ability to successfully fertilize gam. Figure 6–19). ous red algae (up to 29 mol%) and echinoderms (up to 17 mol%). nal muscles may allow for differential contraction of the pharynx and tentacles. Flatt PM. and Ralph PJ. Polychaetes are named because of their nu. 2010. tinian coral Fungia scutaria. and direct burial by beach restoration proj. serpulids are not active borers.like. Variation in coral growth rates with depth at Discovery Bay. 2006). ria: molecular. among other members of the same genus the oldest and largest specimens have been found only on dead corals. ed. roalgal communities of the northern Florida reef tract. indicated initially by tentacle buds (f). Anthony KRN and Fabricius KE. et al. its muscular movements. Santiago. to name a few. genic condition of the giant barrel sponge Xestospon. allowing maximum exposure to sunlight.like filaments and is referred to as a capitulum. This group was prominent on mid. zler 2001. Flow is induced by the movement of choanocyte fla. logical and genetic evidence of the vulnerability of the stylasterid hydrocoral Errina novaezelandiae in New Zealand’s fiords. ous Porifera from the Permian reefs of south China. ics. More open atolls with multiple channels are common here. lary pore in the skeleton. and is circumtropically distrib. The rate of lateral advance into host tissue may be only about 1 cm yr–1 . can co. 2001. Zoantharians and Corallimorpharians Zoanthids and corallimorpharians are colonial an. European Journal of Phycology 34: 371–379. Some gorgonians produce a heavily calcified axial skeleton. Bleaching in Amphi. Conversely. a specialized part of the nacreous layer where adductor muscles are inserted and anchored.). Nether. we will use the traditional taxonomic color groupings to discuss the roles of these organisms on reefs. and de Beer D. Loss of Biodiversity. 2005. seven of them appear to be unique to the Western Atlantic (4 Caribbean and 3 found only in Brazil. 2003. Hyella. and Frankignoulle M. Niau. Using radiocar. In contrast. 1997. minifers: Miniacina. Animals that engage in bioerosion are macrobioeroders and can be distinguished by their feeding habits and hab. –150 140 120 100 80 60 40 20 0 –100 –50 0 Time (×1000 years) before present Relative sea level (m) Pleistocene-Holocene boundary An Introduction to the Structure and Formation of Modern Reefs •27 a crest of coralline algae and encrusting corals.book) 1. position of calcareous sponge spicules: a review and comparison to structurally related materials. b) Scanning electron micrograph of Lithothamnion sp. These Addadi L.e.. unequal sides. Nature 441: 714–718.bound compartments (vesicles) that are anchored to scaffolds such as the cell wall or an organelle. sea anemones constitute a large group of uncalci. provide shade. 1989. Rennell Island in the Solomon Archipelago. (Peyssonneliaceae. Figure 7–2: The amphinomid Hermodice carunculata is most often a predator of cnidarians and is shown here feeding on the orange tube coral Tubastraea coccinea (Scleractinia). a subject that will be described later. ample. 2005.producing. the green layer at the top of Bahamian stromatolites is due primarily to these thickly sheathed cyanophytes (Figure 3–15a). Douady CJ. 2002). ment rates and gene flow of broadcast. toke swarming is a classic example of coordinated mass spawning (see Bentley et al.g.rayed monaxons. 1992. Note underlying aragonite crystals (A. Impact of crustose clionid sponges on Caribbean reef corals. and are of environmental significance because of their symbiotic relationship with another group of protists. lated. but some groups that oc. in particular. Precious corals in Hawaii: discovery of a new bed and revised management measures for exist. ergetic reefs developed as low mounds. tion to chlorophyll a (as in eukaryotic plants). Growth lines are often visible in this region. tens of centimeters wide. Kingsley et al. or heterotrophs (e. Goreau et al. 2007). Beuck L. an island. 2009. ample. Titlyanov EA. et al. matically (Furla et al. These shells are not perforated and thus the cytoplasm projects from the aperture. especially filamentous red. Indeed.in. 1997. Thus while turfs are common elements of reef surfaces. bic crystals have three mutually perpendicular crystallo. and are selected for tube construction. In an interesting twist. but simple or branched filaments that are common on various surfaces (e. 2003. Thus. pali and collumella are well developed. siderably according to both taxonomic position and environment. Figure 1–16: Chuuk Atoll (formerly Truk) in the Caroline Islands is a good example of an “almost atoll. The coenenchyme is also the principal site for the production of sclerites. However. Marine Biology 154: 973–984. These may include volcanic activity. Stoddart DR and Steers JA. phy and Marine Biology Annual Review 48: 161–212. 2005. Maragos JE. Lastly. ler M. 1995. or enzymes like carbonic anhydrase (see equation 4–3. both chemical softening agents and mechanical abrasion have been implicated in different sipunculan groups. pete for space among themselves and with other reef organisms. Cohen Y. 4–1a). Spicules referable to the demosponges have been found in deposits from southern China that are ~580 My old. mer. Journal of Experi. in.hunting divers. while they are more often ellipsoidal (Stoddart 1965). cal. These morphs do not become dominant simply by virtue of their low pro. or a shallow oceanic platform.like forms are quite common in shallow waters of the Florida. which mediates the polymerization mass than any other type of sponge (Leys et al. The evolution of mating systems in tropical reef corals. Z. Such serpulid reefs.Bunker Group. cially to the north along the continental shelf. Chicago: University of Chicago Press. as is their construction by individual cells. Memoirs of the Queensland Museum 44: 667–673. Science 221: 1175–1176.500 km.200 liters of water per minute or 72. nar. defensive chemicals are generally considered to be more effective in reducing predation pressure as described below.. Geller JB. or supple. Two other types of atoll structures (raised atolls and almost atolls) are described below. Riosmena.Cuetos et al. 1932. Losses to the system represent de. Hopley D.proton exchange mechanism (two protons for one Ca++ ). and Sharp K. non. All skeletal growth reflects some combination of ex. whereupon morphological complexity. Notes on the shallow water gorgonian. Scale ap. tends northward about 150 km from its northern bound. and Jaap WC. sium. duce their own extracellular polymers. Mexico. there is some overlap among morpho. a group derived from five. Structural biology: choosing the crystallization path less traveled. However. or by competition for growing space. molecular analyses of phylogeny do not strongly support those morphological distinctions (e. and perithallus (per). Coral Reefs 16: 247–259. In contrast. gia. Lifespans and growth patterns of two deep. such as sea urchin spines. astreoides as it appears without X. Leão ZMAN and Dominguez JML. or branches (redrawn from Hill 1972. sity II: Understanding and Protecting Our Biologi. This placement. Fukunaga A. 2008).g. Photo courtesy of Paul Humann. in this case. In contrast. about 8 cm wide.like depression called a lagoon. Calcinai et al. mism. lack flagella and live within the cytoplasm of their hosts where they provide them with nutrition from photosynthesis. nadian Government recently declared these unique reefs “Areas of Interest” for possible future inclusion within a more formal Marine Protected Area (see Chapter 16). but they are most abundant on deep reefs and seamounts where they may constitute much of the three. Discovery of the novel candidate phylum “Poribacte. and Environment 31: 679–694. nite crystals produced at night form a new but lightly calcified series of sclerodermites that result in skeletal extension. Finks RM and Rigby JK.” some authors sought to distinguish prominent reef ridges that grow by active accretion rather than by erosional sculpting. Van Soest RWM.5 Gya. Berkeley: University of California Press. not all nematocysts are toxicants.E. their location on or within the carbonate. hollow cells called porocytes (Figure 5–1) are scattered among the flattened surface cells. Changes in skeletal density over time (carbonate weight per unit volume) are distin. the same species may develop different ecologies that result from combinations of sexual and asexual modes of reproduction.Alsumard T. The relative proportion of each is a function of pH. Darwin C. but also to influence the specific crystal form as well. Cladistic analysis of Medusozoa and cnidarian evolution.” and to the latter by growing “downward. Similarly. Figueiredo 1997). Tomitani A. 1999. The entire field is ~5 µm across. Hiti Atoll is similar. mers that often directs the process of ion accumula. proximately 1. 1983. 1999. 1965. laris species complex. hence the common name “elephant ear” corallimorphs. where such reefs have received the most attention.like branches called stipes (Figure 4–1e). The subtidal forms grow as large columns up to 2 m high in depths of 7–8 m (Figure 3–14b). sive. rational processes are evident in many of the older high islands of Hawaii where fringing reefs are well devel. often including cellulose microfibrils (Nobles et al. which grow using another animal for support (in this case.5 Mya corresponded to significant environmental changes and the appearance of new carbonate. calcification. Environmental Microbiology 6: 1096–1101. Likewise. the matrix appears not only to control crys. Indeed. Enamel in fishes is distinguished as enameloid due to the pe. tropores. Calcareous green algae of the genus Halimeda have persisted for 150 My and now contribute to sedi. stratum on shallow or intertidal tropical reefs worldwide (Figure 7–9). karyotic cell’s membrane. The margins of such islands may also be partly or completely rimmed by mangroves in the intertidal zone. the reef crest is separated from shore by a basin. mostly demosponges. or on deep slopes (e. Chaves. The zoanthid genus Gerardia is one of two groups called “gold corals” that typically occur in waters exceed. Sponges harbor a diverse array of bacterial symbionts including cyanobacteria.Román A. however. in some cases all the way back to molecular nitrogen (Figure 5. 2002. and additional cycles may double the previous number. 2003. each can recreate the specific mineral from a supersatu.shaped structure with two arms (palps) just below the lips called the building organ (Fig. shelf. Proceedings of the 9th International Coral Reef Symposium. Even in the clearest ocean water. The thallus of some epilithic species can also drape over reef edges. Sedimentology 46: 985.specific density of symbiotic dinoflagellates in tropical reef anthozoans. phology and sedimentology of Halimeda bioherms from the eastern Java Sea (Indonesia). tion is another important choanocyte function. get calculations include: • Reef size and development: How much of the reef is covered with a framework? • Growth rate: How much of the gross carbonate production is incorporated into the framework as bricks and mortar? • Sediment production a: How much sediment is produced by reef organisms? • Sediment production b: How much is the frame. Evidence of cyanobacteria. 2007. In addition. Catch Up. Fautin and Allen 1997). This species is often larger than a centimeter in diameter. In general. most of the others protect the cell by absorbing harmful ultraviolet light. logically and structurally significant for reef develop.Tuvalu group). Macroendolithic bioeroders b. and nutrients on the community calcification rate of a coral reef.topped “table reefs. With rare exceptions. various models suggest that the actual number lies somewhere between 300. shallow. 2004). These symbionts migrate to the endoderm of the planula (e. Carpenter RC. 2001). tem may not be an adequate means of assessing species boundaries (Meroz. their habitats are often unprotected from destructive deep. Interspecific aggressive behavior of the corallimorpharian Corynactis californica (Cni.drained platforms (Fabricius and De’ath 2001). ity are shown at right. herbivores. Stoddart DR. Wood R. additional crystals eventually fill the extracellular spaces within the cortex and produce a mature segment.molecular. rotenoids perform a photosynthetic function. Barrier Reef. usually called the tetrasporophyte because of their propensity to form clusters of four haploid spores. nal brooding described below. Chemical warfare on coral reefs: sponge me. French Polynesia. cations that require water retention. 2008. sometimes threadlike suspension. generally reflecting the overall pattern of the utricular surfaces (Figure 4–8 a). Molecular Ecology 17: 1840– 1849. the Bahamas.. with approximately 15. fragmentation is another form of asexual reproduction in which the colony separates and continues its growth as two or more clones. Journal of Animal Ecology 74: 313–321. Trichodesmium. 2006. Guam 1: 713–718. its contribu.Fonnegra A and Zea S. Sea level change through the last glacial cycle. fin and Bradshaw 2000) whose growth may be regulated by that of their hosts. 363: 3293–3308. cialized thoracic glands as calcium carbonate. They are calcareous. cious—that is. Analysis suggests that zooxanthellae in clionaids cluster within the same genetic lineage or clade (clade G) as certain fora.moving (vagile) species.g. Chlorophyta): fleshy vs. Florida International University Patsy Mclaughlin*. Tropical coast of Brazil. 2008. there are no coastal coral reefs known from northern Brazil to Venezuela where the Orinoco and Amazon rivers discharge. bryopsidales): a Cretaceous genus that diversified in the Cenozoic. eastern Pacific reefs are influenced by a complex regional ocean.associated enzymes that mobilize and pump cal. including sea urchins.90. they may develop exten. Foraminiferans constitute a heterogeneous group. Effects of herbivory and nutrients on the early colonization of crustose coralline and fleshy algae. 2002. ing selectivity among herbivorous coral reef fishes. fission. and surrounds a number of inshore fringing reefs. USA). different types of carbonic anhydrases may be located on the cell membrane. mann. Smith GJ. especially the reef crests with their characteristic growth of calcareous red algae (Pandolfi 1996). The lagoon is located within the protection of the reef’s rim. Thus. However. late or AZ.Penney SJ and Racey A. Here along with branched. or they can be overgrown by mangrove forests or by reef growth. let may include them as a significant part of their diet (Horn 1989.gov). b) Pink coral sculpture of Fuku. and to Richard Walter. spectives. but they can vary both quantitatively and qualitatively among hosts. Unlike their tropical counterparts. Based on Nicol 1931. and northwestern Africa (Figure 6–26). which currently sits atop the hotspot. or a chitinized lower lip that rasps tissue. The deep. here and there surmounted by a low verdant island with dazzling white shores. in turn. and more closely resembles plastic than any natural substance. Facies 56: 173–177. However. indicates that their evolutionary origins were separate from other sponge groups and from multicellular animals (metazoans) in general. 2007b). Symbiont bearing. and López. and longitudi. they resemble those of chloroplasts in plants. Physicochemical fac. whose components are produced by specialized glands that open into the building organ. Pawlik JR. Miami 2: 15–21. including branching. Garcia. low water temperatures can occur within the tropics because of the influence of cool eastern boundary currents. Kuguru BL. Anbar AD. Thus. Plants. for example. Three cell regions can often be distinguished as fila. Leys SP. they suffer high mortality owing to a variety of anthropogenic activities. NZ. This polymorph is very uncommon in marine organisms. And thus. as well as bloom formation and the synthesis of toxins. Asexual reproduction in solitary Scler. duce a wide variety of chemi. Correlation of boron isotopic composition with ultra. ral levels of turbidity vary by locality and may decrease the depth at which reef growth takes place. lationship with dinoflagellates belonging to the genus Reef Corals and Their Allies•119 disposition of protons. ously described. and smaller lateral corallites. 2003. sistance is not a feature of siliceous sponge reefs (Chap. but may not necessarily be calcified. a far smaller one surrounding San Andrés and Providencia islands in the southwestern Caribbean (Milliman 1973). with up to 50% of the sponge’s energy budget and 80% of its carbon budget derived from photosynthesis (Wilkinson 1983. 337 16 Reef Resilience. heavy surge and exposure at low tide).like appearance. dant (e. 207 Decapod Crustaceans. Pitlick TJ and Paul V. 1991. as is the case for Aphrocallistes vastus. Bavestrello G. bonate saturation. dition.. fusion and differentiation of archaeocytes. 1982. sponges produce interlocking spicules that achieve the effect of a unified skeleton (see review by Uriz 2006). and turtles (Meylan 1988) routinely prey on them.living cyanobacteria. Thus. In addition. There are several distinct processes by which clones are produced. The film bridges the spaces between sand grains and acts as a sediment trap.developed spur and groove system at lower right. even though some have classified them differently (Fauchald and Jumars 1979). cus spongarium does not fix nitrogen (Usher 2008). Algal size and shape often correspond to ecologi. glandular warts. 2006). or in protected areas such as bays. Free spicules: pinulate hexactine (upper left). However. and mushroom. and the septal ends may fuse in the middle of the corallite forming a colu.30168.crystalline organic envelopes have been noted during the mineralization process in Halimeda by some. Reef Sponges•85 Zooxanthellae are thought to improve the growth rate of these sponges as they spread laterally. Aquatic Microbial Ecology 51: 45–53. Turon X. Serpulids are not physi. Dubois et al. 2007. Hypercalcified sponges. and Dubinsky Z. caribaeorum in the western Atlantic) exhibit growth rates of up to 4 mm per day (Suchanek and Green 1981). These circulate water into and out of the coelenteric cavity and have important roles in the maintenance of internal pres. secrete. coral reefs are the economic engine for many tropical nations (Moberg and Folke 1999). bonate test blocks placed in water 100–275 m deep (Gek. Giant subtidal stromatolites forming in normal salinity waters. Mills MM and Sebens KP. cals with an equally wide variety of functions. they settle on cal. ing into a polyp. bite and actively remove carbonate as they feed. 1983. International Journal of Tropical Biol. 2005. Scully EP. which is then molded into place at its anterior margin. Sessile Foraminifera The abundant forams described above are unattached and free. although gemmules are also produced Figure 5–2: Scanning electron micrograph showing part of a sponge choanoderm. amoeba. Phylogenetic im. 1992. the surface. 2007. the pharynx has a rasp. pora vertebralis. Figure 3–2: Details of the cyanobacterial cell. versity Press. northwestern Gulf of Mexico: controls on the distribution and growth morphology. Coral Reefs 22: 433–446. Sexual Modes There are two general types of sexual reproduction among corals. which are conspicuously bright yellow. and is therefore a US ton plus 12%. Brümmer F. commonly referred to as feather. xanthophylls. and 16 ATP just to reduce one molecule of N 2. Alternatively. and Braga JC. I spent an incalculable amount of time gathering images and seeking permission to use them. A newly discovered method of reproduction in gorgonian coral. Garcia. Neumann AC and Macintyre IG. without fertilization). nicholsoni. More protons are produced (2). tracted individual at right is a species of the genus Fungia. tropical waters have a symbiotic re. Red Algae With 4. or they may be absent if the lagoon is closed to oceanic water exchange. miales) in the upper fore. Bio. Different groups of organisms often form distinct types of matrices. This intimate physical association facilitates the interplay of host and symbiont metabolism. 2007. Soong K and Cho LC. Over 12. tance of frame. and similar amounts may be found on reefs in the Florida. crustaceans. 1998. Piel J. Thus. Achituv Y. Modified after Meinesz 1980. b). the tube of these serpulids is typically covered with living host tissue. S. Woodroffe CD. Rigby JK.magnesium cal. Fiege and Ten Hove (1999) have shown that this genus exists as a complex of 10–12 closely related species groups. 2006). Gorgonin. bioactive fatty acids.than. and permanently attached plants and animals that form the scaffold of the reef system and allow for the attachment and growth of other organisms. tide pools (right). taliina. These fuse to form a zygote that elongates into a “protosphere” with a large cytoplasmic volume before it settles to the bottom and produces fine. still at the grain surface. and many other invertebrates) living in shallow. Rosalie. Skeletal architecture and microstructure of the calcifying coral Fungia. lelochemicals. Indonesia 1: 515–520. These can dominate environ. In some cases. High diversity and host.Org. Centre Scientifique de Monaco Rüdiger Bieler. Certain groups of fishes constitute an important guild of spongivores. bacterium Synechococcus spongarium among sponge hosts. The inside is now alkaline and conducive to cal. Protons generated by the conversion to carbonate can accompany bicarbonate across the cell membrane. larger algae tend to be long. 2005. Thus. Metabolic wastes produced by the capture and digestion of food are not lost as coral resources. a level that is tolerated by few species. Proceedings of the 7th Inter. Soft. 1999. a process called klepto. release of sperm is often distinguished from broadcast spawning as spermcast mating (Bishop and Pemberton 2006). especially in well. chemical as well as life history characteristics. Indeed. while larger objects are rejected and drop into the surrounding water. ceedings of the 3rd International Coral Reef Sympo. in turn. where they formed vast depos. tension and accretion. this structure is used for tube construction. Reef Corals and Their Allies•109 Asexual reproduction is by no means universal. Figure 3–8: Various growth forms of cyanobacteria. 2011. thus contributing significantly to reef sediment production.pdf Murray JW. Pagés J. However. noes called seamounts that often rise from the ocean Shallow spur and groove topographies are pro. ductivity. Evidence suggests that populations of C.” and indeed some Tuamotuan atolls are composites of as many as 300 such cays. Evolu. curs internally by males that engage in spermcast mat. Likewise. Henry RP. turbid waters. Tribollet A. Photo by author. lization. They may be open at one or both ends. Blanchon P and Jones B. The body is subdivided into numerous folds of gastrodermis and mesoglea called mesenteries. Petit M. (Gerardia sp. each of which is truncated to form a hexagonal dipyra. New Mexico. Moreno EC. parchment.point dried SEM preparation of a decalcified Halimeda segment showing the arrangement of the central medullary filaments (mf) and cortical filaments (cf) that become branched as they approach the surface of the segment.axis is la. sion over sexual reproduction. 2007. Much of the confusion is due to the terminology that attempts to distinguish reef. low islands extends about 2. lations often found in sponges are one group of such consumers. oped ~10–11 kya as a barrier reef. which are all obligate anemone symbionts. although other forms that are more complex are found in some species. before reaching volcanic basement rock (Figure 1–15b).. including six. The mouth may be accompanied by paired or multiple feed. Photo courtesy of Paul Humann. Such conditions can give rise to a well. although hidden in caves. only the haploid gametophyte is well devel. produce aragonite needles. Biomass and sec.like sensory structures called cirri. the ratio of organic matter to carbonate is increased. Stud. Tribollet A .g.like core composed of the enzyme silicatein.Org.like. Sponges.positive forms. However. Godinot et al. and Raven JA. zil. Biodiversity and Conservation 11: 851–885. calcareous sponge groups col.HDTV. The Calcarea produce no dis. ily or exclusively of sclerites that are completely fused to one another. An additional type of defense is afforded by the ability of some sabellids to sever the cartilage that supports the radioles if they are stretched or pulled.containing enzyme that serves to reversibly catalyze the conversion of more abundant bicarbonate to carbon dioxide. or in the coenenchyme between polyps. Keats DW. 1995. these do not form natural groups of organisms. organisms that form arborescent or encrusting colo. including the oldest of them. tribute to the cementation process in the construction of Halimeda bioherms and coral reefs.like plate that can be extended like a tongue or rotated internally to grind its food. and settlement. Georgios T. where they occur regularly. creeping across the substratum. High. 2006. “Everyone must be struck with astonishment. mucus.Williams R.like fusulinids up to 8 mm long are abun. eous spicules and a few produce massive. and they vigorously defend the host from competitors. b) “Downstream collection” in sabellid polychaetes. and some within the genus Acropora.fracture. However. Skeletal development in Acro.enclosed space where aragonite deposition occurs (Okazaki et al.dimensional structure of the substratum.Pagès C. Henkel TP and Pawlik JR. These include sclerocytes that secrete mineralized spicules. Weiss MP.. Wainwright et al. all of them died at the end of the Permian 251 Mya dur. funnel. 2006).relief islands are crowned by members of this salt. Endolithic cyanobacteria are also important pri. 2000). feed on species of Age. Fora. linida (Mississippian to Permian.Parker G and D’Elia CF. Because asexual reproduction is energetically cheap. the Caribbean) that may be a reflection of the taxonomic uncertainty in this group. phytes are not significant for coral reefs. Is competition for space between the encrusting exca. In addition. Journal of Phycology 35: 24–34. las sponges reduce depolarization. c) polyps may separate after intratentacular budding as in Dichocoenia stokesi from Florida. Carbohydrate is stored in the form of floridean starch that is more highly branched than other starches and. although it appears that carbonate. Courtesy of the Image Science & Anal. 1963. Zale and Merrifield 1989. Smithsonian Institution US National Museum Florentin Maurasse. line algae. 1987. axonia. The Timetree of Life. referred to as broadcast spawning. including calcare.0. live unattached to the substratum as adults. unlike tree stumps or peat deposits.) west of the island of Hawaii at 380 m.oral prostomium fused to a post. Zooxanthellae may provide several key products or conditions that influence light. Sethmann I and Wörheide G. for example. and in some cases. 2007). which is likewise calculated on a time and area basis.less spermatia.5 m in diameter and up to 1 m high. can adapt to as little as 0. This feature unites the classes Hy. algae. 