Minerals Management Sewice MMS 84-0038 THE ECOLOGY OF THE SOUTH FLORIDA EFS: A Community Profile
U.S. Bepaflment of the Interior Fish and Wildlife Service Minerals Management Service BR \\ a0 FWSjOBS - 82/08 MMS 844038 August 1984
THE ECOLOGY OF THE SOUTH FLORIDA CORAL REEFS: A COMMUNITY PROFILE
Walter C. Jaap Florida Department of Natural Resources Marine Research Laboratory St. Petersburg, Florida 33701
Project Officer
J. Kenneth Adams lvlinerals Management Service Gulf of Mexico OCS Region Post Office Box 7944 Metairie, Louisiana 7001 0
Prepared for
National Coastal Ecosystems Tern Division of Biological Services Research and Development Fish and Wildiife Service U.S. Department of the Interior Washington, D.C. 20240
and
Gulf of Mexico OCS Reginnd Office Minerals haanagemen&Service U.S. Department of the Itlieriot. Metaisie, Louisiana 700 18 ihc oplnlons, findings, conduslori>, or recornrnel~datlonsexpressed In tius report are those of the authors and do not nece\sdrily refle~tthe vlc\lis of the U.S I-~rhand Wlldl~feService Or the Mlnerals Managenlent Service unless so cic51gnGitcdby uthcr '~uthon~cddo~tl~nents I llii coritr,tctual repor"la cdlteti to the standards of the Dlv~slonof Biological Services and does not cc~nfonnc~~tilely to thc tc~llr~i~alcdltlng guidehnes of the Mmcrals 'Managenlent Service. PREFACE
This profile of the coral reef community of south Florida is one in a series of conlnlunity profiles that treat coastal and marine habitats important to ntirrtans. Coral reefs are highly productive habitats which provide living space and protection fro111 predation for large populations of invertebrates and fishes, xnany of which have comnlercial value. Coral reefs also provide an important economic benefit by attracting tourists to south Florida. The information in the report can give a basic understanding of the coral reef community and its role in the regional ecosystem of south Florida. The primary geographic area covered lies seaward of the coast from Miami south and west to the Dry Tortugas. References arc provided for those seeking indepth treatment of a specific facet of coral reef ecology. The format, style, and level of presentation make this syntl~esisreport adapt- able to a variety of needs such as the preparation of environmental assessxnent reports, supplementary reading in marine science courses, and the education of participants in the democratic process of natural resource manage- ment. Comments on or requests for this report should be directed to:
Information Transfer Speciahst Records Management Section (OPS-4) National Coastal Ecosystems Team Gulf of Mexico QCS Region U.S. Fish and Wildlife Service Minerals Management Servlce NASA/Slidell Computer Complex Post Office Box 7944 10 10 Cause Boulevard Metalrie, Louisiana 700 10 Slidell, Louisiana 70458 (504) 837-4720 ext. 25 19 (504) 255-651 I ,ZIIljIIE VIA I IONS
1 1 O\<'IVIC\S 1 2 ( or,ti I(cc.1 f)i--,lr~l)ut~ori I 1 ( orl~rl~~tt!i()I)t\l~~ltution i 4 tildctr~t,iilics~~trt k)l t loridhi ( r~r,tlKccf Kestarch ! 5 t tl)llc)r!uc \i~~!lifl~,ill~*~
4.1 I~~ttt~tluctiun 4 2 21g.i1* ty ilsrlrid iFutn;n 4 ? Sp;.i;tgc~,5;. f; P. Srfin:a!i! 4 4 i'alitiarid '4 4 Other Wcnlirrc i~ioups 5.1 lntroduciion 5.2 Taxononli~Composition and Spatid Distribution 5.3 I)iurnal Migrations 5.4 Summary
CIIAPTE R 6. REEF FISlI by Jarrles 'T. ?'il~nant
6.1 Reproduction and Recruitnlent 6.2 Food Ifabits and Trophuc Structure 6.3 Movcrnent 6.4 Social Organizatlorl 6.5 kcological Aspects of Reef Fish Diversity 6.6 Comrnuility Descr~pllo~~s 5.7 Reef Fish Manage~nent
CHAPTER 7. CORAL REEF ECOLOGY
Introduction Physical-Chemical Envronment Conlmunity Structure Diversity Symbiosis Predator-Prey Relations Productivity Coral Reef Models Natural Impacts
8.1 Human Impacts 8.2 Coral Reef Resource Management 8.3 Management and Mitigation Recommendations
COLOR PL,A'I'ES 'FABLES
Page
( ~rirlrllyr~lalldlidirly'r of rci f-r~ldtcdspecles in Monroe ('ounty, 9980 (NMFS 1981) Morzthly ti:r ti~t~ptratritcz('( ) for Midnil and Key West (YOAA 1981) %f(li~tlxi)i)r~cirutc~r lo11 (nlix) for Cliamr dnd Key West (YOAA 1981) l-for~tiliywintl spceri (krrt/hr) dnd directlor1 for Mla~n~and Key West (NOA.4 1981) Mnior hurrliane5 Lrowng the coral reefs from 1873 to 1966 (Sugg el ai. 1970) Kcef w~wdte~ternperdtures (OC) (Vaughdn 1918) Uottctnl (3 ni) sedwdter teinperdture (OC) dt I. lhhorn Control Reef, Biscayne hdt~oridlI'drk, 197%(fr0111 darly thcrmograph ddta, Biscayne Yational Park) 1 lct'tl ranges for \cverd \outhedst I lorrda rectc (NOAA 1981 ) S,tl~n~tlcsin the i lorlcl,~rezf tract and vlclnity Age and growtlr r,ttc of Recent I lorlda reefs (Shinn et a1 1977, Shun 1980) i,lve hottorn cor;i!s from Schooner Iicef, Uiscaync National Park (four 1-m" quadrants. Jadp 'ind Wtteaton 1977 MS) Octocor,ils at i)onic Kecf (two 20-ni transects, 1977, Wheaton, in prepdratlon a) Octoc orals at ilo~nc('ontrol iiref (two 20-rrl transects, 1977, Wheaton In preparation a) Stony corals at 1)ontc Keet (two 25-m t~ansects,1977, Jaap, unpublished) Stony corals at i)otr~c('ontrol Kaef (two 25-m transects; Jaap, unpubhsfted) ~?itf110utcsof stony coral assoc~dt1011~.Biscayne National Park, based on several Line transccts at cacti reef (8 reefs, 18 transccts) (Jaap, unpubltshcd) blliliorn Heel stony coral farina (lour 4-ri12 plots sarnpled per reef, 1978, Jaap, irn~)ut~lrshcd) 1 lhhrrrn f'oritrol Reef 5tony ~or~ilfauna (four 4-m2 plots samplcd per reef, 1978, J'tit~). ~r~p~t)lishctl) 1 Ikl~ornKccl ~)ctocor~Ifduncl (three 10-111 t~dn~ects,1977, Wheaton, unpubhshed b) I Lklic>rnC'ontrol Kccf o~1ocor;tlf,~un,~ (three LO-rn transects, 15'7: Wheaton, ~~nj~~tt~l~shed1)) ti Ariik Itcrf \tony ~oritltduila .tt 17 rn (trftecr~I-m2 piots, Jdap, unpubl~sheti) in lfarlk rccl tonatron a,~ttcrns i $4 (
llor~tt~lyttllrumuin, nlrdn, allti II~JU~IIUIII\t'.iw.itCr tc~~~pcr'itllre'it Sltlld hey. I. 878-1 890 (V.tughsn 1 " iiY)
Jottrl f'rnt~i.karilp(.'oral ficcf St;itc I'ark ancl Kt-y I argo National Mitrirle Sarlct!t;ir.y C'ross-sectional diagratlr of I.ooe Lcy Kccf. ('ross..srct~onaIili;igrarr~ of Iiircl Ksy IZeef. C1assificattor1 of llirtl Key Kcdf allti Ilry 'I'clrtugas ixinctl 011 al~~incianucof stony corals using pr'ot~p;tvrragc sorting anti (.'zck3r1owshi's cacSficic.nf. COLOR PLATES
Page
j3 satc!lttc pf-rt,topial,ll i.~~~lt~ci~t1 lorlda. Note ldrge passe\ between Islands in 84 ltic fnxdiile K:.!, , 1 1tlnita ~dy:ICS between the rnanniand and the Keys. ii7 1 IcnLii f
5'11 i Greciar, Rocks, Key Largo Xational Marine Sanctuary; elkhorn coral with schools of sergeant major fish (AD~~d~fdz~f~axatilis). Schools of fish at an unnameci reef off islamorada. Southern stingray (Dasyatis at?zcriccrt~a)near the Benwood wreck, Key Largo National Evlarinc Sanctuary. Longjaw squirrelfish (Nolocentrus ascc~~zsiut7is),Molasses Reef, Key Largo National Marine Sanctuary. I'orkfish (A tzisotrenzus ~ir~qinicus)and yellow goatfish (Mulloiriichthys ~narfinicus), Molasses Reef, Key Largo National Marine Sanctuary. Gray angelfish (Potnacunthus arcuatus), Eastern Sambo Reef. French angelfish (Pornucatrthus paru), unnamed reef off Islamorada. Nassau grouper (Epinephelus striatus), Sand Key Keef. Glassy sweeper (Pernpheris schomburglii), French Reef, Key Largo National Marine Sanctuary. Fire coral, Millepora cornplanuta, following heat stress; zooxanthellae expulsion caused bleached appearance. Unknown pathogenic condition (white) in Acropora palmatu. Mobidity-mortality in Diploria srrigosa caused by fishcollecting poison (rotenone and organic carrier). Shipwrecked Captain Allen on Middle Sambo Reef, 1973. Motor Vessel Lola and salvage barge on Looe Key Reef, 1976. Discarded steel from the Lola in the spur and groove tract. Anchor on coral. Boats lied up to anchor buoys at Molasses Reef, Key Largo National Marine Sanctuary. Eyebolt securing anchor buoys to the reef platforrn at French Reef: Key Largo National Marine Sanctuary. Scars on elkhorn coral (Acropora palrnatu) caused by a boat propeller, Elkhorn Keef, Biscayne National Park . Fish trap without a buoy line (ghost trap) on unnamed reef off Islamorada. Steel junk from Carysfort lighthouse on the reef flat. Accumulated batteries from Carysfort lighthouse on the reef flat. Batteries are used in the lghthouse operation. Lobster trap buoy line tangled in elkhorn coral (Acropora palrnata) at Sand Key Reef, September 1983. Zooxanthellae expulsion, Eastern Sambo Reef, August-September 1983. ABBREVIATIONS
Blscayr~eNational Park (I!.%) Coast Guard Ilcpartrnent of knvironmental Regulation (Flonrla) L)cpartrncnt of NdturaI Resources (f lorlda) 1)epartmcnt of Admini~tralion(Florida) I)eparllx~cntof Statc (Fiorncia) (U S ) tr,nv~ronmentalI'rotcct~on Agency (0S ) PIS^ and Wlldhfe S, ervice i [J S ) <;ct)loglcal Survey John Pcnnckaxnp Ioral Reel State Park Key Lalgo Natic>nal Marine Sat~ctuary(orlglnally referred to as Key Largo Coral Reef Marine Sanctuary in cdrly (itneral Mandgemertt Plan dcvclopcd by YO&&) L.KNMS Looe hey N,lt~onalMarinc Sanctuary MMS M~neral.;Managenlent Servlce NMI'S Ndtlondl Mdrlne I.isher~esService NOAA Nationdl Occalr~cand Atnlospheric Adnlinistration N PS hatlon~ii)~;rk Service lJIl<' Sttl. Ilcv. or SI) US.4t.l. Y ill' Years Heforc I'rcscnt CONVERSION FACTORS
Metric to U.S. Customary
Multiply To Obtain millimeters (mm) inches centimeters (cm) inches meters feet kilometers (km) miles square meters (m2) square feet square kilometers (km2) square miles hectares (ha) acres liters (1) gallons cubic meters (m3) cubic feet cubic meters (m3) acre-feet milligrams (mg) ounces grams (g) ounces kilograms (kg) pounds metric tons (mt) pounds metric tons (mt) short tons kilocalories (kcal) BTU
Celsius degrees 1.8(C0) + 32 Fahrenheit degrees
U.S. Customary to Metric inches 25.40 millimeters inches 2.54 centimeters feet (ft) 0.3048 meters fathoms 1.829 meters miles (mi) 1.609 kilometers nautical miles (nmi) 1.852 kilometers
square feet (ft2) 0.0929 square meters square miles (mi2 ) 2.590 square kilometers acres 0.4047 hectares
gallons (gal) 3.785 liters cubic feet (ft3) 0.0283 1 cubic meters acre-feet 1,233.0 cubic meters
ounces (oz) 28.35 grams pounds (lb) 0.4536 kilograms short tons (ton) 0.9072 metric tons BTU 0.2520 kilocalories
Fahrenheit degrees 0.5556 (PO - 32) Celsius degrees
XI ACKNOWLEDGMENTS
bunding for develop~nentand pubhcation of thls docurnent was jointly sponsored by the Fish and wlldhfe Seplce and the hlll>erals Management Service of the U.S Department of the Interior. Marly people had a duect or ~ndlrectrnflucnce in the creation of this document. I he responsibhty for content and format are thore of the prrnclpal author (W. Jaap), M. Ifurn~nwrote the section on algae: G. Schmahl, sponges; R. Johansson, plankton; and J. 'Iiln~ant,fish. Reviewers included I. Bright, G. Bruger, J.H. tludson, J Uot~nsack,G. kiuntrman, 1. Lang, and J. Wells, these lnd~vidualsare thanked for their criticisms and suggestions for m~provernent.Karllyn Jaap and Alan70 t:elder assisted with preliminary typing, organizing the kterature, and cditlng. The 1:lorxda 1)epartrnent of Natural Resources Bureau of Marine Research, St. Petersburg, aided and contributed in the areas of library resources, unpublished data and graphs, and rise of photographic files. ?'he follawmg individuals are thanked for thelr assistance I:. Joyce, C. butch, K Steidinger, W. I,yons, I) Camp, J. Wheaton, W. Marovec, !,. Truby, M. Krost, and L. 'fesler. l'he U.S. Natlonal Park Service is also tharlkctf fur the frcedotrl to use field data from the Biscayne National Park coral reel study. The concepts and foundation for this docurnent are based on experience and most of all the cross flow of mformatlon from a peat number of it~cl~vdualswho llave studied Florida's coral reefs. The following people directly or indirectly providcd stimulation arid informat~onfor thls document; John Wells, Robert Ginsburg, tugene Shinn, J.N. Itucfson, Jenmter Whcaton, J. Morgan Wells, David Oleson, Peter Better, Lowell Thoinas, Norm Blake, Phlhp l)ustan, John fialas, Gary l)avis, rimid Yosse Loya. Ken Adarns (formerly with the U.S. bish and Wilrlhfe Servlce, now w~ththe Minerals Management Service) B expressly thanked for his patience and prodding. IIe saw the need for this report and hsefforts have ;rirlecl the author In resolving problems and gett~ngtlie work completed. Gaye Farris (l1.S. Flsh and Wildhfe Scrvrcc editor) was resportsible tor the final editing whlct~greatly improved the coherence drtd readability of this report Betty Mrotly perfortncd the Ilerculean cffort of proofreading. Wc tflar~kthe Minerals Mdnagement Sewlce, C;ulf of Mexico Regional Office, far printing this docu- racrrl wltlr color plates which xncreasc the scicrltific usefitlness of tltc report. Larry kfandley, Carla Langley, I)ch\~~cMiller, Kutlr h4onlcal. (:ynttl~ii Nxcholson, I'atricla l'haggard, Cathy Cartwright, Mardyn Parker, and hllfllt C;r~fflttof that office cotlt~ibl~tedyearnan efforts to the corrlpletiun of this pnbhcation, XEktb puhlicatlofi 1s cfcdlcatcd to all who seek the trutti, especially Taylor .4lexander; lost friends Li~l Brown artti i rank Wcscott; and espcc~allyKar~lyn. CHAPTER 1
CHARACTERIZATION OF THE RESOURCE
1.1 OVERVIEW low-diversity coral assemblages, but important fishery habitat. Grouper find refuge, feed, and breed in and near Coral reefs are highly complex and diverse these structures. From Stuart (St. Lucie Inlet) to near communities of biota, a phenomenon of the tropics Palm Beach is a transitional community of Oculina bank and subtropics limited by such factors as suitable sub- flora and fauna and hardier elements of the tropical reef stratum, temperature, light, and sedimentation. In biota. This region is characterized by the convergence of simplest terms coral reefs are concentrated complexes the temperate and subtropical climate zones. From Palm of corals and other organisms that construct a limestone Beach southward to Miami (Cape Florida) elements of structure in shallow water. In the initial building process, the tropical coral reef biota become increasingly impor- a set of primary framework builders set down the first tant in a north-to-south gradient; however, the building structure; later colonizers add to the volume. Skeletal of three-dimensional reef structures does not occur. This breakdown by physical and biological actions creates area is characterized as an octocoraldominated hard- carbonate sediments, which are recycled by other biolog- ground community (Goldberg 1973a). The two stony ical processes or are cemented to the reef framework corals most responsible for reef building (Acropora through biological or geochemical processes. paimata, elkhorn coral, and Montastraea annuloris, star The coral reef complex found off southeast coral) rarely occur here, and currently do not actively Florida represents a mosaic that exhibits extreme build reefs. Acropora palmata was once an important variability in all parameters used to evaluate biological reef builder in this area, but it ceased building reefs communities. Coral reefs provide a wide spectrum of about 4,000 years before present (YBP) (Lighty 1977; vocational and recreational activities. Many important Lighty et al. 1978). Today only a few isolated colonies fisheries are directly tied to these reef communities; are found north of Fort Lauderdale. a reef's principal resource value (economically) is as a highly productive habitat: it concentrates marine protein The region of maximum coral reef development in a localized area. Coral reefs also play a significant is restricted to south and west of Cape Florida, offshore role in the tourist industry of southeast Florida. of the Florida Keys archipelago (Figure 1). This small While the level of reef usage is increasing as chain of islands extending from Soldier Key to Dry southeast Florida experiences rapid population growth, Tortugas exhibits a diverse pattern of hardgrounds, management of these reef communities tends to lag or patch reefs, and bank reefs from 25 m to 13 km off- is unresponsive to the problems described herein. shore. This is the only shallow water (< 10 m) tropical Scientific and lay Literature has reported real coral reef ecosystem found on the Continental Shelf of and potential threats to southeast Florida coral reefs North America, and has been refened to as "The Florida (Straughn 1972; Voss 1973; Davis 1977a; Dustan Reef Tract" (Vaughan 1914a). This discontinuous 1977b; Bright et al. 198 1). Impacts have included vessel assemblage of reefs forms an arc paralleling the Keys' groundings and sinkings, oil spills, anchor damage, beach coastline in a general southwesterly trend. Landward, renourishment dredging, fishery activities (lobster trap the reefs are bounded by the Keys and a series of shal- recovery), tropical fish and invertebrate collecting, low embayments (Biscayne Bay, Card and Barnes shipwreck salvage, and diving-related activities. Individ- Sounds, and Florida Bay); seaward of the reefs are the ually these acts do not greatly affect the resource Straits of Fiorida and the Florida Current. The Florida vitality, but the chronic and synergistic nature of some Current, a subsystem of the Gulf Stream, plays an of these acts is cause for concern. important role in the existence and maintenance of coral The goal of this document is to serve as a refer- reefs off southeast Florida. It modifies the environment ence for those interested and concerned individuals by moderating winter temperatures. The current's source responsible for environmental management of the 1s tropical; hence, its waters are significantly warmer resource, as well as those seeking a better understanding than resident shelf water masses during the winter, of Florida's coral reefs. thereby modifying winter thermal conditions such that offshore reef development is not hindered by extremely 1.2 CORAL REEF DISTRIBUTION cold weather that occasionally occurs when cold fronts intrude into southeast Florida. Inshore patch reefs, Although the tropical coral reef communities however, are more vulnerable to cold extremes caused found off southeast Florida (Figure 1) are the emphasis by winter weather. The current system is dynamic, and of t!t:s report, 3 brief surn:nary of coral reef distrihutiooq eddies or meanders bring considerable volumes of water throughout Florida will aid m understand~ng this re- into the reef environment. This brings plankton, a food source. From the Georgia border to near Fort Pierce on source, and new recruitment from nonresident popula- the Atlantic coast, in depths of 15-50 m, Oczdinu (pret- tions to the reefs. The significance of the Florida Cur- zel coral) bank communities are the dominant coral rent cannot be overestimated when considering coral community (Avent et al. 1977; Reed 1980). These are reef existence off southeast Florida.