2001. ered in 1956.depleted waters. both of which play a role in shaping limestone coasts above the tide (Schnei. The boring microflora in modern coral reef ecosystems: a review of its roles. Plant. Invertebrate Biology 125: 177–194. Figure 1–18: Differential effects of subsidence and tilting can produce different reef types in the same geographic region. plex and toxic brominated compounds found in the ge. This process (sometimes referred to as fission) is particularly com. Biogeosciences 2: 61–73. arborescent skeleton. et al. and Gosline JM. Perry C.Moghrabi SM. Rougerie et al. Cyanobacteria can produce up to 50% of the sponge’s energy budget and 80% of the sponge’s carbon budget through photosynthesis. form. Ecological Research 20: 529–535. 1983.Accumulation of coralline debris B a s e m e n t ro c k 20•Chapter One growing on the protected sides of these islands are often comparatively well developed. tophyllia reesi. Alternatively. some groups produce calcite crystals only externally. shallow. 2000. ing their formation (e.” and others that protect larger sexually reproductive individuals be. Photo courtesy of NOAA Ocean Explorer. Druffel ERM.Water Corals and Ecosys.offs in resistance to competitors and predators.like galler. Although the sponge induces peripheral folding of the colony (Figure 5–9a). Many geo.shaped Onotoa and V. merous. Rinkevich B and Loya Y. Earth Science Reviews 58: 163–231.like prisms. perhaps 1–2 kilometers wide or less. their biological characteristics are poorly known. where they found reef material that could have grown only in shallow water. More highly integrated colonies can respond to stimuli at greater distances from the source. although the geological origin of these islands is not clear (Clouard and Bonneville 2001). 1991. ates and planktonic phototrophs. per. microscopic sclerites. Wörheide and Hooper 2003). (1992) has classified such matrices into four categories. These cells may form a continuous layer inside the sponge forming Sponges play many key roles on reefs. Márquez et al. Bleaching and stress in coral reef eco. all of the polychaete feeding types described above fall into this category.Roubaud et al. Figure 4–16a).water coral ecosystems. silica sponge spicules may ac. Richter C. a b Figure 4–20: a) Encrusting Homotrema rubrum tests under coral. either by accumulating sediment near attachment sites. tions of the cortical spaces.2 mm) thick can be deposited and trapped in one 24. Zoologische Verhandelingen Leiden 345: 313–336. Environmen. 2006). Many of them are distinctly neither U.water areas (e. and the “average” or baseline symbiont density for corals under normal environmental conditions is widely accepted as ap. struction (Figure 1–1a). is derived prin. Mya. 2001.Fine et al. and was produced primarily by coral calcification. However. 88•Chapter Five is surrounded by a star. Photo courtesy of Xavier Pochon. alline algae on the Great Barrier Reef. ously from supersaturated solutions of ions. the clownfish Amphiprion clarkii associates with all ten giant anemo. may favor colony fis. rophytes. Part 2. although the cues that might induce settlement en masse have not been identi. A single chromosome is typically present as a long. From what little is known about the reproductive process. This speci. Photos courtesy of Paul Humann. Likewise. 1985. curs below the surface due to a succession of bacterial communities. Clode P and Marshall AT. However. NASA AQUA satellite image. tending from east central Florida to as far south as south. the cause of the rare but serious paralytic dis. southern Japan. especially in reef environments that have be. The third way: spermcast mating in sessile marine invertebrates. and many others. These distinctions are among those that warrant place. these will ultimately give rise to a second and independent sporophyte plant. Archaeocyaths: a history of phy. The venom from such con. and patches (yellow dots). shoreward wall is protected from waves and strong currents. atolls and barrier reefs on continen. not only does calcification cease in the dark. Most scleraxonian colonies are upright and branched (e. tiated into regions that resemble vascular plants. Rivers are a triple threat to corals. especially dur. Calcified Tissue Inter. During adhesion. 2004).density) bands that mark annual growth (Figure 6–14). Goniastrea retiformis and Stylaraea punctata. A few of these produce a toxin that is injected along with the bite. 2002b. However. wave energy. Wilkinson CR. For ex. a b 86•Chapter Five may be involved. Even on reefs where corals are diminished because of marginal conditions. Volcanoes in the Sea: The Geology of Hawaii. The first of these is composed of six primary septa that extend from the corallite wall close to its center. They not only dilute surface waters. Caribbean Journal of Science 41: 492–507. velopment and antiquity of animal pattern formation.dimensional structure. crabs. For the most part. existing topography and grew seaward into the incoming waves. However.. Coenothecalia): a common Indo. the recipient’s eggs are fertilized internally through a complex system of sperm capture and recognition. spiny hexactine typical of the genus Heterochone. thus sug. Up to 75% of the dry weight of some demosponges is composed of silica spicules. The species problem in Millepora. the evidence of coral reef formation in the tropics is not often destroyed by advancing ice sheets. Heterotrophic bacteria are also abundant in sponges.. biont density is expressed. Dispersal. and Golan Y.Pacific reefs from the intertidal to considerable depths. are characteristic of river water and are detrimental to corals and associated organisms. tery of the filamentous boring stage had to be solved in the laboratory before Porphyra could be grown com. Cold. and Littler DS. 1999). Bundgaard I. produce six primary mesenteries and six (and only six) unbranched and non. consider a reef as any rock that might pose a hazard to navigation. 2001. 2005). Stromatolite/mi. eda. The entire system functions as a natural breakwater that dissipates incoming storm waves. ing sponges: lessons from Pione cf. Reefs and Related Carbonates: Ecology and Sedimentology. and Willis BL. noid form. Transactions of the American Geophysical Union 29: 855–860. Modern crustose coralline algae date from the Early Cretaceous period about 145 Mya (see geological time scale. dant and occurs as two common crystal varieties. and can fix nitrogen. thus enabling the recognition of the appropriate host species as juveniles using olfactory cues (Arvedlund et al. 2000. Similarly. Materials Science and Engineering C 23: 473–477. ing the development of Ningaloo Reef near 22°S latitude.E. 2008). Coral Reefs Tropical coral reefs are unequaled in the marine en. and Duarte LF. b) An epicurean harvest of palolo epitokes from American Samoa. Targino CH. In the subtidal forms. Some species may even use them as a lair from which to ambush prey (Ruppert et al. However. Boller ML. O’Neal W and Pawlik JR. The body posterior to the thorax is typically dif. and finally atolls (right). nobacteria). gonite is more susceptible to chemical breakdown than calcite or magnesium calcite. ara. itzka 1982). sea pens.circulated lagoons may produce steep. The skeletal framework is derived from multinucleate sclerocytes that form a syncytium rather than individual cells (Leys et al. and are com.defined shelf edge reef. and *Note on time: In this text the units Gya. and clover. 2007. Other tubicolous polychaete families are more typi. releases oxygen.like caudal region that terminates with the anus. NASA. Heterocystous cyanobacteria may also form enlarged. With the exception of a few small groups that exist only as medusae. and the form of the thallus (Figure 4–1f). Coral growth and reef growth: a brief review.bodied. Fragmenta. ing on a continental shelf. Some foraminiferans eat oth. 2002). reefs. gions. However. Tahiti 4: 505–526. als from the Great Barrier Reef. but nonetheless may participate in nitrogen cycling through different pathways. and they can be particularly prominent on the hardground communities of Fernando de Naronha. including the mode of fertilization (internal vs. but here are distinguished as internally brooding species to set them apart from exter. edulis and many other bivalves is smooth owing to its construction by flat. Hagan et al. some unicellular algae Reef Minerals and Mineralization•37 fibrous materials are typically associated with a soluble component containing high concentrations of acidic amino acids whose negative charges are thought to bind calcium and concentrate it sufficiently to induce nucle.like calcitic structure that is characteristic of many coral reefs (Chapter 4).5202. Coral Reefs 4: 1–9. Mi. Thus the length of the Great Barrier Reef with. 2007). In: Birke. and its sedimentary importance.g. as can members of the family Primnoi. Walters LJ. Coral Reefs 25: 287–295. it is certainly not true for all of them. A vision for regular.g. 1990. and other food appli. trations of nitrogen that accompany fish farming activi.Decapod Communities. a b –c +c –a 2 +a 2a 2 –a 2 +a –a 1 +a 3 –c +c +a 2 –a 2 –a 3 +a 3 –a 1 +a 1 +a 1 –a 3 Reef Minerals and Mineralization•35 introduced in certain brachiopod shells (Chapter 9). Walters LJ and Smith CM. It might also be noted that the plate motion associated with reef development brings these islands into warmer water closer to the equator where vigorous coral growth occurs. Coral Reefs 4: 1–9. Bulletin of Marine Science 33: 132–151. Spongivory in hawksbill turtles: a diet of glass. This escape autotomy (release of a body part as a defensive measure) may be used by sabel. This flow pathway. et al. and the apparently infrequent occurrence of photosynthetic symbionts. rounding a central lagoon that lacks an island landmass (Figures 1–9 to 1–12). ern Australia (Figure 1–6a). 2006. Marine Ecology Progress Series 443: 265–283. In addition. 2009. The Ecology of the Indonesian Seas. and are thereby distinguished from more selective browsers. the presence of zooxanthellae does not always severely limit the depth distributions of this group. Adey WH. and Motokawa T. However.Mortensen 2005) and made complicated by environmental factors including depth. International Review of Cytology 74: 127–162. Feldstein T. ate production estimates the potential gain to the system. the group also contains a distinct cnida called a spirocyst (Figure 6–2e) composed of a thin but fibrous capsule that houses a spiral structure composed of mini. billionths of a second. 2004.g. 2008 and references therein). These inquilines may incidentally exclude others by virtue of their sheer numbers. Figure 5–15d) and became especially prominent along with sphinctozoans on Permian reefs in southern China (Rigby et al. sess chlorophylls a and c. discharges a mean volume of 200... ployed to fill spaces. 2b.celled species that do not have heterocysts. • Production of structural calcium carbonate or sedi. Razak T and Hoeksema BW. Geological Ap. New Zealand Journal of Marine and Freshwater Research 19: 467–475. fluorescent mate.water groups. The thylakoids contain chlorophyll a. 1982. These forams live for several years and reach reproductive ma.700 years old. ing photosynthesis. Aquatic Conservation: Marine and Freshwater Ecosystems 14: 75–94. Charleston: National Oceanic and Atmo. Quan.concentrating mecha. whereas tests resulting from gamete fusion form smaller ones. Journal of Phy.Fine E. Li C. Thus. Rock and Reef.” including those found on coral reefs. the more broadly applicable histological description of “syncytial tissue” will be used here. Geometric changes over time are referred to as rates of skeletal extension. In addition. Gaspar H. Scolecodont jaws are some. In some groups (e. lene triangles. 2002. oysters (Chapter 8) whose shells are fused or otherwise adhere. where most of the cyto. The biology of reefs and reef organisms / Walter M. ing. 1997. meters in diameter and dozens or in some cases even hundreds of meters high. a photosynthetic perithallus. 2005). ganisms on coral reefs through time has been questioned (Macintyre 1997). Thylakoid Thylakoid membrane membrane Thylakoid membrane Phycocyanin Allophycocyanin Chlorophyll a Chlorophyll a Chlorophyll a Phycoerythrin Figure 3–5: Structure of the phycobilisome. Graham EM.shaped (e. Prochlorophytes are grouped into three genera. posed of a tough and resistant thread. How many Pacific hotspots are fed by deep mantle plumes? Geology 29: 695–698. detected cyanobacteria bloom in the southwestern tropical Pacific: implication for oceanic nitrogen fixation. and when such atolls are removed. suggesting a degree of independence between boring and photosynthetic activity (Zundelevich et al. 2004. Almost all of them are confined to the damselfish genus Amphiprion. Pocillopora damicornis. Kvennefors and Roff 2009). 2006. Coral Reefs 7: 51–56. along with somewhat more complex ones that oc. Galloway SB. 102 Overview. tion and conservation of precious corals. 2006. Sedimentology 35: 805–819. Weiss MP. Steep fore reef slopes may allow reef development to occur to a depth of 100 me. they grow very slowly and leave a record of that growth in the form of bands that may provide a means of as.West Pacific. nal of Sedimentary Petrology 60: 335–360. et al. and similar to seaweed. Reef Algae and Foraminiferans•71 heat. ated ecosystems in shallow. and Stuckenrath R. In: Reisser W. 372 5. ent places. and extends along the body wall or column to a basal aboral epidermis. Coral Reefs 2: 101–110. the association is more compli. Up. The Caribbean’s western. mono. Photo courtesy of Bob Stone. Gilmour 2004). and Atkinson M. a common shallow. ognized from the fossil record since the Pennsylvanian epoch about 300 Mya (Kirtley 1994). Gambi C. ing Fossils. and movement of new cells into the epithallus (e. and organization are often poorly defined. Lower Cambrian cavity.feeding invertebrates including sponges. Photos A. cified connections that solidify the areas between cells. and South Af. Heart of glass: spic. tain symbiotic zooxanthellae and are referred to as “zoo. stromatolites found nearby in seagrass com. Paleoecology and evolution of marine hard substrate communities. der varying turbidity. An overview of biominer. tecture and microstructure of the reef. are unique among animal taxa. ganisms and Their Descendants Exclusive of Animals. crevices. Coral Reefs 27: 601–604. Prouty et al. 2011.like growth of calcareous red algae called the algal ridge. and metamorphosis into a polyp. However. However.like clusters. or more recently as the work evolved. Hagan AB. 1999. tection from storms. or attach to sandy substrata by single or mul. gastropods (some of which consume their hosts). 1985. ranging from violet at wavelengths of 400 nano.g. 2009). 2005. cal Bulletin 190: 125–138. 2007. whereupon each palp is wiped across the mouth. 2002). as are branching corals and topographic high spots on reefs (Lara and Gonza. and as cold as . often making those attributes unreliable for diagnostic purposes. Whether mineral is produced in the intracellular or extracellular environment. Indeed. Wörheide 1998). 2009. subsidence. Figure 3–8f) that appear granular due to the presence of numerous spores. leased during photosynthesis. Algal Structure Most multicellular algal groups are benthic seaweeds that may resemble more complex marine plants like sea. isms do not belong exclusively to one functional group. Their skeleton was calcareous. more heavily calcified Halimeda segments and aggressively compete for them when foraging in groups (Overholtzer and Motta 1999. and a 3). Hoffman. asexually reproduced larvae. and decreases in water quality that may be conducive to high sipunculan density time is a bit complicated and depends on geography. growing on the skeleton of gold coral.1 My Oahu 3. tal transmission of symbionts could imply that hosts are freely capable of selecting from among a smorgasbord of free. 2008). drial and nuclear genes suggest that stony corals are monophyletic but most families of stony corals are not (order Scleractinia. Larval settle. All of these structures are fused with the corallite walls and floor and may exhibit rough or smooth sur. Indian Ocean). among many others—are almost entirely composed of these structures (Appendix 2 a. udoteal. Proceedings of the 8th Inter. identification courtesy of Kristian Fauchald. and Bergman B. Microbor. gers or predators. a b Figure 6–25: a) Sula reef complex. Conversely. A gradient of wave energy often determines which group is likely to become dominant. 1955. Biomineralization Whether under direct biological control or not. and that the initial colonization by corals resulted in a simple fringing reef. Daly M. 2008. Marine Ecology Progress Series 232: 105–114. b) Slab of P. Other anthozoan groups called corals (e. otic dinoflagellates in reef coral communities from Hawaii. nizational efficiency of their aquiferous system. 2003. 2007). both focus on shifts among the components of dissolved inorganic carbon (DIC). the primary tube that is secreted during larval metamor. College of Wooster. Al. 1997. and Westpha. outer prismatic layer of calcite. Figure 4–11). In: Warme JE and Shanley eds. Acta Geologica Hispanica 37: 61–72. Boschma 1948. persal. 2004. et al.. cal defense of marine invertebrate larvae.Australia. This is the only species widely recognized among several that have been given dubious names. Some anthozoan groups develop specialized elongated.associated. welling can occur locally in many places. there are no living forms that resemble them. especially in tropical. Journal of the Marine Biological Association of the United King. fertilization is internal and the fertil. some hosts pro. or they may simply “overtop” and shade photosynthetic organisms. a b Reef Corals and Their Allies•123 ferred orientation optimizes flow through the fan. a large number of endemic species (found only in a defined area) have been discovered. This area is dominated almost exclu. cal polyps.restricted habitats. Croix. crozones. 2000). and the figures quoted here over sandy or muddy substrata. Häussermann V. Services provided to their hosts may include many of the functions of free. actively growing reefs. tailed examination of bioerosion is given in the chapters that follow. clear water where green. the relatively greater contribution is due to the accumulation of microscleres or megasclere fragments. Fieseler L. and ecology of the blue coral Heliopora coe. and allows the colony to bend rather than twist in stronger current or wave ener. In fluid dynamics..photosynthetic. Algal ridges of the Caribbean Sea and West Indies. bonate. These morphologies are most abundant in the shallow. It is suspected that large snapping shrimp popu. Langdon C. many “barrier. The thallus may be erect and attached by a single large rhizoidal mass. The amount of light reaching a reef coral is addition. tary components of reef structure since Cretaceous time over 150 Mya. Knorre H and Krumbein WE. For an in. Brazil. and central Pacific. In: Dubinsky Z.gov/stories2008/20080116 _hawaiicoral. it can then be used as a nitrogen source by S. interpreted as a red alga. The Huon Pen. similar to Halimeda (Clifton and Clifton 1999). cially in the Pacific.rich zone called the inner (shallow) fore reef (James and Macintyre 1985). 1999. Horn M. Jokiel PL. c) Scripps Institute of Oceanography global topography v. nels in skeletal material or carbonate rocks. Species may possess more than one clade. Clifton and Clifton 1999). 229 polychaete worms. b) Fringing reefs as well as other types may develop coralline but. dreds of years old (Fabricius 1995) and can produce large amounts of spiculite at their bases. and occasionally ex.long chain on the eastern side of the Great Bahama Bank (Figure 3–14a). and the result has been a considerable list of terms describ. cite crystal may be plate. which have different species of Acropora.0–8. Hughes TP. The usual ex. and biotechnological potential. Amphinomid worms (Family Amphinomidae) have many of the structures typical of mobile worms. called “vital effects” by geochemists. aprica–C.” which consist of filamentous and short. Sedi. Miles MS. and their relationship with the living tissues of calcifying organisms.20). It follows that the Caicos Bank (southeastern Bahamas archipelago) would then qualify as the second largest atoll. texture. depending in part on local hydrodynamics. while commercial exploitation of red and pink corals has flourished in the Mediterranean and in the Pacific since the 1960s.e. Sponge: effect on the form of reef corals. Azooxanthellate? Most Hawaiian black corals contain Symbiodinium. 2008. to the SE in winter. Development of microbial endolithic communities: the influence of light and time. However. In contrast. binsky Z. and strong wind and wave activity is frequent. Journal of Phycology 39: 937–947. ered as “subsections” or may be grouped into five orders (Castenholz 2001). tially kept pace with sea level. and out through the exhalent surface composed of relatively large oscular tubes. Diversity and Distributions 12: 165–178.1% of light reaching the surface of their carbonate substrata (Shashar and Stambler 1992). Amemiya S. nels. ians have a flattened disc with many small tentacles that contain numerous and potent nematocysts. but contrasts with the upstream collecting system of certain sipunculans (see below). which are often colorful and spiraled (Figure 7–4a). Marine Ecology. which can sometimes be seen swimming within the aquif. a fringing reef develops on part of the east coast. Zoological Journal of the Linnean Society 139: 419–437. Botanica Marina 39: 395–399. Char. Biological Factors Affecting the Abundance of Halimeda Halimeda is abundant owing to high growth rates and short lifetimes. ter renewal time for classification of atoll lagoons in the Tuamotu Archipelago (French Polynesia). Goreau TF. Certain large macroalgae may become erect. Science 239: 393–395.Johnson Space Center STS068–252–42. algae that produce erect branches and tufts form a distinct downy or velvet. thus giving corallimorpharians a competitive advantage (Chadwick 1987. often many leagues in diameter. This includes income from tourism. which is called the prismatic layer. sphinctozoans (c). Brooding and Symbiont Inheritance With some exceptions (e. Despite a nematocyst. most of the Hawaiian Islands. showing both known and suspected locations extending from North America to what is now eastern Europe. Tsounis G. Nitrogen fixation also occurs in the water column and is another source of nitrogen for sponges through the heterotrophic uptake of particulate organic matter. mica et Cosmochimica Acta 70: 4617–4634. Bra. Photo courtesy of Malcolm W. The seaward walls form a shield. the hallmark of the phylum. rine Ecology Progress Series 301: 129–134. and Gennin A.P. but both appear to begin life as ordinary epizoic zoanthids. Figueiredo MA de O. 2007b). rubrum may be dislodged from their attachment sites and become incorporated into the sedi.com). 1984. and sediment transport. 1997. spreading >10 mm yr–1 . which in turn de. cil Centre of Excellence. and other environmental insults can result in the expulsion of zooxanthellae from the host. organic sheath. but more commonly this occurs during f. It broods its fertilized eggs. chaetae. and Weiss KM. 1984.Alsumard 1999). sis. These often form encrustations. Such massive. which are converted by carbonic anhydrase to bicarbon. and where surveys have been made using modern statistical methods. SW Pacific. Walters LJ..000 m).like and encrusting corals may dominate. When irritated. 1: an ultrastructure study of thallus development. spongia muta are primarily monaxons held together by spongin fibers. among other factors. Magnino G and Gaino E. the genus Halimeda (family Halimedaceae) is one of the most im. Photos by author. Scoffin TP and Bradshaw C. Proliferation of algae on coral reefs is part of a larger problem of reef decline that will be addressed in Chapter 15.wcmc. skeleton. Sponge de.. Both of these op. Journal of Geophysical Research 112: 1–14. or that formed by certain marine algae (e. From the perspective of the host. 2005. 2007. ute significantly to the three. tend beneath the surface. See Maldonado and Riesgo (2008) for a general review and further details of sponge reproduction. ice growth. After these corals are undermined. thor. not all of them are “coral friendly. in more exposed locations it may develop complex to. 1988). and Schneider J. are included in an order of about 4. cultures of the leafy gametophytes are grown with clamshells to facilitate the settlement and growth of carpospores (see http://www. also usually tube dwellers. Most atolls in the western Pacific and Indian oceans (and therefore most atolls) grow on gradually subsiding volcanic plat. boring sponges (Chapter 5) appear at the surface as a thin layer of tis.like con. sidence and deepening sea levels.like growth of skeletal carbonate. Schlesinger Y. row). sion.Pacific spur and groove systems are seen as primarily erosional. Tribollet A. and others are capable of burrowing as deep as 5 cm into carbonate rock (Chughtai and Knight. Porst M.dimensional structures. nauer JL. In: Carr NG and Whitton BA. like bees in a hive.building coral. Vroom et al. ters predatory fishes (Paul and van Alstyne 1988b). tematics 34: 661–689. 2006. thoracic regions bears stout. Macmillan. although off. as shown in Figure 6–35. Some corals are capable of adjusting their heterotrophic carbon intake according to plankton availability. a process in which superficial and internal boring may remove hundreds of grams of carbonate m–2 yr–1 . Atrich No shaft or barbs. b) Polished section of grains from a Bahamian stromatolite in plane. only portions of the body containing oddly.000 m along the Gulf of Mexico. Growth of the coral skeleton at the microscopic level occurs through the production of magnesium. The bent Hawaiian. 1990. However. Like many deep. ings of the 3rd International Coral Reef Symposium. zooxanthelle per host cell is typically not the way sym. riers. Mortensen and Buhl.g. and a fourth vertical line called the c. The thallus exists in a variety of forms. These compounds may have calcium. Schröder HC. cement that welds adjacent grains together (Figure 3–15b). Secretion of organic acids that result from carbohydrate metabolism Figure 5–10: The boring sponge Aka coralliphagum. riodic influence of river outflow and lowered salinities (e. in turn. Habitat. Risk MJ. lowed by overharvest. All of these algal groups display physical. and Duffy EJ.mortar schematic depicted in Figure 1–1a. Photo courtesy of Lau.like network of reticulopodia used for feeding. Paleoceanography of coral reefs in the Hawaiian. indicated a contemporaneous decrease in tem. some attaining volumes of a cubic meter or more. tion is enhanced by light or photosynthesis. Benolt M. courtesy of Paul Humann. rounded by 5–7 smaller (~0. drecht: Kluwer Academic Publishers: 123–139. 2006). However. but in addition to cross. 2007. The western coast of Australia lacks a true eastern boundary current because of the Indonesian archipelago. contains approx. Figure 6–33: a) Broadcast spawning of the hermaphroditic Carib. 2006. New colonies are produced by spontaneous colonial fission. ure 1–13). where their abundance and diversity often exceeds that of cor. windward reefs. Frank U. 2004). 2003).developed reef crest and are rimmed by low islands typically built 2–4 m above sea level by the accumulation of coralline debris. within 1–2 days of gamete formation. spective 32: 162–173.CS1. 2000. and prominent polyhedral bodies that contain some of the enzymes required for carbon fixation (Figure 3–2). including com.000 species are grouped in two taxa. eds.to. face layer that surrounds the gastrodermal canals. 1999. Miami 1: 477–483.like ribosomes are also found in most mitochondria and in all plant chloroplasts. line algal turf with white calcified tips. 3. but because it forms a (Weiner and Dove 2003). antipatharians are not typical of shallow water environments.. These and other important reef octocorals with more limited distributions are described below. and attachment of small fragments. Richter et al. Journal of the Marine Biological Association of the United Kingdom 87: 1721–1726. shape. In the unexploited condition. Photo courtesy of NOAA Ocean Explorer. Others may form a suspension. Unlike most sponges.g. but they are formed in separate bundles between layers of porous. Growth rates de. cific.org. ied conceptacles are usually protected from herbivores and other sources of physical damage. However. Figure 1–14a). sulting from various biological and physical forces.like gametophyte tissue within the fe. photosynthetic capacity varies even within the same colony during the day as a function of the light environment. 1986. a marine photosynthetic prokaryote of global significance. Pacific Science 55: 291–299. and many develop an enlarged cortex with thick cell walls that contain highly elastic polymers or are strengthened by cellulose or calcification.” or mushroom. 1995). Fish with generalized diets prefer chemically unde. Coral Reefs 19: 221–230. bell 2001). Sponge Species Most sponges produce a number of spicule types. McDonald et al. and Ilan M. Tuamotu Archipelago. The function of sur. although they may also be mul. and Guam. allowing some degree of development after fertilization. In. Lobban and Harrison 1994). tigaciones Marina y Costeras 35: 259–265. 152 Sipunculans. ties. Corallimorpharians are a small group of hexacorals whose polyps are typically flattened. 1995. Morphol. Wagner D. James Cook University. and Pawlik JR. cold. this restricted time frame occurs only at night or the early morning. Oceanography 9: 23–27. ian coral Galaxea fascicularis. some forams produce a milky “porcela. Reconnaissance geomorphology of Rangi. 1985. 2008. 1999. Barrie JV.. several centimeters thick and >1 m across (Lang 1974). both fishing and dredging create large volumes of sediment that in. Thus. 1997. ing the larger axial corallite at right. WD. There are well. 1980. or antibiotic properties. and LaRocha CL. nus Agelas (after Bickmeyer et al. cur in hemispherical stony corals (e. separates the main tissue layers and forms anchorage sites for the development of nerves and muscles in these two layers. however. the volcanic peak should disappear and leave a concave lagoon in its place. ficial and the original needles are preserved just below across the cell membranes for photosynthesis.like structure that is often enlarged in cultured material. though there are a few calcite. more than any other river in the world. Perera N. or in cyclosystems.” a term that is not always appropriate but is applied to several fleshy octocoral families that form massive. In contrast to mobile poly. Koehl MAR. central Pacific. Simple sclerites often form in the mesoglea of the polyp. The others are epilithic. seus and Medusa revisited. with special reference to microorganisms. Tarduno J. 1977. 2006). which are sometimes accompanied by another set around the oral region that close over it and the tentacles during contrac. sity Press. cally estimated as gross carbonate production. Advances in Marine Biology 52: 1–145. chemical or biological processes turn sediment into sedimentary rock through a modification of its mineral. Skeletal archi. Danovaro R. Morphological and cytological aspects of algal calcification.green eggs. Falcón LI. Free University of Brussels. Proceedings of the National Acad. However. Journal of the Ma. perhaps better known for its place as the last major battle between the United States and the Imperial Japanese Navy during World War II. Jamaican coralline sponges: their morphology. Clionaid sponges generally dominate the boring sponge community and account for 90% or more of the total borings in most coral heads (e. especially in the face of global warming. including the tissues of many reef sponges (Chapter 5) and numerous other groups. New York: Springer. but in others.energy. liphagum (after Grube et al. 1970. ways true and in such cases the generations are said to be heteromorphic.and deep. boxed area = costae.. ing the outer micritic layer (m) and larger crystals of aragonite (boxed area) below. although smaller sizes may be em. Diversity of coralline red algae and extinction patterns from the early Cretaceous to the Pleistocene. far fewer studies have focused on millepores and their zooxanthellae compared with stony corals. Neumann and Macintyre (1985) refer to reefs that grew in concert with sea level rise as in depths of 5 m or less (Figure 1–22 a. The high level of calci. cally requires a competent larva may occur for days or weeks and is followed by settling. and hexacti. 1999. 1°N. ing around within their tubes to defecate. which is able to close off the opening to the burrow. rather than through pores. phosphate. Such deposits. Nitrogen fixation by cyanobacteria provides a source of nitrogen for the scleractinian coral Mon. retain their eggs within the polyps rather than release them. a region re. Blooms of certain benthic species are opportunists and can blanket the bottom and dominate areas once covered with seagrass.reef growth: a dynamic balance. or assist in locomotion along the bottom. and may dissolve some propor. sabellid trochophores are all nonfeeding leci.poor waters found on most reefs and is presumed to be nutri. cal lifetimes (Bachar et al.M. Microbial Ecology 11: 461–498. as well as brittle stars and others that are epizoic and may simply use them as a perch (e. meters to a meter or more below the surface. among other unique fea. and the adaptive significance of the bleaching response. fied from Krautter et al. Wunsch M. ductive effort in the brooding coral Pocillopora dami. including large Pacific island groups such as the Tuamo. Ecosystems of the World. cify. theless. Corallinales) on a coral reef. clude green (Chlorophyta). Marine Drugs 2. 2008. while being tough and elastic owing to a tightly layered construction. which provides cyanobacteria with some degree of shelter from grazing animals (see below) and from desiccation during periods of exposure from low tide.. 1995. In contrast. tan reef. Mo. Science 240: 1737–1743. tecture referred to as a microbial mat. Coral Reefs 22: 109–120. Indeed. This group is chemically defended. lids that can exchange oxygen through their body wall and do not rely heavily on their branchial crowns for that function (Kennedy and Kryvi 1980). bonic anhydrase is found in high concentrations in the calicoblast cell membrane as well as in the organic matrix. Therefore. lite from the marine sponge Didiscus oxeata. fied from Allemand et al. Pasternak Z.building corals have requirements for clear. Little is known about the biology of either of these genera. 2003. Sánchez et al. Others are formed by the sand. However. Shifting roles of heterotrophy and autotrophy in coral energetics un. Mature cnidae. development occurs on the colony surface in a mucus cocoon. eastern Papua New Guinea. a region where many more undoubt. and become bioerod. and Garrity GM. cool and deep oceans are thought to be the most limited environments for the production of this mineral. Such beds are widespread and occur in protected shallow. were unable to sustain their growth during rapid rises in sea level and remained submerged in relatively deep water. this genus can dominate the substratum. tures are not always cemented into a framework.shaped. the more tropical St. and Management. in Chapter 4). On rare occasions. even though some of these patches are quite large and may exhibit a coral cover of >50% (Spalding et al. ants of this group often distinguish shallow and deeper water hosts. The entire system covers a distance of about 2. in turn. ment to the mucus and protein components (Bonar 1972). Vol. ish Columbia and northern California. 387 vii Many people have been instrumental in the develop. Milleporids are pantropical (found in all tropical regions. and Protula bispiralis.Pagès et al. Other examples of biological control in. hemispheri. barrier reefs (e. and elsewhere. ecol. Pollution in Tropical Aquatic Systems. which in some cases may develop a prominent buttress and chan. lower left. but ancient reefs contain little or none of it. The radioles are composed of ciliated units called pinnules that trap and sort organic matter by size (see Figure 7–6). Porites lutea. c) NOAA researchers display a large colony of the scleraxonian gorgonian Paragorgia arborea (also called “bubblegum coral”) collected by submersible vehicle. rous polychaetes. 1999. and Wa.specific interactions with structural refuge. Gorgonian corals typically have two different types of skeletal support: (a) layers of sclerites at different levels within the colony and (b) an axial skeleton. P. In addition. gen fixation in symbiotic marine sponges: ecological significance and difficulties in detection. ditions (Dullo et al. Both are holaxonian gorgonians. gesting that aging or compromised cells are consumed. Rosenheim et al. 2001. proximate. containing the photosynthetic and reproductive cells (conceptacles). or by rupture of the body wall. light levels may be too high and thereby suppress photosynthetic bicarbonate uptake. for ex. Although the intertidal zone often contains high lev. 1977. especially those with muscle systems composed of epi. Once fertilized. Coral Reefs of the USA. rent Dabouineau. and within about 50 My.water corals from deep sea ancestors. López. mounds. Most Atlantic examples are composed primarily by Lophelia pertusa. Demosponges All other members of the Porifera are composed of distinct cells and are placed in a separate subphylum. Lambrechts DYM. 2007. the symbionts remove heterotrophic waste products and function as metabolic kidneys. 1997. male and female are separate and produce eggs and spermatia in cavities called conceptacles. pearance as reef builders. 2008. Phototrophic nutrition and symbiont di. ogy 155: 159–171. 1990. Rhodophyta). Marine Biology 132: 691–701. ography 21: PA 4217. b) Drilling through coralline carbon. ronment. Charpy. 2003. disease. with the hope that they might be that way again someday. 2003). the Gulf of California. 1997. phyte plants. a diploid sporophyte and a haploid gametophyte. Carbon utilization may also fail to take into account periodic growth. most widespread. the pH is reduced to about eight.L. An early Holocene reef in the western Atlantic: submers. A third crystalline form called vaterite is relatively rare among marine organisms. which began 11. and other colonial cnidarians whose coelenteron is divided by eight unpaired mesenteries and eight corresponding tentacles with characteristic feather. Matsumoto 2007) and food supply (Risk et al. Autotrophy versus heterotrophy: the origin of carbon determines its fate in a symbiotic sea anemone. Rock borers may also have a thickened caudal shield at their posterior end that can aid the process of boring (e. analgesic (pain re. tion from adult thalli. tions: pre–Hurricane Allen conditions in Discovery Bay. Legentil et al. tertidal zone. whereas others may be hermaphroditic (Simpson 1980). buds may begin within the tentacles. 1994). per. Mohamed NM. while their neighbors form atolls with a typical lagoon (Spalding et al.term response of light.Paige et al. equal to 1. However.1%–0. the percent cover of coralline algae and corals.water reef corals (e. 375 Index. the c. sev. Journal of Experimental Marine Biology and Ecology 175: 105–120. this tissue is comprised of cells with junctions that constitute an im. Uncomfortable with the resulting terminology of “living spurs. De Goeij JM. where such “pendant” growth may reach a length of a meter or more. 2000).Moving Polychaetes Polychaetes either are freely mobile. rier and fringing reefs that grew in terraces.. Excavating animals actively bite and remove carbonate as they feed. and especially glycerol (Muscatine 1990). ceans form colonies with a rubbery or leathery consis. as global positioning system (GPS) technology and deep. Even so. As a result. often a coral colony. The largest continuous Lophelia construction known thus far is a complex 35 km long and 3 km wide called Rost Reef. while several zoanthid genera grow epizoically. which traverse the mesohyl. like canal systems. Indeed. the skeleton was reputed to serve double duty as a love charm (Hickson 1924).shaped bend between the Hawaiian and Emperor chains occurred relatively quickly between 42 and 50 Mya and is thought to represent a shift in the direction of the hotspot. 1988. mation. energy requirement. Of the 77 islands. St. d) external skeletal spines. perhaps as a means of dissuading generalist predators. a gorgonian or antipatharian coral. Oceanogra. and steroids with similar functions (Figure 5–22). have been identified and are collectively referred to as cyanosponges (Usher 2008). eventually rotating through 360°. Late Mio. Left to them. or adapt to changing conditions that favor them at the expense of corals and other more slowly grow. Figure 4–2: Turf algae are typically less than 10 mm in height. 2001. coral settlement. barger 1978. Because of its vast expanse. c b Chaetetid Pillar Lamellae Stromatoporoid Sphinctozoan dInozoan a Figure 5–15: Fossil hypercalcified sponges showing the structural variations of distinct Paleozoic clades including the chaetetids (a).shaped spicule bundles in the ectosome that transmit light deep into the sponge interior. Some groups of anemones are spermcasters in which fertilized eggs are brooded by the female and are held until larval formation is com. giving it the longest history of any coral (Colgan 1984).1 m3 (100 liters) may be capable of pump. In general. 2007. the calcium salt (CaCO 3) is the most abun. eastern Indian Ocean following the Sumatran earthquake and tsunami. In: Eldredge N and Stanley SM. Figure 5–17c). Bayer FM and Owre HB. mesohyl. ranking alongside kelp forests and sea. Razak and Hoeksema 2003). Nova Southeastern University. ure 5–17a) are typically small. Coralline algae typically occupy the most energetic zones. bean coral Montastraea cavernosa. although not necessarily a chemi. personal communication.brown color and colonies can occur in a wide variety of forms. Ayre DJ. but the mys. and Reefbase.IOC Register of Ma. ment. However. Marshall JF and Davies PJ. Discovery of a “living dinosaur”: globally unique modern hexactinellid sponge reefs off British Co. ric changes over time. 1986. Morato T. and Addadi L. Miller KJ and Mundy CN. even within a spe. Riul et al. similar to more complex photosynthetic eukaryotes including seaweeds and vascular plants. a type of collagen. 1979. most were solitary and true frame builders. Many are suspension feeders with a specialized prostomium that forms an elaborate crown of highly modified. 1989. Planulae are considered fully developed when the oral pore connects to the endoderm. unlike in other algae. Meibom et al.Paula JC. gen. These include the carotenes. However. Chemical defenses.driven calcium. It is also noteworthy that while zooxanthellae are common symbionts in many tropical hosts. era Halimeda and Udotea are distasteful to many herbi. Hooper and van Soest 2002). composed of tubules situated at the folded anterior end of a distinct thoracic region called the collar (Figure 7–4b). Pedal laceration by the anemone Aiptasia pallida. STS093. Adey 1998). Phase shift from a coral. 2003. Ma. leaves. tralian coastal environments. sion in sea anemones: integrative studies of life cycle evolution. Conversely. rises 153 m from its base at 789 m off the coast of South Caro. Marine Chemical Ecol. and the effect of climate change. These structures serve as points of anchorage for the body wall musculature. although branched.secreting organisms as depicted in Figure 1–1a and further described below. Reef Corals and Their Allies•127 are azooxanthellate. This sponge ge. Kingsford M. 2006. Moderate. verrucosa. Such large colonies may be 60 to >100 years old (Mortensen and Buhl. The basal polyp region in colonial forms is attached to its neighbors and to the substratum by tube. particularly other polychaetes and crustaceans (Wilson 1971. Pentagonal Atafu (Tokelau.12303. Trophic Dynamics and Ecological Interactions The Aquiferous System and Sponge Feeding Although there are some deep. However. and Mekong rivers. Coral Reefs 14: 131–140. Colgan MW. Panama 2: 1581–1586. ing activities of cyanobacteria. especially in the intertidal. Journal of Ex. tion of Madracis fragilis n. Botanical Review 72: 330–366. Some species endolithic. ate as the adjacent coral reefs. The spionid tube is U. tannins and chlorophyll) tend to absorb light energy. Coral Reefs 24: 160. ranging 1–12 months. Santa Barbara. and scrape carbonate as they do so. fjords. timated that a single Halimeda plant can reproduce itself by vegetative growth in a single month. Barque Canada Reef (South China Sea. The first is the so. 2001. Efferent mechanisms of discharging cnidae. a zinc. glassy. the epitokes.. Molecular Ecology 17: 1304–1312. All three are identical in their chemical composition. Burns BP. These layers are deposited at an extremely slow rate. 4–2). 2008. Townsville. 1999. with numer. calcite. These in. et al. b) Bioeroders are commonly categorized by size. day old juvenile showing radioles (r) and operculum (o). some of which rise thousands of meters from the sea bottom. New York: Academic Press. Charpy L. In: Dubinsky Z.energy environments where waves and currents pump seawater through the framework and sediment. Western Australia. Some polychaetes. Proceedings of the Biological Society of Washington 10: 100–109.armed and expendable “scouts. 1996. b. tors. 2003. times called submerged atolls or bank atolls. low). If the basic dihexagon (Figure 2–6a) is bent at its equator.” The latter group is common in the North Atlantic and SE Alaska to the NW continental United Figure 6–24: a) Colony of pink coral Coral. 25 distinct and often ornate microscleres are produced by various members of this group (Figure 5–5). PLoS ONE 3(9): e3222. Dipyramids in the hexagonal system can have six or twelve faces. distribution. Blair SM.. tant group of mostly colonial forms that are common on coral reefs. which may attach to or insert itself into the substratum. Santos UM. Steneck RS. Biomineralization. 2008.like structures called stolons.H. New Georgia Island in the Solomon Archipelago). The southern.3. faces. Hedley 1956). cretory cells. which give these algae their characteristic pink or red color. national Coral Reef Symposium. bonate density. However. and even within the same zone under differing hydrody. In: Culver SJ. nobacteria. Williams et al. Homotrema. a distinctly different pattern is obtained when the bud arises within the whorl of tentacles. especially on the windward side where broken coral gravel. While examples of red algae can be found in both shallow and deep water. and fossil rela. or some combination thereof (Liao et al. 2008). and to Mary Corrado. 2005. similar to certain zoanthids. tant groups of marine protists include heterotrophic cili.dependent equilibrium as shown in Equation 4–1: At seawater pH (8. imaginary lines of reference that connect the crystal faces or the angles between them. Aquatic Conservation: Marine and Freshwater Ecosystems 17: 195–215. inhibits overgrowth of sea. Three. and Ginsburg RN. balances heavy grazing by herbivores. McMurray SE. Thus. closely related species. Conversely. Pro. Clavico EEG. shelves. tion to reef carbonate appeared to exceed that of corals (Hillis. Drechlser C. Should conditions that cause bleaching become extreme in the short term. Taphonomy of coral reefs: a review. pounds. including oyster aquaculture operations. and in some cases can extend to more than 200 years. phycoerythrin. Cell. including the familiar sea whips and sea fans found on coral reefs worldwide. whereas others form crystals internally and then exocy. become filamentous. These are based on DNA sequence variations that code for the large and small subunit ribosomal RNA. Tribollet 2008). the encrusting black cyano. 2005). Gittings SR. Most reef corals begin showing ill effects from summer temperatures that exceed 30°C (described more specifi. Senckenberg Museum. In calcifying species where high calcium levels may occur in the extracellular or intercellular spaces. 2000) and are best known for their reproductive habits (described below). Literature Cited Reef Sponges•97 reef sponges. A number of factors. Coral Reefs 26: 617–633. Species of Spirobranchus are commonly associated with scleractinian corals. which lack a medusa stage. but others perform the same shift using carbonic anhydrase (CA). Marine Ecology Progress Series 207: 273–281. Walters LJ. paulkayphotography. charted seamount. as well as many members of the Anthozoa.sized sand grains. Stanley and Helmle 2003. photosynthetic nutrients are excreted from the rhizoids into the sediment where fermenting bacterial populations create strong reducing conditions around the “rhizosphere. fungi and bacte. biogenic reefs are carbonate buildups result. and even developed a chan. and Palaeobiological Ap. glossan opisthobranchs: taxonomic trends and evolu. Coral bleaching. For ex. called the prostomium. creases its spatial complexity. ducers in dead carbonate substrates of Hawaiian coral reefs. often containing a rich assemblage of fish and suspension. sues. 2003. supply. the cells actively control the pH and movement of key ions into the mineralizing site. vae go? Adult genotypic diversity conflicts with repro. Such reefs may have been “shot in the back by their own lagoons” (Figure 1–23b) and are a type of give. Sherrell RM. erate an upward current. munities that stabilize the bottom. and in the in. Hirose M. Grigg 1997). contribute to enhanced growth rates. is left behind in its burrow to regener. Arvedlund et al. All three stages and both types of plants are isomorphic— that is. several species are eco. review by Lewis 2006).412. 2008. 2005.compounds necessary for their growth. their ability to withstand harsh conditions (e. Panama. cretion. 1994. “low islands” are called coral cays. One of the most obvious distinctions between the two groups is in tube construction. Northern Mariana Islands. niferans are placed in the suborder Miliolina. poraneous approaches using cores.like protrusions from the body wall are about 20 cm long and 5 cm in diameter (Figure 5–12). Bioeroding Sponges Boring sponges are carbonate excavation specialists that have been recognized since the early Cambrian (Ko. fucata. Phototrophic sponges are often flattened.living species. sils did not produce spicules. These structures contain large numbers of zooxanthellae and are extended during the day. A simple and diffuse nerve net occurs in some taxa. Panama 1: 263–274. Alácran Reef. mental Marine Biology and Ecology 367: 61–64. nor do they possess true vas. Reefs. not be reestablished (e. parchment.Hoyes JS.: Octocorallia) from Chatham Rise (New Zealand).g. dae). the archaeocytes (Müller 2006). c a d e b Figure 6–29: Asexual bud. and pH are also important determinants of Mg. The Free. logs: biological and ecological effects. Interactions Explored. Major bulk dissolved organic carbon (DOC) re. they are the result of huge numbers of individual larvae. land of Hawaii. Biological control of calcification in symbiotic corals has two components that appear to be important in modify.versus live. underwaterexplorer. even within the tropics coral reefs may be restricted by extreme temperatures or by river outflow with its deleterious effects of sediment deposition. a recent analysis of Vaceletia’s ribosomal DNA suggests more specifically that these hyper. and overfish. Beneath the tissue. Calcifying tissue may fill the spaces remaining between corallites or they may share walls with no space between. However. are induced within an extracellular organic envelope called a biofilm. Photo courtesy of Ian Macintyre. Macintyre IG and Marshall JF. Coral reef benthic cyanobacteria as food and refuge: diversity. indicating that the largest mounds could be approximately 3. One such sponge.Pacific counterparts Figure 1–9: Atoll shape varies considerably ac. as is a similar pigment in certain other sessile foraminiferan groups (Krautwig et al. Krautter.up reefs off southeastern Florida. dia: bristlecone pine of the deep sea. ted against the range of photosynthetically active radiation. Marine plant diversity. Wakefield TS. a hexagonal crystal has two identical hexagonal faces that make up the top and bottom. snapper. Palaeogeography. dophyta) primary production due to deposition of a thin sediment layer. tion in cyanobacteria. meristem (mer).energy environments.g. The second largest linear barrier reef in the world is located off the coast of Belize in the Caribbean. rescent octocorals described below. Photos by author. structurally and ecologically significant organisms that produce signifi. more heavily calcified seg. and Larson DW. including Bora Bora. and displays limited develop. Tube growth requires the serpu. 1990. cally produced during asexual reproduction. Paraplexaura. Ecological studies of black corals in Hawaii. 2009. Thus. but also differs in the way it is assembled (De Nooijer et al. nutrient. Redwood of the reef: growth and age of the giant barrel sponge Xestospongia muta in the Florida Keys. clonal populations can recog. The group is widespread in sandy or muddy substrata and are common on reefs. and are withdrawn at night (Figure 6–8a). vading newly exposed carbonate substrata within a few days. hunmathi and Miladhunmadulu atolls (two names. Proceedings of the 6th International Coral Reef Sym. sug. ments. Lewis JC. Sedimentology 20: 574–614. Samoans and other Polynesian groups. sub. and San. Modi. and although most of them do not calcify. ride sheath elongate as the cell divides and bores into the grain. ganisms. rapidly growing macroalgae over crustose corallines may shift the structure of coral communities in complex and subtle ways. The initial deposit is a network of micro. Coral Reefs 4: 125–134. Figure 3–4: The structure of . this proposition does not always account for the proportion of sexually vs. c) skeletal cross section showing laminated growth rings and hollow center. 358 Are Reserves Enough?. Photo courtesy of Sally Leys. The specific conditions and activities governing stromatolite growth are complex. Reed JK. six. 1948. The deposits can be massive with 85% or more of the thallus weight as carbonate. 2007).water reefs do not conform to the framework concept yet produce cemented structures composed of a thin veneer of small corals growing closely together as a carpet (Riegl and Piller 1999). Hine and Steinmetz 1984). les 1998). Characterization of mac. Atoll Research Bulletin 458: 1–20. Tai. not reincorporated into the reef structure). rowing by having a U. Both of these pigment complexes are blue and give these organ. all of which are organized into distinct layers.. Capricorn. This class of about 6. fringing reefs may gradually or rapidly drop into deeper water. and Manfred Krautter. and Ferrier. careous sediments of the Great Barrier Reef. ATPase production requires a source of energy in the form of ATP. ed. Lesser MP. ative phosphate acquisition in giant. instead of the calcium. They are often biologically complex. The bor. son of biocalcification processes in two coralline dem.g. The form of organisms that dominate the reef crest may vary according to the strength of wind. and other photosynthetic pigments that occur in zooxanthel. mental nutrition. there may be a series of organisms and complex biochemical transfor. oped. thotrophs (Rouse and Fitzhugh 1994). tional Museum Todd LaJeunesse. 