The Keys or islands act as barriers to cross shelf reefs the sea bottom (where it has been investigated) water transport from shallower bays and sounds. These consists of sediments, sponge habitats, and rubble, with bays are very shallow, hence much influenced by mete- occasional rocky outcrops that generally parallel the orological events. Temperature, salinity, and turbidity basic reefs. can be significantly affected. Heavy rainfall, drought, The coral reefs, seagrasses, and mangroves are all and winter cold fronts are the major influences. Coral integrated into the coastal ecosystem of southeastern reef distribution patterns reflect the extent of water Florida. Motile species and trophic energy webs are exchange between the bays and sounds and the Atlantic. integrated and interrelated among the three communi- The larger northern Keys (Elliott and Largo) have exten- ties. sive reef development off their coasts; the middle Keys, which are smaller and more separated, have numerous 1.4 HISTORICAL RESUME' OF FLORIDA CORAL channels communicating with Florida Bay (Plate la) and REEF RESEARCH exhibit less reef development than the upper or lower Keys. The influence of Florida Bay on water quality Arbitrarily, coral reef research can be chrono- negatively affects reef developnient in the middle logically placed into three periods: early (to 1900), Keys area. middle (1900 to 1950), and recent (1950 to present). Major bank reefs off the Florida Keys include Early research was descriptive and taxonomic, and was Carysfort, Elbow, Key Largo Dry Docks, Grecian stimulated by the need to provide safe navigation for Rocks (Plate 3a), French (Plate 1b), Molasses, Alligator, marine commerce. National security also had some Tennessee, Sombrero, Looe Key (Plate 2a and b), influence in that the Caribbean Sea prior to 1860 was Eastern, Middle, and Western Sambos, American Shoals, frequented by pirates and European naval powers. Eastern and Western Dry Rocks, Rock Keys, and Sand During the age of discovery, the Florida coastline was Key. Dry Tortugas is studded with various coral reefs crudely mapped by the Spanish and Enghh. Agassiz (Davis 1979, 1982). (1852, 1869, 1880, 1885, 1888), LeConte (1857), Hunt Off the west coast of Florida, tropical reef (1863), and Pourtales (1863-1869, 1871, 1878, 1880) development is nonexistent. Ledges and outcroppings provided the earliest reef descriptions and details about are a special rocky habitat which supports an association the coast. They also initiated systematic description of of hardy corals and other biota; however, they do not the many organisms found in and near the coral reefs. construct three-dimensional reefs. The Florida Middle During the middle period, establishment of the Ground (a fossil reef formation about 157 km northwest Tortugas Laboratory on Loggerhead Key by the Car- of Tampa Bay), while exhibiting higher diversity in coral negie Institution was the most significant factor bene- species, is not an active coral reef comparable with those fiting cord reef studies during the first half of the found off southeast Florida. These areas in the eastern twentieth century. Alfred G. Mayer and his colleagues Gulf of Mexico, however, are critical habitats that conducted much fundamental coral reef research from should be provided with rational management, especially the Tortugas Laboratory. Mayer (1914, 1916, 1918) since important fisheries are found in association with determined the thermal tolerances of tropical marine the gulf live bottom communities. organisms in general and reef coral in particular. T. Wayland Vaughan was the most prolific Carnegie reef 1.3 COMMUNITY DISTRIBUTION researcher, reporting on the geology of Tortugas, south Florida, and the Bahamas (Vaughan 1909, 19 10, 19 12, The Florida Keys possess coral reef communities 1914a, f 9 14b, 1914c, 1914e, 1915a; Vaughan and Shaw similar to those found in the Caribbean and other 1916). He studied the taxonomy and growth rates of tropical Atlantic areas. The reefs are bathymetrically scleractinian corals (Vaughan 1911, 19 14d, 191Sb), distributed from less than 1 to 41 m deep. Large-scale summarized seawater temperatures at reef tract light- synoptic mapping of the Keys' reefs has been recently houses (1918), and reported on the Tortugas current completed (Marszalek et aI. 1977; Marszaiek 1981, structux e (I 935). Other students of scieractinian corals 1982). Caution should be exercised when interpreting include Duerden (1 904), postlarval development in these maps, as they use mapping units based on geo- Siderastrea radians; Matthai (19161, systematics; Bosch- logical and biological criteria, and in some cases errant ma (1925, 1929), postlarval development in Manicina interpretation could occur due to their large scale and areolata; Wells (19321, thermal tolerance knowledge; criteria used. and Yonge (1935a, l935b, 19371, M. areolata aute- The basic pattern of marine conimunities m cology, taxonomy of Siderastrea spp., and mucus southeast Florida (where commercial development has production in stony corals. Other reef-related research not occurred) is one of a shoreline dominated by man- from the Carnegie laboratory includes work by Cary groves, rocky intertidal, or sedimentary environments. (1914, 1916, 1918a, 1918b), Bctocordlia; Taylor From the intertidal zone to Hawk Channel 1s a mosatc of ( 1928 ), algae; deLaubenfels (1 9361, Yorifera; and environments including sediments, seagrass exposed Longley and Hildebrand (1941), fish. Another first for Pleistocene rock, and some patch reefs. Seaward of the Tortugas laboratory was that the first underwater Hawk Channel, patch reef abundance and frequency color photographs of coral reefs were taken by Longley increase and bank reefs begin to occur. Seagrass is often at Tortugas. Dole (1914) and Dole and Chambers (191 8) abundant and sediment is coarser. Seaward of the bank reported on salinity at Tortugas and off Fowey Rocks 3 ). lighthouse. L.eCon~pte ( 1937) described some 7 Oi tt~gd~ Jameson (198 1 Florida Current plankton was reported reefs. Welk (1933, unpublished) contains ~cm,!t!erdt>fe on by Bsizarai~ (1957). Emery (1968) discussed coral informati~non the Tortugas reefs. A fire dc:,iioy~dthe reef plankton. Coral predators and boring and rasping biota were reported by Robertson (1963), Ebbs (1966), Tortugas Laboratory m 1937; it was never rebullt. Verrill (1902), Smith (1943, 1948, 19711, Antonius (1974b), IIein and Risk (1975), and Hudson Vaughan and Wells (1943), and Wells (1956) are the (1977). Physiology of reef corals, particularly the relevant scieractlnlan taxonomnc and systematic refer- symbiotic relationships of corals with zooxanthellae, was ences. During the recent penod (1950 lo present) there studied by Kanwisher and Wainwright (1967), Kriegel has been a great proliferation of scient~fichterature (19721, Chalker (1976, 1977), and Chalker and Taylor dealing w~thFlorida coral reefs. Growth rates of coral (1978). Reports on the impact of human activities specles were detailed by Broecker and Thurber (1965), include McCloskey and Chesher (1971), Hubbard and Shinn (19661, Jaap (lc)74), Landon (1975), Exniliani et Pocock (1972), Straughan (1 972), Voss (1 973), Anto- al. (I97$), Dodge ( 19801, and Hudson (1981 ). Popu- nius (1974b, 1976, 1977), Courtenay et al. (1974), lation dynamics of scleractinian corals were reported by Griffin (1974), Jaap and Wheaton (19751, Manker 1)ustan (1977a). Brooks (1963), IIalley (1979), Davis (1975), Shinn (1975), Chan (1976), Britt and Associates (1982), and Porter et al. (1982) added to Tortugas (19771, Davis (I977a), Dustan (1977b), and Bright et al. mforrnation. (198 1). Coral reef growth rates in Florida were reported L~cologicalstudies and reviews include Smith et by Noffmeister and Multer (1964), Shinn et al. (1977), al. (1050), Voss and Voss (19551, Jones (1963, 1977), and Shinn (1 980). C'rlynn (1973), lludson et al. (19761, Hudson (19811, Echinoid distributions in John Pennekamp Coral Cloldberg (19733, 1973b). Antonius (1974a1, Klssling Reef State Park were studied by Kier and Grant (1965). (1975). and Shinn (1975). Coral reef fish studies in- Miller el al. (1977) reported on diatoms in the park. cluded Springer and McErlean (1 962a, 1962b3, Starck Janleson (198 1) reported on the biology, geology, and (i1)68), Starck and Davis (19661, Bohlke and Chaplin cultural resources in deeper regions of the Key Largo (1968), X:~ncry (1973). and Jones and l'hompson National Marine Sanctuary. Workshop and symposia (1078). Rccf morphotogy and physiography were literature includes Stursa (1974) and Taylor (1977). reportcct by Stephenson and Stephenson (1950). Shinn Shin11 (1979) detailed collecting-pennit requirements for (1966), Kissling (1065), Starck and Schroeder (1'165), biological and geological sampling in the Keys' area. St.irck ( 19b0). Straugtian (1978), 'i'urmel and Swanson (1Wl), Weeks (19721, llannau and Mock (1973), and 1.S ECONOMIC SlGNIFICANCE Avent el al, (1077). (;cological htcraturc includes Ciinsburg (1956), Exploitation of Florida's reef resources began Rruoks (1963). ME anti Brooks (1965). iioffrneistcr and with the Caloosa Indians, who harvested marine protein Muler (1 9681, Duane and Meisbcrgcr (I 969), bnos (fishes, lobsters, and conchs), shells, and coral for (10701, MuXter (iY7 1), Raymond (1 972). Ginsburg trading. Excavated middens occasionally contain coral (1975), kms (19741, Ciieason (19741, lloffnaclster artifacts. Durlng the mid-17th to late-19th centuries, the (19741, I:tios (1977), Lighty (1977), Lighty ct al. Florida reefs posed a significant navigational hazard to 119781, %ant3 (19X0), Mitchell'i'ripping (198 I), and huropeans and Americans. Today, salvagers recover gold, Cituolrf and 14ncts(1982). Geographical research includes silver, and artifacts from the numerous shipwrecks work. by (;~lrsburg artd Shinti (1641, Mars~alekel al. adjacent to the reefs. Many reefs are named for ship- (lC177), hlarsmick (15181, 1982), Kecd (1980), and wrecks. Looe Key Reef is named for the 44-gun British Sloct tiart and Fosberg (1 98 I ), Frigate, the IiMS Looe, which was wrecked on the reef Culdcboc>ksor general references Lo t:londa reefs the niorning of 5 Febn~ary 1744. Molasses Reef is tncludc ftoflmckstcr et ai. (196.4). (;msburg (1972a, named for an unknown vessel laden with molasses that 1072b). Xuller (19741, Voss (I976), Greenberg (1977). foundered there. ('iltxn lit)7Xa), drrd Kapian (1982). tiurricane ~nrpacfs011 Early Florida Keys' settlers had a thriving enter- i~lor~darccf's hdvc bcen documented hy Springer and prise of luring unsuspecting ships onto the reefs with hfctarlcan (1962a), Ball ct al. (l967), I'erklns and Enos false beacons. They then claimed salvage rights on the t i ObXj, and Sillrin (19'751 'Thc effect on cord reefs by wreck, salvaged the cxgos, and auctioned them off. other ntrtevrolug~catphenonicna was studied by Jaap Wood from the vessels was also salvaged. Many of the {XY79), Walkci (1981 1, and Roberts et al. (1982). older homes in Key West are constructed with salvaged i)frysica1 and chcrn~ca1oceanography ad~acerrtto the reef ship timbers. In an effort to reduce the shipwrecks, early tract includes studies by liela (1 952)., Chew (1 954). coral reef work was directed toward mitigating the Stortnrrkei (1 9591. Wcnnckens (1 9S9), McCaUum and hazards. This resulted in the lighthouse construction Stockrnan I 1.964f. Gardun and I)era ( 1969). i.$ansoil and between 1825 and 1886 on the most dangerous reefs. Polrnifexler (1972). Lzu (1Y75), Lcc arld Maycr (1977). These lighthouses significantly changed the nature of the Lee and Mouer-s (1W7), and Lelnillg (1979). Keys' economy. Fishing, cigar making, sponge har- 'The ~~tocor~lffiawere reported on by Goldberg vesting. and agriculture replaced shipwrecking as the (IY73a, 1973b). Opresho (1973), Cairns (19773, and major economic endeavor at Key West. Wheaton (I981 1. tlriticrseas parks were reported on by Commercial sale of coral began in Key West V~sset af. (19b91, S~?~tfz(1972), Tzilnoulis (1975), and around 1830 and remained a poorly organized cottage Industry unt~l 1950. Durmg thst period collectors Table 1 used cither grappling hooks fro111 boats or hand liar- Coniliierclal landings of reef-related species in Monroe vested while reef diving. 'Fhe industry changed with the County, 1980 (NMFS 1981). advent of scuba diving and the increased interest by the general public in the rnarine environment. There was increased demand for coral by tourists as well as for Weight Value export to northern markets. No quantitative data exist Species (lb (S) on the nlagnitude or economics of the coral harvest. It is suspected that commercial coral harvest at no time Ballyhoo employed more than 20 individuals working on a part- Jacks time seasonal basis. In 1973 and 1975 Florida enacted Dolphin statutes making it illegal to collect, sell, or damage stony Grouper & scamp corals (Millepora and Scleractinia) and two species of sea Ziogfish fan (Gorgonia) within State waters. In 1976, the Federal Jewfish Covernrrient (Bureau of Land Management) wrote Shark regulations under the authority of the Outer Continental Snappers Shelf Lands Act to protect corals and reefs in the area Lane undcr federal jurisdiction (beyond the 3-mi limit in the Mangrove Atlantic). 'The Fifth Circuit Court of Appeals, however. Mutton ruled that these regulations can only be applied when Red active mineral or petroleum exploration or production is Vermilion occurring in the immediate vicinity of the coral. Current- Y ellowtail ly, the Gulf of Mexico and South Atlantic £.'ishery Triggerfish Management Councils are preparing a nianage~nentplan Warsaw grouper for corals and coral reefs in the region bctwccn North Spiny lobster Carolina and the 'Texas-Mexican border. Spanish lobster 'Today, coral being sold is foreign. From 1977 to 1979, 200,000 pieces of coral were imported with a dockside value of S3 1,500. Retail niarkup would place the value of imported coral at about $95,000. Most of this coral came from the Philippines, where collecting and selling coral is illegal, but enforcement is difficult because of the thousands of islands belonging to this in Monroe County devoted entirely to tourist diving. For nation. the most part this activity is a nonconsumptive form of Economically, Florida coral reefs directly or reef usage. Most tourists corne to experience the reef indirectly generate an estimated $30 million-$50 million environment firsthand and to observe fish. Some divers annually within the Monroe County region. 'These do spearfish and catch lobsters. Spearfishing is banned in monies come from all aspects of fishing, diving, educa- some marine parks and sanctuaries, i.e., John Penne- tion, and research. Commercial fishing in particular kamp Coral Reef State Park, Key Largo National Marine depends heavily on the coral recf habitats. Most of the Sanctuary (JPCRSP-KLNMS), and Ft. Jefferson Nation- sought-after species spend all or part of their lives in the al Monument. Much of the tourist diving is concentrated reefs. For some species, the coral reefs are a nursery area offshore of Key Largo in JPCRSP--KLNMS, off Big where juveniles mature into adults. Many species breed Vine Key at Looe Key National Marine Sanctuary and/or feed within the confines of the coral recf. Kesi- (LKNMS), and off Key West. dent fish populations may only seek shelter and refuge Besides dlving, there are glass-bottom boat tours in the reef and feed in the nearby surrounding grass that allow the nondiver to enjoy the reef firsthand flats or the open sea. 'The life history patterns of indi- without getting wet. Charter airplanes also fly tounsts vidual species vary, but the reef is a critical link to the over the reefs. Tourist gift shops market many reef- success of these species. Table 1 presents commercial related souvenirs, from colorful T-shirts to postcards. landings of reef-related species in Monroe County for All levels of education, from elementary to 1980. graduate school (including youth organizations, scouts, Diving as a sport and hobby attracts more than a sea camps, and diving schools), bring students to the million people to the Florida Keys annually. 'These coral reefs to supplement classroom experiences. Special divers rent and purchase equipment, charter tours to the pubhcations, documentaries, and movies about the coral reefs, and purchase food and lodging. Tourists come reefs are produced. These are an economic and educa- from as nearby as Homestead and Miami, and as far tional benefit to the Nation. away as Europe and Canada. A 1979 Skindiver magazine Economic impacts of applied and bas~c~esearch survey indicated that the Florida Keys was the most on coral reef communities include equipment rentals, air popular diving location in the United States among fills, and lodging. Potential commercial applrcalions traveling divers. The survey reported that the average of this research will benefit pharmacolog~ (4~ntl-c;ancer diver spent about $7 18 per trip. There are 40 businesses compounds from vat-ious reef organisms are being te\tcd), mcdlcsnc, (art~tlciali,rirrc\) c-t~i~ig~i,rccl fl>h- an item of cornrncrce rhelr I~ahitatvalue and attractror~ enes m,rnagcrr~ent,aquarjd. rii~nin~turc~e~hnlyues, dnd to divers are worth far more than as an item of com- arci~acolugy merce. Whde coral reefs ren~runon the seafloor, they are Keverluc fmnl all thest. 'ictrkitie.; reenters the like a good investment they continue to generate sotith I:lortcia ccononiy ~ndgenerates employment for monies through the marine protein harvested there and many 0th~~.people ln the SCNICC seciors. The corals' the dlvers who come to enjoy them. As cur~os,they gretitcst V&C, however, m as a kvlng resource, and not as bnng a smaller dividend to a fewer number of people. CHAPTER 2
THE ENVIRONMENT
2.1 CLIMATE
The climate of southeast Florida is characterized as subtropical marine in Miami and tropical maritime in Key West (NOAA 1981). The region is heavily influ- enced by the adjacent marine environments; the Florida Current, Gulf of Mexico, and Atlantic Ocean affect the terrestrial and marine climates during different seasons. Daily average air temperatures in Key West range from lows of 18.8' C during January to highs of 31.9~C in August (NOAA 1981). Compared to those of Key West, JFMAMJ JASOND Miami air temperatures are measurably lower in winter Figure 2. Monthly mean air temperature for Key and slightly lower during summer (Table 2 and Figures 2 West and Miami (NOAA 1981 ). Data base is 29 years and 3). There is a dry season from November to April in for Key West; 38, Miami. Miami and November to May in Key West. Key West averages 1,007.4 mm of precipitation annually; Miami, 1,s 18.9 mm (Table 3 and Figure 4). Key West's weather 301 0 MIAMI station is within 0.5 km of the coast and is more repre- sentative of the reef situation. Throughout most of the year southeast and east-southeast winds prevail, and velocities normally range from 10 to 20 km/hr; wind velocity is less during the summer (Table 4 and Figure 5 ).
2.2 HURRICANES j210 1 Severe hurricanes are common meteorological JFMAMJ JASOND phenomena in southeast Florida. A formal hurricane Figure 3. Monthly minimum air temperature for Key West and Miami (NOAA 1981). Data base is years for Key West; 38, Miami.
Table 2 Table 3
Monthly air temperatures (OC) for Miami and Key West Monthly precipitation (mm) for Miami and Key West (NOAA 1981). (NOAA 1981). Miami -Key West Miami Key West perioda Range Mean Range Mean ~onth~ mean mean
January 54.6 42.4 January February 49.5 47 .O February March 52.6 39.6 March April 91.4 55.1 April May 155.4 63.8 May June 228.6 115.6 June 175.5 104.4 July July August 170.7 113.5 August September 222.0 186.4 September October 207.8 141.5 October November 69.1 67 .8 November December 41.7 38.6 December
Yearly 14.8-32.2 24.2 18.8-31.9 25.7 Yearly total 1,518.9 1,015.7
"C~inatedata base for Miami = 38 years; a~limatedata base for Miami= 38 years; Key West = 29 years. Key West = 29 years. 7 D KEY WE57 seasor, exlsts fronl i June to 30 November. Thirteen 0 MIAMI 1 major hurricanes have passed across the reef tract and struck the Florida Keys since 1894. Criteria for a major hurricane include one or more of the following: 200- km/hr wind speed; winds reaching out 160 km from the hurricane eye; barometric pressure of 716.28 mm of mercury or less; and/or tide surge of 2.7 m or greater. The Florida Keys has a greater probability of hurricane ol.lir:~impact (one in seven) than any other Florida coastal area JFMAMJ JASOND (Florida Department of Natural Resources 1974). Facts pertaining to hurricane impacts on coral reefs will be Figure 4. Monthly mean precipitation for Key detailed in the reef ecology chapter. Table 5 summarizes West and Miami (NOAA 1981). Data base is 29 years major hurricanes that have crossed the reef tract since for Key West; 38, Miami. 1873.
2.3 SOLAR RADIATION Table 4 Peak solar radiation occurs between 0900 and Monthly wind speed (km/hr) and direction for Miami and 1500 hr in Florida (Barnes and Taylor 1973). Trans- Key West (NOAA 198 1). mittance through the water column is greatest at solar noon; albedo (reflection) is significant during early Miami Key West morning and late afternoon. According to Hanson and Montha Mean Direction Mean Direction Poindexter (1972), maximum solar energy was expended speed speed at the air-sea interface from 1000 to 1400 hr; most of (kmlhr) (km/hr) the energy was expended in heating the water column. Gordon and Dera (1969) studied solar radiation atten- January NN W NF, uation between Key Largo and Great Abaco Island, February ESE SE Bahamas, and found that in the surface layers (0-5 in) March SE SE the attenuation (diffusion coefficient) was 0.1 1-0.57 April ESE ESE Kd/m between Miami and the Florida Current. Hanson May ESE ESE and Poindexter (1972) reported that the amount of solar June SE SE radiation impinging on the bottom at 13 m ranged July S E ESE between 5% and 17%; zenith angle, cloud cover, wave August SE USE action, and turbidity all influenced transmittance. September ESE IISE Kanwisher and Wainwright (1967) reported that Florida October ENE EN E reef corals requlred between 200 and 700 footcandles November N EN E (fc) of solar illurnlnation for autotrophic self-reliance llecenlber N NE (compensation point). On clear days these values cor- related with a depth of about 30 m. Y carly 14.8 ESE 18.2 ESE 2.4 SEAWATER TEMPERATURE
"~lirnatedata base for Miami = 38 years; 'The most extensive data base for reef seawater Key West = 29 years. temperatures appears in Vaughan (1918). These data came from several lighthouses and Ft. Jefferson, Dry 1 ortugas, and are summariz,ed in Table 6 and Figures 6-10. More recent data from a patch reef in Blscayne National Park are presented in Table 7 and Figure 11. Temperature extremes are from 14@to 38' C; most annual ranges are from 18@to 30' C. Mean values are above 18@C, the threshold temperature generally ac- cepted for structural reef development by reef bullding corals (U'ells 1956). Temperature variability (range and standard devration) is greater during the winter. In recent viintcrs (rice 197 6 5, polar a2i masses have cooled coastal waters, causing fish kills and coral mortalities. Areas that have suffered the greatest harm from thermal JFMAMJ JASOND stresses are off Loggerhead Key, Dry Tortugas (staghorn Figure 5. Monthly mean wind velocity for Key coral thickets), and patch reefs off Plantation Key (Hens West and Miami (N0.4.A 1981). Data base 1s 29 and Chickens and The Rocks). Cold water masses are years for Key West: 38. Miami. creatcd in Florida Bay during the winter passage of polar Table 5
Major hurricanes crossing the coral reefs from 1873 to 1966 (Sugg et al. 1970).
Hurricane Date Wind speed Tide height name 13ay Year (km/hr) (m
18-30 September 1894 167 test.) - I896 161 (est.) 27 August 1900 - - 1 1-22 October 1906 - - 6-1 3 October 1909 - - 9-23 October 1910 177-201 (est.) 5 2-1 5 September 1919 177 (est.) - 1 1-22 September 1926 222 2-4 22 September-4 October 1929 3 1 August-7 September 1933 201-225 - 29 August-10 September 1935 322 5-6 30 October-8 November 1935 121 - 3-14 October 1941 121-198 - 12-23 October 1944 193 2 -4 11-20 September 1945 175-315 2-4 5-14 October 1946 129 5 9-1 6 October 1947 - - 18-25 September 1948 196 2 -6 3-1 5 October 1948 161 2 Easy 1-9 September 1950 117 - King 13-19 October 1950 196-241 2 Donna 29 August-I 3 September 1960 225-322 4 Cleo 20 August-3 September 1964 177-217 2 (est .) Betsy 27 August-1 2 September 1965 266 (gusts) 2-3 Alma 4-1 4 June 1966 201 - Inez 21 September-1 1 October 1966 2 1-150 2
Table 6
Reef seawater temperatures (OC) (Vaughan 191 8).
Reef Tortugas Sand Key Carysfort Fowey Rocks (1 879-1 907) (1 878-1 890) (1878-1899) (1 879-1 9 12) Month Range Mean Range Mean Range Mean Range Mean
January 19.4-24.8 22.1 17.9-24.1 21.8 18.2-25.9 22.5 15.8-26.2 22.2 February 18.7-24.4 22.1 18.3-25.3 22.8 20.6-24.8 23.0 15.6-24.5 22.5 March 19.6-25.4 22.8 20.4-27.3 23.8 21.1-25.6 23.1 18.7-27.4 22.9 April 17.9-25.8 23.6 22.5-28.5 26.1 22.4-26.4 23.9 21.0-29.4 24.4 May 2 1.9-28.3 25.6 25 3-29.9 28.2 24.0-27.9 25.5 21.4-29.2 26.2 June 23.6-29.3 27.2 27.7-32.2 29.8 25.4-29.4 28.8 21 -5-29.6 27.2 July 24.?-31.1 28.8 29 9-3 1.9 31.1 27.1-30'2 30.0 23.4-30.9 28.3 August 24.2-30.9 29.3 29.1-32.2 30.7 26.5-30.3 30.0 23.5-31.2 28.7 September 24.1-30.5 28.8 28.7-3 1.2 30.3 27.2-30.1 29.6 22.9-30.7 28.3 October 22.5-29.4 27.3 24.5-30.1 27.6 23.8-29.1 27.6 22.3-29.4 27.0 November 2 1.4-28 .O 25.3 22.3-27.1 25.1 23.0-28.7 25.5 20.0-28.4 25.0 December 21.3-26.4 23.2 18.4-26.0 22.5 21.0-27.3 23.3 15.8-27.9 23.1
9 2 20 MAXIMUM i- A 0 MEAN I 0 MINIMUM 12 1, , , , , , , , , , , , .~.,~.,..,,. JFMAMJ JASOND JFMAMJ JASOND
Figure 6. Monthly minimum, mean, and maximum Figure 9. Monthly minimum, mean, and maximum seawater temperature at Fowey Rocks, 1879-1912 seawater temperature at Dry Tortugas, 1879-1907 (Vaughan 19 18). (Vaughan 19 18).
28 - 26 - 2 24 - m -1 22 LU - 5 20- 18 - 0 CARYSFORT 0 DRY TORTUGAS 14 A FOWEY ROCKS 12, , r r . , , , , , , O MEAN JFMAMJJASOND Q MINIMUM Figure 10. Minimum monthly seawater temperature 10 , , , , , , . , , . , , JFMAMJJASOND at Carysfort Reef, Dry Tortugas, and Fowey Rocks. Figure 7. Monthly minimum, mean, and maximum seawater temperature at Carysfort Reef, 1878-1899 (Vaughan 19 18).
Table 7
Bottom (3 m) seawater temperature (OC) at Elkhorn Control Keef, Biscayne National Park, 1978 (from daily thermograph data, Biscayne National Park). Standard Month Range Mean deviation 40 - January m February 2 30- i March -il 0 April 4 E 20- May MAXIMUM June Ci MEAN July 0 MINIMUM August tO.~~.~.?~.stqSeptember JFMAMJ JASOND October Figure 8. Monthly minimum, mean, and maximum November seawater temperature at Sand Key, 1878-1890 December (Vaughan 19 18). water depth during spring low tides when shallow reef flats may be near emergent. During the summer, if wind speed is low, heating of the water column may cause hyperthermic conditions such that zooxanthellae (sym- biotic algae within the coral tissue) may be expelled 0MINIMUM (Jaap 1979; Hudson, in press). During the winter the 0 MEAN reef flat water may be hypothermic, again causing 16 D MAXIMUM thermal stress (Hudson et al. 1976; Hudson, in press). 121, , , , , , , , , , . , A trend of decreasing mean levels and ranges is J FMAMJJASOND noted as one approaches the Gulf of Mexico. Tidal cur- Figure I 1. Monthly minimum, mean, and maximum rents for the reef area are not presented in the Tide seawater temperature (3m) at Elkhorn Control Current Tables prepared by the National Oceanic and Reef, Biscayne National Park, 1978 (from thermo- Atmospheric Administration. There are significant tidal graph data, Biscayne National Park). currents between Florida Bay and the Atlantic.
2.6 SALINITY air masses (Roberts et al. 1982). Offshore (into the The reef tract area experiences oceanic salinities Atlantic) transport is the effect of tidal pumping, as summarized in Table 9. The region adjacent to Bis- northerly winds, and density gradients. Communities cayne Bay is an area where heavy rainfall can reduce adjacent to tidal channels suffer the greatest impact salinities for short periods. Dole and Chambers (191 8) from this thermal stress. Conversely, during the summer, reported that heavy precipitation in Miami almost the Florida Bay water may become hyperthermal always was followed by temporary reduction in chlorin- (> 31' C) because of solar heating, and communities ity and salinity at Fowey Rocks 24 hr later. Flat low near tidal passes are more affected. land, porous soils, and distance offsliore minimize the Seawater temperature reflects the annual climatic effect of rainfall on salinity in the reef tract areas. cycle: seawater temperatures are lowest from December Rarely, entrained Mississippi River spring runoff through March. Temperature peaks in August or early is carried along the inshore side of the Florida Current. September and cools throughout the fall and winter. This water mass has salinities of 32-34 ppt, which is A comparison of seawater temperature with air temper- within the tolerance limits of reef corals. ature (Tables 2, 6, and 7) for Miami with the recent Diurnal salinity fluctuations at Margot Fish Shoal Biscayne National Park data shows that mean seawater ranged from 37.8 to 37.3 ppt because of evaporation temperature is warmer than mean air temperature and precipitation (Jones 1963). throughout the year. During hot summer periods, density sinking on shallow reef areas is common. Evaporation creates dense 2.5 TIDES hypersaline surface layers whicP sink and mix poorly with subsurface cooler waters. Swimmers can feel and The Atlantic coast from Miami to Key West see this phenomenon. exhibits a semidiurnal tidal pattern. The area close to Key West is influenced by the Gulf of Mexico, which experiences semidaily or daily tides. Table 8 presents 2.7 DISSOLVED OXYGEN mean and spring tide ranges for several reefs. The major effect of tides on reef communities is the reduction of Jones (1963) and Jaap and Wheaton (1975) provide limited information on dissolved oxygen. 'I'he water column ranges diurnally from 90% to 125% Table 8 oxygen saturation. Daily maximum values zre attained between 1400 and 1600 hr (Jones 1963). Tidal ranges for several southeast Florlda reefs (NORA 198 1). 2.8 TURBIDITY
Transparency (equivalent Secchi disc depth) for Mean several stations off the Florida Keys was reported by Location tide level Mean tide Spring range Wilhams et al. (1 960). They reported thar the equivalent (m) (m) (m) Secchi distances ranged from 4.5 lo 35 rn in this area. Variabibty of water clarity is considerable. Following Fowey Rocks 0.4 0.7 G .? storms thc water may be nearly opaque. Plankton and Molasses Reef 0.3 0.7 0.8 suspended matter also affect water clarity. Alligator Reef 0.4 0.6 0.7 Water clarity and sedimentation rates for a Sand Key Reef 0.2 0.4 0.5 dredge operatxon near Basin Wills, Key Largo, were Garden Key, 0.2 0.3 Not gven reported by Griffm (1974). Ambient or background Dry Tortugas suspension concentration in the water column ranged 11 Sa':i~~iuzs11: the Florida reef tract and vicinity
Salinity Location Dates range fppt) Source
Dry Tortugas 1913 35.2 - 36.1 Dole 1914 Fowey Rocks 1914-1916 34.2 - 38.6 Dole and Chambers 19 18 Soldier Key 1945-1946 33.1 - 37.1 Smith et al. 1950 Key West area 1953-1954 33.3 - 37 .O Chew 1954 Margot Fish Shoal 1961-1963 36.8 - 37.3 Jones 1963
from (2.5 to 3.7 mg/l at a nearshore patch reef. Suspen- Bank) imply that no near-surface formations older than sion concentration within the dredge plume ranged from upper Miocene are extant. The Florida pennisula south 18 to 2 12 mg/l. These levels are probably comparable to of Lake Okeechobee has been a carbonate depositional the upper limits on the reef tract following a hurricane region during portions of the Mesozoic and throughout or major storm. the Cenozoic. Recent carbonate sediments are grossly similar to older material. Two major strata dominate the 2.9 CURRENTS southern Florida coastal areas from Miami southward along the keys. The Key Largo formation is a reef facies A warm water current (the Florida Current) formed during the Pleistocene and will be discussed in flows through the Straits of Florida. Wust (1924) more detail later. The Miami formation is also a Pleis- calculated that 26 m3/sec pass through the constricted tocene stratum that is composed of oolite and bryozoan area between Florida and Cuba. The current's velocity is skeletons. in the magnitude of 150 cm/sec. It is composed of two water masses in its surface layers. The eastern core is 2.1 1 GEOLOGIC HISTORY AND PROCESSES composed of Caribbean water that flows into the Gulf of Mexico through the Straits of Yucatan; the western During the past 20 million years, major change or nearshore portion of the current is composed of has occurred in western Atlantic coral reefs. Previously, water that flows from the Gulf of Mexico (Wennekens a cosmopolitan reef biota was found throughout the 1959). 'The Florida Current comes closest to Florida off tropics. Twenty million YBP, a land barrier emerged Palm Beach, where the central axis is about 15 km off terminating water movement between the Indian Ocean the coast. Off Dry Tortugas the current is 124 km south and the Mediterranean Sea. Approximately 7 million of the islands. The current is very dynamic and meanders YBP, the Central American Isthmus developed, separat- a good deal. Generation of eddies off the main body of ing the Caribbean Sea and Atlantic Ocean from the the current brings current waters onto the shelf and to Pacific Ocean. What was previously a circumequatorial the reef environments. As noted earlier, the current tropical zone had thus been isolated into separate moderates winter temperature and brings plankton of biogeographic provinces. During the late Oligocene and Caribbean origin into the reefs. Maul (1976) presented early Miocene, 20-30 million YBP, the western Atlantic the variability of various components of the Gulf Stream area experienced its greatest proliferation of reef build- system (Figure 12). Schomer and Drew (1982) recently ing. During the Pleistocene (at least 1 million YBP), revicwed the physical and chemical environment of major environmental change extirpated the Pan Atlantic- southern Florida. Pacific biota. Glacial periods occurred during this time; during each period, sea level was drastically reduced and 2.10 GEOLOGICAL SETTING the marine climate was cooler. Pacific genera of Sclerac- tinia that were eliminated from the western Atlantic The following information is paraphrased from a include Stylophora, Pocillopora, Goniastrea, Goniopora, field gmde to south Florida sediments (Ginsburg 1972a). Pavona, and Seriatopora (Newell 197 1). The Florida-Bahamas region is a carbonate platform part In Florida, a major coral reef community devel- of the Atlantic and Gulf coastal province. The platform oped during the last major interglacial period, San- is dissected by deep-water channels. The most important gamon, 100,000-112,000 YBP; it was killed during colrsideratlon here is the Straits of Elor~da.wkch E the last glacial advance (Wisconsin) because of the nearly 550 m deep. Marine portions of this platform are reduction in sea level and the cooler climate. This generally shallow, less than 16 m deep in most places. Pleistocene reef, known as the Key Largo formation, Deep test drilling in the area indicates the platform has extends from Miami Beach to at least Dry Tortugas and had a history of continuous subsidence and deposi- seaward to the Straits of Florida. It varies in thickness tion of shallow-water carbonates anri evaporites. Maxi- from 23 to 61 m or more; the basement has not been mum depths of d~3penetration (5,486 m at Cay Sal reached in several cores. Wherever its base has been lo- catcd, the farmation was found to rest atop calcareous- ranged from 0.65 to 4.85 1ni1,000 years (.Fable 10; quartz sands. From near Mlarnl to Big Plne Key the Key %inn et al. 1977). For comparison, Adey (1977) Largo formation n found near the surface, but from Ulg reported reef growth off St. Croix, U. S. Virgn Islands, Pmc Key to Key West is overlain by the Mlarnt Oollte was I5 rn/ 1,000 years (this is the upper lirnit for Carib- formation, whrcll may be upwards of 12 m thick (lioff- bean reef growth). Shinn et al. (1 977) reported that the rllclster and Multer 1968). ffoffrnelster and Multer base of recent reefs was I1leistocene Key Largo Reef, (1968) and fioffnlelster (1W4) reported that the ex- fossil mangrove peat, and cross-bedded quartz fossil sand poseci portion of the Key Largo formallon in the Elonda dunes. Keys 1s represcntatlve of a low-wave-energy patch-reef More recently, an extensive harrier reef existed cornmunlry. Ille rrialn cvldence for thlis 1s that certain off the Ft. Lauderdale area, but it was extirpated about sclcract~nian corals, in particular ilcru[>ora pnltnata 7,000 YBP (Lighty 1977; Lighty et al. 1978). This reef (elkhorn coral), are absent in the fossil record. Presum- was a shallow-water Acropora palrnata community. The ably a series of events occurred such that the seaward demise of this reef was attributed to environmental part of thc Kcy Largo formation did, at one time, change caused by increasing sea level (Lighty et al. posses high wavc-cnergy corrl~nun~t~esaitriilar to today's 1978). Recent coral reef growth off the Florida Keys reefs; however, as sea lcvel rose tiuring the Ilolocene started from 5,000 to 7,000 years ago. transftrcsuon, these fossll cornmunltles were appar- 1)evelopment of a coral reef integrates biological, ently eroded away hy wave action. Iloffmc~ster(1974) geological, chemical, and physical processes. Soon after repurteci that a core made neur Looc Key Reef recovered the first coral colonies settle and start to grow, the ira~mentsot ti pu!rrlar~l froirt I8 m hclow the surface. breakdown of organism skcletons by biological and lhc ilolo~enetran~gression of sea lcvel (kigurcs plzysical agents occurs. Sediments crcated by tftese 1 J und 14) tnd~catcsthat I0,OOO years ago the sea level activities become a part of the reef. was &bout 30 rtt lowcr thitn today (Ligl~tyet a!. 1982). Finer sediments filter into voids and borings, and Stlinrl tat 81 (1977) cored several recent reefs frorn Mlamt the coarser fractions fill the interstitial space between tleaclt to Ilry l ortupas arrd dated 1n1Cial g~owl h frorn the reef framework. Reef tract sediments are carbonate 5,250 to 7.X0O YUI' Rccf growth ctr accrctlor~r;rtcs and are do~t~inatedby algal and coral skeletal material
EUSTATlC CHANGES IN SEA LEVEL YEARS x103
2 4 6 8 10 12 14 16 18 20 22 24
ENVELOPE ENCLOSES
CURRAY & SHEPAHI) - FAlRBRiOGE 1960
MlLLlMAN & EMERY
URRAY 1965
Figure 13. Sea-level change during tlle 1.loIocene Period (Gmsborg, ('o~npsrativeSed~~rtentology Laboratory, Ullrvcrsrty of Mumi, Florida ).