1: reef habitats.energy trade winds north and south of the equator (Chapter 1).bottom communities. tered into eight distinct genetic lineages or clades that are distinguished by letters A–H (see Appendix 3). at least in shallow water (Tak.g. Lastly. Försterra et al.noaanews. The state of a reef can generally be described as a net value of carbonate addition and subtraction due to re. nia (a clade formerly designated as a subclass) in which most of the skeleton is composed of a proteinaceous cor. Bulletin of Marine Science 33: 132–151. In: Birkeland C. cies of Porphyra are economically important. multicellular algae and protists in general constitute an artificial grouping that is in the process of deconstruction into many “kingdoms” or kingdom.shaped burrows to procure food. and Bümmer F. Schuhmacher H. especially against other cnidarians. Some authors have suggested that these clonal larvae develop from eggs parthenogenetically (i. and stems. Ecosystems of the World. in contrast. Ellison AM. 2001. University of Victoria. especially in regions where typhoons and hurricanes are common (e. Demo. the modern view is that fringing reefs develop in a complex variety of ways. forams are single.West Pacific range. Structure. but especially the tentacular epi. Figure 3–8f). Marine Biology 108: 137–144. cystous. while others pro.g. 1968. over time and exposure to fresh water. Folds in the calicodermis reflect the structure of the septa. A framework of A.1.temperate waters. land provide a storm. Not surprisingly. inviting detachment from the substratum. epidermis and endoderm vs. creased losses due to storms and wave action (Wulff 1997). which projects as an oral cone.feeding polychaetes.C. ville: Australian Institute of Marine Science.like biological glass. Invertebrate Biology 125: 281–287. Neves E and Johnsson R. Siphono. 2001. is surrounded by a barrier reef system with a circumference of >1300 km (Spalding et al. ment of atoms (or ions) forms a series of two. American Zoologist 30: 23–35. microscopic examination is required to distinguish them.g. Marine Ecology Progress Series 397: 53–61. nobacteria. and its dependence on yolk or other modes of larval nutrition. Ulstrup KE. ters of genetically identical or closely related colonies. Figure 1–8: NW New Caledonia barrier reef and atolls. Map by J. and may likewise play a role in fixing nitrogen as well as cycling of different forms of it. diversity “hotspots. especially on barrier reefs and atolls on which they are well developed. or filamentous microalgae. ferred to as the middle lamella (Figure 4–16a). Cnidaria). depth. and can occur in depths of 10 m or more (Ross 1977). In contrast. especially encrusting and excavating species (e. Others have found that some reefs. When the absorption spectra of the major cyanobacterial pigments are taken into account.poor) central trop. some cyano. Chemical defenses in sacco. tion of the carbonate as well. gists often considered the group as sponge. 1989. Ecological Economics 29: 215–233. caudata (also referred to as P. The boring mechanisms of cyanobacteria (and other microbial endoliths) are unknown. tion. 2007. nitrogen fixation in such environments may exceed that produced on and in the reef itself (Shashar et al. Photo by author. erated away from the oral surface even though food is subsequently collected on the oral side. they can substantially alter the compo. posium. meandrina. for assembling the disarticulated bones of a bumphead parrotfish skull shown in Figure 12–8. lae (Schuhmacher and Zibrowius 1985). and provide nucleation sites for mineralization. it is less clear that the host benefits significantly from N. et al.” some of which may be 20 km or more across. Figure 5–3). Henkel TP. zooid) and canal system. presenting a wide oral disc and numerous. some of which can become quite large. and reduc. 2000. and another that is fatal called sponge orange band disease (SOB) and is accompanied by the total collapse of host (Cowart et al. the Hawaiian chain continues as a se. Indonesia 1: 23–27. cludes species that are not necessarily threatened with extinction. cies. succes. An Introduction to the Structure and Formation of Modern Reefs •15 (Milliman 1973). land C. one of the hallmarks of red algae is the production of a second spore. While the latter ac. binders. where on a local basis the tests may cover as much as 65% of the substratum (Jindrich 1983). Molecular Phylo. flattened discs and dome. and it has Figure 5–21: Agelas conifera produces distasteful alkaloid com. In: Gupta BKS. and morphology (Chapter 5). An Introduction to the Structure and Formation of Modern Reefs •9 especially on the windward side (Figure 1–5). Associations between Caribbean sponges and photosynthetic organisms.” Riding (2002) distinguishes these as “segment reefs. Marine Biology 145: 875–884. Chapter 10). Amaral et al. b). The Scleractinian Skeleton Most scleractinian reef corals form an aragonitic cor. its windward side. the north Pacific coast. posits. Raised reefs are further described below (see Uplifting and Tilting). however. insula include successive fringing and barrier reefs that have been uplifted. there is no definitive evidence for any of these interactions. nected to its neighbors by a thin. ———. Some species form tests composed of aragonite whereas others do not form their own carbonate and produce an agglutinated test by embedding and cementing foreign material (e. Broadcast Spawning Sexual reproduction among corals involves complex and variable mating systems. Larkum AWD. perate region (Birkeland 1997). Fietzke J. br. despite its common use. although at one. This process. Anderson PAV. each of which is joined by a separate shelf (Figure 6–23a). accelerating at >5 x 106 times gravity (approxi.Williams et al. photosynthesis.g. plete. mar 2005). The production of various microbialites is an important. capable of in. can be quite challenging because of the types of boring and calcify. doi:10. doi:10. American Scien. ed. 144 Free. daria: Hydrozoa: Athecatae). stratum. certain members of the genus Tubastraea are capable of forming large fan.like structures called blades and these may be attached to one or more stem. a range of 250 to >900 g m–2 yr–1 may be deposited in some locations (Weinbauer and Velimirov 1995). Biology of marine herbivorous fishes. and as a cure for rheumatism and other pathologies (hence the term “anti.. New paradigms for supporting the resilience of marine eco. one that is cyclical and has no long. Bunge H. niidae). the average gross carbon. Helbig U. Nervous systems at this level also may be associated with the development of more advanced muscle tissues that include smooth and striated fibers. Surface. Amsterdam: Elsevier Science BV. line. Micron 39: 209–228. the larger (older) size classes of exploited black corals have declined ow. nerves.0 (e.third of all scleractin. Report on certain hydroid. pillar. Hy. typically blade. Kinsey DW. however. (2005) found that process appears to be endogenously controlled (i. brate Paleontology. Perhaps for these reasons. and because the relationship with zooxanthellae is obligatory for these forams. Cabioch G. Two living demosponge genera produce stromatoporoid. tually making the structure as a whole less porous and more resistant to hydrodynamic forces. and are particularly well known in fresh. the tropical Pacific. Extraordi. In other cases. Chi KJ. Atafu (center) has well.like structures. Roopin M. calci. The cell walls of red algae are typically composed of cellulose fibrils. 2001. Pacific. In quiet muddy settings such as inshore lagoons. allel to the cell membranes. Pho. Johannes RE. is also the dominant light. McClanahan TR. These calcisponges are typically small. an anatomically complex Caribbean alga that overgrows corals. Integrative and Comparative Biology 45: 615–622. Stanley GD and Helmle KP.g.cell communication. Bicarbonate is presumably trans. Figure 5–6: Spicules from the Caribbean barrel sponge Xesto. 2010 http://topex. This degree of toxicity is nearly equivalent to bacterially produced botulin. Palaeoclimates: the first two billion years. while unimportant commercially. pographies with moats.known group of species (Young and Henriksen 2003.g. Asso. 2005. Elbert DC. mollusk shells. Norton TA. Paul VJ. are they are released through a corresponding small ampul. sometimes as large as 5 cm in diam. Biology and ecological functioning of coral reefs in the Main Hawaiian Islands. In others. The Free. 1995. and can be im. The resultant external fertiliza. Canada: controls on growth and development. slowly growing species. two types of bleach. which is often an actively feeding planktotroph. plains some.energy conditions. 354 Stemming the Tide: Sanctuaries. Indonesian P. Coral Disease and Health Workshop: Coral Histopa. Within the last 2. Makatea. Vol. Seddon S.water gorgonians Prim. Marine Geology 3: 369–383. While many of them (calcare. 1983). carbonate to CO 2. In: Geldsetzer HHJ. Neves and Johnsson 2009). tacular budding patterns. Figure 5–18b). These formations may contain at least as much carbon. Diversity and distribution of reef organ. Atoll Research Bulletin 459: 1–46. Some marine species. fertilization (Miller and Ayre 2004). These demosponges are present worldwide and play an important role. 2006.E. A comparative demographic anal. Invertebrate Biol.and. The lamellar.Horani FA. and Green EP.ocean diving submersible vehicles are used to find and remove these slowly growing corals. and Macdonald KS. In contrast. Duffy JE. The dark spots within the sand grains and within the crust (e. Indeed. scribed. Depth limit for reef building corals in the Au’au Channel. Together. Such conditions and high. A gelatinous connective tissue. Leys SP. south Ber. As described more specifically in later chapters. tercellular spaces tend to become more heavily calcified. This amphinomid even sequesters palytoxins from Palyt. and de. Coral Reefs 21: 375–390. tritional state of the host. http://www. and Martens CS. siding volcanic platforms are fringing reefs initially (left). While not all cyanobacteria are capable of nitrogen fixation. 1981. However. chaetae. Seamounts: Ecology.g. Chondrosida. although aragonite is produced by a few taxa. lik 2003). Zooxanthellae are packed within host gastrodermal cells. Daly M. Inquilinism. ily recognized by its distinctive yellow. “high islands. Dullo W. Ayre DJ and Grossberg RK. Manila 2: 679–684. Burton PM.com (BoraBora/ pictures/5d_9716. newly formed calcium carbon. season. Clayton WS Jr.building coral Fungia sp.swept cliffs or canyons. Such resistance may also be a factor in the pres. tive implications. The small amount of tidal exchange. The range of this species includes the western Atlan.are found commonly in tropical reef environments throughout the world.like reef. Materials Science and Engineering C 23(4): 473–477. In some cases. Cronin G and Hay ME. which could decrease pH by producing protons thereby increasing carbonate solubility (Ehrlich et al. and Irvine LM.like cells. nes feed only occasionally (Fautin 1991) and thus may rely heavily on their symbiotic zooxanthellae. Treatise on Invertebrate Paleontology. Eggs are brooded within female colonies where they develop into planulae that escape through ampullary pores. Sethmann I. Bergquist PR. cellular organisms became diverse and abundant. and phospholipids. although Reef Algae and Foraminiferans•67 production of structural carbonate or sediment. are tracks of the same hotspot that is now under the island of Hawaii. S. LaJeunesse et al. Biological Bulletin 136: 130–139. Facies 51: 33–48.000 to 17. The primary pigment. Panama 1: 725–730. Summons RE. The soft coral community on a sheltered reef quadrat at Laing Island (Papua New Guinea). a. Oceanography 21: 108–117. Jour. However. Their nematocysts deliver a toxin that produces a painful welt and is responsible for Millepora’s common name. and Cay Sal Bank (see Figure 3–14) between Florida and Cuba. position.water species from the Caribbean. sabellariids use particles 125–500 µm in diameter for tube construction. ———. Natale F. The first attempt only penetrated to 30 m. and Systematics 8: 23–43. In contrast. are all in this group. calcite and aragonite. The open. 2007. Golubic S. hollow. 1977. Highsmith RC. islands and sandbars can close the lagoon so that exchange is limited Figure 1–7: Belize barrier reef and its three adjacent atolls as labeled. ability to destruction by storms. Photos courtesy of Paul Humann. These medullary filaments then divide and form a white. lipids. Webb GE. These give rise to the rhizoids or holdfasts and the initial segments. 2008. or by sexual reproduction that begins on a core of foreign material.Shallow fore reef with buttress and channel system 15 m . many corallines periodically shed all or part of their epithallial layer along with accumulated sediment and epiphytes.induced turbidity following sea level rise (Adey et al.tolerant group and are referred to as “mangrove cays. width. Relatively large colonies can dominate the benthos.g. non. The free.” The meristem may contribute to its own growth by cell division.MacDonald and Leys 2008).pone. Abby Collier was instrumental in smoothing the path toward finalizing the manuscript. The end of it marked the warm interval that defines the present Holocene epoch. most (~75%) of the nitrogen fixation and most of the cyanobacterial biomass occurs intertidally. and Hoegh. chloroplasts of cyanobacterial ancestry were the forerunners of modern eukaryotic plants. crustose corallines from several genera can form unattached. as shown in Equation 4–2: Depending on the species. This genus is widespread on Indo. Western Australia. 1988.diving depths are still collected by trophy. Holo. Gorgonian extracts have been shown to deter feeding by fishes and by some inver. imately 25 species of red or pink corals known for their value as jewelry. tre. rapid assessment of the tropical northwestern Atlan. and Coll JC. as do certain foraminiferans (Chapter 4). and Allemand D. Coral Reefs 28: 547. however. gametes are broadcasted from the gametophyte and fertilization takes place in the water. gamete or mucus production. Limnology and Oceanography 52: 907–911. The solution effect of fresh water ponds and groundwater rich in CO 2 from decaying vegetation is thought to be particularly important in the pitting and scarring of limestone surfaces. 1–21a) provide a continuous record of reef growth dur. including a continuous supply of sediment that that can be sorted during turbulent re. which form limited refuges for corals and associated reef organ. Vol. and Yonge CM. peptides. Fadlallah YH. They can also change their orientation to take advantage of favorable current flows (Dubois et al. each ending in a protuberance where nema. Zoology 2: 1–248.Furman 2004). Verbruggen H and Kooistra WHCF. and carrageenan is used as a gelling agent in ice cream. ther calcify nor produce spicules (Wörheide 2008)! Inozoans were a fossil group of sponges with dense calcareous skeletons. The choanocyte flagella may change direction every few seconds. Seaweed susceptibility to herbivory: chemical and morphological correlates. ment of both the written and illustrative portions of this book. but appear to have followed a different evolutionary pathway.600 years ago (Grad. Depending on the complex interplay of these factors with environmental variables. Both of these methods can extend growth horizontally. epizoic serpent stars curled around the branches.specific bacterial lineages appear to have co. clude that vigorous conditions promote reef accretion. New York: McGraw. ever. or they may be separated from them by a sandy terrace or trough (Figure 1–14b). This species is highly sought by commercial trawl fisheries between Brit. nities. vised description of in situ “zooxanthellae” genus Symbiodinium. Nuñez CV. Four southwestern Caribbean atolls: Courtown Cays. Cairns. By contrast. orange. Johnson Space Center ISS004. fied segments as contributions to the sediment (Figure 4–5 a. the light. 2003. sociated with the surfaces of certain key species. and the form of the me. feeding can be continuous.consumer balance in intertidal biofilms. Ex. erations are prominent in sponges. Pueschel CM and Saunders GW. or the medullary filaments may occupy part of the surface and interrupt the cortical layer. The type of tube. poorly circulated lagoons are particularly susceptible to dramatic physical changes. Thus. Figure 6–15: a) Scanning electron microscopy image of aragonite crystal layers from the septa of Fungia sp. With these observations in mind. 1988. pink. However. and Tartinville B. Andria JR.5 Gya such mats became significant reef builders. er’s viewpoint. 1997.000 years ago when Uplift and Tilting As observed by Darwin. sea urchins (Birenheide et al. and tintinnids: ultrastructural character. “the symbiotic association permits rapid growth because the coral can invest in flimsy scaffolding at night with the certainty that the bricks and mortar will be available in the morning. 3 Reef Growth and the Carbonate Budget. and Systematics 38: 255–273. When noncarbonate rock substrata (e. gastrodermis (see discus. Few studies have been devoted to reproductive biology in this group. especially in regions that do not experience strong upwelling. Deep Sea Corals and Coral Reefs Although scleractinian corals are best known for their diversity in warm. Goreau TF and Hartman WD. perhaps when the filaments be. The entire structure that extends to and above the surface (reef crest plus islands) is generally referred to as the rim of the reef. although often in distinctive patterns and thick. Species with ag. personal communication) called “soft cor. In: Connell DW and Hawker DW. while some have suggested that shedding alone is sufficient to maintain CCA populations intact (e. H. Conversely in less harsh subtidal regions. A recent assessment based upon molecular genetics and cell structure has placed the Chlorophyta in a “supergroup. and lasted for only about 10–15 million years. 2001. 1993). 2005. However. Coles 2003). ifera. Bali. even in areas of intense graz. Sherman CDH. This is the haploid gametophyte stage (1n). nel and buttress system.1072. Nitrogen Fixation and Trophic Relationships Cyanobacteria play a key role as nitrogen. bafflers slow water flow and facilitate sediment deposi. sponges are typically placed in their own sub. especially those in the intertidal zone and other space. western Pacific) in 1896. Verte. and may not receive adequate nutrition from them (e. These organisms are aggres. flat. 2006. covering 40% or more of the available substrata in some areas. “naked corals”) from which modern scleractinian corals may have evolved (Stanley 2003. secrete substances that play an important role in promoting the settlement and adhesion of coral larvae. Raiatea to the south and Tahaa to the north. often restricting the larger crystals to the interior por. Elder. Figure 6–11: Colony of the antipatharian Antipathella fiordensis showing white tissue and attendant. 1997). A radiation reflecting the appearance of modern segments are often a dominant component of reef sediments. the northwest end of the Soci. a number that can vary in some species from 6 to 11 (e. Calvert P.rich carbonate granules 1 micron or less in diameter. 2006). and in some cases for attachment. 2001.. Many of these compounds are toxic to cancer cells. pursuit. especially in shallow. structural characteristics of the scleractinian coral skeleton: a biocalcification proxy. None. scribed below. eas. Cyanobacteria. Barrier reefs and atolls develop complex topogra. These are connected to the siphonous system so that cytoplasm can flow into the gametangia from the segments to which they are at. Fink and Rigby 2004). Paul VJ and Hay ME. and the histone proteins associated with the complex structure of eukaryotic chromosomes. coral reefs (frame reefs) are generally restricted to tropical and subtropical regions that are no more than 30 degrees of latitude above and below the equator. cial reference to diversity and distribution of deep. is 79 km across and 90 m deep. ever.sized sediments as shallow reefs are converted to terrestrial and semi. Hexacorals are represented by six orders including anemones. A reappraisal of the chemical and physical defenses of Caribbean gorgo. and they occur erratically over time and space within the reef. into the spongeocoel.gathering devices.” They produce 20 different types of distinctive megascleres. ing environments favorable for microborers may play a role in preserving the segmented. Zooxanthellae Digestive cell Neuron and muscle layers Spirocyst Nematocyst Cilium Microvilli Epitheliomuscular cell (EMC) Epitheliomuscular cell (EMC) Mucus cell Mesoglea Epidermis Gastrodermis Coelenteron Figure 6–1: Model of anthozoan epidermal and gastrodermal tissue layers. wildlifeland. or acute and fatal. or T.water Lophelia pertusa coral reefs off the southeastern US: Blake Plateau. However. Biodiversity and Conservation 19: 153–167. ing colonization by boring organisms and other sessile competitors where the hosts would otherwise be vulner. 1986. phosis is required as a foundation for further growth and is a structure that cannot be replicated by adult tissues (Pernet 2001). and they contribute to a complex pattern of carbonate lithification (infilling. the intrusion of cool. Müller WEG. glossa) as antipredator defenses. Figure 6–30: Detail of Caribbean coral skeletons with extraten. Studies of stony corals and other zooxanthellate an. However. 1860) (Cni.. For example. Cyanobacte. Underwood AJ. cal to the sponge because water flow provides a renewed source of oxygen.rayed forms called hexactines (Figure 5–4). 2004. nile fishes: species.density bands typically correspond to warmer sea surface tem. The Belize Barrier Reef was designated as a World Heritage Site in 1996.induced turbidity and sediment deposition. 2008. sophical Transactions of the Royal Society B.Rivera and Paul 2000). In: Margulis L. pollutants. four other regions of the body are often distinguished. chaetes. bonate in the tropics. and new partnerships could be forged with the more toler. karyotic plants. Thus. Types of Reefs Reefs are usually categorized by their size.bearing segments are formed and it is referred to as a chaetiger larva. Habitat use by sponge. 1990. ticeable at the bottom of steep reef slopes where corals with eroded bases may accumulate in a heap after rolling mud by definition is composed of fine particulates. these grains become welded around the periph. members of the genus Madracis) develop a single cycle of 8 or 10 septa. The Evolution of Modern Reefs a. Indeed. 1993. or banks. http://www . zooxanthellae enhance calcification rates and yield an overall growth of 8–22 mm per year. participate in a mass spawning event in which gametes are shed synchronously for a few hours during a few days. The most spectacular of these coral spawning events occurs on the Great Barrier Reef (GBR). 1988). They live in protected habitats that may include mud.lived alcyonaceans have also been Figure 6–21: Alcyonacean form: a colony of the stalked genus Sarcophyton from a reef in Vanuatu. Regardless of growth form.9 GA) barrier reef complex. 1996. collectively created a rimmed carbonate shelf and slope on these ancient reefs (Wood 1998). Polychaete worms can be significant contributors to reef communities by their trophic habits. and the Figure 6–16: Scleractinian zooxanthellae in culture. the Great Barrier Reef and the Belize barrier reef systems) are not associated with volcanic platforms at all and are among those referred to as non. suggesting that they may be more pervasive in carbonate environments than is currently appreciated. anything that increases the mortality of frame. Coral Reefs 24: 67–73. eventually becoming capped with algae and other calcareous organisms at sea level. wave re. This suggests a primarily physico. Warraber Island. als: autotrophs or heterotrophs? Biological Bulletin 141: 247–260. SJ. bind.water corals from the Pacific are typically associated with clade C. cornis. and are subse. Zundelevich A. brates. these may be restricted to openings in the rim called passes or channels. Decho AW. an average of 11.77'89—dc23 2013003715 o This paper meets the requirements of ANSI/NISO Z39. high turbidity. these pools may be exposed only briefly and may harbor certain hardy coral species in ad. 1998). Thus. the dinoflagellates. elongated rhizopodia that are typical of a number of protists. which is ~0. ———. ings of the 5th International Coral Reef Symposium. Photosynthetic pigments (Table 4–1) include chlorophylls a and d. gram.Johnson Space Center. the sea hare Stylo.bearing females. and Dodge RE. ing have been described. thesis using chlorophyll while producing molecular oxy. Skyring GW and Bauld J. they may feed more efficiently than gorgonians. This can occur owing to bioerosion alone. Frame. Ber. Indeed. Phylo. In addition. 1973. or specifically control it. temperatures may exceed 35°C. Kroeger KF. rier reef tract.sea corals..sized tests Regardless of test size and the number of chambers. Extracel. Paul VJ. The largest of them formed mounds several hundred meters across and tens of meters high.building Hexactinellida found in waters off British Columbia reaches a height of 1. predation pressure by fishes is thought to drive the association because the sponge pro. 1988b.pdf. some low. Rützler K. Likewise. duce funnel. meters (nm) to red at 700 nm. bacterial symbionts in sponges. ture and the stellate surface features of Astrosclera are consistent with stromatoporoid fossils (Wörheide et al. ects (Dubois et al. isms. Flow cytometric analysis of virus.g. ries 22. Proceedings of the 3rd Inter. Titlyanov et al.term research. ct. However. which might be expected as an intermediate stage of reef formation. Sexual reproduction among anemones varies con.lived particulates that stay in suspension for long periods cause turbidity. lonic tissue becomes embedded in a colonial epidermis and a few hydrozoans produce a heavily calcified skel. Such high popu. 2000. gonematales). 1997. Integrative and Comparative Biology 45: 369–376. oc. Additional accessory pigments form phycobilisomes (Figure 3–5). Deep.violet and red light. Sponges. ers. A whiff of oxygen before the great oxidation event. the most common of which is longitudinal fission.like branches superficially resemble certain growth forms of Millepora and are dotted with small pores that are con. 205 Overview. their characteristic pigments are phycocyanin and especially phycoerythrin. Paleozoic and Mesozoic chaetetids often occurred on shallow reefs and in the open. 1999. Spirocyst Long. ciated community with the zoanthid Palythoa carib. cent stromatolites. Varying growth rates in bamboo corals: sclerochronology and radiocarbon dating of a mid.com. support very large populations of a fanlike species that occur in such exceptional densi. or density. Bo M. or may adopt a “sprawling” morphology and attach by multiple rhizoids. Coral Reefs 13: 135–136. or diminishing over time. Klumpp and Polunin 1989). eton from this material. the pinacocytes. often less than 1 cm in diameter. Family Clionaidae D’Orbigny. High pH environment Ca2+ Ca2+ 2H+ Ca Ca2+2+ 2H2H+ Ca2+ 2H+ CaCO 3 HCO 3– CO 32– HCO 3– H + HCO 3– CO 2 Low pH environment ATP-driven Ca:H exchange Nucleus Chloroplast Chloroplast Chloroplast Starch granule Starch granule Starch granule Pit connection High magnesium calcite crystals Protons for photo. Townsville: Australian Institute of Marine Science.producing metazoans formed enormous reef complexes. it is the world’s larg. the region around the anus is thickened by cuticular material composed of hardened collagenous protein. Bidirectional sex change in mushroom stony corals. 305 15 Reefs Now and in the Next 100 Years. Their boring habit may be related in part to feeding activities. suggesting that the adaptive mechanism may have maturational or other unknown components (Fautin 1991). especially in wave. heat. Boca Raton: CRC Press: 147–166. called the nacreous layer (Figure 2–3). calcite. 1990.swept and exposed intertidal zone. 2006. Structure and diagenesis of recent algal. the result is harmless and each mutually benefits from the increases in surface area and feeding capacity.38478. Partitioning among grazers of food resources within damselfish Adey WH. Although there is a diversity of form within the group. tivity by heterotrophic bacteria. thus dramatically increasing the Figure 6–9: a) The colonial anemone Playthoa. Biomineralization. where colonies can reach 3–4 m in height and serve as substrata for many small invertebrates and shel. the known geographic patterns of algal diversity may be counterintuitive. 2004). ented on the surface of the thallus and are arranged in concentric bands. 1995. and formed in layers that are oriented perpendicular to the shell surface (Figure 2–2) giving it a rough texture. Scale approximate. Biochemical Systematics and Ecology 36: 283–296. Charpy. Mortensen PB and Buhl. This species produces a dense aragonitic skeleton along with siliceous spicules at the living surface that become incorporated into the ara. When released from the parent. all of these are plants. Chlorophyll b–containing oxygenic photosynthetic prokaryotes: Oxychlorobacteria (Pro. Science 279: 879–882. The largest of the group is Rangi. This incremental and discontinuous growth is especially evident in large hemispherical or massive corals that may grow to sev. the corallum is essentially a layered sequence of dead corallites and internal dissepi. Similar to psammophiles. Witter A. Marine Biology 154: 547–556. Pfannkuchen M. Rasser M and Riegl B. vironment and competitive success in the Corallimor. New York: Chapman and Hall: 411–435. the crystals are organized in concentric bands. The influence of wave exposure on the ecological zonation of Caribbean coral reefs. Scale approximate. tary Geology 159: 113–132. Scott L. cific precious coral harvests are often unregulated and be moderately common locally within their Indo. veyed by the tentacular cilia to the lips where organic matter is removed. 1998). Isotopic temperature records. However. removal of protons from the periphery of the cell walls creates an alkaline environ. Apparently.water sediment found behind the reef crest. Many produce leaf. 1986. Coral Reefs 263–270. ordinate corals. epilithic. and Marine Protected Areas. Coralline algae also form part of this community. but also opened up his considerable library of underwater photographs to me. The extent of this physical and physiological connected. tesy of Frank Parrish. 114•Chapter Six distinction. riaticum. including extreme fluctuations of temperature and sa.g. the egg begins dividing quickly and be.html. lans they are 45 times larger. tesy of Image Science and Analysis Laboratory. There are apparent differences among host preferences within the same geographic region (e. However. Thus. cated axial skeleton (actually an “exoskeleton” produced by epidermal tissue). Golbuu Y and Richmond RH. the photosynthesis to respiration (P:R) ratio expressed on a daily basis can be greater than 1:1. Molecular evidence for host. 2006. we might conclude that hermatypic corals are always large and zooxanthellate.secretory glands. The first requires core drilling and ex.flow pathway in coral reef food webs. In the opposing view of biological control. Instead. out nutritional quality—“junk food” as Davies (1984) put it. Lazar B. Philo. Cohen et al. or to track changes that have occurred within them. Their tubes. they do possess numerous epi. nobacteria.magnesium calcites are those containing more than 4 molar percent (4 mol%) of the total car. a) Axial pol. 2006. and reef development over the last 34 million years. and mollusk shells. cal Resources. Wörheide G. Table 4–1: Characteristics of Algal Divisions Division Approx.Johnson Space Center ISS004. Figure 5–3: Cloud. shore islands can provide a refuge (e. Figure 7–1: Anterior segments of the jawed worm Nereis sp. 56% of living corals had been invaded to varying degrees by 12 different species of boring sponges (Nava and Carballo 2008).Cambrian. and in Australia. Thunell R. The Hague: Martinus Nijhoff. Bonaire. The Flower Garden Banks. some of which were massive. All are burrow. US Virgin Islands 117–127. the fire corals (family Milleporidae). Reece. and mangroves. Photo. The framework was composed of a particular massively branched coral. This material was collected from the lower intertidal of the Exuma Islands (Bahamas). it is in the path of the cool Canary Cur. are welded by coccoid cyanobacteria. Allemand D. cells per square centimeter is the measure of choice. Earth Science Re. for production of “nori” used as a garnish for noodles and as a wrap for sushi.. remaining static. Inozoan calcare. derived from palps. NDF Work. and survival. but deeper. cilaginous sheaths are produced. Figure 6–8: a) Caribbean anemone Lebrunia danae with pseu. A cluster of the Indo. cialized ventral sac and is extruded from it as a string that is molded by complex movements of a muscular col. Indeed. and many of these unusual compounds have been isolated from sponge. Growth is dynamic and is influenced by grazing her. Feeding habits differ considerably as described more fully below. many organ. the Tuamotu group region has the highest density of such reefs in the world. Carbonate Production Under favorable conditions Halimeda species ex. Anthony and Fabricius 2000). plankton from water passing over them (Reiswig 1971. bodies against the organic matrix in scleractinians: a new tool to study coral biomineralization. ing current along the aboral side of the radioles (Fig. Calcified laminations are formed in association with bacterial biofilms that. In some cases. Figure 5–21). Indeed. pared with the intertidal zone. logical term for resource sharing (e. Another of the Maldives. Macintyre IG. als. but everyone else’s as well. gae may overgrow. cipitate calcium carbonate in place. talline or other mineral deposits may form spontane. lia). 2008. Algae and Fungi. ies may protrude only part of the way from the body wall and do not contact the coelenteron.density bands are thought to form during the cooler months of the year. However. making them more susceptible to damage by strong currents or storms. 52 Overview. that lies in close proximity to the skeleton as it forms. Chicago Deron Burkepile. Montana. Factors that favor the colonization of fleshy. Most new growth occurs at night (e. cluding some nudibranch mollusks (Todd 1981). however. recent work has shown that radial growth is only a few microns per year in specimens grow. rine Biological Association of the United Kingdom 83: 37–39. the Calcaxonia and the Scler. Other reef sponges produce terpenoids. 2008. 1997. or other consumable material (Magnino and Gaino 1998. View from left to right is 5 m.dimensional structure that is elevated above and is distinct from the surrounding. Marine Ecology Progress Series 423: 101. but it can also oc. and Wagner M. Eakin CM. and their abundance further reduces the area available for tures with active growth on all sides. Sabellariids often grow as individual mounds up to several meters across and a meter high. are composed of two tissue groups that are derived from corresponding embryonic layers of the planula larva. but small species (in contrast to serpulids) of. a narrow cul. but not assuredly annual). or transferred within a reef to build islands. Combosch DJ. in. haps as an adaptation to capture prey more efficiently (see Goldberg and Taylor 1996). Abbott AT. 1999. thotrophic (Fadlallah 1983). Figure 4–1: Algal functional groups and comparative structure. the smaller cilia guide them into a series of “food grooves” along the oral side of the radiole arm where they are sorted by size and conducted toward the mouth.7 My Maui 1. Tompkins. suggesting a predominant wave and current pattern in this shallow. the coelenteron. Effects of intermittent flow and irradiance level on back reef Porites corals at elevated seawater temperatures.shaped portion of their body which then de. The pioneers include a diverse assemblage of heterotrophic bacteria that produce a polymeric film at the stromatolite surface. or elongated as pencil. Some sponge groups are significant agents of bioerosion. drecht: Kluwer Academic Publishers: 37–56. Herrmann H. and symbionts. Field biology of Halimeda tuna (Bryopsidales. and c). and the only member of the octocoral family Tubiporidae.htm.like blow. and each of them gives rise to nu.000 km farther northwest where they have developed into atolls. Carnegie Institute of Washington 27: 117–124. nite crystals are often flattened and plate. Raven JA.6–2. Thus. Jones 2008). and are chemi. benthic trawling was legally suspended in 2002 (Conway et al. respiration in most free.dimensional repeated solid will result in the growth of a crystal. but lack the complex porphyrin structure (Figure 3–4). and are aherma. sue and the underlying coral carbonate. incompletely un. tion. but there are also variations by species. Because these struc. Ammar MSA. the flavor has been described as fishy.g. fully established itself in other estuarine environments around the world. eds. Aoba T. Journal of Phycology 42: 453–463. Aquatic Conservation: Marine and Freshwater Ecosystems 13 (supplement): S5. The ecology of the sponge larva. proach to underlying biomineralization principles. Irrawaddy. Crabs not only seek refuge in sabellariid reefs.g. Fukunaga 2008).” This microenvironment is fa. open form of H. Atoll rims are sometimes ring. ing the incorporation of extranuclear bacterial genes. Scharf FS. They lack symmetry and true tissues (see below) and can regenerate all cell types from a single group of stem cells. Thus. giant anemones. 2003.25°N is the northernmost such structure and is 25 My old (Figure 1–17d). ity of the neuromuscular system in certain cnidarians is suggestive of mesodermal construction found in more advanced. et al. ing symbiotic bacteria and sponge archaeocytes. Among these. water habitats. b) De. including some that are unable to survive without them. Wahle CM. 2002. in turn. 84•Chapter Five The Structural and Functional Roles of Sponges on Reefs Reef Growth/Recovery: Binding and Stabilization A number of sponges serve as “living glue. and Palm Beach Counties. centrated weaponry in certain cnidarians may interfere with nerve conduction and cause immobility. ifera: applications and paleoenvironmental analysis. 2008. Micron 40: 169–193. Some sponges are enormously efficient at capturing and removing organic matter and yet never achieve large sizes. most sponges produce two size classes of spicules. Trichodesmium is also one of the most prominent of all nitrogen. In certain types of green algae. and trophic interactions. Lifetimes are correspondingly short. Pro. ment is removed from the base of the tube in a similar fashion. Printed in the United States of America 22 21 20 19 18 17 16 15 14 13 1 2 3 4 5 ISBN.like layers. also appear to favor the successful invasion of coral tissue by these organisms (Rützler 2002b. ———. the island track is very short.like backbone that includes proteins similar in composition to silk. The shape of the colony and its spines are important features that are often used to define species.g. or have become incorporated within the sediment. a pro. The polychaete head is conspicuous and large relative to clitellates. World Atlas of Coral Reefs. Reef cor. Siberia. ranging from sand to silt. Si(OH) 4. Engel S and Pawlik JR. Incidence and iden. Indeed. Risk MJ. ing crustose coralline algae (Rhodophyta. b). Reefbase. Coral Reefs 1988. Most groups of symbiotic cyanobacteria fix nitrogen. The radioles often possess photoreceptor cells or even compound eyes that make them very sensitive to light changes. among others. 2001). While there are modern examples of reefs that are formed by sponges. 1993. global sea levels began rising from glacial meltwater about 18. and lustrous ap. 2001). Courtesy of the Image Science & Analysis Laboratory. and Fitt WK. Marie D. This constitutes a distinct sporophyte generation that proliferates as it bores within bivalve shells and other skeletal and nonskeletal carbonates.bearing foraminifera: har. but are typically reduced or absent on segments associated with the head and the anus. Coral Reefs. asexually produced diploid carpospores. where the eggs are fertilized internally. a starch that differs in the way the glucose units are bound to each other. a hidden species without photo.shaped forms often had double. and exist as dynamically interactive communities of protists.226. and less elaborately constructed spic. tinct from antipathin (the antipatharian skeleton). ganisms that are capable of deep carbonate grazing with specialized mouthparts.edge reefs and large. 25. Marine Biology 139: 991–998. 2003. ganic matter they produce. Of these. Canese S. Titschak.. ganic acids from respiration (e. The process and timing of larval tissue differentiation is complex and may vary according to life history characteristics. Vol. however.30167. for example. Some polychaetes (and other consumers) do not fit nicely into typi. sperm alone are expelled and are brought by currents to eggs that are retained by gonochoric or hermaphroditic females. some important groups form calcar. and worm reefs (Chapter 7). pora tenuis) and planula. Thus.” and again. Fission is far more com. and were abundant in the sediments of Figure 4–21: a) Rice. Their other anatomical features are very similar to those of stony corals (described below). 2006. Guest JR. muda. Krautter et al. trophic bacteria within coral. primary production. called transverse fission. Nontji A. Heidelberg: Springer Dordrecht. vironment in which associated organisms interact and thereby produce a complex community. Biological Bulletin 205: 319–330. Harrington L. is common in the oligotrophic (nutrient. Miles 1991). tion of these cements are numerous. als may form high. graphic aspects of the soft coral Sinularia flexibilis leading to local dominance on coral reefs. is formed as cytoplasmic granules that are not associated with the chloroplasts. but then become highly branched. However. and in the presence of high calcium levels. Photos cour. In the South Pacific. sion may provide larvae with a portable food supply and a capacity to travel long distances. about 100 Mya. Indeed. Thorington GU and Hessinger DA.200 m of shallow. et al. portant element of sponge bioerosion. 2003. More than 100 genera and 5. similar to certain black corals. Large benthic forams were once voluminous contribu. Al. Relation. Matsuda S. ment. Pisera A. De Beer and Larkum 2001). and nerve tracts or rings. and Kitazato H. Bioeroders are often categorized according to their size. Maldonado M. Geology 5: 749–754. and may have a life span similar to that of Gerardia. Models of this system suggest that a membrane.living. In: Freiwald A and Roberts JM. filter and clarify water from terrestrial sources. Tambutté et al. tress and channel systems in both shallow and deep water as depicted here off the north coast of Jamaica and the windward portions of Grand Cayman Island. and Ilan M. and thus colonies are often referred to as “precious” red or pink corals. portant nitrogen. The result is typically a range of growth forms that occur within a taxon. Atoll Environment and Ecology. Members of this genus oc. Some groups develop ex. 2007). Therefore. forming radial compartments like spokes on a wheel when viewed in cross section. 1–1 Coral Islands Several authors have described different types of islands associated with coral reefs. cene growth history of a reef complex on a cool. While certain corals may exhibit a capacity to become autotrophic under certain conditions. Zea S. As the name suggests. doi:10.edge reef architecture around Grand Cayman. Nonetheless. line as they do in the Red Sea. However.. appearing in the fossil record before the Cambrian period. Love GD. which ini. ment worldwide. There may be as many as 200. and are thereby likely to be under direct cellular control. 1988. Members of the suborder Fusu. Rodrigues LJ.rich environments. Meade RH. Davies 1991. Teleostei) dwelling among the tentacles of sea anemones: a cleaner fish with an unusual client? Journal of Experimental Marine Biology and Ecology 329: 161–173.dissolving mechanisms may be localized in the leading end of the filament. 220 11 Reef Echinoderms. nai Peninsula.000 years (Macintyre 2007). ganism’s metabolism and its environment. While it occurs prominently below low tide. Individual species are seasonal and only a few remain abundant throughout the year. Calcification begins when the utricles fuse and form an intercortical space that is isolated from seawater. tory relationship between self. and octocoral sclerites include some that resemble spicules.fws. Sorting sponges. Simple symbiont.g. covered daily by the tides. There are six crystal systems. metic fusion produces the diploid sporophyte generation. far below the depth where they could have grown. Journal of Experimental Marine Biology and Ecology 335: 167–176. These specialized polymorphic polyps may include nonfeeding individuals that defend the colony with one or more stinging tentacles. Interestingly. New York: Academic Press: 4: 59–105. Photo courtesy of Dieter Fieger. Chemical variability within the marine sponge Aply. Russian Journal of Bioorganic Chemistry 27: 2–16.2148. cations for dispersal. Reef Corals and Their Allies•125 referred to as “antihermatypic” (Schuhmacher 1997) be. Fosså JH. der and Le Campion. 2007. 2007. The thorax consists of two segments beginning immediately below and behind the mouth. operculum.building corals are said to be hermatypic. NASA. and Kauai. Many such formations create a complex three. Hydrobiologia 471: 101–110. although two families of planktonic species are abundant in the open ocean. cessful coral settlement (Kuffner et al. including a reef crest that is often exposed at low tide. sea levels. Farmer MA.water runoff. patch reefs.g. or chronically with hu. and Hine AC. and even these may be covered by the absorption of carotenoids and allophycocyanin (not shown). occurring only as symbionts in certain groups of tunicates described in Chapter 9. shop Case Studies 1: 1–25. volume. and while that class of minerals is a large one. Likewise. posed a large part of the reef framework. A Little Crystallography Crystals are formed by minerals when they be. Geochemical perspectives on coral mineralization. Third. The other 11. removing as much as 1 kg m–2 yr–1 on some reefs. However. maining on the fringes of SCUBA. However. Coral reefs may occur uncommonly at salinities as low as 25 ppt (Coles and Jokiel 1992). Baja California Mexico. 2003. Gerardia and Savalia completely engulf their hosts and then begin to secrete their own very distinctive skeleton (Parrish 2007. another epithelioid called the choanoderm. but serve to point out the gaps in our knowledge of these important reef organisms. The result is often a massive ridge. Management notwithstanding. One of these symbiotic Soritidae. bivores (see below) that limit the development of the calcareous thallus on one hand. Postglacial growth history of a French Polynesian bar. spontaneous division of the body into one or more parts. This raises the question. Hydrozoan polyps are measured in millimeters. but there are distinctions among algal groups (Chapter 4). Some are so heavily armored they cannot contract completely into their calices (singular = calyx). Science 177: 270–272.defined pharynx. but this chapter emphasizes those that are sessile and live in tubes or in the reef framework. Wilson DE.specific bacterial group known so far (Usher 2008). These and other fea. Westfall JA and Elliott CF. sclerites are con. Dubois J. Browsing animals target their food specifically and have little or no effect on the substratum. Heliopora’s range is very similar to that of Tubipora musica (Veron 2000) and both species are Figure 6–22: Large (~8 x 10 m) encrusting Sinularia colonies from the Great Barrier Reef may be hundreds of years old. ment in eukaryotic plants. However. Rossi S. Colony defense and behavioral differentiation in the eusocial shrimp Synalpheus regalis. Arvedlund M. 2003.. ments replace forams on reefs in deeper water (Collen and Garton 2004). Despite the limited diversity. anthozoans are sepa. New York: Plenum Press. Yahel et al. Figure 1–19). the degree of development within the female. Gautschi G. Photo courtesy of Jane Prince. recruitment. proximately spherical) ranging from 0. In contrast. reefs on offshore islands are fairly common in this region. Thus. 2. The Biology of Some Other Calcareous Chlorophytes Many other members of the Bryopsidales are present on or near reefs and contribute aragonite to them.like or proteinaceous external covering and are referred to as “naked forams. the Niger and Zambezi rivers pre. 1997. tion to the head.dwelling sponges. tions: pre. The column is often more complex than that of other anthozoans. nematocyst types. and Negri A.. The form of mature larvae can vary. Andréfouët et al. There are 34 species that are currently recognized (see Verbruggen and Kooistra 2004). P a cif c p la te S o lid d e n se ro c k Zone of magma formation Hotspot H aw aiian ridge Kauai 5. Science 299: 1846–1847. there are no species known to have evolved a symbiosis with zooxanthellae. Hsu C. Coral reef sponges: do predatory fishes restrict their distribution? Limnology and Oceanog. Members of the imperforate family Soritidae are impor. However. 79 tion. tos courtesy of Paul Humann. Kramer PR. emy of Sciences. and Sporadotrema. Bioeroders vary in their be. Habitat and fauna of deep. Similarly. lium sp. ported to the organic matrix where high pH facilitates its conversion to carbonate. Weiner and Dove 2003). 2003). kaline and enriched in carbonate. acteristic laminated appearance. Kelaher BP. 1998. Cold. and rock or coral crevices. some Figure 6–35: Reefscape at Punta Galeras. when considered together. squid. ing mechanism thus appears to be primarily chemical. coral. likely as an intracellular endosymbiont. terrestrial environments. or those engaged in poorly regulated markets for the curio or jewelry trade. Nitri. Bayer FM and Grasshoff M. bon and energy flux in reef corals. This erosional feature is caused by endolithic organisms among which cyanobacteria are prominent. the an. Adl SM. how. 2004). However. Scale courtesy of Peter Glynn. umes of hydrocarbons have been discovered amid their deposits offshore of Tunisia and Libya. thellae. Chondril. herms (Chapter 1). Ball EE. ized eggs remain within the female or female portion of the plant where they are housed and perhaps nourished during development. der stress (e.co. In type 2 matrices. which facilitates feeding by the polyps. 1997. Tropical sponges in the genera Agelas and Dysidea. shrubs. These include the Atlantic genus Penicillus and the Figure 4–11: Life cycle of Halimeda. and there is no evidence of persistent hotspot activity (Clouard and Bonneville 2001). Element partition. The ecology of nudibranch molluscs. Griffin S. gener. com. leothermometer in paleoenvironment analysis and paleoceanography (late Miocene. Coffroth MA and Lasker HR. though once shallower and farther south. 132•Chapter Six follow egg release during mass spawning. and Coates AG. Sidri M. mucose. synthesis 68•Chapter Four As a group coralline algae are more common in reef areas with clear water. eastern Indian Ocean. Lewis and Von Wallis 1991). or different combinations thereof. and may have been capable of life spans of up to 500 years (Wood 2000). vival dynamics of scleractinian coral larvae and impli.energy en.Johnson Space Center STS046–77–31. The thallus of these and certain other macroalgae is further differen. Fishes with established symbioses are unaffected by host nematocysts owing to their mucus coating. Facies 44: 265–282. allowing them to tunnel deeply into coral skeletons. rent.foraminiferal reefs. The Order Gorgonacea Gorgonians. Column buds can produce small radial polyps from a larger axial polyp. (b) removal of protons or other calcification inhibi. Croix: its implications with respect to late Cenozoic coral reef development. and cells within the colony (Gateño et al. which are abundant in reef waters with a con. most of which lack zooxanthellae. rDNA): finding the molecular and morphological gap in Caribbean gorgonian corals. Construction of early Proterozoic (1. The material has a marble. Note that in this scenario. The filaments. Zhao et al. but selected and stored by a specialized ventral sac. Ma. plications for the physiology of the hexactinellid con. the Octocorallia) produce a skeleton composed of microscopic calcareous elements (sclerites). 2006). However. marily by coralline algae. eds. Many of these sponges were globally important in reef construction. The extensive sheaths left behind by Solentia are subsequently fused by microcrystalline micrite. Nitrogen fixation or dissolved ammonium up. Moreover. branched. 2000). rington 2008).. 2007. The absorption of light by chlorophyll causes the ejection of electrons that are shuttled through a complex electron transport system. ceedings of the 8th International Coral Reefs Sym. tion of nitrogen from benthic sediments by three spe. However.up reef (Neumann and Macintyre 1985). Effect of car. low rocky surfaces and may include phototrophs with low light requirements. Blanchon and Jones 1997. the pinacoderm and choanoderm are syncytial networks of cytoplasm that al. nated by coral growth owing to generally lower wave energies and a more consistent rise in sea level over the last few thousand years. not all of which are simply explained (see below).like “reefs. Geister 1977. Determinate growth and modularity in a gor. Endolithic microflora are major primary pro. Instead.dimensional framework (Figure 7–5) that supports a relatively rich and diverse assemblage of algae. Zooxanthellae are known to take up from their hosts significant quantities of vital nutrients including phos. Boulder: Geo.Horani et al. growing in place). and (c) the ability of the coral community to respond to those forces.depth description of these topics. lae. Papua New Guinea. ed. Coral Reefs 26: 53–68. ments from this layer and is typically the thickest region.g.photosynthetic nitrogen.1371/journal.effort ratio. and ascidians (Chapters 6 and 9). The two shorter axes are desig. Chen CA. Philosophical Transactions of the Royal Society B: Biological Sciences 363: 2641– 2650. Phyco.000–6.shaped crowns and the conspicuous red and blue pigments found in some ser. Quinn RJ. cated than revealed by a simple trophic relationship. 