14 BURIED REEF
PLEISTOCENE SURFACE
PLEISTOCENE SURFACE
INCIPIENT REEF
Figure 14. Sea-level change in a Bahamian shelf reef system during the last 10,600 years (Hine and Neuman 1977).
Table 10
Age and growth rate of Recent Florida reefs (Shinn et al. 1977; Shinn 1980).
Base age (Y BP) Accretion Growth rate Reef (with confidence limits) (m) (m/1,000 yr)
Long Key 5,630k120 5 .O 0.65 Cary sfort 5,250B5 7.3 0.864.85 Grecian Rocks 5,950k100 9.5 6-8 Bahia Honda 7,160335 4.6-8.2 1.14 Looe Key 6,58030 7.3 1.12 Bird Key 6,01730 13.7 1.36-4.85
(Ginsburg 1956). Halimeda and other codiacean algal et al. concluded that cntstose coralhe algae and stony plates are the most common algal skeletal material. The corals (nine spec~es)annually fixed 163 metric tons of sedimentary material becomes incorporated into the reef CaC03 on a reef platform with a surface area of 10,800 framework through the process of being bound to the m2 (9 kg CaC?o3/m2/year). For comparison, nonreef platform by crustose coralline algae and through the depositional marine environmenrs in southeastern Hor- geochemical processes of in situ cementation by high ida are reported to produce 0.25-1 kg ~a~~~/rn'/~ear magnesium calcite cements. Ginsburg and Schroeder (Stockman et al. 1967; Moore 1972). (1973), among others, detailed the processes of marine Ghiold and Enos (1982) studied CaC03 produc- cements in coral reefs. tion in the brain coral Diplorza labnvri?zfhifurmts from Steam et al. (1977) presented the most recent several patch reefs in John Pennekamp Coral Reef State quantitative budget of calcium carbonate (CaCO,) Park (JPCRSPJ-Key Largo National Marine Sanctuary within a coral reef ecosystem (F~gure15). This was (KLNMS). Production (CaCO,) was I I .8W.3 kg based on the study of a Barbados fringing reef. While the ca~O~/rn~/yearof area occupied by the colony. Porites magnitude of individual components may differ, the astreoldes annual CaC03 production ranged from concepts are valid and applicable to Florida reefs. Stearn 13.63.5 to 14.023.1 kgjm2/year at Middle Sambo Reef I5 klgurc 15. ('aicium carbonate flow model bascci on a Barbados fringing reef (Stearn et al. 1077).
near Key West (Kissling 1977). hiontastraea aruiuhr~s years, which is within the magnitude of Holocene reef has derrior~stratedan~lual CaCOB production of 20 kg growth in Florida (Shinn et al. 1977). ~'a('~)~/fi~~/~uarin Barbados (Stearn et al. 1977). Frost et al. (1977) and Taylor (1977) contain C;hiold ant1 Kilos (1982) extrapolated production inany papers about coral reef geology. values to vcrtical reef accretion rates of 2.2 in/1,000 CHAPTER 3
CORAL REEF COMMUNITY TYPES
3.1 INTRODUCTION by octocorals and a sparse number of stony (hard) corals (lable 11) The llve bottorn conl~nunitynlay vary in size A biotic community as defined by E.1'. Odu~n frorrl a sinall drea of tens of square meters to one of (197 1) is any assemblage of populations living in a pre- several hundred square meters. The stony corals most scribed area of physical habitat. Four discernible coral co~nn~onlyfound m the nearshore associations Include community types occur off southeast florida.' Gener- S~dcrustt-curatfluns, Por~respontes, Y. asfreozdes, Manl- ally in a seaward progression are found live bottoms. crna urt~olata, Solt7tzasrrea Ilycrdes, and S bourn on^. patch reefs, transitional reefs, and bank reefs. 'I'he Along with these specles, I)rplc)rra ckvusa, Mrllepora relief (height above bottom) of these communities and alt rt orlais, and D~chococ~tirustellurrs are conimonly the dominance of stony corals as a structural element found in deeper coniimun~t~esI'he region surrounding increase in a similar progression. All of these communi- the hve bottorn coxnmutut~es is sedlmentary, seagrass, ties can be physically characterized as shallow water, rock. or sponge Lhe seagrass comxnunity 1s treated m wave-resistant, three-dimensional carbonate accretions detall In Zlen~an(1982). constructed by limestone-secreting organisms (prin- Stony coral species are commonIy found m the cipally corals, algae, and bryazoans) on a pre-existing seagrass and sedmlentary environments adjacent to the hard substrate. This basic structural component is reef communltles. These are nfa~zlclnaartwlata, E'or~tes augmented by other community members, sedentary porlter, C'ladocoru at buscuiu, and Sidf~rastrearadrans, and mobile, permanent and transient. hardy tolerant species thdt do not attaln great size.
3.2 LIVE BOTTOM COMMUNITY 3.3 PATCH REEF COMMUNITY The live bottom cominunity, also known as hardground, is generally found closest to shore, e.g., in Patch reefs (Plates I la, 1 lb, and 12a) are the tidal passes, under bridges, and short distances seaward second major type of coral community, and they are a of the intertidal zone. It usually occupies exposed fossil most conspicuous element in this region. 'Their distri- reef formations, limestone, and other rocky substrates. bution is n~ostlyseaward of Key Largo and Elliott Key; The faunal and floral elements are not consistent, and however, they also occur off Big Pine Key, Key West, the assemblage is usually visually dominated by octo- and Dry Tortugas. They are usually found seaward of corals, algae, sponges, and smaller hardy stony coral Hawk Channel, but a few very nearshore patch reefs species. exist. Two examples of nearshore patch reefs are (1) an These co~nmunitiesdo not actively accrcte or assemblage of patch reefs just to the north of Caesar's build massive structures. 'I'hey support diverse inverte- Creek (BNP) and (2) a small reef just off Cow Key brate and vertebrate communities arid provide an impor- Channel, Boca Chlca Key. Both of these reefs present tant nursery area for cornmcrcial and sport harvested the general impression of survival under marginal con- species. Live bottom habitats are scattered from St. dltlons. Lucie inlet southward to Dry 'Iortugas in depths ranging A patch reef characteristically has upward of 3 m from less than 1 n1 to beyond 30 m. While these descrip- of relief and is dome-shaped. The surrounding bottom tions may imply a single comrnunity type, the nearshore may be sedlmentary, seagrass (Plate l2h), or rock, Most and offshore types differ greatly in species composition. patch reefs off southeast Florida are found in 2- to 9-m In either case the octocorals (soft corals) often dominate depths; their upper surface may be nearly emergent at in terms of numerical abundance and density. An low tide, exeniplified by Basin Nlll Shoals off Key example of an offshore live bottom conlmunity is Largo. Patch reefs are roughly circular in outline and the reef (Schooner Reef) at number 2 buoy in Biscayne vary in slze from about 30-700 m In diameter. Because National Park (BNP, Figure 16) off Elliott Key. It exhib- of different ages of the nuxnerous patch reefs and their its little relief with the exception of a srllall ballast pile local environmental conditions, generalizations fail to from an old shipwreck on the north side off the reef. adequately characterize them. Jones (1 963, 1957) The platform is li~nestonewith small ledges, solution presented information about their physical and chemical holes, and pockets of sediments. It is heavily colonized environment near Margot Fish Shoal off 5,lliott Key and the dynamics of developmental stages. Smith and Tyler (1 975) and Jones f 1977) presented the view that the patch reef cornrnunity is slrnilar to a super organism, in 'Darwin (1 842), based on his experiences m the Pacific that it goes through severaI Efc stages. The nature of the and Indian Oceans, defined three types of coral reefs: community reflects the reef's age. While the coral fringing, barrier, and atoll. Although many attempts community goes through its cycle af development, the have been made to extrapolate these forins to Atlantic fish assemblages change according to nlche availability. coral reefs, Darwinian-defined reefs do not presently An assu~neddevelopmental sketch of a patch reef would occur In Florida. include the Eoflowmg: 17 Figure 16. Coral reefs in Eiscayne National Park.
18 Table 11
Live bottom corals from Schooner Reef, Biscayne National Park (four 1-m2 quadrants; Jaap and Wheaton 1977 MS.).
Percent Mean 0.1.b No. of of total no, of Species Typea (ranking) Frequency colonies colonies colonies/m2
Plcxaura lzonzotn alla 0 73 4 18 9.63 4.50 f'orites porites S 73 4 18 9.63 4.50 Gorgonla verttalina 0 70 4 14 7.49 3.50 Density Pseudopterogorgia arnericarza 0 7 0 4 14 7.49 3.50 Rrlareuin asbestinum 0 67 4 10 5.35 2.50 Mean colonies/m2 = 45.75 b'unzcea tournefortz 0 65 4 8 4.28 2 .OO Standard deviation = 1 5.46 Muricecz atlai?tica 0 6 5 4 8 4.28 2 .OO Range = 25-61 Pseuilopfexauru porosa 0 63 4 6 3.21 1.50 Plexaurella f usi,feru 0 62 4 5 2.67 1.25 Eunicea succlnea 0 5 8 3 25 13.37 6.25 Mean species/m2 = 18.50 Plexa uru jlexuosa 0 5 1 3 11 5.88 2.75 Standard deviation = 4.53 Murtceop~lsIluvzda 0 5 1 3 10 5.35 2.50 Range = 13-21 Porires astreoides S 47 3 6 3.21 1.50 Pl~xuurella~lichororna 0 47 3 3 1.60 0.75 Millep91-aa/c~cortzis S 34 2 7 3.74 1.75 Diversity indicesc - Pseudop/emura flagellosa 0 3 2 2 4 2.14 1 .OO Euttlcea lacinzuta 0 3 2 2 2 1.07 0.50 H ' = 4.27 Agur~czrrugerzcifes S 3 2 2 2 1.07 0.50 H'max. = 4.91 Pl(axaurellagrnsea 0 3 1 2 2 1.07 0.50 J' = 0.87 Eutllcea nzamn~osa 0 3 0 2 2 1.07 0.50 Pseudoplexuura wagenaarr 0 29 2 2 1.07 0.50 Drchocoertiu stellar~s S 16 I 2 1.07 0.50 Pseudopterugorgia acerosa 0 15 1 1 0.53 0.25 Eunicca calyculata 0 15 1 1 0.53 0.25 Eut?icea fusca 0 15 1 1 0.53 0.25 f:avla fraguin S 15 1 1 0.53 0.25 Ps'srudupterogargia kallos 0 14 1 1 0.53 0.25 P~eudopterc~gurgzablpinnata 0 14 1 1 0.53 0.25 S1dc7rasr1-encrderea S 14 1 1 0.53 0.25 Mon tastrueci caveriiosa S 14 1 -1 0.53 0.25 Total 187
a~ = octocoral, S = stony cord. b~.I. = Biological Index, McCloskey (1970). '~iversit~computed with log:,. (1) Initial pioneering settlement of coral larvae have a less diverse, nearly monospecific fauna. Patch On appropriate substrates. rhese specles might include reefs exhibit extreme variability in coral abundance, Porites porites, Manlcitia areolata, and Favla fragurn. density, and diversity (Table 14). Macrobenthic algae They would be preparatory colonizers which would constituted from 0% to 7% of the linear biomass, sponge eventually die and their skeletons would become the 0% to 3.6%. Octocorals constituted a numerically domi- hard substrate that the major framework corals would nant element; however, their linear biomass and macro- settle on. If the bottom were rocky, this stage might be habitat potential are less than many of the stony corals, unnecessary. especially the massive framework builders. Octocorals do (2) The settlement and growth, both upward and occupy much of the space on the mature older patch outward, of the primary framework builders. While reefs. There appears to be a competitive exclusion by other coral species are conimon in the patch reef com- octocorals in the interior of older patch reefs. A repre- munity, Siderusrreu siderea, Mvntastraea annularis, sentative patch reef is the reef at buoy number 4 (Dome Diploria strigosa, D. kibyrinthiff~rmis,and Colpophyllia Reef) in BNP 16. It is about 250 rn in diameter. The nafans build the massive frame or three-dimensional greatest relief is about 2 m and is found on the north- structure of most reefs. Shortly after the framework west side. Tables 12a and b and 13a and b present element has settled, the boring and rasping fauna starts the octocoral and stony corals found in this reef based the production of reef sediment. Sediments fill inter- on several transects. Reef organic linear biomass based stitial space and arc incorporated into the reef frame. In on line transect surveys from four patch reefs in Bis- time, some coral deaths occur providing space for cayne National Park shows that mean stony coral cover settlement of coral larvae, an essential element for was 25.7210.6% in the stony coral-dominated zones growth of the reef. Tlze newly developing reef 1s a focal (Table 14). In' the interior octocoral-dominated zones point for attracting the diverse flora and fauna conlrnon the organic linear biomass for stony corals was 14.3% to the coral reefs. Some of this occurs because of larval 29.7% (Table 14). There is usually a halo of barren sand set tlement, especially for tile sessile specles. Mobile and reef talus (rubble) with sparse grasses and algae sur- knvertebratcs and vertebrates (sorne residents, some rounding the periphery of most patch reefs (Plates 1 la temporary refugees) niovc to the reef as niches favorable and b). The rubble, composed mostly of dead coral to tlieir requlrernents beco~rrcavailable (I'lates 13a and colonies that have been swept away from the reef by b). storms, is potential substrate for colonization. Outward (3) I'hc maturing stage. I)uring this stagc the rcef reef expansion is presumably dependent on the reef talus .grows upward and outward, providlrlg surface area for in sedimentary environnlents for substrate creation. secondary colonizers. 'l'he major franlework builders tialos around some patch reefs resulted from the black attam 2 m in diarncter and greater. I'he boring and sea urchin (Diadema antillarurn) feeding nocturnally on rasping fauna excavate considerable material from the algae and seagrasses surrounding the reef, according to i~asalstrrfaccs ctf the corals, creating a labyrinth of cavas Sa~nmarco(1972), Ogden et al. (19731, and Sammarco that become occupied by a crypt~cbiota In t~me,these et al. (19'74). Randall (1965), however, reported that caves and tunnels ;ire enlarged, and larger f~shand herbivorous reef fish were responsible for patch reef irlvcrtcbratcs lake refuge in thcrn. I he new rilches within halos. In either case, the herbivorous consumers graze thu interior of the reef are colonrzed by a wide taxo- away the flora adjacent to the reef, creating a barren r-ionuc spcctrunr of shade-loving organisms. zone or halo. (4) 'i'lic fully mature stage. Because the prltnary I'atch reefs are important habitats for many reef framework-buildt~lg corals have approached sea level, fish, permanent and transient (I'late 13a). They provide upward growth IS limited. 'l'hc undersurface of thcse shelter, food, and breeding ground for mobile fauna. The corals is cavernous, and in tune the framework bccorncs spiny lobster (I'anulir~csargus) (Plate 19b) utilizes patch so weakcnCtl that the upper surface may collapse ulward reef habitats during part of its life history. forming a n~bblt.or houlderhkc su~facc.'1 his 1s usually Like most reefs, patch reefs show a temporal irregularly flat and, 111 many cases, 1s dominated by change when storms or temperature extremes disturb the octocorals. iiaystackhhe colonies of ,tf. or~tzulur~rappcar communities. S~nallercoral colonies are dislodged and I:, lzavr large dead areas that are colonized by other transported from their growth positions. If the new spccics (Millcporu ulctcorrro and Gorgvtiiu ~*c,rzralltlu). position is favorable, they may continue to grow; if not, Ifthe recf cornpietcly collapscs from a major storm or they may die. This is especially true for octocorals and vessel. grormtiing, the former rriajor relief patch rcef encrusting colonies of Mil!epora that are on unstable becomes a low-rehef plie of rubble that is usually doml- substrates, principally octocoral axes. These colonies nated by octcrcorals and nonframework building stony may be swept conlpletely off the reef by heavy wave corals. In this growth and decay of the patch reef surge. There appears to be high mortality due to this cornrnrrnity rntu the live bottorn cominunity, geo- stress; if so, intense recruitment usually prevents large chemical processes play an important role 1n ce~rienting open areas. and binding sedin~cnts. The association of coral found on any particular 3.4 TRANSITIONAL REEF COMMUNITY ptch reef LS most probably governed by random chance. Same patch reefs have very diverse stony coral faunas The tern1 transitional reef is used to describe while otlrzr reefs, only a few meters or kilometers away, those reefs that have the rudiments of bank reefs (see Table 12a
Octocorals at Dome Reef (two 20-m transects, 1977; Wheaton, in preparation a).
Percent B.La No. of of total Species (ranking) Frequency colonies colonies Plexaura flexuosa 3 9 2 4 1 19.33 Plexaura humomaIIa 3 6 2 2 9 13.67 Pseudoplexaura porosa 3 6 2 2 8 13.20 Pseudopterogorgia acerosa 34 2 29 13.67 Pseudoplexaura flagellosa 3 1 2 12 5.66 Gorgonia ventalina 3 1 2 12 5.66 Pseudopterogorgia americana 3 0 2 17 8.01 Briareum asbestinurn 29 2 10 4.7 1 Plexaurella fusifera 26 2 7 3.30 Eunicea tourneforti 25 2 6 2.83 Plexaurella nutans 22 2 2 0.94 Eunicea fusca 15 1 4 1.88 Eunicea succinea 14 1 3 1.41 Muricea atlantica 12 1 4 1.88 Eunicea laciniata 12 1 4 1.88 Eunicea calyculata 10 1 1 0.47 Plexaurella grisea 10 1 1 0.47 Muricea elongata 10 1 1 0.47 Eunicea marnrnosa 10 1 1 0.47 Total 212 a B.I. = Biological Index, McCloskey (1970).
Table 12b
Octocorals at Dome Control Reef (two 20-m transects, 1977; Wheaton, in preparation a).
Percent 3.1.~ No. of of total Species (ranking) Frequency colonies colonies
Plexaura homomalla 40 2 49 19.76 f'seudoplexaura porosa 36 2 3 0 12.10 Plexaura flexuosa 35 2 32 12.90 Pseudoprerogorgia americana 35 2 26 10.48 Pseudopterogorgia acerosa 34 2 2 5 10.08 Gorgonia ventalina 27 2 10 4.03 Briareum asbestinum 26 2 12 4.84 Eunicea tourneforti 26 2 9 3.63 Pseudoplexaura flagellosa 26 2 12 4.84 Eunicea calyculata 26 2 12 4.84 Eunicea succinea 2 2 2 4 1.61 Plexaurella fusifera 2 2 2 4 1.61 Plexaurella grisea 22 2 4 1.61 Muricea elongata 22 2 4 I .61 Eunicea fusca 2 1 2 3 1.21 Muricea atlantzca 2 1 2 3 1.21 Muriceopszs flavzda 14 1 5 2.02 Eunicea laciniata 11 1 2 0.81 Plexaurella nutans 10 1 1 0.40 Eunicea clavigera I0 1 -1 0.40 a~.~.= Biological Index, McClaskey (1970). 2 1 S 5.0 EE'E L8'0 = ,I SS'O EE'E ZE'E = .xew ,H Z6'B L9'9
88'2 = IH EL'B 19'9 EL'Z L9'9 ,sa3~ptq1Cps~aya EL'Z 19'9 09'OE EE'E I E- 11 = uo~$e~nappzepue~s OZ.8 00'0 I 00's 1 = .ue.r)/sa!uo103 ma^ 61'2 19'9 8B' I 1 EE.E I d$!suaa 69. I E EE'EE
~aao3 (w3) saruo1os sapoios huanbasd (%u!yuel) sarnadg ~elolJO la~ol) TEJolJo $0 'oh; q'I'B ~ua~ad ~ua3rad C1 -- r-1 .(paqsgqndun 'deer : ~~61'S33asuell ut-SZ OM$)&aaX folluo3auroa je sIeros Auo)S ~EIa1qe.L
- - 99'0 = ,I 2110'0 S I 9S'Z f 1 6 I aronlt,> a~io~old?~ OZ'Z = -xew ,H LEOO'O S 9s-Z I I 6 1 ur~~UAJ' uz't nd OZ'Z = I H LEOO'O S 9S'Z I I 6 t: sapjoayso Y,>-IT~O~ L8b0-0 S.9 9S'Z I 1 6 1 x C~AJ~~i~~.itjcfoj~n.ijg sa3:puj Al!siah!a St00.0 0 1 9s'Z f I 6 7 sriu[lafsnzitaooo yn~ 4 ELZI'O OLI 9s'; 1 I 6 l rzrm nir nzll.iydoii,w:) ;['r = uo!le!nap plepuels Zl I0.0 S I El's z z 9E salldnd ~,JPJ(,~ 0s-61 = 'sue1~/sayuolo3uva~ SZZO'O OE If'S ?r Z 9F vanap!s aanls~uap~~ 690 1'0 OBI I S'OZ P Z 8E jlriro313yn u~oda~lzjq Apsuaa 2659'0 088 _CgVES I Z Z Ob siraltlrruo n~nn~sn~uojg .ran05 (m~) sa~uofor, samo[os huanbarg (2ur~ua~) sar3adg ~eloz30 "ahon ~elo~$0 30 -0% e 18 lua3~ad $tra3lad .fpaysgqndun 'deer :LL6[ 'smasue13 ur-sz ornl) jaax auaa $e sp.703 A Attributes of stony coral associations, Biscayne National Park, based on several hetransects at each reef (8 reefs, 18 transects) (Jaap, unpublished). Stony coral associations Transitional Attributes bank reefs Patch reefs Livebottom E2,3;EC 1,2,3: St 1,2;StC l;D 1,2; E 1; Sc 1,2; ScC 1,2; 5 transects DC 1: 6 transects StC 2; DC 2: 7 transects Dominant species Acropora cervicornis Montastraea annularis Porites porites Acropora palmata Millepora alcicornis Porites astreoides % of colonies 48.2 Unique species Millepora complanata Stephanocoenia michelinii, Diploria lahyrinthiformis Siderastrea radians Colpophyllia natans, Mycetophyllia ferox, Mycetophyllia Iamarckiana No. of species No, of colonies Diversity indices b H 'n H 'c H' rnax. J 'n 0.78 0.72 0.75 J 'c 0.72 0.54 0.76 % cover (5 f SD) Algae 0.3 rf: 0.3 1.7 k2.7 2.0 rf: 2.5 Sponge 0.7 f0.8 1.1 5 1.4 2.7 5 3.3 Octocoral 25.2 k 12.8 23.2 f 11.4 38.3 k 15.3 Stony coral 28.9 k 15.1 25.7 f 10.6 14.3 rf: 9.7 Abiotic substrate (sand, + rubble + 46.7 k 9.3 48.4 f 12.1 42 k19.0 rock) a~:Elkhorn Reef D: Dome Reef Sc: Schooner Reef St: Star Reef C: Control Reef, e.g., StC; Star Control Reef. 'Hh, J'n computed by abundance. H'c, J'c computed by cover. 2 3 Section 3.5) but are not as fully developed. They can steel ship sank during U'WII and has numerous corals and be thought of as ernbryonic bank reefs, or a series of other sessile growth. The fish fauna is smilar to those coalesced patch reefs, or welldeveloped hardgrounds of nearby natural reefs. The "French wreck" is a similar with relief. The marginal bank reefs found inshore of phenomenon off Loggerhead Key, Dry Tortugas. Arti- i'acific Reef and continuing south to Pennekamp Park ficial reef substrate is also provided by ballast stone or boundary in BNP are examples of the embryonic bank large blocks of igneous rock that were lost in shipwrecks. reefs. 'rhey possess a well-developed reef flat with some These are also colonized by reef organisms, notably trend of spur development by Acropora palmata on the hlillepora sp. A number of large barges and ships have seawarti fringe. 'The reef at number 1 buoy (Elkhorn been purposely scuttled in waters of 30 m and greater as Reef) at BNI) (Piate 18a) exe~nphfiesthese reefs. fish havens; these also develop natural reef growth. ElkIlor11 Reef is about 500 m long in its northisouth axls and some 200 ni long in its eastlwest axis. The reef 3.5 MAJOR BANK REEF COMMUNITIES flat is a rocky platform with undercut solution holes and ledges and sedimentary environments seaward and Bank reefs are typically elongated and form a landward. Off the northeast reef flat there are several narrow, linear, discont~nuousarc from Miami south and large haystack colonles of ilfontastraca annuluris (Plate west along the Keys to the Dry Tortugas. They are 5b), which are exiensively excavated on the under- located near the abrupt change in bottom slope, which surface. Ttteir upper surfaces are dead, in some cases marks the seaward edge of the Floridan Plateau and they stipporlii~g tlic sea fan, Ciorgonia vvetltulina. The area occur mostly between the 5- and 10-m depth contours. seaward of the rcef flat platform is visually domi- The distinctive features of these reefs are the nated by oelocorats and occasional large clusters of occurence of Acropora paltnara, the coral zonation by f'rjriter pontrs. There is a slight slope to the reef flat that depth (Table 18 and Figure 181, and the seaward spur rn sotnr places has dense clusters of Acroparu palrnatu, and groove formation. cfkhortl coral. A stirnmary of coral species, abundance, Major bank reefs vary in morphology and species and density is presented in Tables lSa, 1 Sb, 16a, and composition. Some, such as Long and Ajax Reefs I bh. (Figure 16 and Table 17), off EUiott Key, are sparsely Acro~~~~rtrpaitnata colonies on the reef flat deveIoped in thelr shallow zones, but below 12 m there interface are situated with major branches oriented into is growth on antecedent reef platforms. Better developed prevailing seas, east/southcast. On the rcef flat small, bank reefs exhibit a zonation which varies from reef to lrlorc utiifor~nlyoricntcd colorlies in less dense aggrega- reef. Reef age, relative position, hydrodynamics, and lions art. conirmon. They arc associa.ted wilh colonies of underlying topography are the major influences on hf~llcprtractrrrrplartclfo, Gorgonla vcntalctra, and Diploria differences in reef morphology and zonation. The ahvosa. l'ilrs cclral otten propagates or spreads by vege- general pattern is presented in 'Table 18. Specific details tative "fragments'" (Plate lob). Broker1 fragments for individual reefs are given below. scvtLle In the nearby boeeo~a and in a short time are rollilly ;ittacltotl. 'lhey grow upward Eorining new eolo- Carysfort Reef ntes. I'hc ctustercii cistribution of this species is proba- bly relstcd to lltc tact that fragtnents are not trdnsported Carysfort Reef (Figure 17) is the northernmost great distances frorrl pascnt colonies. Mergner (1977) reef in the KLNMS system. A large lighthouse is near rcpurlcd that hf. corrirtlrinutu (Plate 3b) is an indicator of the north end of the reef flat. The reef flat has an ticavy wavc surge. It is cotnniarm in the reef flats and spur extensive concentration of Acropora palrnatu on its anti grox)ve lract arcas wilere wavc surge is greatest. seaward flank. The spur and groove formations are in an ,~(.i.i~po~rrc-iLrvrc'orrrr.c (Plate 4b), the other colnmon early stage of development. Seaward portions of the reef ilorly coriri. occur:, ul small patches. It 1s not firrllly flat grade into a nearly barren area that separates the A. attach44 to thr Zwtton~and is rnoved about a good deal pal~rzatw colonies from the buttress zone. The buttress ciur~ni: slorrnr I+urrfc.s asireot~ic~~and P. pontes are also zone has extensive areas of low relief residual spurs and terY corilrrlclrr on the reef flat. 'I'lie former is firmly grooves that have coral develop~nent. Haystack size ;~ttJtll~d,ti.hilr* the latter 1s $lot. colonies of hlontastraea a~zrzularu and fields of A. 011 OCC~S~CI~,CIislo~ipcd it. CC~~VICOJII~Scolonies cervlcornis are common. The reef platform terminates at (01131 zii co111acf wit11 -4. p(11ttiufu colonies, a gaIl or fuse about 27 m. Table 19 presents density and abundance Is foflklcd i?jacttoll r~aCtlOnwilere the t~suesof the lnforrnatron for the stony corals in the 14- to 15-m t'olonlrs krrcet. l he tapering cylindrical bnnchcs of depth range. A broad expanse of rock separates the *1 L.cr~!I'(v?~i~~grot% upward entwlnerl wlfh the flat reef platform proper from hard-substrate deep reef frondose hranttles (sf 1. pahrrntu .-I pairneru overgrows communities. These deeper communities were described L'~'~~lc.of-trr.\w11e.n they are in contact (J. C. l,ang, by- Jaap (l98lf arid Wheaton (1981). Outlying com- I!nlversity uf .Iexa$, Aushai; personal observation). inunities continue to a depth of about 41 m. 'The stony -Anotfier type of transitional reef develops on coral composition is similar to that of the deep reef; aftifkid substrates found tlrroughout the reef tract. Agaricla larnarck~,A. fragilis, and Nehoseris cucullata are 'ffWattract reef' biota and in titile become very recake. the dominant species on vertical faces, while Szderastrea 'fhey include shipwxecks and other mater~als,A good siderea, rM. cavernosa, and Madracis rnirabilis are com- Cxa~n~Ieis the Bcrrctuual wreck in KLNMS. large mon on the horizontal surfaces. 2 4 Table 15a Elkhorn Reef stony coral fauna (four 4-rn2 plots sampled per reef, 1978; Jaap, unpublished). Percent B.I.~ No. of of total Mean no. Species (ranking) Frequency colonies colonies colonies/m2 Porites astreoides 315 16 60 54.05 3.7500 Density Acroporu cervicornzs Diploria clivnsu Mean coloniesjm2 = 6.94 Porites porites Standard deviation = 2.35 Acropora palrnata Range = 3-1 2 Siderastrea siderea Miliepora alczcornis Diversity indices Palythoa sp. :Ilillepora complunata Mean species/m2 = 3.3 1 Favia fragutn Standard deviation = I .54 Diploria srrigosa Range = 1-6 Dichclcoeniu stellnris fl' = 2.99 'Total H'max. = 3.59 J' = 0.64 ".I. = Biological Index, McCloskey (1970). Table 15b Elkhorn Control Reef stony coral fauna (four 4-m2 plots sampled per reef, 1978; Jaap, unpublished). Percent B.I.~ No. of of total Mean no. Speci~i (ranking) Frequency colonies colonies colonies/m2 I'OPI~CSa~frcoldes 260 14 6 8 59.65 4.2500 Density ~llilleporaalcicori~ts 146 8 9 7.89 0.5625 Drplorzu clrvosa 115 6 7 6.14 0.4375 Mean colonies/m2 = 7.13 Acropora cervicurnis 96 5 6 5.26 0.3750 Standard deviation = 2.35 Pontes porifcs 94 5 7 6.14 0.4375 Range = 3-9 Agarzcln agarzclles 76 4 6 5.26 0.3750 Sidevustreu sicfereu 5 6 3 5 4.39 0.3 125 Mean species/m2 = 3.19 t'rrlyf hoct sp. 38 2 2 1.75 0.1250 Standard deviation = 1.22 D~ckocoeniu$fellarls 20 1 1 0.88 0.6250 Range = 1-5 Mlll~poracompianntu 19 1 1 0.88 0.6250 Poritt~sbrawneri 19 1 1 0.88 0.6250 Diversity indices IGvzu f ragurrt 19 1 1 0.88 0.6250 - H' = 2.22 Totaf. 114 N' max. = 3.58 J' = 0.62 P aJ3,1. = Biological index, IvicCloskey (1970). Table 16a Elkhorn Reef octocoral fauna (three 20-m transects, 1977; Whealon, in preparation b). Percent No. of of total Specics B.I.~ Frequency colonies colonies C;org:otjta vc>ntuhnu Eurucea SUCC~~CU Pk*xuuru jlexuusa kcudoptcro~orgiuarn encunu Pkxuura t~orntrtrtullu Muricea utluntica f'6~cudoplcxauroporosu d"seudoplc#uuurcr jlugtrilcrsu Euriiccw tournefi~rri iJ!~*xuurellaju~ifercl Pscuclopfrrogor~~uuctroJu f:utriccu 1ur.r nicrft~ Iisrudr>rttcrogurgia kullos Af uriccopsu Jluvldu iYrrnicbtlncnl.vru kara ITtexanrt~lludlr'iro tc?fr~u 'I'able 16b I IkIt~)rnC'ontrt)l Rccf octocoral fauna (three 20-rn transects, 1977; Wheaton, in preparation b). Percent No. of of total Specws 13.1." Frequency colonies colonies I'.~i~uti~~~~~i~ro~r:t~~gt~~urn crrrr.trttil 54 3 39 13.27 j "j3.1. = Ulological Index, McCluskey ( 1970). 26 Table 17 Ajax Reef stony coral fauna at 17 m (fifteen 1-m2 plots; Jaap, unpublished). Percent ~.l.~ of total Density Species (ranking) Abundance colonies Mean Std.dev. D.I. b ilfontastraea annularis 32 52 24.19 3.47 2.92 C Siderastrea siderea 32 50 23.26 4.17 2.86 C Millepora alcicortlis 2 5 32 14.88 2.13 1.96 R Montastraea cavernosa I8 2 1 9.77 1.40 1.55 R Stephar~ocoeniamicheliizii 10 14 6.5 1 0.93 1.75 C Madracis riecactis 9 9 4.19 0.60 0.74 R Porites astreoides 8 8 3.72 0.53 0.64 R Agaricia agaricites 5 5 2.33 0.40 0.51 R Dichocoenia stellaris 3 3 1.40 0.20 0.56 R Porites porites 3 3 1.40 0.20 0.77 C Eusmilia fastigiata 3 3 1.40 0.20 0.41 R Agaricia lamarcki 2 2 0.93 0.13 0.52 C Siderastrea radians 2 2 0.93 0.13 0.35 R Diploria labyriiztlziformis 2 2 0.93 0.13 0.52 C Meandrina rneandrites 2 2 0.93 0.13 0.35 R Acropora cervicornis 2 2 0.93 0.13 0.35 R Mycetophyllia aliciae 1 1 0.47 0.07 0.26 R Madracis nzirabilis 1 1 0.47 0.07 0.26 R Mycetoplzyllia lamarckiana 1 1 0.47 0.07 0.26 R Helioseris cucullata 1 1 0.47 0.07 0.26 R Colpoph~~ilianatans 1 1 0.47 0.07 0.26 R Total species Total colonies Density, square meter Mean Std. dev. Range Species: 6.13 2.36 1-9 Colonies: 14.33 6.59 1-27 a~.~.= Biological Index, McCloskey (1970). b~.~.= Dispersion Index, Elliott (197 1): C = Contagious (clustered), R = Random, U = Uniform. John Pennekemp Coral Reef State Park Hawk Channel ---'-- Figure 17. John Pennekamp Coral Reef State Park and Key Largo National Marine Sanctum. Table 18 Bank reef zonation patterns. Depth Zone (m1 Conspicuous organisms Back reef/rubble area 0.6 - 1.8 Porites astreoides, Favia fragum Reef Rat 0.6 - 1.2 Diploria clivosa, Porites astreoides, (Plate 14a) Crustose coralline algae Shallow spur and groove 1.2- 2.4 Miliepora complanata, Palythoa sp (Plates 14b and 15a) Deep spur and groove 2.4 - 4.6 Gorgonia ventalina, (Plates 15b and 16a) Acropora palmata Buttress or fore-reef 4.6 - 30.0 Moniastraea an)zularis, Diploria (i31atcs 16b and 17a) strigosa, Colpophyllia natans ffclioseris cucullata, Agaricia fragilis, Madracis mirabilis Spur arid Groove and Sediments LOOE KEY REEF W~stanca[m) E3gure 18. Cross-sectional diagram of Looe Key Reef. 28 Table 19 Carysfort Reef stony coral fauna at 14 - 15 m (ten 1-m2 plots; Jaap, unpublished). Percent l3.1.~ of total Density Species (ranking) Abundance colonies Mean Std.dev. D.I.~ Montastraea annularis Acrvpora cervicorrzis Agaricia agaricites Siderastrea siderea Porites porites Stephanocoeniu michelinii Porites astreoides Colpophyllia natans Mycetophylyllia aliciae Mycetophyllia lamarckiana Millrporu alcicornis MeIioseris cucullata Total species Total colonies Density, square meter Mean Std. dev. - Range Species: 3.70 1.06 2 -5 Colonies: 12.50 8 -44 4 -31 a~.~.= Biological Index, McCloskey (1970). b~.~.= Dispersion Index, Elliott (197 1): C = Contagious (clustered), R = Random, U = Uniform. French Reef Grecian Rocks French Reef, KLNMS (Figure 17; Plate lb) Grecian Rocks, KLNMS (Plate 3a), described in is farther south and is a popular reef for dive tours. It detail by Shinn (1963, 19801, 1s located somewhat has a widely separated reef flat and poorly developed inshore of the main reef he of Carysfort, Elbow, shallow spur and groove zones. The deep spur and French, and Molasses Reefs. U&e most bank reefs, groove zone is well developed and has cavernous tunnels Grecian Rocks has no fore reef or buttress zone. The through many of the spurs. Acropora palrnata and M. zonational pattern includes five zones based on Shinn's annularis are the dominant stony corals on the spur (1963) terminology (Table 21 ). It is about 600 m Iong formations. The spurs are usually 5-6 m deep on their and 200 m wide with the long axis in a northeastfsouth- tops; this limits the firecoral Millepora complanata and west trend. The reef is surrounded by sediments and has the yellow mat zooanthid Palythoa sp. from this reef a narrow bathymetric range of 1.5-7.6 m. zone. Table 20 presents information on the abundances and densities of stony corals from the deep spur and Key Largo Dry Rocks groove zone at French Reef. The deep spur and groove zone merges with a well-developed fore-reef buttress Key Largo Dry Rocks is near and similar to zone. Seaward outcrops similar to those at Carysfort Grecian Rocks, but has greater depth and more defined Reef are also present seaward of French Recf. A deep- spur and groove development Orientation of A, palrnata reef survey seaward of about 40 m characterized the branches are toward the prevailing seas, east to east/ bottom as sedimentary with occasional sponges, tilefish southeast. burrows, and occasional outcrops of limestone with Key Largo Dry Rocks received much study epibenthic reef biota (Jameson 1981). A few solitary following Hurricane Donna in 1960 (Ball et at. 1967; corals (Paracyathus puchellus) were collected. Algal Shinn 1975). Ae that time the charts referred to Grecian nodules were found between 33 and 45 m (Shinn 19811. Rocks as Key Largo Dry Kocks and vice versa. 2 9 Table 20 French Reef stony coral fauna at 6 m (twentyseven 1-rn2 plots; Jaap, unpublished). Percent B.I.~ of total Density b Species (ranking) Abundance colonies Mean Std.dev. D.I. A cropora cervicornis Millepora alczcornis Acropora palrnata Agaricza agaricites Montastraea annularis Siderastrea siderea Porites astreoides Dichocoenia stellaris M.vcetophyllia sp. Coipophj~lharlatans f'orztes porzres Montustraea cavernosa Favia fragum Diploria clivosa 'Total species Total colonies Ilensity, square meter Mean Std. dev. Range Species: 3.37 1.71 1-6 Colonies: 12.48 8.74 1-36 a~.l.= Biological Index, McCloskey (1970). b~).~.= Dispersion Index, Elliott (1971): C = Contagious (clustered), R = Random, U = Uniform, Table 21 Grecian Rocks zonation pattern (Shinn 1963, 1980). Depth Conspicuous organisms Lone (m) and remarks Back reef 0 -0.9 Nonoriented Acropora palma ta, A. cervicornis Reef flat 0 - 0.9 A. palmatlr Acropora palmata 0-1.2 Oriented A. paltnara Weak spur and groove 1.2 - 1.8 ~Woiztastraeaarznularzs, M. cornplaizata Seaward rubble 1.8 - 2.4 Mostly coral rubble, few Szderastrca siderea and M. annularis 30 Since the hurricane, staghorn corals (A. cer- M. annulark just seaward of the deep spur and groove vicornis) have overgrown a great deal of Key Largo Dry zone. Table 22 provides information on the densities and Rocks previously unoccupied by reef corals. The slower abundances of stony corals in the spur and groove zone growing star corals in the reef proper are also being at Molasses Reef. The investigation of seaward zones overgrown by the more rapid growing staghorn corals. with a submersible revealed rocky outcrops and sedi- Shinn (1 975) reported on this phenomenon and has mentary environments similar to those described for followed up the situation with annual inspections of a Carysfort and French Reefs (Jaap 198 1). particular set of colonies that are being encroached upon by A. cervicornis. Looe Key Reef Molasses Reef The zanation and morphology of Looe Key Reef, Looe Key National Marine Sanctuary (Plates 2a Molasses Reef, KLNMS (Figure 17), is the and b) are presented in Figure 18. Table 23 lists the southernmost of the 03shore bank reefs in the park density and abundance of stony coral fauna on Looe system. It is the most popular reef with scuba diving Key Reef. The major significant difference in this reef tourists and has a large lighthouse that makes it very from others is the wide area between the buttress zone easy to find. This reef fits most of the patterns defined and the dzep reef. In the northeastern part of the reef, in Table 18; however, its reef flat is located a relatively the deep reef does riot exist; however, in the south- long distance from the spur and groove zone. The western end, the deep reef community is evident in buttress zone is quite narrow, but has large colonies of patches between sedimentary deposits. Shinn et al. (in Table 22 Molasses Reef stony coral fauna at 5 - 6 m (twenty-five 1-m2 plots; Jaap, unpublished). Percent B.I.~ of total Density Species (ranking) Abundance colonies Mean Std.dev. D.I. b Acropora palmata Acropora cervicornis Montastraea annularis Millepora alcicornis Siderasfri?a siderea Millepora complanata Agaricia agaricites Montastraea cavernosa Porites astreozdes Favia fi.agum Diploria labyrinthiformis Madracis mira bilis Mycetophyllia lamarckiana Colophyllia natans Dichocoenia stokesii Meandrina meandrites Mycetophyllia aliciae Total species 17 Total colonies 325 Density, square meter Mean Std. dev. Range Species: 3.36 1.55 1-7 Colonies: 13.00 6.7 1 4-29 a~.~.= Biological Index, McCloskey (1970). b~.~.= Dispersion Index, Elliott (1971): C = Contagious (clustered), R = Random, U = Uniform. 3 1 Table 23 Looe Key Reef stony coral fauna at 1 - 27 m (fifteen 1-m2 plots; Jaap, unpublished). Percent B.I." of total Density Species (ran king) Abundance colonies Mean Std.dev. D.I.~ Porires astreazdes Milkpura comr~ianata Aganeia ugarzcttes i'orr'tes porites Ftavla fragum Siderastrea siderea A cropora rervzcurnzs Madracis decactis Sfcphanoccreniu michelznit Sidernstr~aradians Morrtastraeu cavernosa Mycc~rophylltalamurchcaria Milleporii alcicornls A rropora palrna fa Montustra~aannularts Mcaniirtncr muundrrtcs 'I'otal species 'I'otal colonies Density, square rrleter Mean Std. dev. Range Species: 4.06 2.19 1-8 ('oionies: 12.19 9 .29 3 -34 a~.l.= Bictlogical Index, McCloskey (1970). "IS.I. - Oispersion index, EUiort (1971): C = Contagious (clustered), R = Random, U = Uniform. press] reported that net sediment transport was to the Bird Key Reef and that sediment was slowly covering the reef's deeper areas. Bird Key Reef, Dry Tortugas National Monu- ment (Figure 19), is unusual in that it does not have a Eastern and Middle Sranbn, Eastern Dry Rocks. Rock spur and groove zone in shallow water, and A. palrnata is Key, and Sand Key Reefs not found at all in this reef community. The major coral accretion is in waters deeper than 10 m on an antecedent These, reefs follow the general bank reef pattern, spur and groove formation that apparently developed They have wellitevelopcd seaward platforms that are during a lower Holocene sea level stand. Shinn et al. sparated from the main reef structure and that have f 1977) reported that cores bored on the shallow areas of .&dficant relief. Again, these platforms axe separated Bird Key Reef revealed that the reef was not founded on from the main reef body by sediments. a solidified platform but rather on considerable uncon- 32 Figure 19. Cross-sectional diagram of Bird Key Reef. solidatcti sediments overlying a Pleistoceile basement. 1982); reef age or stage of development (Shinn et al. The shallowness of the reef and severe winter temper- 1977, 1981 ), and relationships with sedimentary en- atures probably prevent A. palmato froin occurring on v~ronments(Ginsburg 1956; Shinn et al. 1981). Com- the shallow reef at Bird Key. munity structure patterns reflect the stage of reef development. For example, spur and groove construc- Summary tion requlres long terrn dense aggregations of Acropnra palmata (Shinn 1963; Slinn et al. 1981f. Development No universal pattern exists; each reef has dif- of the spur and groove habitat in turn creates micro- ferent features. Major controls include geographic con- habitats (niches) that are necessary for the existence of figuration of the Florida Keys archipetago (("~lnsburg other reef organisins. For example, the Milleporo conr- and Slnn 1964), winter cold fronts chilling Florida plarzata association is found in the turbulent shallower Keys water masses (Hudson et al. 1976, Walker eC ai. portions of the spur and groove habitat (Mergner 1977). CHAPTER 4 CORAL REEF BENTHOS 4.1 INTRODUCTION Earlier references for the study of benthic algae found on Florida reefs and the western Atlantic in The benthos (bottom-dwelling organisms) within general include Harvey (1852, 1853, 1858); Vicker a coral reef is complex, diverse, and for many taxa, (1 908); Collins (1909); Borgesen (1 9 13-1920); Collins poorly known. Benthos in a coral reef constitutes the and Harvey (1917); and Howe (1918, 1920). critical biota or foundation species. In particular, the Contemporary studies of great value include corals and crustose coralline algae are most significant in Earle (1969, 1972b3, which list virtually all algae known niche creation for the other multitudes of species. to occur on Florida's coral reefs. Complexity is shown by the infaunal boring biota; Grazing on algae of coral reefs is discussed by reports of species within a single coral head ranged from Randall (1961a), Mathieson et al. (1975), Wanders 30 to 220 and the number of individuals ranged from (1977), and Brawley and Adey (198 1). 797 to 8,267 (McCloskey 1970; Gibbs 197 1). Most orga- Productivity of algal components within coral nisms were polychaetes, a group that is poorly under- reef communities, including those that bore into lirne- stood, Because it would be literally impossible at this stone and the coral symbionts, is treated by Marsh time to compile a total benthic community profile, this (1970), Littler (1973), and Vooren (1981). chapter will cover those groups which are critical to Colonization and succession of reef algae were the community: algae, sponges, and Cnidarians (e.g., reported by Adey and Vasser (1975), Wanders (1977), corals). Since corals are major elements of the reef, and Brawley and Adey (1981). Algal zonation at emphasis will be on the Milleporina, Octocorallia, and Curacao was described by van den Hoek et al. (1975). Scleractinia. Algae that bore into coral and limestone were Volumes 2 and 3 of Bcology and Geology of studied experimentally by Perkins and Tsentas (1976). C'orol Reefs (Jones and Endean 1973, 1976) contain Coralline algal ridges (Note: these are not found in infornlation on many benthic groups, but the major Florida) are reported by Adey (1975, 1978). The coral- emphasis is on Indo-Pacific reefs. Although the infor- line algae were recently treated by Johansen (1981). mation can he extrapolated in many cases to Florida, the The only recent reef algae study from Florida species are for the most part different. Recently, Rutzler reefs is Eiseman (1981), who made diving observations and Macintyre (1982b) published a fine volume on the and collections during the KLNMS deep reef survey biota from the barrier reef at Carrie Bow Cay off Stann using a submersible. The study began at a 30-m depth Creek 'Town, Belize. and reported 60 species from reef and lithothamnion cobble habitats. The algal species from these two habi- 4.2 ALGAE tats were virtually exclusive. (by Harold Humm, Department of Marine Science, Urtiversity of South Florida, St. Pet ersburg) Algae in Coral Reefs Historical One of the characteristics of coral reefs the world over is the apparent paucity of benthic macroalgae on Knowledge of the benthic algae found on coral and around the reefs, at least to the casual observer. reefs of southeast Florida is based largely on inference "Where are the plants?" might be an immediate question froni work done on coral reefs in the West Indies, of a biologist who is familiar with temperate or boreal especially in the Virgin Islands, Yuerto Kico, Jamaica, seas and views a coral reef for the first time because Curacao, Barbados, Guadalupe, and Martkque. There rocky substrata in clear, shallow water would normally are many scaslde research facliiticS open to visiting support a dense stand of large benthic algae. scientists in the Canbbean area but only one in southeast A significant biological control of algae on coral Florida, and that with limited facilities for visitors. Thus, reefs is the competition for space with other epibenthic for these and other reasons, studies of the benthic algal sessile organisms. The diversity of organisms on a coral flora of Florida coral reefs are sparse, reef probably exceeds that of all other marine eco- William Randolph Taylor of the University of systems. The sessile animals occupy space that would Michigan was the first to make a significant contribution otherwise be colonized by benthic algae. On recently to the knowledge about the species of benthic algae exposed Limestone substrates, especially following found on Florida coral reefs, after he sped the summers physical impact, benthic algae may be the first colo- of IS261 926 a1 the C'arneae Laboratory, Dry Tortugas. nizers, but they are usually replaced by sponges, turn- Taylor's work at Tortugas was a general floristic survey; cates, corals, and bryozoans after a short period. An- his field data provided information on species associated other major b~ologicdcontrol on benthic macroalgae with reef environments. These data are included in is grazing by the herbivorous invertebrates and fish. Taylor (1928) and Taylor (19601, Marine Algae of the Of all shallow-water marine ecosystems, none are more Tropical and Subtropical ~'olrstsof the Americas, still profoundly affected by grazing than coral reefs. available from the University of Michigan Press. The effects of grazing on coral reefs have been demonstrated experintentally in many reef studies, al- the experiment at Curacao. Wanders concluded that though apparently not in Florida, by placing a barrier grazing does not reduce the primary productivity per over a selected site and observing the response of the unit area in Curacao, but only affects species compo- algae that are protected from most grazing. Randall sition. (196lb), working in Hawaii, found a significant differ- Brawley and Adey (198 1) observed the effects of ence between the algal cover inside an enclosure and that another category of grazers on coral reefs (other than of areas outside it after 2 months. Inside the enclosure fish and macroinvertebrates such as urchins) that might the algae grew to normal height and breadth (to 30 mm); be referred to as micrograzers, crustaceans of the Order outside the enclosure, the algae averaged 1 mm in height. Amphipoda. In a coraI reef microcosm in the Smith- In Lameshure Bay, St. John, Virgin Islands, Randall sonian Institution, Washington, D.C., they observed that excluded fish, but allowed herbivorous sea urchins a tube-building amphipod of the genus Amphithoe (Diadema antillarum) in the closed area. Results showed grazed selectively on filamentous algae. They suggested that the fish were the major grazers. While the urchins that this micrograzer may help reduce competition for also fed on algae, their action did not affect the algae as the encrusting coralline algae. severely as did the fish. Randall also showed that panot- While grazing by fish is a major factor in the fish and surgeonfish graze on seagrass adjacent to the paucity of fleshy algae in coral reef habitats, there are a reef, thereby causing the sand halos around patch reefs. few coral reef fish whose activity has the opposite effect. Research conducted during the submersible habitat One of these is the three-spot damselfish, Pomacenrrus study known as Tektite 11 at Lameshur Bay, St. John, planifrons. It establishes a territory, especially in the Virgin Islands, confirmed and expanded Randall's proximity of Acropora cervicornis, A. palmata, and observations on grazing. Fish grazing was found to be a Montastraea annularis, and then bites segments, strips, or major factor influencing biomass and species diversity of patches of the coral, killing the polyps. These dead coral benthic algae on coral reefs. In addition to cage exper- areas are colonized in a few days by a variety of erect iments, algae was transplanted from elsewhere to the -growing fleshy algae. The damselfish, by its unusual reef; the algae was rapidly eaten. Grazing pressure was aggressive habit of defending its territory, keeps oat algal intense to a distance of 30 m; beyond that distance it grazers and thus maintains extensive patches of fleshy gradually tapered off (Earle 1972a; Mathieson et al. algae on the reef where normally they would not exist. 1975). Detailed observations on the damselfish and their algal Ogden (1976) reported on algae-grazer relation- lawns were reported in Jamaica by Brawley and Adey ships on coral reefs and reported those algae that grazers (1977) and in other Caribbean localities including tend to avoid. Prolific algal growth occurred only in Jamaica by Kaufman (1977). The threespot damselfish areas inaccessible to grazing herbivores, such as wave- occurs on Florida's coral reefs as do all the algae species washed surfaces and beachrock benches. Ogden and recorded and listed by Brawley and Adey from Discov- Lobe1 (1978) summarized the role of herbivorous fish ery Bay, Jamaica. The exception is Halimeda goreauii. and urchins in coral reef communities with special reference to reefs around St. Croix, Virgin Islands. Algal Groups A comprehensive study of the effects of elim- ination or reduction of grazing by means of cages was Benthic algae of tropical coral reefs can be reported by Wanders (1977), who workcd at Curacao. categorized into four major groups, all occupying areas His results are applicable to Florida reefs. fie found that of the reef itself or areas under the influence of grazers crustose coralline algae depend upon grazing for protec- inhabiting the reef. They are crustose coralline algae tion from competition by erect fleshy algae. Under a that encrust corals, reef rock, and other Limestone cage that covered a patch of coralline algae, the fleshy skeletal material; filamentous and fleshy algae, which algae soon colonized the surface of crustose species, occur as sparse vegetation and dense vegetation; algae resulting in the death of the crustose species. The on unconsolidated sediments, which are erect rnacro- crustose forms were penetrated by microscopic species algae of the order Siphonales and mats of bluegreen that bored into the limestone. When clean artificial algae; and excavating or boring algae. substrates were placed under cages on the reef, a succes- sion was observed as the fleshy algae colonized it. During Grustose Coralline Algae the first 6-8 weeks t$e colonizers were principally filamentous brown and green algae of the genera Crif- The most distinctive and characteristic algal fordia, Cladophora, and Enteromorpka. After 10-15 group of the coral reef is the crust-forming coraILine red weeks these were replaced by larger filamentous and algae Rhodophyceae, order Cryptonemiales, family parenchymatous species, in particular, the filamentous Corallinaceae, subfamily Melobesieae. These algae form a red algae Centraceras cbvularum and Wrangelia argus; a thin or massive crust with or without erect branches and small, erect-growing coralline red, Jania capillacea; the are calcified throughout. When Living, they are usually a larger red algae Spyridia filamentosa, Pterocladia arner- shade of red in low light, but may be yellow-brown nn icana, and Laurencia microcladda; and the flat, dichot- surface light. They are chalk-white when dead, but soon omous brown algae Dicfyota dichotoma. A11 of these are become greenish as a result of the establishment of the common species on Florida coral reefs and can be found green and bluegreen algae that bore into limestone and where succession on new substrates would be similar to lend color to the upper few to 5 mm of the skeleton, Crustose corallines form small, scattered colonies The mat-forming bluegreen algae community is or large, distinct patches. On Florida reefs, there may be composed primarily of filamentous species (sensu some degree of development of incipient algal ridges, a Drouet 1968), e.g., kficrocoleus lynghyaceus and Schizo- formation found extensively on coral reefs in the eastern thrix calcicolu. Other filamentous species, including Caribbean Sea (Adey 1978). CoraUine algae are common 130rphyrosiphon notartszi and Schzzothrix arenaria, on the underside of corals such as Acrupora spp. and and several coccoid (spherical) species are usually colonize dead coral fragments that hreak off and fall to associated with these mats. Ginsburg (1972b) reported the seafloor. Large patches are found on the reef plat- that the filaments of mat-forming bluegreens grow form limestone, the living surface layer covering former- upward during the day and selectively trap sediment ly living veneers of the same species. Minute species that particles at night; horizontal growth of another species form small, thin crusts are often epiphytic upon larger, results in binding of trapped sediments. fleshy algae, seagrass leaves, octocoral skeletons, mollusk Erect-growing anchored green algae in sediments shells, and hydroid colony bases. Crustose corahes are are found within the genera Hakmeda, Penicrllus, Udo- best developed in shallow, turbulent areas where Light tea, Rhzzocephalus, Avralnvillea, and Caulerpa. They are intensity is high and fish grazers arc part~allydeterred by usually scattered and scarce immediately around the wave forces. reefs, but become progressively more abundant away Colonization, succession, and growth rate of the from the reef proper as grazing pressure is reduced. The tropical. cruslose coralline algae were unknown until the dense cluster of rhizoids that anchor these plants stabi- study of Actey and Vassar (1975) in St. Crolx. Their lizes the sediments and absorbs nutrients. Since these results probably apply in Florida, whch has fewer algae are coenocytic, transport of nutrients to the tops species, for April through Nove~nber. Adey and Vassar and of photosynthetic products to the rhizoids is carried (1975) reportcd that the margins of a crust grow 1-2 on rapidly in conjunction with cytoplasmic streaming. min/month and that accretion rates arc 1-5 mmjyear, They are a unique group of h.ghly evolved green algae, as depending on herbivore grazing, especially parrot- advanced among the algae as the flowering plants on fish. land. Members of this group also contribute signif- Filamentous and Fleshy Algae icantly to calcium carbonate production and sediment on and around the reef. This is especially notable among E'ilanxcntous and fleshy algae arc uncalcified the species of ffalirncda. coral-reef macroalgae that colonize coral rubble and reef litnestones. This group is most adversely affected by Excavating or Boring Algae grazing. '1 wu subdivisions are recogl~i~edthe dense and sparse vegetation. 'I'hey con~prise the same species, Anlong the least conspicuous and most often although 111e denvc comniunity, rf not spatially lkrnitcd overlooked algae are those possessing the ability to bore to crcvices OF six~all patches, extubits greater species into limestone by dissolving it as they grow. To the d~vessity. 'l'hc' sparsc cornrriunlly occurs in arciis re- unaided eye they are visible as a greenish tinge or dis- cdving the heaviest grazing pressure, resulting in cropped coloration at the surface of dead coral, mollusk shells, Furnrs often 1-2 tt1111 Itigh. There IS a transition OF sparse dead coralline algae, and other Limestone material. cornmunlty on ur near the reef to the dense contn~unity Boring algae belong to three taxonomic groups. Most are located away from the reef, resulting fro~na gradient in bluegreen (Cyanobacteria), some are green (CNoro- gl-azxng pressure. phyta), and the remaining one (Xanthophyta) has no common name. Unconsolidated Sediment Algae Representatives of three families of bluegreen algae are known from Florida coral reefs: Entophysalis Pew algae have the furlctiollal alxlity to anchor dfusta, family Chamaesiphonaceae; Schtzorhrix cal- ln tinconsolidated sediment. For this reason, the sea- ci~ola,family Oscillatoriaceae; il.lastzgocoleus testarum, ~.~asscs,ruotcd fornts tirat cvolvcd on land and radiated family Stigonemataceae. The green algae Gomontia mto the nxarine environnlent, dot~~inateshallow sedi- ployrhlzu and the xanthophyte Ostreobzum quekettiz mentary habitats, especially in troplcal regions. There are known from Florida. arc, huwcver, a few specialized Jlgae on loose sedirucn- 'I'hese algae are collected by dissolving a small tary environments; these are cspecially well represented sanlple of greenish Limestone in dilute hydrochloric acid in coral reef habitats. and examining isolated filaments. Keys to the species are There are two major groups capable of colonizing found in Ilumm and Wicks (1980). A comprehensive sandy or muddy bottom areas. I'he bluegreen dgae experimental study of Limestone-boring algae is found in (Cyanobacteria) form mats and penetrate the sediment Perkins and Tsentas (1976); the study was from St. to some extent. Green algae, belonging to the order Croix. They reported five bluegreen and three green Siphonales, erect coenocytic plants (lacking cell walls) algae species. One of their bluegreens, Ca1othri.x sp. having a dense cluster of root-like rhizoids at the base (Calofhr~xcrustacea; sensu Drouet 1968), has not been that provides a firm anchorage in unconsotidaled sedi- reported as a boring alga in Florida; however, it is ments. Mathieson et al. (I 975) included an underwater abundant on Florida reefs and must be presumed to be a photograph of this community. borer. Algae that bore into limestone contribute signif- 36 icantly to the dissolving of limestone, adding calcium common sponge species have characteristic shapes, and bicarbonate to the mineral pool in the reef environ- colors, and habitat preferences which allow them to be ment. identified confidently in the field, at least within a specified geographic area (e.g., Florida). 