2006. but the most pervasive is the process of budding. Brominated pyrrole alkaloids from marine Age. such as their role in reef formation (baffler. After Veron 2000. 1999. Interspecific hybridization and re. Pro. called zooxanthellae (family Symbiodiniaceae). One species in particular.Hurricane Allen conditions in Discovery Bay. trum. the list becomes a bit shorter (Appendix 2c and d). These include unicellular forms that divide by simple fission (order Chroococcales). Larvae of many species avoid bright light or do not survive in turbulent environments where colonial tissues can be shredded (Grigg 1965). or as the product of multiple cells or cell layers. 1999. and Saunders JB. 2004.500 named species (Norton et al. located parallel to the coast of northeast. 2003. for example. In the protection of the crest. Domart. can become ex.g. the zygote that forms within the fe. Miake et al. Japanese god of happiness. The evolution of inorganic carbon. 2008. these algae are generally un. cies and have evolved elaborate biochemical pathways that detoxify the allelochemicals produced by their prey which may involve some form of ejection. often as branched or more typically as dome. and cementing agents. and are controversial because they often do not leave evidence of the cellular processes that produced them. parathorax. Weaver DC. Coral Reefs 20: 69–76. Proceed. nomer (e. Many groups were cone shaped. and the shelf atolls of Belize (Gischler 2003). nisms (reviewed by Raven et al. gies. turf algae are almost always the dominant form. In: Lee JJ and Anderson OR. Archaeocyatha. Marie D. Sloan and Irlandi 2008). Members of the family Terebellidae are often tubico. even though little is known about its role in the growth of the host or its symbionts. Marine Pollution Bulletin 51: 570–579. Density and habitat of three deep. ed. and/or the slowing of volcanic and metamorphic outgassing as large continental landmasses became established (Kasting and Ono 2006. 2007). it has been suggested that perhaps clonal planulae are produced un. quent swells.. Figure 5–22: Sponges pro. Some of the most diverse and ecologically dominant chlorophytes found on tropical reefs are members of the order Bryopsidales. Biologically induced miner. Sponges are such efficient suspension feeders that reefs with large populations of sponges are capable of depleting pico.. Mutualisms among species of coral reef sponges. Truncated pyramids are formed at opposite ends away from the axis of symmetry. Sustainable management guidelines for stony coral fisheries. serpulids may pro. a few that are benthic and feed on detritus. Fixed nitrogen is exported to adjacent reefs by compounds that are re.like sheath). Margino. Badger MR. nected throughout the colony surface by canals. ture of Galaxea fascicularis exert septa: a high resolu. eralogy and Geochemistry 54: 151–187. fied tropical seaweed genus Halimeda (Bryopsidales. an Evolutionary Per. causes energy expenditure from the necessity of sediment re. shell fragments) within the ma. Hydrobiologia 501: 149–166. Resistance.Alsumard 1999. 2002. Macintyre IG and Reid PR. thus delivering water to the spongeocoel. like colonies on reefs. ological state of the cell. Colonna M. tains reef. Marine Fisheries Review 64: 13–20. Clade differences among symbionts in different hosts may account for some of these differential responses (Erwin and Thacker 2008b). ary Perspective 2: 273–293. by convention. b) Polyps may be closely packed and share a common wall or c) arise farther from the parent and become more clearly separate. Courtesy of Springer Science and Business Media BV. a division of the Alcyonacea in which the polyps are connected to others by canals called stolons (Fabricius and Alderslade 2001). ferent types of megascleres. Photo courtesy of André Freiwald. 1989. New fission processes in the zoanthid Palythoa caribaeo. cial reference to the Great Barrier Reef. Less subtly. Density. Coral reefs in tropical regions that exceed these limits may favor only the hardiest of species and are likely to exhibit low diversity and poor reef development. Indeed. vent reef formation along considerable lengths of the tropical coasts of west and east Africa. An Introduction to the Structure and Formation of Modern Reefs •7 obligate symbiotic relationship with photosynthetic cells called zooxanthellae that affect the nutrition.Paige CA. The anemonefish symbiosis: what is known and what is not. Photo from West Papua. and when decay rates are calculated or detonation dates are known. wealth. damage that is quickly repaired by meristem. casionally even some of the tentacles. and other factors. Figure 1–10: Many of the Maldive Islands (right) are atolls that are composed of miniature atoll-like islands called faroes (left). and west Africa. but also acted as an attractant for further brittlestar coloniza. Helman Y.. often conforming to its geometry as the term “fringe” implies. the bundles more often do not remain integrated for long and break apart within some minutes after spawning. depending on their properties. In addition. For example. if present. the abundance of Halimeda may gain from an overall resistance to herbivory. Photos courtesy of the Image Science & Anal. Most of their lagoons are well circulated and open to the sea with more than one channel. life can be rigorous. Chang et al.226. Prepared with liquid nitrogen freeze. the thallus is also readily fragmented by water motion. The origin of palytoxin is unclear. Evolution. Scripps Institute of Oceanography Joe Pawlik. Such a sponge might grow to double its surface area. Bignot G. The genus Halimeda. daria. protein content. This keeps the pH and calcium levels in the organic matrix high. 362 Appendices.West Pa. antipatharians. which operates at high levels in calicoblast cells during the day. 2007. fleshy macrophytes are to some degree inversely related. rine Ecology Progress Series 121: 211–216.209. 1986).. ally augmented by other functional groups composed of carbonate. could also be considered a cementing agent. Agar is used as a growth medium for microbes. minifera promote calcification by elevating their intra. ———. Eakin 1996).. day off British Columbia are essentially unchanged from those Jurassic reefs. Koutsoukos PG. These miniature atoll islands called faroes (Figure 1–10) are unique to the northern Maldives and compose a large proportion of the roughly 1. Photo courtesy of Jere Lipps. while retaining nervous and digestive tissue connections among them. posium 2: 1255–1258. Lateral growth is inhibited at the base by sand motion on the channel floor..cyanosite. brooder categories. variations in local populations. is also quite variable within and between species groups that have been rec. rounding seawater and are referred to as low microbial abundance (LMA) sponges. these endoliths can have a significant impact on both the amount of carbon fixed within living and dead corals and the amount of carbonate removed from coral reefs (Tribollet et al. the larger macrogamete is referred to as the egg. including the “Poribacteria. New York: Academic/Plenum Publishers: 173–185. a testament to the general acceptance of Darwin’s model. d. Adey WH. et al. Frame Reef Formation and Structure The framework of any reef requires the presence of one or more foundation species that define its physical structure. San Diego: Academic Press: 157–220. However. stylasterines rarely occur in large numbers except in unusual environments. and even black can be exhibited. Origin of atoll la. University of Technology. 1998. In addition. 2008. and a deep layer next to the substratum (in encrusting species) or around the inner skeleton described below. 1969. often composed of a tough and resilient protein. as well as erosion. 2006. or use a combination of the two during feeding (e. nivorous because they are capable of capturing relatively large plankton. Budd AF. fying groups of hydrozoans will be considered here. such as the tiny volcanic island of Uvea (northeast of Fiji and west of Samoa). 1991. ken during storms.g. In addition. and its rim develops 20–40 km from shore. ally merging with others during extension and accretion. Palaios 25: 754–763. Leys SP and Eerkes. However. blue corals can be quite abundant and can form colonies several meters in diameter. Indeed. and the behavior of the occupant vary considerably among groups.g. Global biodiversity patterns of ma. quieter water than A. Aquiferous systems and pathways of water flow are shown in yellow (after Bayer and Owre 1968. 1987. Wulff JL. perhaps by the acquisition of dissolved organic matter (Graham et al. Australia 1: 533–560. Coral Reefs 6: 251–262. eds. 1990. boring spionids as a group are regarded as substantial in their bioerosive effects on coral reefs (Hutchings 1986). calcifica. ergy budgets of shallow. Polydora villosa. or they may be hollow and deliver venom to active prey (Figure 7–1). Siliceous sponge spicules in coral reef sediments. competency. Heterochone calyx. Phosphorous (e. Co. ation. surface.. includ. it is often considered a limiting factor in coral nutrition (Yellow. Pho.P and Dauphin Y. cyanobacteria symbioses: a comparison between host. Octo. uted as a cryptobiont. sium. and Cappola VA. Goh NKC.forming organisms. pounds.g. A few genera are abundant rock borers and are of interest here for that reason. Even groups with rudimentary tentacles capture prey or detritus using mucus nets or mesenterial filaments. ate into a complete colony. they are released at night at a predictable month and phase of the Moon..6695. ed. ositional environments modern and ancient. An alarming threat to the red organ pipe coral Tubipora musica and suggested solutions. cervicornis formed much of the carbonate at Alá. Association of the sponge Tethya orphei (Porifera.lived Caribbean barrel sponge Xestospongia muta. Effect of grazing on coralline algae in seasonal. Scavengers feed on carrion. Devonian stromatoporoids on reefs in Western Australia. lonial cnidarians (e. brian period. Sperm clusters fuse with the walls of the exhalent canals and are then released. pacts in Aceh Province and North Sumatra. Thus. upright tufts that form a canopy less than 1 cm high over the sub. b) Hamelin stromatolites at low tide. duce calcified shells and are heterotrophs. NASA. Steindler L. Logan JB. eds. often consists of 24 septa. 2001. Grajales A. Solandt JL and Campbell AC. mid.diversity reef systems that are found scattered along the west coasts of the Americas ranging from Ecuador to Mexico (Glynn and Ault 2000). Fabricius KE. and Wörheide G. The roles of carbonic anhydrase in photosynthetic CO 2 concentrating mechanisms. Barnes DJ and Crossland CJ.like crystals composes the myostracum. they are overgrown by algae and are more extensively bored. Second. McFarland FK. Kvennefors ECE and Roff G.000 km long. but are also known to occur in hosts with a low microbial abundance (Southwell et al. nies.associated or. Baha. lata) corals. Eusociality in a coral reef shrimp.4. merous spaces between the crystals (i. menting agents. allum (colonial skeleton) composed of individual cup. although it is also known from at least three Caribbean species (Erwin and Thacker 2007). but it may best be referred to as a “fringing barrier reef” (Kramer et al. Riul P. and is facilitated by a combination of favorable wind conditions and the presence of relatively weak eastern boundary current systems. liceous spicules and skeleton frameworks in sponges: origin. Skel. 146•Chapter Seven These feather. Williams and Hallock 2004). In: Gorb SN. haps using them as optical fibers to maximize the use of available light (Gaino et al. That is because area changes with the square of the length (12 ). about 0. The crests of its exposed reefs and their characteristic biota were constructed at sea level. These are specialized as digestive tissues. Any of the island categories described here can be additionally categorized as vegetated or unvegetated. and can occur in a variety of habitats in the Indo. Larger foraminifera and sedimentation around Fongdale Island. ch. in. Species richness of recent Scleractinia. regeneration. In. Norwegian shelf. tentacle. Journal of Materials Chemistry 7: 689–702. Huston 1985). 1997). cesses. for example. Reed JK. bleaching. part of French Poly. Unson MD. and even annelids (Hickson 1924). Usher K. Elsevier. Vertical rock faces in the intertidal zone are of. and Sazima I.. Australia.like tubes that compose the unique bright red skeleton of Tubipora musica. viri. The head may be modified according to the type of diet. Limnology and Oceanography 34:896–912. This is the largest island barrier reef system in the world. take.” similar to a traditional kingdom. the calcium carbonate contributes to the fine fraction of the sediment.shore reef system. Hine AC. which is employed to fix carbon into an organic form like glucose. Marine cyanobacteria: a prolific source of natural products.aboral axis. cal. Certain noncalcareous rhodophytes (e. The clearest examples of bank reefs are those that Figure 1–12: Detail of Rangiroa Atoll. and intertidal algal “cup reefs” around the island of Bermuda (Ginsburg and Schroeder 1973). Competence periods are short.. The role of zooxanthellae in the nu. produce a calcareous skeleton primarily in the form of magnesium. it has also been proposed that Indo. stromatoporoid. are quickly replaced *The nanoscale is a diminutive that refers to a billionth (10–9 ) of a unit. Evolution 52: 379–393. While the exact mechanism is unknown. Indonesia. Indeed.Bard C. but more often. 1988. including staghorn corals.08–0. and like them. for example. Photos courtesy of Katha. gations of corals surrounded by sand or seagrass. Burke RB. shrimps. Dordrecht. Red algae are also included in this supergroup. The cells may continue this process. 2006. Scyllidae). 1997. The same operation is thought to elevate the calcium supply within the cali. and families. alkaline waters. hydrodynamics. such as diatoms. and Roncador Banks) to recommend marine.g. chemical.pigment complexes that absorb energy at different wavelengths and act as light. ally significant broadcasters. Pile A. but the extent of their overall effect on reefs is unclear. Annual Review of Systematics and Ecology 29: 179–206. In: Boone DR. Jompa J and McCook LJ.like cen. Hexactinellid reefs are currently endangered. Cockell CS. plex compared with gram. The abdominal region constitutes most of the body and is easily recognizable by segments with distinct chaetae. Coral Reefs. Shinn EA. ———. grooved.building corals at Green Island. quisitely ornate shields or thorny scales that may differ in complexity according to their location on the colony (e. tant tropical representatives. 2005b.HDTV. 2006. and some formed massive or chain.sensitive cells. The lithobiontic ecological niche. which places it in the orthorhombic system. The group became extraordinarily abundant and diverse in shallow Paleozoic seas. a. Darwin’s subsidence model has been modified as a neo. genus Synalpheus. including mound. the granules at the density) and dark (high. Brooding may continue for several hours or days within the egg. However. All types of reefs are riddled with organisms that bore into or otherwise remove carbonate (Table 1–1). many groups are a rich source of al. alline algae in Abrolhos. The only group of note is the dictyotacean genus Padina (Figure 4–14). and Bates NR. 2002b. that is not al. or changing from a bushy to a more planar growth form (e. crobial diversity of extant stromatolites in the hyper. Goldberg WM and Taylor GT. Having extended the definition of coral this far. Likewise. Geological Society of America Bulletin 113: 837–854. at least in the short run. brooding coral species transmit their zooxanthellae directly to the eggs. The incorporation of magnesium is a complex phe. the provision of settle.rayed forms called triradiates (Figure 5–8) and single.5 times the length of the modern Great Barrier Reef (Ghiold 1991. or in the cytoplasm. fishes (Wulff 1994). tation during storms. Chapter 8) fall into this category. Photo by author. Likewise. chaetes are ecto. Light inside sponges. the anterior sterile portion. represents the group and clearly separates them from the distantly related Milleporidae. cent Corals. the proportion of carbonate will increase.curcuphenol. 1881. Sammarco PW. 2007). especially certain members of the wide. Ultrastructure and embryonic development of a syconoid calcareous sponge. and there may be as many as 18. leothermometry. There are two additional clades within this group.containing mesenterial filaments. Phycologia 19: 110–138.92537.dimensional framework attributable to the production of a scaf. Essex: Longman Group UK. and Cunningham CW. Internal conversion will occur in species that cotransport bicarbonate + H+ across the cell membrane. known since at least the Upper Ju. As the chloroplasts mature. In ad. The process is also thought to have a biological component. often only 2–3 days.P. or even as a food source (Randall and Fautin 2002. and survival of some western Atlantic reefs during the Holocene Transgression. 118•Chapter Six Symbiodinium. particularly the larger members like those in the genus Spirobranchus (Kupriyanova et al. sometimes propagating dense clusters of geneti. Eisenhauer A. Blooms of Trichodes. Figure 1–1b). ronmental information (Druffel et al. Ehrlich H. ments. tremely common in dead reef substrata on both Atlantic and Pacific reefs (Moreno. Dark calcification and the daily rhythm of calcification in the scleractinian coral Galaxea fascicularis. there are three that are unequivocal and vital: (1) they influence the calcification process. Sa. Loya Y. coupled with the increased density of the cooling volcano. otic phototrophs. editorial director of sciences of the University of Chicago Press. as could chelating agents. calyx over a dense bed of Farrea occa (Fo). growth is rapid. Most of the losses were in the form of sediment derived from bio. Nonetheless. ever. centration of up to 12. However. Some of these.thick green bands they form in coralline aragonite. These can produce injury to non. turbulent. C. though they account for only about 0. Calcareous sponge spicules. eds. Some specialists refer to reefs that are produced by small skeletal pieces or sand grains that are cemented together as bioherms or “reef mounds. These are the only known examples of living sponge reefs in the world (Conway et al. The phenology of sexual reproduction by green algae (Bryopsidales) on Caribbean coral reefs. velop muddy bays or mangrove forests. making the term “center Figure 6–14: a) X. Collen JD and Garton DW. Marine Ecology Progress Series 303: 133–144. 2001. Lang JC and Ginsburg RN. In addition. Cabioch J. and López. ———.2 % of the Earth’s surface (Reaka. Spur and groove formation on the Flor. Gastrovacular circulation in an octocoral: evidence of Reef Corals and Their Allies•139 Stylasteridae) in shallow water of the Patagonian fjord region. ate either incidentally or superficially. courtesy of Andrew Eksner. mental Marine Biology and Ecology 14: 231–249. Age. cludes both filamentous and single. hydroxyapatite (HA). Tahiti 3: 95–103. 2003. on coral reefs.like jaws or other specializations that allow them access to blood or other fluids. like the apple that does not fall far from the tree. The coralline algae (family Corallinaceae) are among the best known of these and are composed of 25 genera. Hardground coral communities are present. sometimes in a conspicu. Lavik G. The forealgal crust facies of the deeper fore reefs in the Red Sea. More commonly. glutinated tests can be traced to the Ordovician and perhaps to the Cambrian period.loving).Daryan B.like network of interconnected surface canals called astrorhizae (Figure 5–14 inset). Biological Bulletin 204: 146–154. tion. The geologic time scale. or storms) and still survive. To emphasize the respiratory function. These agents of bioerosion include a wide variety of organisms that are sometimes classified by their size. Australia.sand patches. fied. Free. like” reefs are present. The diagenetic cements that fill reef spaces by binding sediment are also high. USA 106: 15374–15378.” and lastly. Some of these unattached coralline reefs are threatened by nearby commercial fishing (e. and Weis VM. tralia. dition.0. Photo by author. typically create a network of fine cavities or pits within carbonate substrata. Ox. New Perspectives in Sponge Biology. 1975. some. algae. Fernando de Naronha. lite wall and floor.. the relationship between corals and their zooxanthellae can be variable and may not always explain their reef. www. The epidermis includes the oral region cover. The shape of atolls.Legentil S. The innermost cell layer is composed of flagellated collar cells. Chloro. we would also rule that corals without symbionts (also called azooxanthel. or to mucus. egory. Changes in light availability. plankton. coccus. 1981. sociated microbial abundances impact marine demo. The family Spionidae is a large one. tions.like parapodia found on each body segment. and shoals. Science 292: 679–686. 2–5 cortical layers and utricles may develop in the new segment. c) Wind. Puglisi et al. persal.29046. although some species are burrowers. Hexactinellid Reefs While modern demosponges have important roles to play on reefs. ing (Figure 6–34b). and Life Histories of the Eukaryotic Microor. Indeed. rather than the more typical calcite. the largest in the Tuamotu group. b) Cross. Glynn 1983. The Maldives consist of 26 such structures. soft corals may constitute nearly half of the living surface of some reefs there (Tursch and Tursch 1982. Sánchez JA. this hypothesis has become highly controver. a number of algae display anatomically simple forms including single eukaryotic cells. cies of bamboo corals (family Isididae. This latitudinal limit is imposed by temperature decreases. tated as randomly oriented crystals on the surface of the thallus. A new bag is put into position to replace it as frequently as every five minutes. et al. These polychaetes are often found in areas that are environmentally rigorous. lin: Springer: 601–617. a phenomenon called coral bleaching. Some tubeworms are capable of turn. Mor. Bellwood DR. 2003. violet. brown algae reflect a completely different lineage and are placed in the supergroup Chromalveolata. 2005). 1999). which adds an additional 750 reefs in an area of about 6. Filamentous cyanobacterial species may be restricted to the ectosomal surface or ramify throughout it. The first is Ca. tion. Wilkinson CR. Kerswell AP. Muller. Tanaka Y.induced cellular calcium elevation. McConnaughey TA and Whelan JF. less. mentary Geology 185: 205–213. trogen fixation (acetylene reduction) on a coral reef. Boero F. ogy and Sociobiology 51:488–495. most reefs along the same Western Australian coast are the Houtman Abrolhos. increased rain. or tunnel. three. volcanic material) is colonized by reef organisms. ans. By contrast. and Phipps CV. is found in the Indo. and Hentschel U. reef growth is likely to be relatively slow inside the lagoon. Diversity and expression of nitrogen fixation genes in Hentschel U. although some groups maintain an obligatory symbiotic relationship with endosymbiotic algae. Indeed. corals. Uriz MJ.000) belong to the Polychaeta. GinkoMaps Project. 2005. meaning that for ev. The crustose coralline alga. brooding of larvae can no longer be assumed as an ex. Thus. These may produce sediment. Hottinger L. ribbean to depths of 30 m or more (Hillis. having devel. Some cyanobacterial species also contain a red pigment. Many such reefs result from abiogenic processes includ. and ky refer to billions. Cabioch G . Algae can be organized into 3–4 often co.g. branch mollusc Elysiella pusilla. The distinctive pink. and unusual island was declared a National Monument in 2009 (see Chapter 16). Light. and Dejardin P. ten abandon old tubes and construct new ones (Rouse and Pleijel 2001). Manila 2: 201–206. and before the evolution of skeleton. and complex interactions. which in some cases are present in sufficient quantities for commercial use. 1997. or persist chronically for several weeks. ria.Antarctic (Ramos and San Martin 1999). polychaetes are often among the first to in.unb. making this group among the deepest. are mostly small mobile species that are common components of coral reefs where they prey on cnidarians and other sessile invertebrates. 1999. Some. 2006. These two terms may apply to any reef. composed of a large sporophyte stage and a microscopic gametophyte stage. This self. buds refer to small protrusions of tissue from the parent that form new polyps. Loss of cyanobacterial cell populations or their pigments can result in the phenomenon of sponge “bleaching. Con. When carpospores are released. Journal of Ex. orthorhombic. Anabaena. sponges exhibit low diversity owing to the dominance of certain polychaete worms that use their hosts for food. Palaios 3: 233–242. rokuju. xanthellate. Paleoceanography of coral reefs in the Hawaiian. both groups exhibit a tendency for specialized morphologies. ognized thus far (see Ten Hove and Kupriyanova 2009). and then transported to the mouth along a ciliated groove. ogy 50:1–55. part 1: reefs. ians.West Pacific region where large island groups—the Tuamotus. they are also exploited as a commod. However. logia 530/531: 433–441. rated from host cytoplasm only by their cell walls and of calcification” misleading (Cuif and Dauphin 2005 a. Nonetheless. These are characteristic of the Indo. also caught up to sea levels that stabilized after the mid. Inter. In: Teichert C. oped. Ef. 2006. a scaffold of poly. Photos by author. longitudinal and trans.Rivera E and Paul VJ. they typically require hard substrata and often exhibit rapid growth. Jones RJ. neath the tentacle surface. 2007. 2000).000 years old. As further adaptation to the tubicolous habit. Thus. Thus spur and groove topography in the region is seen as primarily but not exclusively due to erosion (Gischler 2010). 2005. Stanley GD. Photo courtesy of Paul Humann. duced exhibit various degrees of buoyancy. egories (reviewed by Stoddart and Steers 1977). Figure 6–31: Caribbean coral skeletons displaying intratentacu. ates. associated microbes (see Golubic et al. Still other corals in temperate regions dramatically increase the numbers of their symbionts during the warm summer months. Nonetheless. bryozoans. II: allelopathy. even within the tropics. Opresko DM and Sánchez JA. 2003).000 m and most are 0. epidermal structure. typically co. Both Udotea and Penicillus have a broadcast repro.5 meters in diameter (Figure 6–21). Toyofuku T. actinia) from Chile: characteristics of the biocoenoses.. cific coral reefs. especially with cor. Hillis. Note the black zone. Assessment of coral reefs using herbivory/ nutrient assays and indicator groups of benthic pri. Cnidar. Radax R. AZ. Exuma Islands. a low. jacent cell walls and membranes are discontinuous and are connected by a proteinaceous plug. choric.calcite. Coral reefs: algal structured and mediated ecosystems in shallow. It has 7–8 small islands. One of the largest and most prominent of these. The nervous system is composed primarily of neurons with two processes (neurites) emanating from the cell body. ment: differential localized dispersal and recruitment patterns in Great Barrier Reef corals.like particles. 