4.3 SPONGES Of the more technical taxonomic literature on (by G.P. Schmahl, South Florida Research Center, Caribbean sponges, only a few were based on collections U.S. National Park Service, Homestead). primarily from Florida. An early work that described some Florida reef sponges was that of Carter (1885). Sponges (Porifera) are an important component Later Florida studies on the sponges of the Gulf of of the benthic fauna of Florida reef. Although not Mexico by de Laubenfels (1953) and Little (1963) also usually dominant, sponges are common in most reef included some reef species. The most complete work of zones and can be especially abvdant in certain situ- sponge taxonomy for Florida reef sponges was made by ations. Substrate analysis of the benthic fauna on de Laubenfels (1936) in the Dry Tortugas. Although selected upper Florida Keys patch reefs indicated a now somewhat out of date in terms of the present sponge component ranging from 1.2% to 9.2% of the interpretation of sponge classification, this is an impor- surface area sampled (Jaap and Wheaton 1977). tant work in that all known Caribbean sponge families were listed and their taxonomic characteristics des- Taxonomy cribed. Since the sponge fauna found on Florida reefs is decidedly West Indian, taxonomic works from other Sponges are grouped into four classes. The largest areas of the Caribbean are sufficient for most Florida is the Demospongiae, which account for 95% of all species. Recent studies of this type include those from recent species. Virtually all common shallow water reef the Bahamas (de Laubenfels 1949; Wiedenmayer 1977), sponges are demosponges with most of the remainder be- Bermuda (de Laubenfels 1950), Jamaica (Hechtel 1965), longing to the class Calcarea. The Demospongiae are and Curacao (van Soest 1978, 1980). The studies by characterized by skeletal components consisting of Wiedenmayer (1977) and van Soest (1980) introduced siliceous spicules that are supplemented or replaced by new classificatory interpretations. Taxonomic works on organic spongin, which forms fibers or acts as a ce- burrowing sponges likely to be found in Florida include menting element. The skeleton of the Calcarea, as the Pang (I973a, 1973b) and Rutzler (1974). name implies, is made up of calcareous, usually triradiate A definitive list of Florida sponge species does spicules. A third class of sponges, the Sclerospongiae, not exist. de Laubenfels (1936) described 76 species secrete a compound skeleton of siliceous spicules, from the Dry Tortugas, but 5 were dredged from 1,047 spongin fibers, and calcium carbonate. Sclerosponges can m and 9 others were colIected from sites ranging from be an important structural component of some deep fore 70 to 90 m, which is out of the range of most typical reef environments (1,ang et al. 1975) but have not been Florida coral reef growth. Wiedenmayer (1977) des- reported from Florida reefs (Dustan et al. 1976). The cribed 87 shallow water species from the western Ba- fourth class of sponges, the Hexactinellida, are mainly hamas in a work that reevaluates the validity of scientific deep water species and are characterized by hexactinal names applied to many common reef species. Careful (six-rayed) megascleres (type of spicule). reference to synonymy must be taken into consideration Classification of the sponges is based primarily when comparing Wiedenmayer's (1977) lists with those on the size and shape of the spicules (megascleres and of previous workers. Given the proximity of his study microscleres) and the organization of the skeletal matrix site, it is reasonable to assume that most of the 87 of spicules and organic fibers found within the various species Wiedenmayer (1977) described can also be found species. Gross morphology, surface texture, color, and in Florida. Fiftyseven species of sponges have been arrangement of the incurrent and excurrent ostia are also recorded from Bermuda (de Laubenfels 1950) and considered important. Recently, studies in the areas of Jamaica (Hechtel 1965). Of the species treated by comparative biochemistry (Bergquist and Hartman Wiedenmayer (1977), 39 listed are from the Dry Tor- 1969), reproductive life history characteristics (Levi tugas and 22 from the Florida Keys. These lists were the 1957), structure and function of sponge cells (Simpson result of a literature review and not based on field 1968), and ecology have contributed much to clarify studies, so the actual number of species in these local- taxonomic organization. In spite of this, classification of ities is open to question. the Porifera is still in a state of change and confusion. A brief survey of sponges of the Dry Tortugas by However, the widespread use of scuba, in situ color National Park Service investigators revealed 85 species, photography, and standardized collecting techniques only 43 of which were recorded by de Laubenfeis have all helped in the field identification of major (1 936), and 57 of which were among the 87 species sponge species. Field guides are now available (Voss described by Wiedenmayer (1977). The low overlap of 1976; Colin 1978a; Kaplan 1982) which offer to the species Lists izdii-ates a spcclcs pc,3! rnucb larger than nonspecialist some information on Caribbean reef reported by any one study; the number of species sponges. Of these, the identifications given in Kaplan present is seemingly correlated with the amount of time (1982) are the most up-to-date. Inconsistencies in spent looking. It is mportanl to note that the above nomenclature found in these publications reflect the studies deal w~Phall sponge species encountered in the confusion that surrounds sponge systematics. Most area, lncludlng those common in lagoonal areas, man- 3 7 groves, and inshore pilings. Those sponges that dominate not be seen undel an ordinary light microscope and Was in reef areas are a more or less distinct group and, with thus postulated to be of colloidal (quasi-particulate) some overlap, are substantially different from those nature. Many sponges depend on ambient currents to aid which are abundant in lagoonal areas (Wiedenmayer in water transport through their tissues and to decrease 1977). An extensive study of the patch reefs of Biscayne the amount of energy expended on pumping activities National Park (upper Florida Keys] reported the occur- (Vogel 1977). Thus, hydrodynamic regimes are impor- rence of 98 species associated only with inshore reefs in tant in shaping sponge distributions. In tropical areas, less than 10 m of water (Schmahl and Tilmant 1980). where reef boundary layers are typically low in nutrients Additional species have been collected from deeper and organic particulate matter, increased water flow by water on the outer bank reefs. From the evidence so far, currents can be essential for the survival of many large a conservative estimate of the sponge species occurring species. on or around Florida reefs must be at least 120. Light intensity can be important in shaping sponge communities for various reasons. In one respect, Autecology reduced light intensity increases sponge abundance due to decreased competition from reef corals that depend Sponges exhibit both asexual and sexual repro- on light for survival of symbiotic zooxanthellae. Sponges duction. Asexually sponges may be regenerated from typically proliferate in deep fore reef areas below the fragments, gemmules, and reduction bodies. Sexually, zone of maximum coral growth. Some sponges, however, the group has both dioecious and hermaphroditic have been shown to contain species of symbiotic blue- species. Fertilization is usually internal and both ovip- green algae (or Cyanobacteria) which have been impli- arous and viviparous species are common. This fact cated in the distribution of those species, restricting forms the basis of the division of the Demospongiae into then1 to shallow areas where light is abundant (Wilkinson its twu main subclasses: 'I'etractinornorpl~a (oviparous) 1978). and Ccractinomorpha (viviparous), although exceptions For a good general review of sponge biology, are known. Viviparous species usually incubate paren- consult Bergquist (1978). Several recent collected works c hy mula (solid) larvae, while dcvcloprnent in oviparous have been compiled as the result of symposia on the forms is usually external. biology of sponges or in response to the need for cohe- Amphiblastula (hollow) larvae arc exhibited by a siveness in sponge research. These include Fry (1970), third subciass of I)emospongiac, the IIomoscleromorpha, Harrison and Cowden (1976), Levi and Boury-Esnault and sornc calcareous forms. A promising method that (1979), and Hartman et al. (1980). rrkay bc uscd lo access the extent of asexual reproduc- Quantitative ecological studies of sponges on tlon ur an arca IS the ability of sponges to recognize Caribbean reefs are few. The most complete series of tissue of like genetic cornposition ("self"). Experiments studies of Caribbean demosponges to date were those itlay Isc carried out whereby strains can be recognized carried out by Keiswig (197 la, 197 lb, 1973, 1974) on lhrtxr~ltlt tliffcrcntial acceptance of tissue from other three cornmon species found on Jamaican reefs. These ~nd~virluaisin an arca (Kaye and Ortiz 1981). Kepro- studies set the standard for ecological methodology of dudwn In sponges was reviewed by f;ell (lY74), but sponges and their information is generally compatible tl~e~vI! much yet that ~sunknown about that process m with Florida populations. I-Iis findings emphasized the rrrost specles. A recent, review by Sunpson (1080) variability displayed by the different species in regard to pointed out the many areas open to research. pumping activities, feeding, life history characteristics, L,arval ccology and behavior is fundaeneiatal to population dynamics, and respiration. Ecological studies tllc il~str~lmtionof adult forms. Most sponges have sex- on entire sponge communities are rarer still. Alcolado ually prcxfuced larvae that are free-swinilning, at least for (1979) investigated the ecological structure of sponges a short period, although some species have been shown on a Cuban reef, and data were given in Wiedenmayer to prodoct: larvae that do not swirn but: creep over tho (1977) stressing synecological relationships of sponges in subst~atc(Ncrgcluist 1978). 7'hree physical attributes his Bahama sites. The difficulty of carrying out such influcn~-..etlrc swimn~ing arxd settlement of sponge stud~es1s reviewed by Kutzler (1978) who also gave larvae. light, gavity. and water turbulence. The com- some methods for quantitative assessment of sponges paritive rlzorpiiology and behavior of some Demosponge on coral reefs. larvae have been revicwed by Bcrgquist ct al. (1979). From these studies and from qualitative and Sponges are filter feeders and must take great incidental observations, sponge communities are known cluantities of water fro111 which they remove plankton, to be partitioned along habitat and depth gradients on bac;t-erk organics, and other nutrients from the water coral reefs. Habitat preference, reproductive strategies, column. It has been estimated that the abundance and growth form, and competitive ability are important pumping activity of sponges on the fore reef slope of biological agents that influence sponge distribution. Discovery Bay, Jamaica, arc such that a volume ettuiv- Abiot~cfactors, some linked directly or indirectly with alenr to tire cntiro water coluxiln passes ~hroughthe depth (e.p., light mtensity, temperature, intensity of population every 24 hours (Reiswig 1974). Sponges are wave and surge action, and sedimentation) are also major capable of removing extremely sniaU organic particles. co~ltroliingforces in shaping sponge assemblages, In a study by Keiswlg (1971b), 80% of the organic Massive sponges are rare on reef flats where small carbon removed by three species of sponges could or low encrusting forms are predominant (Reiswig 1973; 38 Alcolado 1979), presumably because of the scouring enlarging coral base to grow on, while the coral is aided action of waves and the increased sediment load caused by protection from boring organisms that usually gain by turbulence. Many species cannot tolerate high sedi- access through the coral's undersurface. The coral ment loads. In Verongia lacunosa pumping rate is re- possibly benefits also from greater access to hetero- duced under increased sediment conditions (Gerrodette trophic energy sources carried in the spongecreated and Flechsig 1979). Species with high sediment toler- water currents. An obvious symbiotic relationship in ance, such as Tethya crypta, can be common on the reef Caribbean sponges involves species of Zoanthidea flat as are encrusting species such as Spirastr~llacunc- (Anthozoa) which grow on the surface of some sponges. tatrix and Tfzalysias juniperina. Species of the genus 'Taxonomy of the sponge-associated zoanthids has been Cliona are also common, infesting dead coral skeletons given by West f 1979). Aspects of the ecology of sponge- and other suitable substrate, In deeper areas (> 7 m), zoanthid associations have been investigated by Crocker where conditions are more predictable and favorable, and Reiswig (1981), who found a high specificity be- massive sponges increase in abundance. Common species tween species of host sponges and their associated found on the outer reefs of the upper Florida Keys zoanthids. Lewis (1 982) studied some functional aspects include Amphimedon compressa, Iotrochota birotu- of this relationship and suggested that the zoanthid lata, Ulosa ruetzleri, Ircirzia strobilina, Ectypoplasia presence may have a deleterious effect by decreasing the ferox, Callyspongia vaginalis, Niphates erecta, N. digi- host sponge's pumping capabilities. talis, N. amorpha, Cliona deletrix, and Xestospongia Another important role of sponges in coral reef rnuta (name designations after van Soest 1978 and ecosystems is that of providing shelter for other reef 1980, and Wiedenmayer 1977). organisms. Tyler and Bohlke (1 972) listed 39 species of fish that associate with sponges, at least 5 of which are Synecology obligate sponge dwellers. The interior cavities of certain sponges are inhabited by numerous invertebrates (Pearse Sponges are major competitors with other epi- 1932; Westinga and Hoetjes 1981), mostly crustaceans, benthic organisms for space and other resources in reef of which the alpheid shrimps are the most prominent. habitats. Sponges have the greatest overgrowth capa- The residents also include polychaete and annelid bility of the major groups of organisms encrusting worms. Reiswig (1973) reported that the sponge Veron- undersurfaces of foliaceous corals (cheilostome ectop- gza gigantea supported large populations of Haplosyllis rocts, crustose algae, and other algae). Results varied spongicola in the canal system; densities of 50-100 with depth, but sponges were the superior overgrowth polychaete individuals per ml of sponge were found. competitors in 77% of the interactions monitored Haplosyllis spongicol~is also a frequent inhabitant of (Jackson and Winston 1982). Overgrowth of Caribbean the sponge Neofibularia nolitangere. corals by sponges has been observed for Chondrilla Sponges serve as a food source for other or- nucula (Glynn 1973), Ectyoplasia sp., and Plakortis sp. ganisms, predominantly coral reef fish and some marine (Lang 1982), and has been demonstrated for Antlzo- turtles. In general, however, predation is not intense. sigmella var. f. incrustans (Vicente 1978), which was Only in 1 1 of 2 12 fish species (5%) studied by Randall found to have a high level of competitive superiority and Hartman (1 968) did sponge remains comprise over compared with corals and other sponges. Some sponge 6% of stomach contents. Angelfish of the genera Holo- species have been shown to exhibit toxicity to corals canthus and Pornacanthus and the whitespotted filefish which they overgrow, causing necrosis of the coral tissue (Cantherhines rnacrocerus) were the major sponge along the line of contact (Bryan 1973). Such a mech- predators. Abundance of mineralized sclerites, noxious anism of competitive interaction presumably concerns chemical substances, and tough fibrous components have the production of species-specific allelochemicals (Jack- been identified as reasons for low predation pressure on son and Buss 1975; Buss 1976), which can lead to most coral reef sponges. The toxicity to fishes of many intricate competitive networks and act to allow high common exposed (noncryptic) Caribbean sponges has diversity in areas of low disturbance. Sponges have also been shown by Green (1977) through forced feeding been shown to enter into complex epizoic relationships experiments. He also noted that, in general, as latitude (living on one another) with one another (Rutzler 1970) decreases, the incidence of toxicity in sponges increases, which may be in response to competitive pressures. presumably in response to the high diversity and density Sponges are involved in symbiotic relationships of associated fishes in the tropics. Predation, therefore, with other reef organisms. Many sponges exist in sym- is not usually considered a direct force limiting the biosis with bluegreen algae (or Cyanobacteria), which are distribution of coral reef sponges. This is not true for all mostly free living within the mesophyll and may consti- sponge ecosystems, as was illustrated by Dayton et al. tute up to 52% of the cellular material of the sponge (1974) in an Antarctic community. (Rutzler 1981). Nutrient translocation of algal products Sponges have been regarded as a major force in to coral reef sponges has been demonstrated by Wilkin- the bioerosionai process on coral reefs (Goreau and son (1979). Sponges can, therefore, supplement their Iiartman 1963; Rutzier 1975). The boring sponges are energy requirements through this relationship. Goreau classed mostly in the family Clionidae (genus Clzona), and Hartman (1966) noted that the Caribbean sponge but species of the Adocidae (Siphonodzctyon) and the Mycale laevis has a symbiotic relationship with some Spirastrellidae (Sphecrosporzgia, Anthoszgmelltz) have stony corals. The sponge benefits from a protected and also been shown to excavate coral Limestone skele- 39 tons. They weaken the substrate, causing the collapse ecological studies. As William Beebe (1928) wrote: and dislodgement of some corals and creating cryptic ". . .when, in the Iliad, Homer described a sponge as 'full habitats in the interior of coral skeletons. In a study of of holes,' he expressed about all the knowledge which Cliona lampa in Bermuda, Ncumann (1966) found that mankind has possessed until comparatively recent as much as 6-7 kg of carbonate detritus could be gener- times." Although there have been some advances in ated from 1 m2 of sponge-infected substrate in 100 knowledge of sponge biology since 1928, there is still days. Iiudson (1977) reported six species of boring much to be learned about the physiology, ecology, and sponges that were principal in the bioerosion of Man- evolution of sponges. rastraea anizularzs (star coral) at Hens and Chickens Reef in Florida. In contrast to the erosional effects of the boring species, many other species contribute to the structural The phylum Cnidaria (Table 24) includes jelly- integrity of coral reefs by binding unconsolidated reef fish, sea anemones, corals, and hydrozoans. Although frame material together and thereby increasing rates of extremely variable in appearance, all members have a carbonate accretion (Wulff and Buss 1979). radially symmetrical body plan. The saclike body has Sponges play a major role in the ecology of a central stomach cavity with a single opening that is Florida coral reefs, but have been greatly neglected in usually surrounded by foodcapturing tentacles. Stinginp Table 24 Classification of major reef benthic Cnidaria, ORDER MILLEPORINA: fire corals OKDER STYLASTERXNA: hydrocorals CLASS ANTflOZOA SIJBC'LASS OCTOCORALLIA (ALCYONARIA) sc~suBayer 1961 ORIIER STOLONI12EKA ORI3flR 'T'ELES1 ACEA ORllfiK ALCYONACEA: fleshy soft corals OKIjER COKCONACEA: sea whips, sea feathers, sea fans. sea plumes, other gorgonian corals ORDHIZ I'ENN.ATULAC:EA: sea pens SUBCLASS %OANTflARlA (IIEXACORALLIA) ORDER ACTINIARIA: arlenlones ORDEK COKALLIMORPWARXA: false coral anemones ORDER LOANT111DLiA: carpel anenloncs ORDEK CEKIANTHARIA: tube anemones, often parasitic in other organisms ORDER SCLERrXCTINlA (MADREPORARIA). true stony corals ORDbR ANTIPATHARXLZ: black or thorny corals capsules (nematocysts) on the tentacles narcotize prey zooxanthellae in protein synthesis. The calcification rate before they are drawn into the mouth, and sometimes and colony growth are greatly enhanced by this sym- can inflict powerful stings on humans. biotic relationship. Plankton, captured by the dactylo- The two classes of Cnidaria in which major zooids and digested by the gastrozooids, probably colonial reef forms are found are the I-Iydrozoa and provide the micronutrients to the polyps. Anthozoa. Within the Hydrozoa class is the order Growth rate data for Millepora are limited or Milleporina, containing the fire corals. The Anthozoa nonexistent. After settlement the growth seems to be class has two subclasses. (1) Octocorallia or soft corals, very rapid, probably upward growth approaches 10 cm including those in the order Gorgonacea (gorgonians), annually (author's subjective estimate). Encrusting forms and (2) Zoantharia, containing the order Scleractinia are transitory, instability of substratum usually leads to or true stony corals. The following will describe the early mortality for the colony. fire corals; the soft corals, especially the gorgonians; Wahle (1980) reported that Mzllepora colonies and the stony corals. detect octocoral colonies from stimuli present in sur- rounding seawater and redirect growth to reach the Milleporina octocoral colony. Mrlleporu then grow over the surface of the octocoral, thus gaining new space. On Florida reefs, the Milleporina are represented Millepora complannta has a limited bathymetric by a single genus, Millepora. These fire corals are quite distribution. It is generally restricted to the reef flat and common throughout the western Atlantic tropical coral shallow spur and groove zones (0-5 m). On Looe Key reefs. A colony consists of a calcareous skeleton with an Reef (Table 23) it was second in abundance and frequen- internal meshwork and external pores through which the cy, with a mean density of slightly greater than three polyps retract and expand. The two kinds of polyps are colonies per square meter. Mergner (1 977) reported that the dactylozooid and the gastrozooid. The dactylozooid, M. cornplanata was an indicator species of photopl-Lilic the defensive and food-collecting polyp, has potent and rheophilic environments; its ecological niche is ap- stinging apparatus, the nematocyst. The nematocyst parently Limited to the shallow, well-illuminated and tur- contains a small hypodermiclike structure. This is a bulent waters found in shallow windward reef communi- capsule with a coiled barb, flagella trigger, and a strong ties. neurotoxin. Upon stimulation the trigger releases the barb that is shot into the prey or predator by hydraulic Octocorallia (Soft Corals) pressure, and the poison is released. The gastrozooid contains a mouth and digestive enzymes. Dactylozooids The Octocorallia, commonly called soft corals or and gastrozooids are distributed in cyclosystems of five octocorals, are conspicuous in coral reef communities to seven dactylozooids around each gastrozooid. off southeast Florida (Table 25). They have various Two species of Milleporu are found in Florida. M. shapes ranging from inconspicuous encrusting mats to alcicornis is a digitate branching form, and M. compla- large sea fans. Nearly all types possess calcareous spicules nata (Plate 3b) is a truncated bladed form. In many embedded in an organic matrix. Specific species charac- cases, it is impossible to render a specific determination. teristics include colonial morphology, branching pat- Thin encrusting veneers of reef rubble, the skeletons of terns, and morphology and configuration of the spicules. other organisms, and jetsam prevent recognition of the The fundamental morphological character is the polyp, specific characters. The western Atlantic species, M. which has eight pinnate tentacles. squurrosa, is a nominal species. Stearn and Riding (1973) The Gorgonacea (gorgonians) are the common- and DeWeerdt (1981) showed that M. squarrosa was an est octocoral in southeast Florida reefs in depths less ecomorph of M. complanata. Boschma (1948, 1956) dis- than 30 m. They include the sea fans, sea plumes, sea cussed the systematics and taxonomy of Millepora. feathers, sea whips, and sea rods (Plate 4a), which are all Miilepora's life history reflects hydroid meta- verv flexible and attached by a holdfast or base to the genesis or alternation of polyp and medusa generations. reef platform. Branches of gorgonians possess a horny, The polyp represents the benthic stage. It asexually solid center, while other groups have a solid or caicar- produces planktonic medusae, which develop gametes eous axis. that, when fertilized, produce planktonic larvae. The The taxonomy of the major categories Is cur- larvae settle and metamorphose into a juvenile Milleporu rently being revised. Bayer (1961) is the most complete colony. Recruitment can also come from fragment single source for identification of the shallow water propagules. Following storms or physical impact, the octocorals found off southeast Florida, and Cairns broken pieces have regenerative powers to grow into new (1977) is a useful field guide. Deichmann (1936)report- daughter colonies. ed on the deep-water octocorals. iMilleporu is a functional autotroph and hetero- Octocoral autecology, including environmental troph. It has very dense concentrations of zooxanthellae tolerances sun~marizedby Bayer (1956). is simrlar to (microscopic, symbiotic dinoflagellate algae) in its that of the stony corals (see next section, Sclzractima). endoderrnic tissues, discussed in detail in Chapter 7. The Octocorals exhibit polytrophlsrn, obtaining energy from zooxanthellae are autotrophic and provide the host multiple sources; plankiivory and autotrophlsm are the animal tissue with carbohydrates and some proteins. two major sources. Polyp nitrogenous wastes and C02 are used by the Reproduction is generally dioecious. Colonies (sohi) suhstrdtc Juvende \sexually ~rnniature)character- nstics art' not sgn~fl~antiydifferent from the adult's. Octocoral fauna In shallow (<30 111) southeast Elorida Greatest mortahty occurs riurlng larval and juvenile reef commur~ities(Bayer 196 1, Opresko 1973; stages. Growth proceeds by asexual buddmg of polyps Wheaton 198 1, in preparation b) and 2s determinant. Octocoral growth rates have not been intensely studied. Kmzie (1974) reported that the black sea rod (Piexaura hornornuNa) exhibited colony Species Patch reef Bank reef growth of 1040 mm annually. The study also noted that sexually mature colonies were 25-35 rnm in height. Kinzie's study was in the Cayman Islands but would Urwrclciri urbcrttnurn x s generally apply to Flonda populations. Icilrgorglu jc'hrarnrn I x x Octocorals suffer high mortality from storms Erj~thropc)li~utni'urrt~ucorton x x when wave surge is too great for the holdfast or the sub- I'lrxuura h~trnottzullu x s strate itself becomes tlislodged. The colony 1s often car- I' jlrrtcosrr s x ried off the reef proper and recovery after dislodgement I%rcudtrplc~~ritirupnrosu x Y IS frequently unsuccessful. 1'. j7ilgttu<)$li X .Y [Iens~tyof octocorals ranges fronl very dense to P. \~fuge~i~xurt x Y sparse, dependent upon the habitat. vanabhty is quite i', sructs Y .Y high. Work at Blscayrle Ndtional Park, for example, IAr rr zcc.