2009.5 to 3 µm) and is com. Schwarz et al.” The result of this condition may be mild and inconsequential. tion of coral reef sponge populations.most algal ridges in Cozumel. It is thought that high. thus maintaining a char. Colonization. shaft and hollow tubule readily distinguished. perimental Marine Biology and Ecology 96: 127–146. US National Park Service. most of which are 20–50 mm long at maturity. Overview Algae and Foraminiferida are part of a diverse group of organisms that have been traditionally excluded from other eukaryotic groups (animals. Risk KM. However. pora) species (Coelenterata: Scleractinia).g. Johnson CR and Mann KH. Zooxanthellae are incorporated into the egg and are increased by division within the larva and thereafter. 1999.g. Some cyanobacteria in turfs produce toxins (Chapter 3) that kill live coral tissue on contact or otherwise reduce suc. Clanahan TR. This makes it a losing proposition for a single layer of choanocytes to pump out an increasingly large volume of water. as protection against sorcery. ton they consume. 54 Brown Algae.000 m3 (200 x 106 liters) of water every second. and physiology (e. 1994.0002429. and the chemistry of crustose coralline algae (Rhodophyta. The biological processes that give rise to one form or the other are not understood. typified by stromatolite reefs described in Chapter 3. and were highly diverse morphologically. sp. although unipolar and multipolar neurons have also been described. a substantial amount of it may not be accounted for (Edmunds and Davies 1986).g. both regional and local. Comparative study of the chemis. A few scleractinians exhibit as many as 8 cycles and 768 septa. synchronized releases. 2008). the area covered and un. 1998. Caribbean shallow. Penicillus is quite distinctive and is commonly called shaving brush algae. and community levels in coral reefs: a review on interactions and control by carbonate chemistry. sperm typically arise from choanocytes that cluster into cysts. Ward. Cementation oc. 1999). These relationships are evolutionarily significant and will be considered further in Chapter 14. Marine Ecology Progress Series 323: 107–117. 79 Overview. Prouty NG. Rinkevich and Loya 1984).Jones 1988. a volcanic center 100– 200 km across where hot magma (molten. Symbiosis 10: 23–46. lar arrangements and therefore their physicochemical properties are distinct. while spur. which are of. 2005. giganteus generally appears to be an obligate coral as. Some species can simply pinch off a teardrop. Holocene coral reef rubble and its binding agents. Ecosystems of the World. demosponges may be the largest and most prominent benthic organisms. ria that live in close association with the cyanobacterial sheath. but the lay. Tiny Rose Atoll (American Samoa. Some poly. and Scrapers. 2006). Bayer FM and Macintyre IG. lite structure of Montastraea cavernosa: Note that 12 primary and 12 secondary septa are serrated and costate (shown in boxed area). 2008.or plate.. 2002). eds. smother. Lepore E. Bulletin of Marine Science 1972. Indeed.rays.West Pacific locations.dwelling shrimp are eusocial organisms. Ferguson and Davis 2008). Almost all members of this family form a calcare. is initiated with ring 6 counting from the initial growth ring (I). perhaps not the best candidates for commercial exploitation. and the endosome with its aquiferous system. thus encouraging sediment deposition (as well as larval settlement). Figure 6–18b).fao. Tambutté E. Paerl HW. described below. For example. and susceptibility to herbivory of diploid vs. but bristle with knob.000 years when sea level rise slowed and the reef developed Figure 1–22: a) Living Caribbean staghorn cor. An overview of biominer. and Pawlowski J. rhizoids may possess attributes that are analogous to the roots of vascular plants and function as “pseudo.water reefs generally and coral reefs in particular. They are also among the most an. and may provide the zooxanthellae with either ammonium or organic nitrogen. ing those that are functionally male and female at the same or at different times. and in some cases produce large patches (e.shaped bur. grasses. Chang WL. They must be composed by organisms that secrete. Yellowlees et al. atoporella nicholsoni. Kauai (= 5. typically near the outer mesohyl (Figure 5–18a). 2010.g. 2006. Likewise. the basal region may also con.Dade. Antipatharians Black corals are sometimes common on coral reefs.g. especially during the winter months. 1997. or it may be partly absent as the result of grazing by a variety of animals with scraping mouthparts. cording to the island’s geological history and its biological response to local and regional conditions. and a few are capable of swimming. a b c d e Reef Sponges•91 cyanobacteria are especially well represented. However. Bank. New York: Springer. b) Leeward side. and likely involves the morphology of the collar gland. regulation of nutrient transfer. 2008). 1989. two different genera and several species may do so.Publication Data Goldberg. Composite of STS093. ration. this compound smoothes the spines and produces a shiny exhibit an extremely large range of spine morphologies and appear to form several distinct clades according to rDNA analysis (e. 352 Synergies and Loss of Resilience. holes. especially during the Paleozoic. the Symplasma. Florida International University * deceased. are scattered throughout the cytoplasm and are typically prokaryotic in both size and structure. particularly from the capture of zooplankton. Smithsonian Institution US Na. and their ratio to fleshy macroalgal abundance. compact branches Hemispherical Figure 1–14: a) Shallow reef zones on a fringing reef (redrawn from US Geological Survey: http:// pubs. Extra. crystal. the heterotrophic bacteria pro. angular symbiosis between the host and its autotrophic and heterotrophic symbionts may serve to increase nu. In contrast. mercially valuable. Journal of Biological Chemistry 283: 25475– 25484. showing the red skeleton where the polyps have contracted. coral.like aggre.g. 2003). neous” test composed of tiny needles of high. order Chroococcales. guished seasonally or annually in the form of distinct layers or bands that reflect a regular alternation in car. loading. Reviews in Min. Goldberg 2002b). From the host perspective. Popp BN. lectively. Several cell types are found here includ. branching corals. the na. and crystal growth. isms. namely S. fication of the cellular site of polychlorinated peptide biosynthesis in the marine sponge Dysidea (Lamel. the outer. 2002. but are especially prominent at depths between 200 and 1.g. and exhibit a syncytial structure. Such organisms are referred to as browsers if they selectively nip off specific bits. Indeed. reefs with high rates of coral mortality often cor. gress Series 175: 285–288. Darwinian reefs. width. netic variants that could represent tens to hundreds of species. desmium form extensive mats of pink to red. Stromatolites still exist today.containing ring compounds typically derived from amino acids) are common in sponges and are distasteful to many potential preda. biosis? Marine Ecology 5: 9–27. Krautwig DWH.phylogenetic and paleontological per.term effects. Initially. dino. a common component of nematocyst toxin is the same material found in bee and reptile venom (Nevalainen et al. microph. Broadcasters with larger eggs tend to produce larvae with a yolky endoderm and thus have been referred to as leci. an island NW of Bora Bora. producing two individuals or sometimes multicrowned individuals that then separate (Hand and Uhlinger 1995). However.. Fiore CL and Jutte PC. in this case. often infiltrated with acidic polysaccha. tondo 1983). Calcite deposition begins at the boundary between the meristem cell walls. Hexacorallia) based on rDNA internal transcribed spacers sequences [sic]. Tambutté E. Compositional heterogeneity and mi. Holo. Macintyre IG. dued but still hydrodynamically active shallow. and violet light is prevalent. barrier reefs. Coral chronometers: seasonal growth bands in reef corals. pendix II of CITES. Diurnal and sea. Schläppy M. Anais da Academia Brasiliera de Ciências 80: 495–513. Nitrogen fixa.. Paul VJ. further described below) form bundles of filaments that produce local anaerobic zones where nitrogenase becomes active. tose the mineral onto the cell surface (De Nooijer et al. the Carolines.” ATPase calcium: proton exchange Calicodermis Oganic matrix Aragonite skeleton CO 2 CO 2 HCO 3– HCO 3– + H+ HCO 3– HCO 3– Ca2+ Ca2+ Ca2+ Ca2+ Ca2+ Ca2+ + CO 32– H 2O + CO 2 Zooxanthellae Seawater in coelenteron Mesoglea Gastrodermis CaCO 3+ H+ H+ H+ H+ H+ Calicodermis Coelenteron P r o t o n c o n c e n t r a t io n g r a d ie n t Calcium concentration gradient Higher Higher Lower Lower H+ 1 2 3 Carbonic anhydrase Equilibrium shift Equilibrium shift Active transport Active transport Protons to coelenteron Protons to coelenteron Active transport Active transport Equilibrium shift Active transport Protons to coelenteron Active transport Figure 6–17: Control of protons and calcium during photosynthesis and calcification: The gastrodermis lines the coelenteron. Kruip J. National Institute of Water and Atmospheric Research. Iwao K.. genetic analyses have not been in agreement Reef Corals and Their Allies•113 tain antipatharian species grow to large sizes and occur in densities of more than one colony per square meter. tions (e. 64•Chapter Four lack a flagellum and cannot swim. and light enhancement of calcifi. Despite its depth. fied species.term records like the Hawaii. corresponds to the temperature at which the skeleton was formed (Goodkin et al. mensional model. tions (described below). Kingsley RJ. Tanner JE. 1997. Both have extensive rhizoid systems that ramify through the substrate and anchor the thallus to the sediment as shown in Udotea sp. Journal of Experi. Micro. western Equatorial Atlantic. I thank Jennifer Myer for the line drawings in chapter 4. tribute to the nutritional well. Indeed. low. there are examples of isomorphic phaeophytes (e. Others are dissected with channels as shown in the boxed area of Fakarava Atoll expanded as c) where there is a transition from relatively large windward cays to small leeward islands. Rezak R. ppt). ceedings of the 8th International Coral Reef Sympo. all of this is exacerbated when combined with decreases in herbivore populations. bility.” which appear to be unique to certain sponges even though they have yet to be cultured and their function is not presently known (Fiesler et al. In Hawaii where the harvest of black coral is regu. Chlorophyta). Samoan P. 2007. Mangeveldt et al. Journal of Sedimentary Research 60: 992–1007. reproduction within the common Indo. 2007). Modern and Ancient Coralline Sponges Coral. crustacea (order Nostocales). Sea level changes are controlled not just by glacial melting. 248 m depth. Proceedings of the 8th Coral Reef Symposium. However. some in the Bahamas). 1995.100 species (S. mentology. 2011). and may be a means of coping with increases in cellular CO 2 and the result. ing in waters >300 m deep. gesting that it too has a vital role in calcification (Al. Tentacles may be limited to the mar. and sea urchins are removed or reduced. Polar Biology 30: 487–492. but each island originates from a single persistent “hotspot” in the Pacific Plate. or even causing the atoll to lose its shape entirely. etc. 2005. yellow. New York: Springer Science and Business Media: 1–8. 2009. Three. ceeded him. there is no equivalent of the Darwin Point for the Society Islands. In: Vacher HL and Quinn T. Cold periods caused formation of ice sheets and the thickening of polar ice caps. a thick. in most cases broadcast.dwelling brittlestars. Courtesy of Image Science and Analysis Laboratory. ter Craig. eral atolls of the Kiribati. Asexual reproduction by transverse fission and some anomalies in the sea anemone Nematostellla vectensis. soglea that interconnects the polyps. cavating caverns up to 12 cm deep within coral colonies (Glynn 1997). ing forms. phytes produce chlorophylls a and b as well as carotenoid pigments. Hickson. they also typically produce a macromolecu. In addition. yet maintain a com. or by competition with other organisms that have a similar taste for sponge tissue. 2002. Type 3 matrices inhibit crystal for. agous feeding refers to the consumption of any form of microscopic prey or particulate food. tropical waters these peaks may become capped with biogenic carbonate and may form the foundation for coral reefs. and by the introduction of a double bond in boxed area b. This listing requires documentation of a 20%–30% decline from an historic baseline. which may be especially important in branched forms that can interlock and resist frequent movement. Raz et al. The physiology of skeleton formation in corals. ysis Laboratory. and orientation. Typically. geologically speaking. Greece). However. nent bonds can be established. Other organisms that require solid substrata. Some species can also produce gigantic siliceous fibers reaching lengths of a meter or more.. Scleractinia and Coral.building. Some sa. This horizon. making them less desir. 2004. 2004. Substratum prefer. when a core was drilled to a depth of 340 meters on Funafuti Atoll (south. the proportion of foram sediment may rise to over 80% of the sand. They also recycle nutrients. tion by organisms is a matrix. Kennedy IR.g.Darwinian concept that ex.3 m long and is common on reefs throughout the tropical Pacific. thetic to structural tissue. Figure 7–7: a) Lateral view of Sabellaria alveolata: ot. Amphinomids may be scaven. Hexactinellida. mones or a combination of signaling molecules and the direct mechanical influence of a single cell on its neigh. nance. fornia Current and northward flow of the Peru Current.step mode of growth in the scleractinian coral skeletons from the micrometre to the overall scale. clude calcareous red algae that precipitate mineral di. This section of the body lacks the well. marine forms often epilithic. The entire range of light used by any oxygenic photosynthetic organism includes all visible light. eds.Victoria and Zea 2005). some have questioned the inclusion of prochlorophytes in the cya. Marine Biology 68: 3321–332. Other polychaetes are a bit more discerning in their feeding and sort the digestibles from the inorganic matter by tentacles and cilia before they reach the mouth. one Hawaiian species appears to be confined to a total area of about 10 km2 where it is the dominant benthic invertebrate (Cairns 2011). although many will invade only carbonate that is not pro. Algal symbiosis and the collapse and recovery of reef communities: Lazarus corals across the K. a tissue now recognizable by its flagellated gland cells. While hexamerous symmetry is the rule. it has a high percentage of coral cover and has been designated a National Marine Sanctuary. 2006. Nonethe. Kinsey 1985). 2006. Grange 1985). 1985. tures. perhaps as a defense (Pennings et al. In: Dubinsky Z. ed. ducers in dead carbonate substrates in Hawaiian coral reefs. new genus. Because the windward reef crest is well developed. a condition of the entire organism that makes them unique among metazoans. tabolites. ing within it. including coralline algae and millepore corals among others. Toxicon 44: 45–51. some hosts that are known to house certain adult clownfishes can kill them as juveniles.13: 978. this species is considered an indicator of Holocene sea levels in the western Atlantic (Lighty et al. Nüchter T.5–0. Martin JM.sized particles (Davies and Hutchings 1983). 2007. eral meters in height and may weigh several tons. et al. the radioles are often referred to collectively as the branchial crown. 2010. a two. and significant vol. phyra. which are used only once. pyramidal. bilaterally symmetrical animals (Seipel and Schmid 2006. With the exception of the very end.armored structures occur on the deep windward reefs of Jamaica. 128•Chapter Six between 300 and 400 meters off the coast of western Norway (Freiwald et al. The sponge surface displayed a star.structure length scales in coral skeleton. Most of these occur as large and small three. in part because they resemble a common growth pattern found in inorganically produced crystal systems that can be precipitated from supersaturated solutions of calcium and carbonate. nobacteria that lack heterocysts. gal forms of Dictyota have become particularly abundant on the reefs in Florida. calcification in crustose coralline algae occurs through the orderly formation of high. Journal of Experimental Marine Biology and Ecology 349: 295–298. Biological Bulletin 196: 70–79. Decrease in Lithothamnion sp. The resulting radiographs show couplets of light (low. Fewer eggs are produced (1–6 per polyp. Most species are held upright by an axial skeleton composed of “gorgonin. 2000.. decalcified coral skeleton taken from the Great Barrier Reef. leaving visible scars on the surface of the substratum. Proceedings of the National Academy of Sciences. and thus provide calories with. There are several orders of these siphonous Figure 4–4: The calcareous green alga Halimeda may be attached by a holdfast to rocky surfaces and form an erect or pendant epilithic thallus.fertilize (i. pletely (Figure 6–30c). Fringing reefs are the most common of the three. Goldstein ST. New NOAA maps show big island has most live coral of main Hawaiian Islands. including certain algae. logica Polonica 48: 497–530. the GBR ex. requiring a large number of protons. Evi. The head often bears a number of eyes. and car. In the calcifying family Corallinaceae. ida Reef Tract. Indo. Wörheide G and Hooper JNA. The sexes of most polychaetes are separate. The high abundance of Halimeda carbonate in tropi. Coyer JA. 1991. The use of the term guild in coral reef ecology and paleoecology: a critical evaluation. Sur. pear to be simply structured. For example. and Titlyanova TV. with a highly organized pattern of 2–3 layers derived primarily from an actively dividing meristem. Calcareous surface similar to onyx. and madreporarian corals procured during the voyage of the HMS Challenger in the years 1873– 1876. Acropora acuminata. There are even “bidirectional” coral hermaphrodites that can change from one sex to the other and then back again (Loya and Sakai 2008). Vermeij MA and Sandin SA. and ecological roles. gastropods. Hine AC and Steinmetz. temporarily causing the adhesion of loose calcareous material to the reef frame. Indeed. downhill. but the volume of the spongeocoel will increase disproportionately. sponse to local and regional conditions. along with respiratory CO 2. alization occurs as the result of interactions between an or. Bretagne Nord. Because many species produce multiple branches. Other notable but smaller barrier reefs occur around Palau (western Caroline Islands). and the Maldives. Community structure of living fossil coralline sponge populations at the 100•Chapter Five Todd C. and Obura DO. among other groups. Framework Large corals Fused oysters Secondary frame producers Sedimentary fll Calcifying algae (Halimeda) Sea urchin spines Mollusc shell fragments Cementing agents Crustose coralline algae Encrusting Millepora Diagenetic cement Binding agents Zoanthids Sponges Bafers Sea fan a b c An Introduction to the Structure and Formation of Modern Reefs •3 within the reef in high.and deeper. but have been uplifted. but also transform and sequester their toxins. and will be an area of focus within each of the chapters that follow. sity to self. 2003). or islands composed of cemented sand dunes rising to 30 m or more (e. 2006). 1997). Berkeley: University of California Press: 515–542. vironments. This change is recorded by growth bands within the axial skeleton (e. Calcitic spicules are for the most part formed in the extracellular environment by multiple cells (Uriz et al. Fora. 2002). This in turn suggests that large col. and those that produce a tough. Cultivation. 2004. Not surprisingly. 1991. Boero et al. Although they lack jaws. 1983). and geographic areas. but few of them present long. Crustose corallines. and Horta PA. the Capricorn. ervation of these sediments in the fossil record. the same technologies are providing a window into their critical role in structur. and in many of these the colony is supported by another internally lo. The genus Phragmatopoma also con. However.like partitions called dissepiments within and be. ter. for example. and Management. rubrum are common on Bermuda’s algal cup reefs (Ginsburg and Schroeder 1973) and the reefs of Belize (Gischler et al. However. Although there are several roles that are played by zooxanthellae.Cheung W and Williams GA. P. Interestingly. Mortensen PB. and detritus) in a mucus ball (Zale and Merrifield 1989. lodysidea) herbacea and symbiotic cyanobacterium Oscillatoria spongelieae by CARD. which are composed of archaeocytes that are enclosed in a protective layer. thotrophic. ous cloud (e. 22: 113–152. and may be found both within and between cells. Chemical Geology 172: 307–329. Reproduction of Cnidaria. Hart T.related alterations to coral reef com. plasm protrudes. Barucca M. and Assmann M. ers of aragonite bundles serve to reinforce the thallus. and may differ in prominence among populations of the same species growing in differ. some brittlestars are thought to prefer certain sponges even if they lack chemical defenses (Hen. these rapidly growing cells add carbonate to the vertical and lateral growth of the crust. Mucilagenoussheath Peptidoglycancellwall Lipopolysaccharide membrane Thylakoidmembrane Chromosome Cyanophycin granule Lipid droplet Polyhedralbody Ribosomes Starchgrains 40•Chapter Three higher. in contrast. In: Van Soest RWM. Blanchon et al. ———. eds. cal functions. Guzman HM. and Cotten J. prevent. Marine Biology 147: 775–788. swim. and Scheitel U.reef environment off Ish. An overview of the shallow. the tube is not Radiole Mouth Ventral sac Collar New edge of tube Tube Pinnules Pinnule cross section Water fow Radiole Food groove Pinnule a b Figure 7–6: a) The crown of large sabellid polychaetes is typically composed of ciliated arms called radioles that are often (but not always) arranged in fan. However. some specific zones produced nearly six times that amount. 1975). cervicornis growth. detritus. fungi. 2009. Little is known concerning the cellular aspects of calcification in crustose coralline algae. although their recovery and growth is strongly dependent on fragment size and physical conditions in the environ. Barnes and Chalker 1990). come a major source of bioerosion.like growths formed from the active accre. Invertebrate Biology 126: 99–112. Lewis JC and Von Wallis E. sentially closed. dense serpulid reefs cover areas Figure 7–5: Biogenic reef structure. and is one of the largest hosts with an oral disc that can reach more than a meter wide. their influence in calcifica. tocorals produce individual calcitic elements called scler. 2004. the chaetiger develops into a juvenile (Figure 7–10). their presence is associated with in. as described in Chapter 15.Medrano 2006). In other cases. However. but likely involves the unusual geometry of their volcanic basements. Storage and remobilization of sus. Bilan MI and Usov AI. courtesy of Elsevier. The reef en. Dor.like oscula.building species in European waters (see Dubois et al. Fay P. The large mounds shown in Figure 7–7c are constituted by Sabellaria alveolata. male’s concave surface and then onto the reef (Lindquist and Hay 1996 and photo in Ritson.. ———. “boom and bust” cycles often occur in which discovery is fol. Revised. Gischler (2010) has suggested that such structures on Atlantic reefs tend to be domi. or halogens that act as hardening agents. There are also more recent examples. Hand. et al. pied by corals. 296 Spawning Mode.quality food resource for intertidal grazers. Rosenheim BE. sponges that differ in chemistry (as well as fiber and skeletal density) are often packed together on reef substrata. and indeed there are penetrat.400 km long. The role of symbiotic algae in car. and is analogous to tree ring analysis (dendrochronology). This diagenetic cement is a particular example of a more general process called diagenesis in which physi. Schöttner SI. Pacific.water Caribbean stylaster. Journal of Experimental Marine Biology and Ecology 125: 145–169. Collins LS. but in addition to ordinary chitinous chaetae. local cur. lar to those that likely contributed to the evolution of the chloroplast. This portion supports lat.like blades. collectively referred to as ecological services. the time required for complete exchange with the surrounding ocean. the whips (center and foreground) are in the family Plexauridae. Indeed.000. Other sets of partitions called secondary mesenter. but experimental work suggests that their adhesiveness is limited and may reduce the utility of these cnidae to the capture of small prey (Thoring. Bioscience 57: 573–583.water species. ther algal growth at the expense of CCA (Belliveau and Paul 2002. Community struc. Even so. eds. Lindquist N. In addition. Acropora millepora. United States National Museum of National History.like secretions from their invertebrate or vertebrate hosts (Martin and Britayev 1998). in New Caledonia. In addition. Kikuchi PRK and Leão ZMAN. 1996. b). In some lagoons. long.initiated.. are unaffected by mixed. among other factors. Vol. ure 5–9b). and Allemand D. Epilithic microborers confine their tunnels and their biomass to the surface.Pacific species are broadcasters (Richmond and Hunter 1990). M. Likewise. and Ripley H. thus eventually weakening and disintegrating the substratum. tion and efficient cycling of host waste products (Chapter 6). Baco A. and temporal characteristics that enable them to com.ATPase. Special credit is also due to Ron Blakey of Colorado Plateau Geosystems for allowing the use of his excellent paleomaps. including four of the named banks located off the coast of Nicaragua in the Caribbean (Appendix 2c). 2001. ISBN 978. choric or hermaphroditic males. ullary cells may have a number of functions in different groups.5199. Most sipunculans are soft sediment burrowers al. 2002. fended. lices dissociate from the colony. 2008. Some clownfishes appear to have an in. or bacteria. alkaline waters. zka 1987. some of which were 10 cm long. as noted above. These eunicids thus play a role in the development of deep. and green species (Figure 4–2). 2001. 2009).000 or more named species (Norton et al. Coral Reefs. 2003. plasty.800 areas displaying spur and groove topography showed that deep spur. Flatt et al. 33 A Little Crystallography. He recognized three principal types of coral reefs on tropical islands and continental coastlines: fringing reefs. including cal. ed. 2005) and may exhibit modest overall vertical rates of growth (e. rolled into a ball. 2005. Romanovicz DK. Proceedings of the Royal So. In addition. Vago R.2–10 cm long.org /2236. 2001. Lambeck K and Chappell J. Thilad. 1992.g. Growth. which eventually fills in the entire cell wall (Borowitzka 1982. it is useful to define the nature of biogenic reefs broadly. Golubic et al. Ross SW. suggesting they were an ancient lineage of coralline algae. This suggests that the symbiotic relationship is autotrophic (i. and Henstock TJ. 1991). Marine and Freshwater Research 48: 589–599. Asexual reproduction does not produce clonal populations Reef Corals and Their Allies•143 Walker TA and Bull GD. rea are also important habitat. The tubes are typically protective. sistence. belong to ancient lineages that produce siliceous spicules along with a massively calcified aragonitic or calcitic skeleton. Schneider J and Le Campion. tist 78: 224–235. a b Figure 4–17: One model linking calcification to photosynthesis in coralline red algae involves active intracellular transport of seawater calcium and subsequent delivery to the calcification site by an ATP. Examples from the western Pacific include wave. In contrast. Their Biology. cient multicellular animals. eggs arise from single choanocytes or archaeocytes. dophyta. courtesy of Elsevier. respiration and energetics of three tropical marine sponges. organ. the rest are brooders. van Oppen MJH. The pharynx also bears a series of hexamerously po.ATPase may actively trans. eds. 2006. but little is known about the specific boring mechanisms in these animals (Rice and Macintyre 1982).g. This movement is also accompanied by subsidence. ate. Johnson Space Center. it appears that the production of amorphous mineral more often serves as a precursor for later deposition of crystalline material. b Figure 3–11: Large stromatolites exposed in Glacier National Park. lites in the Great Slave Lake region of Canada (Northwest Territories) are 1. Journal of Experimental Biology 211: 2827–2831. et al. the skel. cology 26: 6–12. Fragment generation. Although these layers are sometimes closely associated with the living tissue of their hosts. in some 11% of these.. For example. Such populations contribute significantly to bioerosion (e. However. Mechanical design of spicule. The first of these. courtesy of Pål Mortensen. fragmentation. fied anthozoans that are found along rocky coastlines. pearance of the shell.like. b. Dupouy C.” less well. they do not compete well with longer. 66•Chapter Four the details are unclear. Zoccola D. Wood R. Toothpick. or mounds composed of mud or piles of skeletal debris. Because Halimeda can develop 1–2 segments per day and double the bio. Epilithic species.temperature occurrence. come a polyhedral solid with a regularly repeating. Parts a– c show extratentacular budding patterns. 1988. 2004. 2010. Most tubicolous species remain within the tube for life and are sessile.edge Halimeda meadow. 4–2).000 years ago. 90% of which is the lagoon (Figure 1–12). is a branched. abrade. and Chadwick NE. Burrowers including groups of boring bivalves. For ex.S38. and type of crystal formed. a tri. Veron JEN. Climate change and coral reefs: differ. uncalcified algae on the other. but could not continue sustained growth. especially in relatively deep water. roalgae: a physiological perspective. including those in the genera Palola or Eunice. The simplest is a buildup. some groups may become more successful in different reef environments or at different times as described below. Chisholm JRM and Moulin P. Sabel. Current Developments in Bioerosion. especially where turbulent and turbid waters prevail. and that in.g. Col. In one large loggerhead sponge. Science 560– 563. roots. Bald D. ification in coral reefs: some facts and misconceptions.000 m. and even onto their dead skeletons. Rapid sea. Tazoli S. tive 22: 397–414. which control the closing of the shell. t. igaki Island. Ni. 25. up to three times that amount may be dissolved (Zundelevich et al. 2003. Uriz MJ. the polyp then splits longitudinally along the oral. and physiology. Ostreobium spp. Oxford: Black. While not all sabel. in addition to several groups of Archaea (Burns et al. the primary photosynthetic producers for Earth’s ecosystems. Ortega. most Capricorn. Such intermediate cases are not uncommon and are further described below. First. Based on Smith 1955. doi: 10. Bioerosion Bioerosion is caused by the solution and/or excava. Modern members of this group are diverse. flamentous cyanobacterium.and cold.shaped gut or create fecal pellets that are conducted out of the tube through a ciliated. despite their tolerance for these conditions. Thus. point of operational cost and profit. Moreover. the combination of the two may be. and the Aleutian Islands of Alaska. Darwin suggested that fringing reefs on islands were the youngest of all reef types (see below). Science 219: 410. It is unknown if these lower harvest amounts are sustainable from a conservation perspective. tetids. ample. Sydney. However. the most abundant. Such “rhodolith beds” may be one of the Earth’s largest marine communities structured by macrophytes. Lord Howe Island (SE Australia) is in the path of the warm. Most sphinctozoan fos. Figure 4–18: a) Algal ridge from Rarotonga. Coral development: from classical embryology to mo. 2005. Saba Bank in the Caribbean. Palaios 7: 96–107. the actual release Figure 7–10: Larval development of the Sabellariid Phragmato. als. and Loya Y. including multicellular and unicellular forms. and Pomponi SA. 267 Reefs of the Paleozoic Era. 1966. is estimated to cover from 25% to 50% of the available space on Jamaican reefs >50 m deep (Figure 5–14. Demandis KD. and the amount of ambient light. Like serpulids. More often.g. tipathin. the maintenance of turf algae as a dominant form on shallow.” the structure and function of which is discussed further in Chapter 8. The most obvious structural carbonate is the algal ridge. able only microscopically. poridae). or because their de. ferentiated as an abdomen. a sequence that occurs in chiton teeth. Coral Reefs 22: 229–240. Spicules range in size from relatively large and visible. This sequence of events constitutes a form of selective deposit feeding. Visscher PT. These distinctions may also be true of the gametes. Holocene lagoonal development in the isolated carbonate platforms off Belize. However. Daly et al. gregarious or colonial. Cay Sal Bank. Biomineralization: Principles and Con. Huston 1985). Figure 6–6: Branch of Stylaster roseus from the Bahamas showing the arrangement of gastropores (G) and surrounding dactylopores in the form of cyclosystems along the branches. Proceedings of the 9th International Coral Reef Symposium. although different taxa of boring sponges may exhibit different patterns of tunnel and cavity excavation (e.walled aragonite chambers that form con. Canada. bonate do so in the form of calcite. ing the mouth and tentacles. shallow. Allwood et al. lined). or encrusting colo. ford: Oxford University Press. The toxins may be used offensively to obtain prey. Life and Death of Coral Reefs. during the same or different seasons. Within the latter region. 1998.6 µm wide and 10–15 µm long (Figure 4–8 a.” a composite material contain. ogy 11: 97–144. era Sinularia and Sarcophyton on mid. tical. dences for a biochemically driven. E. chains of daughter cells. isms. Goldstein 1999). Endolithic fungi in marine ecosystems. Coral Reefs 20: 399–408. Al. Symbiosis in foramin. Brooding and Other Forms of Larval Formation In contrast to broadcast. deep blue. and others) encounter fre. Little is known about such regulatory mechanisms. http://www. 1996. Allemand D. lecular evidence for multiple lineages in the gorgonian family Plexauridae (Anthozoa: Octocorallia). erosion (i. Ritson. redrawn from Oceanexplorer.noaa. hence the common name “featherduster worms. In contrast.forming species (Figure 6–27c). tinian corals. oir: 13: 30–37. ites are the dark structures between the polyps and around the axial skeleton. spat survivorship.and nano. but it and molecules like it are found in many marine organisms. 2002a. The middle lamella (ml) is typically seen as a thin layer of crystals when viewed between cells (arrow). Examples of how reefs were able or Figure 1–21: a) The tectonically active island of New Guinea is 150 km north of Australia and is divided into two jurisdictions.shell proteins. and dominance by eusocial species among sponge. and on reef. The evolutionary diversification of cyanobacte. gas (Florida). drilla australensis from Western Australia (5 m depth). Ulstrup et al. ribbean angelfish. The coelenteron develops a relatively low pH. is branched and attached to the thy. ———. a flexible. brown algae (Santelices 2002). Catmull J. and Sakai K.g. In a few genera. New York: Oxford University Press: 445–470. Likewise. University of Miami. In : Mc. and van Etten J. come infused with sediments (Birrell et al. 2007). Bahamas: an incipiently drowned Ho. Rasser and Piller 1997). Water transport. or are otherwise biomedically useful for their anti. Fertile segments called gametangia are formed overnight and broadcast spawn clouds of macro.” patterns of richness and ende. Conversely. or those that form such heterocysts but do so in unbranched filaments (order Nostocales) or in V. cyanobacterial colo. and Larkum A.. they can also be quite complex anatomically. though some shrimp used as many as six (MacDonald et al. 1980. phology and hydrology of selected islands of French Polynesia: Tikehau (atoll) and Tahiti (barrier reef). Syllidae) associated with two species of sponges from East Africa (Tanzania. perimental Biology 98: 239–287. trient transfer. The period from fertiliza.7 µm in Reef Cyanobacteria•43 duced. Spirocysts are typically cat.G65 2013 578. Macintyre IG and Glynn PW. Marine Biology 157: 593–602. However. The role of coralline algae in the diets of selected herbivores. Indeed. United States National Museum. since the 1990s rapidly growing macroal. rent and frame reefs do not develop there. ity in the form of jewelry. They occur most commonly in shallow. 1990. 1988. and shelter for the more mobile associates.. curred on ridges behind the old barrier where they grew “keep up reefs” (Figure 1–23a). 5 The Evolution of Modern Reefs. IvoryAndArt. ular mechanisms to environmental control. Others form branched but unjointed thalli. aments (siphons). Cat Island platform. (Rho. with two micrograms being sufficient to cause death in humans. most no time to occupy one host species. Reef Corals and Their Allies 6 Reef Corals and Their Allies•103 by developing stages that are quite abundant just be. especially ectoderm vs. Like. and high levels of wave. tegrative and Comparative Biology 46: 398–406. ineffective venoms. their morphology is dis.like tests and an obligate relation. b). rescent skeleton with characteristic thorny branches.WCMC Biodiversity Se. and nematocyst or zooxanthellae. cyanobacteria) that stimulate or repress the growth of another. depending on coral growth and the topography of the coast. although that alone does not explain the characteristic jaw marks that can be distinguished on the sinuous tunnels they create in rock composed of the same or harder material (Glasby et al. biomass primarily or completely within the substratum Macrobioeroders a. such larviparous development is more common among hermaphroditic species. tube in 4–7 places by a complex of cement proteins. ing the nodes and occupying the central region of each segment. Sedi. Their distinctive calcareous munity appears to be a significant contributor to carbon fixation through photosynthesis. Proceedings of the Biological Society of Washington. Santelices B. Just as in kelp forests. and are generally associated Figure 6–20: a) Sclerites on colonial branch sur. spongy cortex (Figure 4–7a) that is easily distinguished by its widely branched cortical filaments. USA 103: 9096– 9100. sponge larvae are said to be yolk fed or lecithotrophic. stricting organisms living there to only the most tolerant.shaped colonies up to 2 m across. Radtke G. the evidence for nucleation and crystal formation by the matrix is unclear owing to its spatial separation from the site of mineral formation (e. Nature 332: 260–262. hibition of coral recruitment by macroalgae and cyanobacteria. The predominant carbonate in the Corallinaceae is magnesium calcite. an amount that more than doubled between 1999 and 2005. and sediments: complex interac. 6–31c.axis is always oriented perpendicular to the cell wall. Perhaps the best. planula formation. ed. which extends >3000 km farther northwest and represents the oldest and most heavily eroded of these extinct volcanoes (Figure 1–17c). and variable chlorophyll fluorescence analysis of endo. P. Lagoon reefs are complex and may include various types of patch reefs. tex that surrounds a central chord composed of a series of dome. ined.. Thacker RW. about 6 Mya (Martin et al. However.Verlag: 25–31.. 1973. low the surface in shallow water (Figure 4–3). Cnidarian internal stinging mechanism. This arrangement is not unlike the intercortical spaces of Halimeda. Duffy JE. poly. Millepore tissue is typically a golden brown color ow. About 95% of all living sponge species are placed in the cellularian class Demospongiae (Paw. Wood 1998). Note that a metric ton (a tonne) is Reefs and reef. cal one. but not from others.sized particles (Appendix 1). A generic revision and phylogenetic anal. but also by thermal expansion of water. may occur higher up on the buttress wall. rial autotrophs were once classified as “blue. 2005 and references therein). 2009). certain types of calcareous algae. as well as in the population density and types of bioeroding organisms they harbor. is bark.aoml. Wiebe WJ. These characteristics are also relevant to zooxanthellae found in other hosts. karyotes and possess membrane. and organisms that settle and grow on sponge surfaces. Barrie JV. a b c Annelids and Sipunculans•151 Sabellariid reefs are not composed of colonies. ent populations. Macdonald GA. ductive cycle and correspondingly short lifetimes. duce a protective envelope called a sheath (Figure 3–1) Reef Cyanobacteria 3 Figure 3–1: Light microscopic view showing Microcoleus ch. mentology 44: 441–456. pete in this environment due to the considerable wave energy. nes and is the most widely distributed. Metabolites from symbiotic bacteria. Muller. edly remain to be discovered.like appearance. Clearance rates and aquiferous systems in two sponges with contrast. Even though Prochlorococcus species are only 0.derived vesicles inside the cell. 1999.g. a lysosome. 2001. species of Pocillopora become major reef builders in isolated regions such as the eastern Pacific (Glynn and Ault 2000). The former may include volcanic peaks with carbonate margins (e. Towns.forming stages allows the alga to reproduce repeatedly prior to gamete production. Biological control of calcification in reef corals appears to involve two basic components: (a) the function of a specialized calicoblastic tissue and (b) photosynthetic contributions of symbiotic zoo. can allow the formation of smaller individuals that continue their growth away from the parental sponge. They may feed using a muscular pharynx with or without teeth to punch a hole in their prey. and inner nacreous layer of aragonite. sug. but are never associated with zooxanthellae (Stanley and Cairns 1988). 2000. and indeed they may be well represented. Some species may in. ficient and common burrowers that use their protrusible. Ironically. providing nitrogen that is taken up by the zooxanthellae during photosynthesis. egorized separately from nematocysts. This depth was too great and the leeward side lacked the vigorous circulation required for Acropora palmata to produce a framework. Funafuti Atoll. Models suggest that carbon compounds are utilized differentially depending on their source and complexity. however. Trichodesmium A mat-forming cyanobacterium often with unbranched. In addition. an ex. Furla P. d carotenes phycoerythrin phycocyanin floridean starch Reef Algae and Foraminiferans•55 algae. whereas the net carbonate is the amount remaining after accounting for all of the losses re. Adey WH.Pacific. gore. and cause depletion of energy reserves (e. green. especially in the intertidal zone where they may gain protection from predators and desiccation (Pérez et al. millions. As an unbranched coral grows. a brown alga. This could be the result of modern conditions. An Introduction to the Structure and Formation of Modern Reefs •25 erosional sculpting from rain during multiple low stands of sea level. the matrix does not possess specific nucleation sites. How. ern Australia (Collins et al. synthetic capacity of photosystem II in symbiotic dinoflagellates in the Caribbean reef. 2004. are mine and mine alone.feeding currents are more directly channeled directly toward the mouth. However. even. In: Stanley GH Jr.borings created by a unicellular cyanobac.Kerby A. 1982.cz or www . Coral Reefs 9: 231–237. is an indicator of the health and growth of a reef system. ysis of clonal reproduction in a soft coral. 2005. and 32 different classes of crystal symmetry in total. The nature and origin of coral reefs islands. These form new growth and give rise to specialized photo. Comparison of developmental trajectories in the starlet anemone Nematostella vectensis: embryogen. Sponges belonging to this informal group formed small vertical and radial tubules that extended deep into the cortex. along the coast. 2009. 1976. turity after about two years. Apart from the Anthozoa. coral sclerite ultrastructures and experimental ap. Part E. as depicted in many marine biology textbooks. Bristol: Biopress Limited: 111–151. reviewed by Fabricius and Alderslade 2001). and that large colonies may be 2. (b) the history of sea level change around that platform. mary producers (Tribollet et al. These and other long.Pacific (Adey 2005). ment process is distinguished as skeletal accretion. produce a nutrient. Grotolli et al. Figure. and choanoderm of a simple sponge. lation levels have been associated with elevated concen. sition of the plankton in waters they process. Reef Algae and Foraminiferans•63 ments in turn form rhizoids that attach to sand or en.water coral Lophelia pertusa. synthesis). others have found that grazing is the pri. versity of Miami. British Journal of Phycology 21: 217–224.Pacific reefs. 2003. shading this spe. 2008. and Kempf SC. heterotrophic interactions. the older Emperor Seamount chain. Structure and composition of organic reefs and carbonate mud mounds: concepts and cat.third of reef. Many sponges are “living hotels. calci. some members of the genus Madracis may be zooxanthellate or not. some. Florida International University Gautam Sen. and Pawlik JR. An outer (deep) fore reef may be distinguished by species composition and growth form below depths of 15–20 m. Marine biology. Brazil): an atoll built pri. International Review of Hydrobiology 93: 328–341. especially those with broad environmental tolerance compared with many reef organisms. Ecological Monographs 71: 49–71. cal corals may be more common (Geister 1977. ried by strong tidal currents. Rios RE. and Steinen RP. localization. availability of nutrients for the symbionts. Compari. shrimps. ter. and Faulkner DJ.13. 2003. the outer portions of the most heavily calcified segments become increasingly micritized (Chapter 3). D: retracted polyp (dactylo. Calcaxonians include gorgonian families with al.” The wall may persist to depths of >50 m here. The latter group can be functionally male and female at the same time. sive layers (lamellae) grew in the form of vertical. Marine Ecology Pro. While sediment production appears to be a very im. sure to rain that occur here. Penn.3) approximately 89% of the dissolved inor. Proceedings of the National Acad. or as symbionts in several groups of organisms. 104•Chapter Six skeletal surface of millepore corals is perforated with “a thousand pores. 2001). Nature 406: 989–992. 1996). and are surrounded by other specialized “nurse cells” that may contain yolk or other reserve nutrients. Other species with zooxanthellae are always relatively small in shallow water.000 years (Pan. calcification. The algal layer is 5–10 mm below the coral surface. However. there are 24 distinct and ornate. Ma. the absorption of light by pigments found in cyanobacteria shows a potentially broad spectrum of light utilization. mercially. but is flushed by polyp activity and buffered by seawater. Advances in Marine Biology 17: 1–327. Growth rates are not known with certainty for most species. tion 10: 149–154. Fringing reefs off West. the host. position. In its simplest configu. Piel 2004. tology. biological processes. gen as an end product (chloroxygenic photosynthesis). ponents of the digestive process. Perspectives in Plant Ecology. 42•Chapter Three diameter. resistant structures in shallow water. called nitrification. Ghiold J. and Beukens R. 1995). Relict reef system off St. Nor does it explain why some areas (e. is a common “palolo worm” in the Atlantic where it occurs in reef carbonates. Hexactinel. Uwins P. era. Golubic S. 12:1–11. Given high photosynthetic rates by some sponge cyanobacteria. 278 Reefs of the Cenozoic Era. sis. or burrowing. This feature.absorbing complex attached to the thylakoid membrane in cyanobacte. Less commonly. try and structure of gorgonian and antipatharian coral skeletons. specialized cells called akinetes (e. 1978. Figure 4–17). since sandy areas may account for a large proportion of reef. even under these circumstances (e. and bacteria (Bak et al. Two wells were drilled through over 1. Barnes and Taylor 2001). a b Reef Algae and Foraminiferans•69 material for herbivores. eastern Pacific. In: Krupnick GA and Kress WJ. tributed as calcareous sediments after the death of the colony. Smith and Birkeland 2007). manently. tem pores and polyps. Joenje M.coral skeletons. Indeed. but may become threatened if their trade is not carefully regulated. Knight MA. 1: Invertebrates: 257–270. 2008. Hentschel et al. tic oceans. French Polynesia). for example. Sponges (Porifera). similar to the way vertebrate skeletal elements are joined by collagenous ligaments. Others may produce larger. causing them to completely retract and suspend feeding activities (Fautin 1991. Reef Corals and Their Allies•133 pore develops by ectodermal invagination. and mineral composition of sclerites in the alcyonarian coral. Carbonate buildups may include platforms. similar to that exhibited by anthozoan corals (Lewis 1989. ment. ing internal spaces. lographic axes. 2005). the bioeroding community. the growth records of some specimens indicate that rhodoliths may remain in place for periods up to several months. Indeed. After Lambeck K and Chappell J. 1996.. as well as by molecular scaffolds that may play a role in the assembly of crystals. tropical lagoons. Nava and Carballo 2008). Overview Cyanobacteria or cyanophytes are photosynthetic. 2008). are now generally losing carbonate to erosive forces as a result of recent environmental changes (e. but these are typically associated with the parapodia on the body segments rather than the head. For example. 2003.CS1%20BlackCoral/WG9. 2007. tent with the known function of other such tissues. 2005). Ecological obser. zoanthids (Chapters 5 and 6) and certain ascidians (Chapter 9) can hold reef mate. successive uplifts. Marine Biology 147: 497–508. The excellent vector graphics throughout this book were produced by close collaboration with my illustrator.nl. branched. glove manner. although the quantitative importance of these or. Bricker I. Swart PK. Sponge feeding by Caribbean an.lived. Littler MM. segments with a simple cortex and large in. rogate hosts (Marín and Ros 2004). Scleraxonian gorgonians are distinguished by an axial skeleton composed primar. and crustose forms that form an understory across the substratum. and Wei Z.g. structure in the deep. 1997). Nematocysts are the only cnidae known to be capable of delivering venom. The organic phase appears to control carbonate precipitation and crystal growth. The tissue.in. While most of them take a slender and tapered form. they also produce bright white ones that are mineralized with calcium car. Assmann M. rulea (Pallas) in the Pacific. Similarly. Calcite is secreted by a wide array of reef organ. Limnology and Ocean. such processes are integral to understanding the biology of reefs. cyanophytes are prokaryotes. from the Dry Tortu. 2008. Tube construction begins when planktonic serpulid larvae settle and undergo metamorphosis. crests that have grown close to the surface may be buffeted by waves or exposed to air or rain during low tide. the reef would con. 1. tail of stromatolite as in 3–12a showing the internal growth layers.moving polychaetes occurs directly through the body wall. dictidine. van Oostveen MM. 2003. Ecological goods and ser. 1985. cord of the movement of the same hotspot over the last 80+ million years.5 m high. Reef Sponges•99 Grotto. Courtesy of Image Science and Analysis Laboratory. as the process continues the lateral cell walls partly dissolve. that is. plored below. Under these condi. Marine littoral. Seaweeds are traditionally classified into divisions based largely on their pigment composition. undergoes multiple fission producing more coccoid cells. 2011. and thoughtful work. Messing CG. riod. branched. or in sequence (male then female. and fungi). However. eds. tion by more rapidly growing. lagoon. and Calcification in Halimeda A series of interwoven. –a 3 +c –c +a 2 –a 2 –a 1 +a 3 +a 1 Figure 2–8: Aragonite crystals in their simplest form resemble a cereal box with two parallel. although sponges lack the regional specialization and organization required for colonial status. Chlorophytes). De Kluijver A.temperature (e. little is known about these interchanges and the overall ecological importance of Figure 5–19: Section through the Caribbean barrel sponge Xes. Figure 4–9). resulting in the release of mature larvae. crophagous carnivores) to corals and crustaceans. especially those in the form of amino acids. Wentworth Scale (Grain Size). As a result. Atoll Research Bulletin 129: 1–41. Cores have the advantage of viewing net carbonate production. bellids chemically weaken the substrate by the secretion of acidic polysaccharides from glands associated with the chaetae (Chughtai and Knight. and mannitol. Some groups of fishes also invest their teeth with iron. Abou. Sexual reproduction in Acropora (Iso. Potentially damaging ultraviolet light is absorbed by scytonemin.Penney and Racey 2004). Fossil ste. up to 60.. Nature 413: 726–730. the volcanically active is. mor.g. where they cover a large area of the continental shelf. a split occurs between the upper and lower parts of the polyp. Boca Raton: CRC Press: 481–520. Blooms of certain species of Lyngbya can cause skin eruptions on contact owing to a group of allelochemi. Moreover.associated coccoid species. fish Chlorurus sordidus. and carbonate. cles. The order Antipatharia includes about 235 species (Daly et al. which may help explain the existence of “weedy” species with a wide geo. and most of these are hermaphrodites that release eggs and sperm together.building corals and >80% of scleractinian species studied thus far are broadcasters. the symbionts func. pharia. 1940. Both the GBR and Belize reef are continental barrier reefs. ranging from 1.Bunker reefs. Special thanks are due to Peter Glynn (University of Miami) and Chuck Birkeland (University of Hawaii) for the yeoman labor of reading the entire manuscript.87 Gy old and can reach 80 m in length and 20 m thick. Likewise. A Functional Biology of Clonal Ani. Biological Bulletin 213: 227–251. 2003. the population density of this pioneer species may change with time and conditions. Reef fishes may be contributing to this prolifera. lakoid by a related allophycocyanin. cene Halimeda alga. Phycologia 14: 55–69. dark skeleton can be seen through the tissue. a more profusely branched species that prefers deeper. Demosponges typically produce at least 12 dif. More typically. Mills and Sebens 2004). each of which is composed of several classes based on the symmetry and arrangement of their crystallographic axes

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