~~x~lrrr t=r~ Y Y documented a range of 10-50 colonies wrthin a square x E. !UJI!U s meter. Both Wheaton (in preparation a) and Opresko E. Ffi 3 The fireworrn Ncrmod~cecurutlcu- hrriz {Plate 18b) preys on many release sperrrr ~srotke wster ct:ltl:nn: f!owtv=i.cr. !c:tiii;-a- corals including d nuinber of reef tivrr anrf c~tlhryologicaldevc1u~)rnent are rri:erna! Plan- uctocorats. irla larvae are relcast:,i rhrou& the polyp ~troutlr,I'ltosc sp~ciesstudied spaun during the spring, sunlrncr. and 3. f'ypirottru spp. gastropod rnolluscs fall. Recruitment cri nrtv ci~lorlicsresults trurri Iimrval feeti. sornetmles: in groups, on octo- settiernenf Ifc~llowlrxg3 brief planktonic perloti. "xlctd- corals. Gor,qonru vcrtral~t~a1s a rnorphosis occurs after thc farvac scttlc on appropriate favored prey. 4.2 Table 26 5. Nu~~lerousfish have been observed feeding; however, they do not Southeast Florida reef Scleractinia. appear to be obligate octocorali- vores (Randall 1967). ORDER SCLERACTINIA Octocorals are increasingly harvested for human purposes. Many octocorals contain pharmacologically SUBORDER ASTROCOENIINA (Vaughan and active compounds within their tissues. Medical-phar- Wells 1943) macological research requires harvest of prodigious quantities of octocorals to isolate active compounds. Family Astrocoeniidae (Koby) Bayer and Weinheimer (1974) reported on the prosta- glandin compounds extracted from Plexaura homomalla. Stephanocoenia rnichelini (Milne, Edwards, Concern has been registered that this might eliminate a and Ilaime) species population over a wide area if harvest restrictions were not instituted. Another exploitation of octocorals Family Pocilloporidae (Gray) within the area is for live aquaria. Information provided to the Gulf of Mexico Fishery Management Council Madracis decactis (Lyman) reported 5,845 colonies of octocorals belonging to nine M. formosa (Wells) species are harvested annually and sold as aquarium M, mirabilis (sensu Wells) specimens; several are nonreef species. Current State and proposed Federal statutes only protect the sea fan, genus Fanlily Acroporidae (Verrd) Gorgonia. Acropora palmafa (Lamarck) (Plate 5a) Scleractinia A. cervicornis (Lamarck) (Plate 4b) A. prolifera (Lamarck) (Plate 5a) The Scleractmia (stony corals) are a specialized order of Zoantharia, distinguished by an aragonite SUBORDER FUNGIINA (Venill) calcareous exoskeleton. The skeleton (Plate 9a) is composed of a basal plate, external wall, and specialized Family Agariciidae (Gray) internal structures-the septa, pali, and columellae. The group in general expresses radial symmetry set in a hexa- Agaricia agaricites (Linnet) merous mode. Yonge (1940, 1973) and Wells (1957b) A. agarlcites agaricites (~inne') reviewed scleractinian biology, and Vaughan and WeIls A. agarzcites danai (Milne, Edwards, (1943) and Wells (1956) provided a glossary of terms and Haime) and definitions. A. agaricites carinata (Wells) The order Scleractinia is divided into five sub- A. agaricites purpurea (LeSueur) orders (Table 26)-Astrocoeniina, Fungiina, Faviina, A. lamarcki (Milne, Edwards, and Haime) Caryophylliina, and Dendrophylliina. Most shallow- A. undata (Ellis and Solander) water, reef-building corals are found in the first three A. fragilis (Dana) suborders, with the greatest number in the Faviina. With Iielioseris cucullata (Ellis and Solander) few exceptions taxonomic characters are based on the skeleton, especially the septa. Family Siderastreidae (Vaughan and Wells 1943) The fundamental unit of the coral colony is the polyp (Plate 7b), a tiny mass of tissue with a set of Siderastrea radians (Pallas) tentacles and a central mouth. The tentacles and adja- S. szderea (Ellis and Solander) cent connecting tissues are covered with cilia and ne- matocysts. There are three tissue layers: ectoderm, Superfamily Poritidae (Gray) rnesoglea, and endoderm or gastrodem. The endoderm of reef Scleractinia is usually filled with dense concentra- Family Poritidae (Gray) tions of zooxanthellae. The color of coral tlssue, a com- plex of plant and animal tissue, is mostly the result of Porztes astreoides (Lamarck) the plant pigments within the chloroplasts of these P. porites (Pallas) zooxanthellae. Coral tissue covers only the very surface P. porites divaricata (LeSueur) of the limestone skeletal mass. McCloskey (in Muscatine P. porites furcata (Lamarck) and Porter 1977) reported that this tissue complex was p pontes clavarLz (Lamarckl 45% plant and 55% animal by weight. Because coral P. brunneri (Rathbun) tissues contain the producer, and fixed carbon can be passed directly to the coral animal tissues for utilization without a herbivorous intermediary, the association of (contliiued) coral and zooxanthellae is successful in nutrient- impoverished tropical waters. To augment this ability, 1 abil: 1 1 able 26 (continued) '6 (eontinucd SliBC1RI)P H F.AVIIE*;A (Vaughan and Welis 1943) SI!BORI>EK C:ARYOf'IfYLLIINh (Vaughan and Wells 19/13) Superfanlily (*aryophylliidae (Gray) i.arnily i:aryophyllildae (<;ray) SIIBO 1IDL:K I)I.NI)ROl'llYLLIINh (Vaughan and Wells 1943) Family Dendrophylliidae (Gray) I3(1lunc1lilr~llicru/Zoridcrnu (Pourtales) corals have developed other methods to conserve and recycle llrrliting resources such that !ugh productivity is marr~tained.Muscatine and Porter (1977 1 reported that rccf corals are prlrnary protiucers, primary consumers, sccontlary and tertiary consumers, deposit feeders, and saprotrophs, different species have perfected different strategies. 'l'he n~ostuniisual and probably the most impor- trrrlt trophic strategy that is almost universal amortg the rcef-dwelling corals is the prinlary producer activity. 'i'hc autotroph or phototropln 1s carried within the coral tissue as an endosymbiont. Hermatypic scleractinia possess the dinoflagellate %uosutlthellu tnicrc~udr~at~cum (1-oebllch anti Siurley 1979) (also reported as Svrrrbro- ilt~ittir~for (;yrnncxfrn~um) withln their endodernnic tissues Ihe ~ooxanthellae cornplernent the corals by rucychng rtitrogen and ptlosphorus, fixing carbon (I~pids,amino acids, notlamino organic acids, glycerol, organic phosphates. and glucose), producing oxygen, cnlnancing calcification, and removing animal inetaholic products (('02.nitrogenous wastes) Muscatine (1 9731, I aylor (1 973 ), anti Muscatine and Porter (1 977) review- etf the coial host ~ooxanthellaesymbiont relationship. Muscatlile artd Porter (1977) reported that 30% of the casbo~~fixed by ttie tooxanthellae on a clear day would satlsfy all of the coral's carboti needs. t'orals are suspension feeders. Several techniques are used singularly or in con~t>inationto capture zoo- illrisscz ix~liasrrtrtzr~grllti (1)ans) rctrectcd during the day, Uendrugyru cylzndrus is an ;Idi,e,txtoplr3 /Xiit !u?ncirckr~trii(Mrlnc, I'dwartfs, escept~on(Plate ?a). Porter (1974) reported on polyp and 1lzrrrlt.l expanswn.) I'he captured zooplanktors are delivered to .I1 ~kr~larrtlar( fiidnc, I'ilwards, ~ncirialme) the mouth by the tentacles or by reverse beating of the Ai. lt*t Growth rates of scleractinian species from Florida and the Elaharnasa. b Growth rate Species (mm/~r) Location Source Acropora cervicornis Dry Tortugas Vaughan and Shaw 1916' Key Largo Dry Rocks Shinn 1966 Eastern Sambo Jaap 1974 Acropora palrnata Goulding Cay, Bahamas Vaughan and Shaw 1916 Eastern Sambo Jaap 1974 A cropora prolz fcra Goulding Cay, Bahamas Vaughan and Shaw 1916 Agaricza agaricires Dry Tortugas Vaughan and Shaw 1916 Porites porites Dry Tortugas Vaughan and Shaw 1916 Poritcs astreoides Dry Tortugas Vaughan and Shaw 1916 Siderastrea radians Dry Tortugas Vaughan and Shaw 1916 Siderastrea siderea Dry Tortugas Vaughan and Shaw 1916 Pavia fragurn Dry Tortugas Vaughan and Shaw 1916 Diploriu lubj~rintliiforrnis Dry Tortugas Vaughan and Shaw 1916 Diploria clrvosa Dry Tortugas Vaughan and Shaw 1916 Diploria sfrigosa Dry 'Tortugas Vaughan and Shaw 1916 Carysfort Shinn 1975 jlfbtrricinu rrreolatu Dry 'I'ortugas Vaughan and Shaw 1916 bfartlcina oreolu fu rnay nri Dry Tortugas Vaughan and Shaw 1916 MCJ~~US~~UCUcu~~~*rrt(isu Dry Tortugas Vaughan and Shaw 1916 &fon tcrst ru eu unrtulclris Key West Agassiz 1890 Dry Tortugas Vaughan and Shaw 1916 Carysfort Hoffmeister and Multer 1964 Carysfort Shinn 1975 Key Largo area tIudson 1981 O~.uiinada,f&siz Dry Tortugas Vaughan and Shaw 1916 X)ickocuenia st<)kesii Dry Torlugas Vaughan and Shaw 1916 Id@!t(irogyru cj~lindrzts Dry Tortugas Vaughan and Shaw 1916 lsopt~ylliu.sirtuosu Goulding Cay, Bahamas Vaughan and Shaw 1916 l:'crzm~liufusrigiura Dry Tortugas Vaughan and Shaw 1916 "c20ddhg Cay, Bahamas, data were only used when I'ortugas information was unavailable. b~ = increase in branch length, D = increase in diameter, H = increase in height. 6: Vaugl~anand Shaw's multiple values were averaged. l@z turhulcnce S'. crsrtcordes is dominant w~thD. Fragmentation also complicates the issue. During the clrvosu. ill the spur and grooves, Acropora [~ul~iafu(Plate 4-year BXP study, branching species (A. palmata, A. cer- 5a) is dominant (Table 18). In tile buttress zone M. llicornis, and !? pporztes) were frequently fragmented and asrrzubris is most slgnlficant in the reef-buildlng pro- scattered throughout the site causing major change in cesses. 'rabies 17, 19, and 20 present the species ranks, abundance and dominance. What was originally a single abutxdance, detlsitics, and cbispersion patterns for a colony in some cases was broken into 10 or more living number of reefs, note the high degree of variability fragments. The great variability over the period of this withirr and between reefs. In deep reef communities study points out that shallow patch reefs are dynamic (2 30 in) the species that: dominate are Aguricia lam- and that physical controls are significant. Table 28 arcil-I,~Valfracas mirahrlrs, Stepha~zocnrnlatnlchelinii, A. presents a time series of a representative 4-m2 plot at fi.agriis, and M. lormnscz. Ekhorn Reef. Dynamic populatron changes affect the nature of Biological interactions among the Scleractinia are coral associations in shallower reef habitats. Colonies are documented by Lang (197 1, 1973, 1980); Jackson and frequently relocated following storms. In rnmy cases, Buss (1975), Richardson et al. (1979); Wellington they do not die, but reestablish themselves in the new (19801, and Sheppard (1 98 1, 1982). These interactions habitat (Plate 10a). This makes it very difficult to are mostly involved with maintaining and/or expanding evaluate mortality in a time series within a sampling site. space, "Iebensraum" in the reef where spatial resources 46 Table 28 Time series for abundance, density, and dispersion of stony corals at Elkhorn Reef (one 4-m2 plot) (Jaap, unpublished). Abundance B. (total no. Percent of Density Species (ran king) of colonies) total colonies (xb) D.I. b Acropora cervicornis Acropdra palrnata Porites porites Porites astreoides Siderastrea siderea Millepora alcicornis Favia fragurn Diploria clivosa Yearly total Acropora cervicornis Acropora palrnata Siderastrea siderea Porites astreoides Diploria clivosa Millepora alcicornis Yearly total A cropora palma ta Acropora cervicornis Siderastrea siderea Porites astreoides Yearly total Acropora palrnata Siderastrea siderea Porites astreoides Porites porites Acropora cervicornis Diploria clivosa Millepora corn planata Yearly total a~.~.= Biological Index, McCloskey (1970). b~.~.= Dispersion Index, Elliott (1971). C = Contagious (clustered), R = Random, G = liniform, '~otethat a storm during the winter of 1979-80 fragmented a large A. palrnata colony. Survivors established a new set of colonies, thus changing dominance and density. are often a limiting factor. Rapid growth among the 4.5 OTHER BENTHIC GROUPS Acroporid species can shade out other slower growing species (Shinn 1975). Some species have potent external The level of insight into many other coral reef digestive mechanisms whereby the mesenterial filaments benthic groups is poor. Detailed examination will not be are extended and kill the tissues of adjacent corals. Lang attempted here. There are nunlerous diverse habitats (197 1, 1973) documented this digestive mechanism as within a coral reef that provide niches to a multitude of hierarchical among the species of Atlantic Scleractinia. species belonging to nearly every phylum (Plate 8b). Some species compete with this agressive territorial Sedimentary deposits provide habitat for infaunal behavior through sweeper tentacles which keep the organisms. Many mobile forms find refuge under the mesenterial filaments away from the lower hierarchy sedimentary surface and forage for food at night. 'rhe species. It is also probable that alleochemicals are reef rock itself is a habitat for both epifauna and boring important in protecting space. Biological interactions are and cryptic fauna. A single coral head weighing 4.7 kg at probably more significant in deeper reef environments Heron Island contained 1,44 1 polychaete individuals where physical events are less frequent and of an inter- (Grassle 1973); 67% of all organisms from the coral's mediate magnitude. interior were polychaetes. Vittor and Johnson (1977) Coral pathology and disease is an infant research studied polychaetes from Grand Bahama Island and area. Phenomena were reported by Preston (1950) and reported that 84 species were present. Robertson Squires (1965). Laboratory experiments demonstrate (1963), Ebbs (1966), Ilein and Risk (1975), and Hudson that bacteria cause morbidity and mortality in corals (1977) reported on the boring fauna of Florida corals. (Mitchell and Chet 1975 ; Ducklow and Mitchell 1979a, Specific references that deal with the other benthos 1979b). Antonius (1974b) reported that a blue-green include the proceedings volumes from the past Inter- alga, (Isczlkitor~a,was capable of killing corals under national Coral Reef Symposia, the two volumes from the certa~n(laboratory and field) conditions (Plate 19a). The Biology uncl Geology of Coral Reefs, Volumes 23 (1, 2) alga grew over the coral, killing it in a short period. of Bulletin of Marine Science, Frost et al. (1977), Colin (;ladfeller (1982) reported on an undetermined patho- (1978a), and Rutzler and Macfntyre (1982b). Voss et al. gen that caused epidemic mortality in A. pulmuta on a (1969) provided a qualitative account of the biota found rccf off St. Croix, U.S. Virgin Islands. A similar condi- within BNP prior to its becoming a national monument. tion is often seen on Florida reefs; Grecian Rocks The study of most other benthic organisms, save certain appears to be currently infested with this pathogen commercial species (e.g., spiny lobster), is for taxonomic (l'lati: 25a). Pathology and band diseases in corals were purposes. Decapod crustaceans were reported on by teviewcd by Antonius (19818, 1981b). Gore (1981 j. Lyons and Kennedy (1981) and Lyons et al. (1981) reported on biological aspects of the spiny lobster Punulirtis argus. Work (1969) studied West Indian mollusk communities. Kier and Grant (1965) and Kissling and Taylor (1977) discussed echinoderms from Florida reef communities. CHAPTER 5 PLANKTON by J. 0.Roger Johansson Tampa Bay Study Group City of Tampa, Florida 5.1 INTRODUCTION suminarize and critique studies of reef zooplankton and find that, due to often questionable methodologies used, Coral reefs are highly productive marine ecosys- no conclusive data on the relative importance of zoo- tems with a great abundance and diversity of organisms. plankton to coral reef nutrition have yet been published. They are generally surrounded by waters with small Few studies have been conducted describing the plankton standing stocks. Most studies describing coral free living phytoplankton community in the vicinity reef plankton communities have investigated the impor- of coral reefs. Researchers generally agree that phyto- tance of zooplankton in coral nutrition. ?'he results of plankton's contribution to the primary productivity of these studies have been somewhat contradictory. the coral reef ecosystem is small (e.g., Hargraves 1982). The early studies (Yonge 1930 and others) Further, Wood (1 954) suggested that the waters of the generally indicated that there was not a sufficient Great Barrier Reef might have a distinctive reef phyto- amount of zooplankton in the water coluliln around the plankton community. Subsequent studies (Jeffrey 1968; reef to support the corals. Recent studies of coral reef Revelante et al. 1982), however, found a reef commun- zooplankton (Emery 1968; Porter 1974; Sale et al. ity similar to the nearby ocean phytoplankton com- 1976; Alldredge and King 1977; Porter et al. 1977; munity. The following discussion of coral reef plankton Hobson and Chess 1978; Aamner and Carleton 1979; will, therefore, mainly be concerned with the zoo- Hobson and Chess 1979; Rutzler et al. 1980; E-Iamner pIankton community. and Hauri 1981; McWiUiams et al. 1981; and Olllhorst 1982) have shown that methods used in the earlier 5.2 TAXONOMIC COMPOSITION AND SPATIAL studies for capturing the reef zooplankton were not DISTRIBUTION adequate to describe the zooplankton community most nutritionally important to the corals. These plankton As discussed above, several researchers have live in close association with the corals, sometimes described coral reef zooplankton communities in great within the reef itself. detail. However, due to geographical considerations, the The earlier collections used nets which were following discussion will generally be limited to observa- towed or drifted above and sometimes away from the tions of Alligator and Looe Key Reefs in Florida by reefs. The recent studies, using an array of different Emery (1968). The information obtained in these collectioli and observation methods such as stationary studies is somewhat limited due to the lack of quantita- nets, diver-controlled nets, plankton pumps, diver- tive abundance measurements; however, the report gives operated suction devices, emergence traps, light traps, a detailed account of the spatial and temporal distribu- and diver visual and photographic observations (see tion of the different zooplankton taxa found on the Porter et al. 1978; Hamner and Carleton 1979; and reefs. Emery used several different collection methods, Youngbluth 1982 for discussion of sampling tech- including boat- and swimmer-towed nets, suction niques), have found that coral reefs, worldwide, have an devices, light traps, and diver observations, in order to abundant and diverse resident near-reef zooplankton reduce sampling errors of zooplankton living near or community different from the oceanic forms in the within the reefs. Further, he made collections and surrounding ocean. Porter et al. (1978), summarizing observations during both day and night. Iie also subdi- information from mostly indo-Pacific reefs, estimated v~dedthe study areas into several habitats such as grass that 75% of the reef zooplankton standing stock was of beds, lagoons, patch reef, reef tops, caves, and deep benthic origin and that 85% of the biomass was of local reefs. VlsuaI observations by divers, swimmer-towed origin. Parts of this community are, therefore, often not nets, and suction devices produced the most interest~ng considered as truly planktonic, but rather as epibenthic information about the zooplankton on and i.11 close or demersal. Many organisms in the resident comlnunity vicinity of the reefs. are only found in the water column at night and spend Close to the water surface of all areas of the daylight hours on the sediment or within the reef itself reefs, Emery found a typical offshore zooplankton conl- (see Section 5.3). munity, consisting mostly of free living copepods and It is still not clearly demonstrated, however, larvaceans. Visual observations deeper in the water what importance zooplankton play in coral nutrition. column and just above the bottom otten found the reef Emery (1968) states that plankton represent a major plankton in close and dense aggregations or swarms. The source of food lo the reef, while hluscatine and Porter swarms tended to hold together arid move as a unit, and (1977) suggesr that zooplankton feeding does not they also maintained their position agamst the current contribute a majority of either calories or carbon re- and the surge. Emery found four species of copepods m quired for reef corals. Hamner and Carleton (1979) these swarms. Acu~frotonsa, A. spznata, Ozthorza nana, and 0. oculata. He estimated that one swarm of A. were most often captured at night included: fish larvae; spinata contained 1 10,000 copepods/m3. fiamner and large copepods; crab larvae; chaetognaths; ostracods; and Carleton (1979) also reported extremely dense swarms shrimp larvae. Ohlhorst (1982) found that the zooplank- of copepods (max. 3,320,000/m3) on Australian reefs. ton emerged from the substrate of Discovery Bay, Swarming species of copepods, in the Emery study, were Jamaica, at variable rates throughout the night. The peak most often found on the grass beds and reef tops. activity was usually two hours after sunset. However, Monospecific schools of mysids, of one size class, were smaller species of copepods and juvenile members of also found on the the reef tops often inside caves and large species migrated into the water column during the along coral faces. Schooling implies a stronger social day and were least abundant in the water column during behavior than swarming since the individuals that belong the night. She suggested that the reasons for die1 vertical to one size class are longitudinally aligned and move as a migration are many and varied. They include: feeding; unit. Schooling behavior of mysids has also been re- reproduction; escape from predation; and dispersal. ported by other investigators of Alligator Reef (Randall Porter et al. (1977) studied reefs in the Pacific Ocean et al. 1964). On the deep reefs and the patch reefs of and found that the volume of emerging plankton was both Allgator and Looe Key Reefs, close aggregations of greatest over branching coral and least over sand and the plankton did not appear as important. Copepods and rubble. Alldredge and King (1977) and McWilliams et al. mysids were found individually or in looser aggregations. (1981), sampling different reefs of the Great Barrier Emery also observed that the swarming and schooling Reef, also found a positive relationship between the behavior of these organisms was mostly a daytime structural heterogeneity of the substrate and the amount phenomenon. They were often found individually at of epibenthic plankton emerging at dark. The composi- night. Hobson and Chess (1979), reporting on resident tion of the community Alldredge and King found was reef plankton of Hawaiian atolls, found a similar diurnal similar to the one described by Emery (1968), and they pattern for swarms of copepods, mysids, and larval estimated that an average of 2,5 10 zooplankton emerged fishes. per m2 of the reef substrate. McWilliams et al. (1981), Swarming and schooling in dense aggregates are conducting a more detailed study, found not only a important examples of behavior adaptation by resident greater volume of plankton emerging from the coral than reef plankton, which differentiates them from pelagic from the sand but also that the composition of the rising forms. Hamner and Carleton (1979) stated that at least communities were different. The reef bases were transi- seven species of copepods, and probably more, engage in tional zones with a mixture of "coral" and "sand" swarming behavior at coral reefs in three of the world's communities. These authors further found that more oceans. The same authors have suggested that swarming plankton emerged during the summer than during the not only reduces predatory pressure, but also facilitates winter. The annual range of the "coral" fauna was reproduction and permits the plankton to cluster in local 5,030-2,350 animals/m2 and the "sand" fauna 2,720- eddies to maintain a favorable feeding position with 1,150 animals/m2. Also, the plankton samples collected minimum energy cost. In a subsequent paper, Hamner during the summer were generally more diverse in and tiauri (1 98 1 ) related the resident reef zooplank- taxa than winter samples. Hobson and Chess (1979), ton distribution to the reef configuration and the local collecting emerging plankton from the lagoon substrata water motion. They found swarms of copepods and of atolls in the Pacific Ocean, found a considerably less chaetognaths directly upcurrent of the reef face in an abundant plankton community in their traps than entrained water mass, which was subject to less removal reported by Alldrege and King (1977) and McWilliams et than downstream waters. This accumulation of biomass al. (1981). Hobson and Chess (1979) suggested that the and incrcased primary productivity in areas upstream of use of different types of traps may account for some of reefs is one aspect of the well-known "island mass the variations reported between study areas. Youngbluth effect" phenomenon. (1982) tested three types of emergence traps and found that both density and diversity of the zooplankton 5.3 DIURNAL MIGRATIONS collected were affected by the trap design and sampling procedures. He concluded that different mesh size Investigators of cord reef zooplankton who have netting in the traps alone could probably account for taken care to sample the populat~onsnear the substrate much of the variability in plankton abundance and and on a diurnal schedule have found the plankton diversity between different study areas. community undergoes drastic daily changes. Emery (1968) found that nighttime collections contained four 5.4 SUMMARY times greater volume than daytime collections. A suction device with a light close to its mouth to attract the Thc coral reef plankton community can be plankton was used fox thesc collections, and thcrcfare, characterized as follows: quantitative comparisons are not in order. Emery did, however, find a very different composition of the 1. A highly abundant and diverse plankton at night. Many forms, such as polychaetes, zooplankton community is cumaceans, and zoea, which he observed living in caves found on the reefs. It is in and crevices during the day, were swkmmg outside the striking contrast to the open reef in great abundance at night. Other forms which water community nearby. 2. Most of the reef zooplank- 4. The major portion of the resi- ton belong to a resident com- dent plankton community con- munity, which is able to main- sists of epibenthic forms. This tain position on the reef. community generally migrates 3. The community consists partly from the substrate to the water of true planktonic forms living column at dark. in dense aggregates and using the 5. Phytoplankton communities reef configuration and water cur- found over the reefs appear to rents to maintain position. be little different from open ocean assemblages. CHAPTER 6 REEF FISH by James T. Tilmant U.S. National Park Service South Florida Regional Research Center Tropical coral reefs are richer in fish species than new males are rapidly developed from female stock by any other habitat (Marshal 1965; Emery 1973). The fish some indiv~dualsundergoing a protogynous sex reversal. fauna of the Flonda reef tract is wholly tropical Egg productioi~is maximized by this mechanism and a with only seven species that have not been recorded else- greater proportion of energy channeled into population where in the West Indies. Thus, although the study of maintenance (Goldman and Talbot 1976; Smith 1982). coral reef fishes in south Florida has not been extensive, Florida reef specles known to undergo sex reversal considerable information concerning their biology and include most (if not all) parrotfish (Scaridae), many ecology has been developed through studies elsewhere in wrasses (Labridae), and some groupers (Serranidae). the Caribbean. A pelagic larval stage is almost universal among The most complete studies conducted in the coral reef fish specles. Little detailed information is Florida Keys are those by Longley and Hildebrand known about the life of larval reef fish while in the (1941) for the Dry Tortugas, and Starck (1968) for Alli- pelagic phase, although they are frequently taken in off- gator Reef. Field guides to Florida reef fish include shore plankton tows. Studies in which the time of Chaplin and Scott (1972); Greenberg (1977); Stokes spawning and time of period of peak recruitment of and Stokes (1 980); and Kaplan (1 982). juveniles from the plankton have been correlated indi- 'I'hree major reviews concerning the general cate that the larval stage varies from a few weeks to ecology of coral reef fish on a worldwide basis are months depending on the species (Randall 196 la; Moran I:hrlick (1975), Goldman and Talbot (1976), and Sale and Sale 1977). Aging of larvae or newly transformed (1980b). juveniles by counting daily growth increments on the otoliths has revealed that pomacentrids, gobiids, and 6.1 REPRODUCTION AND RECRUITMENT blenniids have larval lives of about 21 days. The larval stage of siganids, acanthurids, chaetodontids, and labrids Most caral reef fish spawn in the water colurnn is 30 or more days (Sale 1980b). As is common among above tfic reef, arid their eggs rcrnain in the plankton many marine invertebrates, many species can probably during development. A few species either lay their extend their larval life until suitable settlement habitat is eggs dernersdly and attend to them or they brood found (Randall 196 la; Johannes 1978; Sale 1980b). thcm by inoutti (Sm~th 1982; Colin 1982). Breeti- The methods by which larval fish are returned to ing aggregations are common to ~aostreef species. These a reef environment have been mostly hypothesized from serve to increase egg fertihzation and release of egg and circumstantial evidence. Generally, it is believed that sperm m the Ice of the reef, thereby increasing disper- current gyres that form in the lee of islands and reefs act sion and reducing predation (Colin 1982). Coral reef as traps to prevent the loss of larvae downstream and, fishes typically produce large numbers of offspring through their rotation, return the larvae to the reef which are then dispersed by means of a pelagic larval (Munro et al. 1973; Leis and Miller 1976; Johannes stage (Ureder and Kosen 1966; Bllrlich 1975; Johannes 1978). Johannes (1978) reported that the spawning 1978; Sale 1980b). The timing of egg release (or hatch- migrations, which bring some species to particular sites ing for those farxlilies with dermersal eggs) is often at precise times, release the gametes into a mass of water tnggrreti by dusk, nlght, or the tides, increasing the that is most likely to return them to the same reef at the ability of newly spawned or hatched eggs to elude end of their pelagic existence. Iie cited the variability in predators (hlyrbrrg X 972, Roberlson and Choat 1974; spawning peaks between various locations, often within Warnex el al. 1975; Bell 1976; Cohn 1976, 1978b, the same region, as possible evidence that spawning 1982, Moycr and Bell 1976: Jaliannes 1978; Lobe1 patterns have arisen as responses to a variable pattern of 1978, Moyer and Nakazono 1978). water circu!ation. The season in which spawning occurs Most species of reef fish spawn several times is not well known for Inany coral reef species, but it is durmg their spawning season. Spawning may be daily, known some species spawn on a regular basis all year biweekly, or monthly (Robertson 1972; Holzberg (Coh 1982). For most species that have been studied, 1973, Warner et ai. 1975; Moyer and hakazono 1978; spawning peaks in the late afternoon and may last from Lobel 1978, Ross 1978) These multiple spawtringq a few minutes to an hour or more depending on the have been explained as a method of ensuring a wide species (Colin 1982). Present knowledge concerning the dispersal of offspring (Sale 19778). reproductive behavior of coral reef fishes in the Carib- Sex reversal is also coinmon In many reef species. bean has been summarized in papers by Smith (1982) Often the adult popuiations are largeiy females (with and Coh(1982). only a small but sufficient percentage of males). If the Seagrass beds are important to the reproduction number of males in the populat~onbecomes reduced, of coral reef fishes m that they act as nurseries for some S species. Ogden and Zieman (1977) reported that at and Costa Rita. This habit was not observed by Randall Tague Bay, St. Croix, juvenile spiny puffer (Diodon), (1 967) during extensive studies in the Caribbean; Finckh squirrelfish (Holocentrus), yellowtail snapper (Ocyurus (1904) in the Ellice Islands; Wood-Jones (1 9 10) at Cross- chrysurus), surgeonfishes (Acanthurus), and numerous Keeling Island; or Choat (1 969, cited in Goldman and wrasses (Halichoeres) were commonly present within the Talbot 1976) on the Great Barrier Reef. Randall (1 974) grassbed. The spotted goatfish (Pseudupeneus macu- did observe coral feeding by scarids in some areas of the latus) and the yellow goatfish (Mulloidicthys martinicus) Pacific, however, and discussed how this habit appears to occur as juveniles in grassbeds off the Florida Keys vary among the same species of fish at different loca- (Springer and McErlean 1962a; Munro 1976). Springer tions. Scraping of massive (head) corals by parrotfish and McErlean reported capturing eight species of juve- does occur at least occasionally on Florida reefs (W.A. nile scarids in Thalassia beds off Matecumbe Key; all but Starck 11, personal comment in Randail 1974; J.T. one, Sparisoma radians, are considered reef species as Tilmant, personal observation). The amount of food adults. These fish remain in the seagrass habitat until taken in this way may be insignificant compared to algae they become too large (> 20 cm) to hide among the ingested by grazing. blades (Ogden and Zieman 1977) and presumably mi- There are few true omnivores among most reef grate to the reef. fishes. The majority of species tend to be substantially more carnivorous than herbivorous. Most herbivorous 6.2 FOOD HABITS AND TROPHIC STRUCTURE fishes have been found to take only a small amount of animal matter. Among the Caribbean species reported Fish constitute a major portion of the animal as omnivorous are the damselfish (beaugregory and the biomass on a coral reef, and they are an important com- threespot damselfish); the grey angelfish and French ponent of the overall trophic structure. Most reef fish angelfish; the scrawled filefish, orangespotted filefish, are carnivores and many studies have shown that carniv- and fringed filefish; and the sharpnose puffer (Randall orous fishes normally represent three to four times the 1967). biomass of herbivorous fishes (Bakus 1969; Goldman Schools of small zooplankton-feeding fishes are and Talbot 1976; Parrish and Zimmerman 1977). This common on coral reefs and zooplankton is an important reversal of the "traditional" biomass pyramid results energy source for larval stages of most reef species. from many carnivorous fishes feeding on invertebrates Among adult zooplankton feeders commonly schooling that form the largest portion of lower trophic levels on on reefs are the herring genera Harengula, Opisthonema, reefs. Benthic invertebrates are of considerable impor- Sardinella, and Jenkinsia; the anchovies (Anchoa); and tance to reef fish populations as energy assimilators at silversides (Atherinornorus and Allanetta). Basslets the planktivore, herbivore, and detrital levels (Vivien (Grammidae), cardinalfish (Apogonidae), glassy sweepers and Peyrot-Clausade 1974). (Pempheridae), and chromis (Chromis spp.) also typi- Herbivorous reef fish are sustained primarily by cally occur in relatively large schools among reef crevices low algal turfs and low-growing filamentous algae and where they feed on plankton. In addition to these adult diatoms (Goldman and Talbot 1976). No phytoplankton planktivores, dense schools of minute larval grunts feeders have been identified in any of the reported (Haemulidae), snappers (Lutjanidae), and wrasses (Labri- studies of reef fish food habits (Hiatt and Strasberg dae) are often quite common hanging above coral heads 1960; Randall 1967; Bakus 1969; Hobson 1974; Gold- or among branches of Acropora where plankton and man and Talbot 1976; Panish and Zimmerman 1977; coral mucus are the only available food. Hobson and Chess 1978). This is in contrast to fish Most of the carnivorous reef fish appear oppor- populations in temperate regions where up to 6% of the tunistic, taking whatever is of the right size and is catch- species may consume phytoplankton (Davies 1957). able within their habitat (Goldman and Talbot 1976). Benthic algal feeders include most parrot fish (Scaridae) This conclusion is based primarily on the variety of food and surgeonfishes (Acanthuridae) and some blennies items observed within the same species at various Ioca- (Blennidae), damselfish (Pomacentridae), butterfly fishes tions. OnIy a few studies of reef fish species have evalu- (Chaetodontidae), and filefish (Balistidae). A number of ated food consum~tionin relation to what was actually studies have shown that the herbivorous reef fish have a available: Jones (1968); Vivien and PeyrotCkusade marked effect on the distribution and abundance of (1974); and Hobson and Chess (1978). aIgae present on a reef. When protected from grazing by Some carnivorous reef fish are highly specialized exclosures, algal communities wiU rapidly increase in in their feeding habits. Among these are the parasite standing crop (Stephenson and Searles 1960; Randall pickers or cleaners (Labroides spp.), juvenile bluehead 196la; Earle 1972a; Montgomery 1980; Montgomery et wrasse, and neon gobies (Cobiosuma spp.), and several al. 1980; Nixon and Brostoff 1981). sponge feeders. Caribbean reef fish reported to feed on The extent to which zooxanthellae is eaten by sponges include butterfly fish (Chaetodontidae), spade- reef fish has been disputed (Randall 1974). Parrotfishes fish (Ephippidae), puffers (Tetraodontidae), and' box- (Scaridae) have been reported to scrape coral in a feed- fishes (Qstraciidae) (Menzef 1959; Bakus 1964; Randall ing manner by Hiatt and Strasberg (1960) in the Mar- and liartman 1968). shall Islands; Motoda (1 940) in the Palao Islands; Talbot Few, if any, reef fish have been classified as detri- (1965) off East Africa; Al-Hussaini (1947) in the Red tal feeders and the utilization and turnover of detritus Sea; and Bakus (1 969) along the Pacific coast of Panama on coral reefs appear to be left primarily to bvexte- 53 brates. Only a few biennies (none occurring in Flofida) nocttlrnal lredcls Inak contribute significantly to the have been reported to be primary detritivores (Cioldman energy budget of the cura! reef through the import of %nd 'Tdbol: 1976). "This number may increase as more organic matter in the form of feces. If so, their contribu- fwd studies for the numerous snraller blennies and tion to reef nutrient lavels and maintenance of Inore gubies become available. Also, much detntal material is sedentary fish abundance may also be important. Meyer ingested by those fish classified as herbivores, and et al. (1983) found what appeared to be higher nutrient detcrnlining actual materxal being foraged is often levels and coral growth rates associated with diurnal difficult (HandalI 1907 1. resting schools of grunts. Verification of these links, Many species utilize the reef only as a refuge. however, requires further investigation. They use adjaccnt seagras beds and sand flats as feeding Use of adjacent grass and sand flats by reef fish grounds (Longley and kfildehrand 1941; Randall 1963; is not strictly a nocturnal phenomenon, but seems to be Starck and lZavis 1966; Davis 1967; Ogden and hhrlich the dominant pattern. Only quite large herbivores, such 1 97 7; Clgden and Zienan 1977). KandaIl(1963) report- as the rainbow parrotfish (Scarus guacarnaia), venture far ed that punts and snappers were so abundant on patch into the grass bed away from the shelter of the reef. Mid- reefs in the Ih-xla Keys that it was obvious that the %zed herbivores are apparently excluded by predators reefs could nor provide food and possibly not even and feed only near the reef (Ogden and Zieman 1977). shelter for all of tttcsa. Both the abundance and diversity Randall (1965) reported parrotfishes (Scarus and Spari- of 1.l;xernulicIs and iutjanids have been reported to he re- soma) and surgeonfishes (Acanthurus) feeding on sea- duced on reefs where acictluate off-reef forage areas are grasses (Thalassia and Syr~rlgodiu,n)closely adjacent to unavailable (Starck and Davis 1966; Kobins 1971; Ale- patch reefs in the Virgin Islands during the day. We vizorr and Brooks 1975, Gladfelter ct al. 1980). Smi- attributed the formation of halos around patch reefs in liarly, whcn recf ehcller ~slacking, what appears to be St. John, Virgin Islands, to this grazing. s~zktublcgssutxctl forage areas may go unusetl (Starck and Davis 1966; Ogdcn rnd Xiernan 1977). 6.3 MOVEMENT In Tggue liay, St. C:rcsrx, V1r6un Isiarids, 79.4'%, of tfrs fii;lres actlveiy feeding in the prdsl?cds at night were Most reef fish seldom change their residence once species tfut scst~giltshlrltcr on ahaccnt coral reefs during they have settled onto a reef and some become quite the day (Knbhlcc atrd Zterx~an,kn pret?aration). Starck sedentary ('I'hresher 1977). Many smaller species, such as and X)avts (1Ybb) l~stedspe~ics of the liolocenrridae, gobies (Ciobiidae), blennies (Blcnniidae), and damselfis11 Lutjrrrnictiic, stid Ilacrrttlbolae families ils occurring d~ur- (E'omacentridae), maintain relatively small territories faally un Alli~torKcaf and feeding nocturnally In aarlja- which they vigorously defend (Salnzon et al. 1968; ccnf sengrass bctl?;, E ypirally both juvenile and adult Kasa 1969; Law 197 1; Myrberg 1972; Vine 1974). 'rag- Zla~rriulitiss~md lutjaizide form hetcrotypic rating sciluoks ging studies have shown that even those larger, wider over prornknent coral. txcudr ($%lostcommonly A croporrt ranging species, such as serranids, lutjanids, haemulids, i?crlniarul arlJ i)c~nfrrytorifrf) OX lit caves or crevices of tllc chaetodontids, and acanthurids, remain, at least diur- rthrl' (f:hrhck artd klutruh 1973; Clyden and ktirhch nally, in a fairly circuxnscribed place on a reef (Bardach 1977). tat Junk ttrcso Crsi~es irrove off the reef illto 1958, Randall 1961b, 1962; Springer and McErlean atij:rr;erit seagiuss bods and sand flats where they feed on 1962a ). Randall (1 962) recaptured some serranids and a~ivsi.iut~ic.mvs.rtcbtrra-rrles (Kaa~drili 1967; Olden and bfirlich luyanids at their initial place of capture up to 3 years Iq73; Mci'usXlttid ct a1 1Y7i?1). At dawn they return to after then release. 'Tagged Naernulon plurniert were t tae reef. repeatedly captured on the same reef by Springer and Starch arrd i)avts ( 196-1) reported that 1 1 of IS hilcErlean (136Za). When the If. plurnicrr were trans- hpacads of haexttuifds frllrnd in dturnrrl rcstmg scttools on planted, they exhibited a tendency to home (Springer Allfg3101" Kesf disilvrsed at z~rghtlo feed. The ltgllter and McBriean 1962a). Ogden and Ehrlich (1977) re- ~010~~4fmlnls (seven spadgsi generally distributed ported on the sucessful homing of If. pplurniert and Ii. titcrnselves alrjtrp, ssrtdy to thln-~rassareas of ?!kc back flrtvvirrieuru~n to original patch reefs over distances as rvef tone Sns$jrcrs (Lutjanitfas) fullowcd a srrvlllar pat- great as 2.7 km in the U.S. Virgm Islands. [err1 with [, gr.tscu.5 itrld I,. synugrrr in spsrst: was to Only a small amount of information is avaikable i~sckrvcf h~hltrtl.At Tagtie li3ay, St. Croix, the nocttirrlai on the usual extent of nocturnal movements of reef dis*trntluCiorn of gux~tsover the pasbeds was qultc sirni- fishes that fecd on adjacent gass beds. As indicated lar to that raportcd in the I'lurida Keya, fklc f:rench abuvc. these fishes potentially can range quite far frorti gunt was nrost dbundatat over thin grass or bare sand their diurnal restuig sites. Lutlat2us grzseus and fiacnzu- w?zile wfutc wrxis were almost always observed by Rub- Ion j7avcrlrnrutuirz ranged as far as 1.6 km froin Alligator blcc ( in preparation ), tmn thick gass . ifc also found tfic Reef (Starck and Davis 19661, while H. plumieri and ntrrrrbcr of ettra: red Fx~!~cs feedulg nosfumAIy over a V fhrotit:egturri typicslfy migrated distances greater 3re:x was posili\-t:lp correlated witli a measure than 100 rn over the grass beds in ?'ague Bay, St. Croix of hab1t;nt coarmpfexity, iri~jtiyrapsoltte for113 of arganl- (Oficrt'i~and Ehrlich 1977; Ogden and Zieman 1977). taiictn <)f tl~f~f~ assemblage whxll: active over the I'he mlgratitzg schools follow a linear path for 2040 m feeding ground. from the reef. and then be& to break up into sinaller It has beem hypoth6~l~cdBy BtUlrlm and lllurlro and snlaller groups in a dendritic pattern (Ogden and f 1974) and Ogden iind Liemail (1977) titdr mwating xianan 1977). I'he distance traveled to feed is undoubt- 54 edly related to the abundance and quality of food avail- groups (Buckman and Ogden 1973 ; Ogden and Buckman able within the seagrass bed and can be expected to vary 1973; Alevizon 1976; Itzkowitz 1977). considerably among locations. 6.5 ECOLOGICAL ASPECTS OF REEF FISH 6.4 SOCIAL ORGANIZATION DIVERSITY Literature on the social behavior and organiza- Only two studies have attempted to fully define tion of reef fish assemblages is extensive (see Ehrlich the diversity of fish species on selected Florida reefs: 1975; Goldman and Talbot 1976; Sale 1977, 1980a; Longley and Hildebrand (1941) and Starck (1968). Thresher 1977; and Emery 1978 for reviews). It has Longley and Hildebrand provided a systematic account been conclusively shown that individuals or groups of of all fishes they captured or observed during 25 years of individuals are not scattered randomly about the reef. investigations at the Dry Tortugas, and Starck listed Rather, their distribution reflects topographic and fishes collected and observed during 9 years of study at biologic features of the environment as well as behav- Alligator Reef. Longley and Hildebrand listed 442 ioral adaptations to disperse (Thresher 197 7). The species, 300 of which are closely associated with coral interaction of ecologically evolved behavior and the reefs. Starck listed 517 species, 389 of which he con- availability of existing resources largely determine the sidered reef species. The remaining species were either social organization within a given reef fish community. offshore pelagic forms, demersal species from deeper Most smaller reef species are either herbivorous water, or strays from adjacent inshore areas. Both lists or planktivorous and are territorial to varying degrees. are from single reef areas and, therefore, probably under- The size of territory and the extent to which it is de- represent the actual diversity of fish species on Florida's fended is a function of resource availability for many reefs. species (Low 197 1; Thresher 1977; Sale 1980a). Thresh- Bohlke and Chaplin (1968) identified 496 fish er (1976) found that the size of area defended by the species within the Bahamas and adjacent waters. About threespot damselfish (Pomacentrus planifrons = Eupo- 450 of these species are known to occur on coral reefs mancentrus planifrons) on Florida reefs varied with and probably approximate Florida's total diversity. the type of resource secured and further that the size of Several less extensive surveys of fish on Florida the area defended against a given intruder was deter- reefs involved visual census, filming, or limited collect- mined by the threatened severity of the intrusion. The ing techniques. One of the earliest surveys was done by size of territory defended against other families, for Jordan and Thompson (1904), who identified 218 spe- example, correlated with the amount of bentkc algae in cies inhabiting the reefs at the Dry Tortugas by using the diet of each species. Thresher (1977) reported that baited lines and various nets. the threespot damselfish in the northern portions of its The high diversity of fish on Florida's tropical range generally defended larger temtories than those in coral reefs is exemplified by a few studies which report- the southern portion, where thicker algal lawns develop- ed the number of species observed within a given limited ed within their territories. area. Bohnsack (1979) recorded a mean number of Differences in social structure have been ob- species ranging from 10 to 23 on isolated natural coral served within a single species at different locations. heads less than 330 x 210 x 150 cm in size off Big Pine Itzkowitz (1 978) and Williams (1978), working at differ- Key, Florida, and 13 to 20 species on small artificial ent sites in Jamaica, reported different social organiza- reefs (160 x 60 x 80 cm) that he established. Alevizon tions for Pomacentrus planifrons. Itzkowitz found and Brooks (1975) observed an average of 14.7 fish males and females occurring together as neighbor- species during 2.75-min samplings during which scuba ing territory holders in the same habitat, while Williams divers took color movies. During the sample period the reported the use of different habitats by the sexes. In diver panned the camera to include most or all fishes other regions, Robertson (1972) at Heron Reef (Great sighted within 4-5 m (Alevizon and Brooks 1975). Barrier Reef) and Potts (1973) at Aldabra Atoll reported At present, there appears to be only six or seven Labroides dimidiatus occurring as males with harems species of reef fish that might be considered endemic to and as mated pairs, respectively. Barlow (1974) found a the U.S. continental reefs. Two of these, Lythrypnus correlation between the social structure exhibited by phorellus and Gobiosoma oceanops, are small gobies; the Acanthurus triostegus and the presence or absence of latter (neon goby) is common. The purple reef fish various other territorial acanthurid species. The extent (Chromis scotti) is also known to occur only within U.S. to which the nature and distribution of food resources waters, although the genus is well represented by at least can predict the form of space and resulting social organi- three other species throughout the West Indies. Ophi- zation used by territorial species was demonstrated by dion selenops, the mooneye cusk eel, is a species report- Thresher (1977) for five species of pomacentrids com- ed by Starck (1968) to occur occasionally within the mon on Florida reefs. Invariably, the organization pre- area of Alligator Reef (near Matecumbe Key), but has dicted closely matched that observed in the field. not been reported elsewhere. Cusk eels are curious elong- The large reef fish species require a greater food ate fishes, highly nocturnal and burrow-ing into mud supply and, therefore, tend to range over a larger area of during the day. At least five species of cusk eels, repre- the reef. These species, although often having distinct senting two genera, are known to occur dong the ranges, are less territorial and frequently form foraging southeastern United States, but the taxonomy of this group is not entirely clear. Two small tropical serranids also influence species diversity, although this is hotly have also been considered endemic to the continental contested (Sale 1976, 1980a; Sale and Dybdahl 1978; United States. One of these, the blue hamlet (Hypoplec- Smith 1978; Talbot et d. 1978; Brock et al. 1979; trus gemma), may only be one of many color morphs of Ogden and Ebersole 1981 ; Sale and Williams 1982). The a single species, H. unicolor, which is common through- controversy centers around whether reef communities out the Caribbean (Tfiresher 1978; Graves and Rosen- have an ordered structure with predictable species re- blatt 1980). The other serranid is the wrasse bass (Lio- cruitment to vacant niches or an unpredictable chance propomu eukrines), which ranges northward along the species recruitment. An unpredictable larval recruitment continental shelf, but is not known to occur elsewhere in would increase local variation in the species present and the Caribbean. The remaining possible endemic species is might well contribute to a higher overall diversity (Gold- the cubbyu (Equetus umbrosus), but its distinction from man and Talbot 1976). Recently some investigators have E. ucuminatus, found elsewhere in the Caribbean, is suggested that differences in recruitment and communi- questionable (Robins et al. 1980). ty structure may be a result of the size of the reef area The reason for high diversity of fish species on considered (Ogden and Ebersole 198 1) or of the time coral reefs is frequently debated and may be related to a interval between which samples have been taken (Bohn- number of factors (Sanders 1968; Goldman and Talbot sack, in preparation). 1976; Smith 1978; Talbot et al. 1978). All biological communities tend to diversify through colonization over 6.6 COMMUNITY DESCRIPTlONS time, and coral reefs exist in an environment generally without major perturbations or great temperature Although fish diversity is usually high on coral change. Most coral reefs have had long and relatively reefs, fewer number of species will appear abundant or stable periods to develop. Within this overall long-range highly obvious to the average reef diver at any one stability of the tropics, intermittent moderate disturb- location. Reef fish, like most organisms, have depth ances from weather events occur. Although we are just and habitat preferences where they can typically be beginning to understand the effects of disturbance seen. Many are active only nocturnally or diurnally, and events (Endean 1976; Bradbury and Young 1981; some species, such as yellowtail snapper and gag grouper Pearson 198 1 ; Woodley et al. 198 1; Davis 1982; Porter (Mycteroperca microlepis), are abundant only season- ct al. 1982; Roberts et al. 1982), they may help to ally. maintain a higher diversity by preventing a resource- A comparison of three Florida reef studies, all of limited equilibrium and the competitive exclusion of which used the same 50-min visual census technique, some species (Connell 1978). shows significant differences in reef fish communities at Coral reefs generally include a variety of micro- each study location (Thompson and Schmidt 1977; habitats related to zones of coral growth, wave exposure, Jones and Thompson 1978; Tilmant et al. 1979). A1- and reef stnlcture (Wells 1957a; Maxwell l968; Stoddart though the top six families comprising the fish com- 1969). This diversity of habitat types allows for an munity in terms of abundance were the same, their rela- increased diversity of fish species. Although habitat tive ranking and individual species members varied relationships have not been extensively studied on (Table 29). Damselfish (Pomacentridae) were the most 1:lorida reefs, few reef fish have been found to be cosmo- common family at Tortugas while ranking third at politan aver all available habitats. At One Tree Island John Pennekamp Coral Reef State Park (JPCRSP) and Recf, Australia, 49% of all species collected were re- second at Biscayne National Park (BNP). Parrotfish stricted to one or another of five major habitats (Gold- (Scaridae) were the most common community member Inan and Talbot 1976). Similar faunal differences among at BNP but were second at JPCRSP and the Dry Tor- habitats have been reported by Gosline (1965) for the tugas. Grunts (Waemulidae) were particularly abundant Hawaiian Islands; Chave and Eckert (1974), Fanning at JPCRSP (most common family) but were third at Island, Sotlth Pacific; Jones and Chase (19751, Guam; BNP and the Tortugas. Of the 165 species observed I larrrtclin-Vivien ( 1977), Tulear Reef, Madagascar; and between JPCRSP and the Dry Tortugas, 1 15 were shared Williams (1982). the Great Barrier Reef. in common. Thirty-one species were seen only in The coexistence of a high number of fish species JPCRSP and 19 were observed only in the Tortugas on coral reefs also implies either that these species are study areas (Jones and Thompson 1978). Overall, fish lughly specialized (occupy finely partitioned niches) or communities at John Pennekamp Coral Reef State Park that there is considerable overiap &I resource utilization. were found to be more diverse and abundant than at the Predators are generally food limited and tend to have Dry Tortugas. more clearly separated niches (Hairston et al. 1960; Seven of the top 10 reef fish species at Biscayne Paine 1966). Keef herbivores such as parrotfishes, dam- National Park were also within the 10 most common selfishes, gobies, and angetfishes are generally believe&to species in fohn Pennekamp Coral Reef State Park. use many of the same food resources (Randall 1967; However, only I0 of BNP's top 20 species were repre- Sm~thand 'Tyler 19 72, Hobson 1974). Both spec~aliza- sented in the 2Q most common species at JPCRSP. tion and resourcesharing appear to occur among various Three major types of coral reefs are recognized reef fish groups and at various life stages within some along the Florida reef tracts: patch reefs, hardground species. live bottom, and bank reefs. Each general reef type Recruitment of juvenile fish to coral reefs may supports a characteristic fish fauna (Table 30). General 56 Table 30 (continued) Reef types and zones Patch Livebottom Bank Species TOP Outer Shallow Deep Fringe Orangcspotted filefish Catztherhines pullus * * Scrawled filefish Alurerus scrrptlts * Slender filefish Mot~nc~utirllustuckcri * Bar jack Cururz.~rli bcr * Uuccn angelfish 110i~curzt1r1~~cllluri~ * Gray nngclfish /20tnuccrnrirusclrcuutus * Kccf I)uttcrfly C%ui rodotf sedrtrruriii~ t*oureyr butterfly (.: cuptsrrrrrirJ Si)oltin butterfly C'. ocelbtur (;oitl\irot gr)i?y (;tiutl~irlcpr~fhotttj).\~trr * f lcvvcfnng gohy ii~gi'io,ired hi~li~t~ili~ * I tcrriilgs t'ililtertiuc * Bermuda chuh hj pirr)sus \c.cturr~li * * Spdnlsh hofiih Ho'ltu~z~l.\YU~IIS * * Slippery dick ilui~chotv-c.t hci~itruru;, * * knellowheaJ wrasse 11. ~urtiofr x C reole wrasse C'iro;icus (?urrul (continued) 58 Table 30 (continued) Reef types and zones Patch Livebottom Bank Species TOP Outer Shallow Veep Fringe I Iogfish Lacht~olaimusrnaxirnus * * Clown wrasse H. macrtlipintza * * * * * Blackear wrasse H. poeyi * l'uddingwife ll. radiatus Bluehead wrasse Thalassoma bifascia turn Schoolmaster snapper Lutjunus apodus Mutton snapper L. atzelis Gray snapper L. griseus Mahogany snapper L. mahogoni Yellowtail snapper Ocj~uruschrysurus Yellow goatfish Mulloidichthys rnartinicus Spotted goatfish I'seudupetzeus rrzaculatus Glassy sweeper Pempheris sclzornburoki Sergeant major A budefduf saxatilis Blue chro~nis Clzrornis cyarzeus Brown chromis C. multilinecltus Yellowtail damselfish Microspathodon chrysurus * Dusky damselfish Pomaceritrus fuscus * Beaugregory P. leucostictus Bicolor damselfish P. partitus Threespot damselfish P, planifrons Cocoa darnselfish P. variabilis * Black margate Anisotrernus surinamensis Porkfish A. virginicus Tomtate Haemulnn aurolineeturn Caesar grunt H. carbonariurn (continued) 5 9 Pa ti!^ livek>ottorri Rlnh Species TOP Outer Stlti!uu Deep r-ringe S~naUmouthgrunt H. chrysarg>lrrum * * French grunt !I. jluvol~~~r~ufutn 4. * I a * Spanish grunt It. ~nucrosforr~itrn * * t* d White grunt W. plumieri r * BlucstripetI grunt El. scwrus * c * Stripcd yarrotf tsh S c4rorc.c+tr,rl!, Ktiinlww parrot ftsh S. grrcjr'urtrtriu Hcd tail p;irrotfisl~ S irr~,iop[~.nart~ * Barred II~JIXL~~ff pt~(j/!u -& Butter Ii,rnslet El 1rrricoi::r * (;rdy\hy f,i.lirtipllc.irts r rui.trrul!ls * fieti grtjnpcs I; ,r!orict * Nahsau grouper i: arriufrrr (cirntin~~d3 60 Table 30 (continued) Recf types and zones Patch .Livebottom Bank Species TOP Outer Shallow Deep Fringe Black grouper Mycteroperca bonaci * * Harlequin bass Serranus rigrinus * e * Lantern bass S. baldwini * Saucereye porgy Calarnus calamus * Jolthead porgy C. bajonado * Great barracuda Sphyraena barracuda * Sharpnose puffer Canthiguster rosrratu * * descriptions of some of the most common fish species provide a varied and cavernous habitat. Along this outer follow. zone larger predators such as grouper, particularly black grouper, red grouper, and Nassau grouper, mutton and Patch Reefs gray snapper, and hogfish can be found. Close inspection of crevices and holes will commonly reveal sharpnose On most patch reefs, fish occur in two distinct puffers, squirrelfish (Holocentrus spp.), small cardinal zones: top and outer. Species commonly seen over the and flamefish (Apogon spp.), soapfish (Rypticus spp.), coral rubble substrate of the reef top are bluehead and glassy sweepers, and, possibly, the green moray (G.~rnno- clown wrasses, the puddingwife, the slippery dick, and thorax funebris). Clinging to coral heads are small neon bicolor damselfish. The dusky and threespot damselfish and masked gobies. The neon goby is noted for its func- staunchly defend small territories of algaecovered rocks. tion as a parasite and mucus picker, or "cleaner," on The cocoa damselfish is also abundant at some localities. large fish species. Gobies frequently establish "cleaning Resting on coral rubble close to a crevice into which stations" into which larger fish will move, become quies- they can dart for cover are redlip blennies, the only cent, spread their opercula, and allow the cleaner fish to species of combtooth blenny (Blenniidae) commonly work freely through the gills and about the head. Juve- found on Florida's coral reefs. Also near protective nile angelfish and the juvenile Spanish hogfish have also crevices are hamlets (Iiypoplectrus spp.) of various been observed functioning as cleaners of larger fish on colors. Scorpion fish (Scorpaenidae) can also be found the reef (Thresher 1980). frequently blending with coral rubble over the top of the Close to the bottom, among the corals along the patch reefs. outer edge of the patch reefs, the puddingwife and yel- Wider ranging species typically seen moving lowhead wrasses become more abundant. Feeding more among the corals within the top zone are the surgeon- in the open along the reef edge are typically saucereye fish (Acanthurus spp.); gray, French, and queen angel- and jolthead porgies, spotted and yellow goatfish, redtad fish, white grunts, and striped, stoplight, and redband parrotfish, and angelfish. Close inspection along grassy parrotfish. Frequently blue tang, ocean surgeons, and areas will frequently reveal the small bucktooth parrot- parrotfish (particularly the midnight parrot) will form flsh, blackear wrasse, and lizardfish (S~rnodu~spp.) At large mixed or monospecific schools while ranging about the base of the corals on sandy substrate, nearly trans- the reef top. Common fish species typically found near parent bridled and goldspot gobies can be seen, as welt as coral heads, branching corals, or other ouicrops on the the harlequin bass. la open sandy arsas i~nmediately top of patch reefs are the bluehead wrasse, sergeant adjacent to the patch reef (but not on it). hoverrng major, tomtate, Caesar grunt, and Spanish grunt. The gobies and yellowhead jawfish (Opzstognathu~aurlfrvrt~) vertical standing trumpetfish and the slender filefish can can usually be seen protruding from their burrows. usually be seen among the branches of the octocorals. The water coIurnn over the patch reefs usullly Around the outer edges of the patch reefs, the does not support a large numbel of fish, but barracuda, water depth is greater (8-10 m), and larger coral heads bar jack (Caranx uuber), yellow jack (C. hartlit~lumtreil, and Mojarras (Gerreidae) are among the most commonly the outer base (mixed hardgrounds) of the bank reefs observed species. Ballyhoo (tfcrnirarnphus braszlrensis) (usually at depths of 20-30 m), large mutton snapper, can often be seen moving about just under the surface. grouper, and porgies (Sparidae) become more abundant. In the quieter waters of the back reef. schools of small The tobaccofish (Serrunus tabucarus) and sand tilefish herring (Iiarrnguia spp.) are common. (Molacuntlzus plurnieri) can easily be found resting openly on the flat calcareous silt bottom in the mixed Live Bottom hardground and sediment zone. In the deep reef zone the bigeye (t'riacanthus Live bottom corninunities are formed on broad arenatus) tends to replace the glasseye snapper (P. areas of limestone outcrops occurring in relatively shal- cruentatus) cornmonly found on the shallower reefs. low water (2 ni or less) within the protected reef zone. Similarly, the cubbyu is replaced by the jackknife-fish, Coral growth may vary greatly in density over the area and hinds and graysby (Epinephelus adscensionis, E. and often is mixed with seagrasses. ?'he habitat generally guttatus, and E. cruentatus) are replaced by the coney has little relief or structure and is composed largely of (h'. fzllvus). ?'he snowy grouper (E. niveatus) becomes octocorals, sponges, and small scleractinian corals. prevalent at depths greater than 30 m. Over these deep Only occasionally are largc colonies of stony corals reefs, the number of species of jacks (Carangidae) and found and these often form an independent cluster. mackerels (Scombridae) greatly increases. Also com- Fish fauna of the live bottom is generally s~rnilar monly seen are the large bar jack (C. ruber), rainbow to that found on the top zone of patch reefs. A few not- runner (Elugatis bipinnuluta), amber jack (Serioh able differences include an increased tendency for tangs dumerili). black jack (Carunx lugubris), bonita (Sarda and surgeonfish to form largc wide-rangng schools, an sarda), and cero rriackerel (Scomberornurus regalis). increasetf abundance of tlre large rambow parrotfish, and Cloud-forming schools of small herrings (Jerzkinsiu and frequently larger schools of white, Spalush, and Caesar Sardinclla) are seen over the reefs at depths greater than grunts. An tntercslirig species incrcase ovcr live bottoms 20 rn during the day. At depths greater than 30 m, a is that of thc lantern bass. I his small bottom-dwelling third species of Serranus, similar in appearance to the bass appears to prefer the broad rocky flats around tobaccofish, becomes common. The chalk bass (S. tor- turtlegrass beds und, occasionally, replaces the sin~llar tugarum) is found around small rubble mounds, coral llarlecluin bass found more frequently on patch reefs. outcrops, and rocks near the base of the reef. At this depth, a short search over the sand plain next to or be- Bank Reefs tween reefs will usually reveal a colony of garden eels (~V.ystacticlitj~shalis). They have long, slender bodies ffnnk rccfs (f.ipurc 1%) form elongated discon- and are found in large colonies where, each protruding tutuous sti~~ctt~resalong, the seaward edge of the reef vertically froin its burrow, they form "gardens" of wav- tract. I lkcy rkse to wlthirl a mctcr or less of the surface ing heads while feeding on plankton carried by the and iravc cxtensrvc reef flats supporting only sparse current. cncti~sllnl: corals over tlieir shdllow zones. Seaward, The number of species described above in no way tikey sltrpt+rapidly to tleptl~sexceeding 20 rn. approximates the total number found within the coral Kcdl~pant1 saddled blenn~es,srnall gobres, and reef habitats of Florida. otf~crsinall \>ottorxl-ltwclh~~pspecies cfominatc the reef flat. Witl~rnthe ckailow q)ur 'ind yrocrvc zonc branched 6.7 REEF FISH MANAGEMENT and f~noctocorals hecvtne much triore previllent. Among dcnse ~~MIIL.~ICSvf staghorn or elkhorn coral Fish resources on the coral reefs of Florida and <,4t.rc~pr~rclspi3.f arc dusky and threespot darnselftsh, elsewhere are lligllly prized economically and are exten- rirlany specLC:, of %runts(Ilrtctriulort spp.), blach margates, sively utilized. Reef-fishing activities commonly include gray anti sc1~~)r~liriastt:rsnappol, scrgeant m;tlors, and co~nnlcrcialhook-and-line harvest, commercial trapping, flcrniuda cllul?. Witfiln the water column ovcr the rcef. sport angling, spearfishing, tropi~alfish collecting, and bar jdchk, hdrractltia, and yellowt,rkl snapper arc easily sc~entlfic collectmg. The importance of reef fish re- seer). sources and their economic potential have been describ- rl~cllsh fauna ut thc dccp spur and groove rone ed in a number of sunlinary reports (Camber 1955, Stru- can be highly vdned tiepcnding on depth anti the arnount sdker 1969, Swinglc et al. 1970, Stevenson and Marshall of reef structure. Welow tl~c,lctoporu coral zone (deep 1974, 13ulhs and Jones 1976, Klirna 1976) In add~tion spur and goovr zone), however, the following become to descr~ptivereports of flslung activities, a number of lncrc prevalent >eflowtarl darnsclfirlr, the rock beauty, exploratory studies and fishing development projects butterfly fish, small groupers (i.c., han~lcts,ldnds, and have been completed for reef fish resources (Brownwell graysby). yeitowtad and retirari parrotfish, largc pudding- and Kainey 197 1; Munro et al. 197 1; Kawaguchi 1974: ives, and large scrdw fed file fish W~thincreasing ciepth. Wolf and Kathjen 1974, Stevenson 1978). Little Infor- there 1s a distutct lIIcTt'dSZ 111 (,'/IIo/~~~s(C'/l:)lliorttis C~U~~CUJ mation, however. is available concerning the lmpact of and C tnulrrlirlc.cilits). the longsnout bufeeitly fish (C' coinmerc~alor recreational use on reef fish populations rrculcurlrc), tnggzrfish (13allrrrs spp arnd C'ftl~ti?t:It~).~ii~s or on rcef fish management. Falrly substantial sectlons Jr~j~tuttroi1, filefish (4hil~'riis app.). jackknife-flsh. of I;lorrda.s coral reefs are now designated as local, schools ctf creole wrasse, and thz blue parratfrsh. ,%lon% State, or National preserves and are being managed for 62 the protection and conservation of their resources. nidae) and groupers (Serranidae). although some tilefish 'The impetus to preserve coral reef resources (Branchiostegidae), jacks (Caran~dae),and triggerfish stems largely from their recognized economic value (Balistidae) are harvested. A complete listing of commer- through tourist attraction and recreational opportuni- cially important reef species can be found in the U.S. ties. But the negative impact of recreational use (i.e., Fishery Management Council's management plans for snorkeling. diving, boating, and fishing) on coral reefs these fish in the Gulf of Mexico (Florida Sea Grant has not been extensively investigated. Interestingly, College 1979) and South Atlantic (South Atlantic although the taking of tropical fish for aquarium pur- Fishery Management Council 1982). poses is prohibited in all coral reef preserves, spearfishing Within Biscayne National Park, a limited assess- is permitted in at least one and hook-and-line fishing is ment of fishing impacts on reef fish populations was permitted in all. Impacts to fish population through conducted by underwater visual surveys concurrently disturbar~ceby boating and diving alone could becor~ie with fishermen's creel census (Tilmant 1981 ). During a significant. On extremely popular reefs, such as Molasses 3-year period, fish catch rates varied inversely with the Reef in the Key Largo National Marine Sanctuary. as number of fishermen using park waters while reef fish many as 20-30 boats with divers are often anchored at populations remained relatively stable. Katnik (1 981 ) one time within a 4-5 ha area (NOAA-OCZM 1979). found that of the reef flats surrounding the Pacific island Specific studies at locations with such an extremely high of Guam, those with the highest fishing pressures level of use are not available. However, within Biscayne showed a marked reduction in the large size classes of National Park, a 5-year study of eight selected patch the most prized species. Also, some fishes of less eco- reefs revealed little disturbance to fish populations nomic importance were more abundant on heavily fished where maximum annual use per reef ranged from 3,400- reefs than on reefs lightly fished (Katnik 1981). Bohn- 3,600 persons diving or snorkeling (Tilmant et al., in sack (1982) reported that he found significantly slnaller preparation). piscivorous predator populations on Looe Key Reef Past research on exploitation of reef fish stocks (near Big Pine Key, Monroe County, Florida) than on can best be described as gaining basic information on life either of the two similar reefs within the Key Largo histories, food habits, growth rates, and basic biology of National Marine Sanctuary. He attributed these differ- important species, and on developing harvest techniques. ences to spearfishing on Looe Key and believed that the Only recently have studies evaluating the ecological abundance of some remaining species on Looe Key Reef impacts of fishing activities begun to be reported (Camp- was affected by the larger predator loss. bell 1977; Davis 1977b; Tilmant 1981; Katnik, in press; In 1977 the United States passed the Fishery Pauly and Ingles, in press; Bohnsack 1982). Conservation and Management Act, which extended The few studies available show that a limited jurisdiction over fisheries to 200 mi and called for estab- number of reef fish are actually sought and comprise Iishing regional fishery management councils to develop most of the sport and commcrcial harvest. Austin et al. specific long-range management plans for each coastal (1977), after interviewing 4,275 recreational fishermen, fishery resource within the Fishery Conservation Zone. reported that only four species groups (grunts, snappers, In response to this legislation, the coral reef fishery dolphin, and grouper) constituted almost the entire resources within U.S. territorial waters are receiving catch. Within the groups, gray snapper and yellowtail much greater attention, and management plans for their snapper accounted for 85% of the harvested snapper; conservation are being developed. The status of biologi- white and bluestriped grunt, 97% of the grunts; and red cal knowledge and application of fishery management and black grouper, 82% of the grouper. Thus, only six principles to coral reef fish stocks was reviewed at a species made up about 80% of the total recreational workshop on reef fishery management sponsored by the catch. The National Park Service has recorded over 120 National Marine Fishery Service in October 1980 species of reef fish caught within Biscayne National Park (Huntsman et al. 1982). The workshop proceedbgs (Dade County, Florida); however, 10 species accounted succinctly summarized the requirements for such for more than 80% of the harvest. management. It is widely recognized that before coral The commercial harvest of fish from coral reef reef ecosystems can be effectively managed, much basic areas is almost entirely directed toward snappers (Lutja- data and many larger-scale multidisciplinary studies are required. CHAPTER 7 CORAL REEF ECOLOGY 7.1 INTRODUCTION 7.2 PHYSICAL-CHEMICAL ENVIRONMENT Wellb (1957d) succln~tlydefined the nature of Ihe subrtratuiri for estdbl~shrnentof a coral reef the coral recf as d "deterintnistlc phenomenon of scden- nlust be rock or consolidated materials. Shlnn et a1 tary organlsinr wit11 high n~etabolisiil hvlng in warrn (1977) reported that the reefs they cored In blonda lraarinc waters wititin the ztrnc of strong illumination. were estabbshed on a basement of I'leistoccne reef fa- 'I'hey arc constructional physiographic features of cles, fossil mangrovc, and llthified crosq-beaded quaitz tropical seas consisting fundarnentaliy of a rigid calcar- (old \and dunes) Ihe Initial settlement of ploneerlng coils tr;lmewitrk made 1117 rnalnly of iritcrlockeii and species 1s followed by successional stages durlng whch c11crtlstetl skclctons of rcef-hurldtng (hcrnmtypic) corals tlie genesis for creatlon of the reef 1s estabhshed Shep- and c:~icareotis dlgac Itic fr~mework controls the pard (1982) discussed the evolut~onof reefs based on c~cc~~~i~ulationof scdrmcnts on, In, dnd around itself data frorn the Red Sea and Ilawau and noted that about 1 hcse sed~nlcntsdrc clcrived f rorn organlc and physical d 50-year period was requlred for a coral reef commun- dcgratldtic)n of Chc frdrue and orgdiiisrns associdted with ity to attam the fourth (final stage) or blndinguf- the recf con$tructors anrl have bulk ten or ntore tlnios as sedirncnt phase I'lor~da reefs are m varlous phases of great ;ts thc fr:brne itself. Ilic coral rect biotol)c 1s a facics develop,nent, some hdve reached considerable develop- of the tuarrno tioplcal hiochorc, and ~tsessential fauna ir~entwhlle others are either juven~leor have suffered rind flora cortsists of cor'iis ant1 calcareous algae, wluch ~rnpactsclcnylng then1 full develop~nent doilllnatc In nn~nhers and vtrlun~c and prov~tic the Most coral reefs off southeast Florlcia are found cctriogical niclic\ cssenti;il to the existence of all otlter In depths of <43 m. Beyond these depths the substra- tum dnd physlcal environments are unfavorable to the existence of herniatypic reef-buildlng Scleract~nla. It should be noted that where the physical environment is tdvordble in other drcas of the Cdnhbean (Bahamas, Behle, Jarndlcd), a set of deep reef Scleractlnla are found ~olonlzlngthe escarpment faces These communities are tilldliy. the IIIC)I)I~C f.~tli~~01 hcnfluc d~id 11eht~)n~ found ds deep as 70-80 m. Off southedst I.lorlda thls sg)oc~cs. Roll1 pliybic,illy ant1 t)~gdn~cally,reefs dre type of cnvrronrnent is, for the niost part, nonexistent. d<>llildt~dtCdstiuct~~re~ rlt~d ,ire tile result of d ncar A deep reef wds recently found in 60- to 90-111 depths Ikifan~z of cotistructiv~ and dcslrucllvc tn~ccs Ilie west of the Dry iortugas. Few details are available at cr~itstrriit~on~lforces, dre Idrgcly orpdnir, tflc accuinula- ti16 ~IIIIC Sclerosponges are also a conspicuous element tion ol tlrc c,ilcdrer)us skcivlons 01 corals, colc~rcous of deep recf coriimunitles, these are not reported from aigL!c, tord~likn~tcr,i,moil~tsLs, ctc , roughly m that ordei f'loridd reefs Dustan et al. (1976) discussed the factors crl ini~~c)rtdncs. . 1)estructivc ilnd tfcgriidational forceil Iiintting sclerosponges from Flonda. tcr~rllny:lo break down thc \tout ficunewc>rkof reefs are Light is probably one of the most slgnlficant ~~dillflllfi~~f111 titc ccd~eii~s~hrcaktlowr~ of coral. dlg;ll. env~ronrnentdl considerations controlhng coral reef de- lilr~llttsc~~n\krtletctn\ 3i>ii reef rock by tlu: nor~rldillfz velopment in general Llght is necessary for the symbio- ai-t~vlticsof ,I wrctc varicly of pertorating, I)onr~g.;tntl t~crelat~onship between the coral and zooxanthellae. dIh\0~~11?~JIc*~~. Sj>(~ll~teS, ~~~(~Xltishil, WOrfYlS. dlld Cifli- 4ny factor wfuch reduces the amount of solar radiation t~o~d\. alcrrc olwiou5, bttt no Inore triiportant, iriipinging on the coral surface will affect growth and dcsl~u~ttvc~~~IICY til wdve ;*C~IC)II . . . [in~lt~di~lg]wlnd- nutrition. Reef corals are almost universally phototropic. rir~u~~~wdvcr, ~tuch dur~i~g hurric;mcs sttihe prodigous Wells (1 957a) reported that species richness at Bikim fldtrznrr*r blow5 oil zref5 " iliolf wds closely controlled by solar *Ilum~nationand f'hdt wirirt~%'ells (1 (3.573) scpoited 1s stiii vdild. indirectly Influenced by temperature and oxygen. Gene- I'setakicti rnvttstipitions into vdrious ecological functioiis rdl solar radiation character~stlcsIn klorida were dlscus- tlav*. rnzredseci i>ur ur~iierst'indri~garid apptecidtiuil of Eed m Chapter 2. Ihe nature of hght is significant Joklel rkte cord1 ri>etsIn ger~cral iddy, I~londa'scoral reefs dre and York (1 982) reported that specialized pigments In pruh,ibly one ul. the n~stvis~ted reef sy\telns In the coral t~ssuesfilter the potentially dangerous ultraviolet world, yet the htc. history of inany spccles 1s S~IHpoorly portions of the spectrum, thus allowlng the corals to underrtviiif f he recent lncredse In coastal devclopr~~er~t occupy the shallow waters wlthout belng harmed The nrrJ q9uli tilvlii.g Irdb g~fl~~dkdc.oI~cc~III dt)olil lhc vl~dllly zuoxantheilde xrt: able to ~onlpcnhdtefor changes ul uf the coral reefs. Slr~cevery iltt!c edtly babelme eco1og1- rad~atronquahty and quantity through changes in the11 cal infcrrrnatr~>rrIS availahlc for sor~panson,qital~tat~ve plgrrlcnts Wethey and Porter (1967) reported that reef ~ubject~r~statrments are often oftcred irt support that corals were able to compensate to a depth of 25 m the reefs arc. detenordtmg, The most srgn~ficantimpacts wxthout Ioss of autotropluc efficiency. Kanwlsher and on tile I-londa cord1 reef? are, however, caused by natu- ~ainwrighf(1967) reported on resplratlon and produc- ra! events over which man has no cuntrol ttvlty in Florida reef corals from a study at Hens and C'l~ckensReef. Wells ct dl. 1 10731 rrpvrtcd that sunle ftdal conti~tions off the southeast coast of corals uzre best cl;issil';ed as produccn bsszd oti f" I.lortdd were preser~tcdIn 'latale 8. All rsrlges are Iess (~rnducinvit>:resplrat~orl) ratlix being gredter tti,zn a,nr thin t 1x1. 1 tddl conditions only posc a threat to the Teltiper%ttureis related to solar radi,itlun illrillxi- sh,~liowerportion:, of tile reef flat u11d spur anti groove @ng on the water column and the ~nflttence of the (lult rows Reef emergence rarely occurs except under Stream or E 1ontl.i C'urr~nr. H'htie rec'f corals heave been syncrgtst~r:n~eteurt)logtcat ~nfltiences. Wl~en spring low described a& ~tel>otherz~~lc(Wells l VZ 7b). Inmy spcclcs L~dcs occur at or near mjdddy tlurtr~g the surnntcr, are rery to1er:tnt of temperature barxdtlc)n (Clayer 1914. thcrrnal tte:~tlngcan occur (Plate 24). 19 16, 13 18 ). Reef COT~~Sfl01111~) ~O~~tftlltllllph>bl~logl- I)~ssoived oriygerk ts depc?.~~ilerrtupoil photosyn- cal cond~tions)best hetween 311'1 29' C'. CI0r,i1 reef tlies~s,irld rcsptraclotl of bcntkric orgniilsrtls. I ernpcrz~ture development is reported to he bantted hy lovl terrlpexii- anil salinit) cctntrol the s,ltur,htiox~level of ouygcn ln the tttres. Illis was proposed by Vatigh.in (1 9 llH for t,'loricla. water. Studies by Vangti,~n (1 01 4d), M,rycr [I 91 8), He reported thiit corals did not huild recfs where tern- Yonpc dntl Ntchoil~(1931 ). inlit11 et '11, (1950). and peratures fell helow the 18' (' !limirnurn for proloxrpecl Jones (1063) reported tltdt oxygen 1s not truly :r tlri~itlrrg periods 1he growth and dernise of stdghorn reefs at the factor far corals ~trtdc'r IHOS~circli~tl~t~trlces. Oxype~i Ilry Iortugas dre related to terupcrature control. I)uring ren'talns In supcrsuturiitio!~durisig rilost of the day. jotles favor.tble penucis tlzesc popiilatlons psohfcrate, but (1l>63) reportc'ti that the d~iirr'tdlrililge was fr~)nl90'r to ~~~asior~ally,I cold w~irterreduces tcrrrpcrature below L 25'; saturatlun durtng she suri~r~ier tulerance lcvcls dnd rrlass nlortnl~tyoccurs (Porter et 31. Water ccrluain pli is ~n!Iuonccil by organic 1982) Wittun a rlr~ade,if lrilld ~onditiorlsprevail, the snct:~bnlrsin. Studies by Juxieb 1 19t~31 hi~td Jaap and pctpulatlons recover lelnperatiire datd In t lgures 6-1 1 Wl~raton(1975 t reported pfl values ot near Q. It is iiot a conlgrared recent d:itit with tht trorli the early pdrt of this cerlturf Ihr ditta ~ntplyno ~n.ijorchange m chnlate, It is apparent thdt tcmper:ittirr is vflc vf the 7.3 COMMUNITY STROC'TURI: rnalot controls of reef cieveloprrlerzt off southe~ststI londa (ttoherts et ai. 1982) ,411ottrer source of c01dw~itc.rstrc.is <'vt,~lrecls otl s~)uttiu,istI lotirl,t have developed is trorl~oc~~rs~u~tdl ~ipwelh~~g troll1 sources hene;ttt~tire over the past 5,000-7,000 ycdls. it 1s prcsulnetl thdt t'lur~dd ( tirrtBnt I~OSCst~:ttll~etI layers oc~~s1011.iIly "surviv,t! slochh" hving 111 witdt 1% ~IUWdeeper water catibc I~tcdhrerlt14t 1\111>.Ilvet-rclated stresses 111 srrilirrler tluring tlte wlsci~t~cir~ire dgth WCIL' the scctl pnpuli~trons urunlly occur cttlrttig midday wltert sprint: low tide5 tor rcj~lrrrislzrrzantoi tllc rrcls as the 5e;k Icv~Irose durtng ca~tnc!de with t.alrrr WIDL~~C)~~I~~OIIY (1'k;lte 2Jbj. I he thc fioloccrlc trdnsgrcsstsn Solnc cv~cterice for tl~ls~s ~Ildli~wreef !ldt\ iuf fcr trunt ther rilal \tresses ~nrtnlfcstcd tound krt tcrr<~cessc'iwa~d of sorne of the slrajor rcefs In ac)uxaritltellde eupulsron Ilitrdviolet ligltt rrijy alsc) he (Mtddic Sarribo 'snti Sdnd Kcy Meets b. I kicsc were invoiverl w~tfi tJus phenoulrnort. ,\s w'1tc.r tlcptfl is i>rotrakiy :tcttvr constructionat rcefs dorrrlg $1 Ivwcr sc:i rcduceri, lllr IAV frytit rnay Elr~tc\irtli~.:etlt strength lo 'tlr Irsvel stdnd wii~c!~werc tirownut1 as sea level rt)se rapitily. tux~cJJ,~ physlc,illy hurrl thrv tis>uc\ (Jokiel and korh lho toss11 harrier roe1 olt Broward arrd lX~lrrtf%edcIi li)K1) ( ourti~cs~lso provtclcs creclrhlf~tyto tlris ttxcory (1 tgtlty X,tltrirt> ii zc~nlrollciiby prcc~pit~~tiorl.cbdpvrd- 1 977 1 hrri~ctltc origltr ot the recent rt~efs,orpdrirslrka tron, .inil ~\rt~r>ft'11rer rti,ijor r,iirtt,ill ovt.1 ,titl,~cetrt I'tncl hd~~'llrl~f \llffl~1~111llltlC 10 <~CVC~L)I)Illtl) 1011tj)~l'hCC)IIl- iliii~iitlc~'~IC tt\tt'ilX> ill tlir OLC;IIIII r'lrlgv 1,1171~~(1 rr~url~lre\i lie) rir.play all level5 of corlrplcuty rrt ttrcir rcportetl r'irtges trclrrt 33 1 to ih 6 pp1 Ic>nt'r tl(l;t>:l \tru~Lurc. tllvcrs~ty, rr~tt.ractrons, uornpcixtivll. and rc~~yrtctl+,crrrlt drtiri1'ii cha~~ge~.111 3 tlritc \crte\ frljttt 11oplli~rt+~,~tlOn\lll~>\ 'Il,rrgrtt L.lsll bhit,il ot I I ll~otthey S'~iirr!t) doc., ntrt Irl tcl ins of ,it~c~ncl'irtc~c.tlotninance, ,inti diversity. nortii'tlty pox '1 tiirc'it to tkrc rccl\ Alter Irurr~i~ttrc v,iria>tt%cItXr~rrtrt.r diipl,ty cr*rtairt cl.i,rr.ittcrrctic jvatterni. r,llns, i~c~wc.rer,tficrc rb 3 p~)tetlti~~Ifog diliitirin ii~xctcr llttb \tA\\rSi- I>c.~ilf~i,s1s lltc nlort stiiblc and I~ebt%uitcd fos runoff f he only reportcti poisit)lo i.lllr~~rk-rt.Idts11 111111. hSTli'> %ttid1~'5f !I<' l~litbls~I>ellfflO?i. plilIlhFt>ll, i)tld phunn~tienirn u,~s tho 1878 iortrrg,ig 't~l,~~kw,rrrr." ncktc)t~r xtii!,ii j~lopcrt~ei.tfadt rlx,ihs if tl~ff~cr~llto mahe WIIILII*a> rt*portcd EO fl~vckilled \lyn~licvntpoj)t~idtio~lz, t1111c \L"IIC'LI io~!~p.iri\o~liJIIV 'r~gl~lflc~t~tccl'r tlldt dl- uf cor,rl> Zl,tycr I lc102) rcyortcd Oldt 4 grcdi ~~iiir~llcttrl Itlolii~itLorIrt* orpntsltii 'ire c%entrnl tts tf~creef. otiier\ ttiy 21 * el__w. ~,~~,.,,~~~ curiirnuntktlies Bruce (l"l7h) and fbtlctrt 11'9761 reporred lWl W!*?illi on asboc,l:stior~sof tlvmg c~traf~and crustacean wntnlen- saltsrtt rt~cotal reeh wrth most of the einphasts on Pacific 8 "*-"--#a?-*-*-- ccel.r; mmE LKtP Orgtnism rnrcr~ctlonsut a cord reef &re also coniplex and lxzghly zlcvetopril. Surrre drr ohligate whll~ 3. - --. -- EL~- ~XXF~F ~tilers &re IJ~OSE~~by cflance. Patton (1976) wrote, "Corals exceed ali, other anvcXrtchra&cgroups in the dtvemcj crf tiwrrrs dnrl muxnhcrs of species to whch they i-d"-*-L-- i-d"-*-L-- tea. 91, 9D. JS, 0. ptdy Itme." kuud ~11t.i S~P'XXCK are tkie nlyor factors LWEt OF SIMILARITY attraetnte tu assoclatm f'kc cord skcietarl offers an i igttrc 21.. Eirttrl,$xaty el cujal launs on t*lvnd;r reels esprtctieily gourf ~~acizefor snlaUcr crrganksrns to escape UZX~IRgitxi~p svrr,agc carrilr~p~nii Ctuhanuwski'~; cueffi- psecistsor* N~frieroti$sp~ngm. ixr53ychaetes, bivalves, and csclnr. atpuncuhds find habta~t &ax ilae base of corals. Otlzer t8 organnms live successfully on the coral's upper surface. coelentric feeding behavior with the exte~~sionof mesen- Sorrie. such as copepods. have developed specialized terial filaments. l'11e rnore "aggressive" species have the fornls to hve in the coral polyps, they are rnore worjn- abil~tyto digest the tissues of less "aggressive" adjacent hke than crustacean (Patton 1976). Sotne organlsnis species. Lang (1973) described this form of interaction. have developcd unique ways to niodify the coral skele- It is not a rapid phenomenon; in some cases it may take ton. The subfamily f'yrogomatinae (barnacles) and the months for the interaction to be noticed. decapod crustacean Dotneciu ncc~t~rhophoraare able to rnod~fy the Scleractinian skeleton. Dortzecia ueutzrhu- 7.6 PREDATOR-PREY RELATIONS phoru lives on A. pult~rutubranches (Patton 1967) and is able to influence calcification such that the coral builds I'redator-prey relationships are far too great and a skeletal structure around the crab. 'I'he process initiates coxrlplex to detail here. One of the more well-known when a crab seeks a position on the branch margins corallivores is the polychaete ficrrncxiice curunculutu where bifurcation has begun; the crab maintains its (Plate 18b), a large marine worm that is documented to position, and the skeleton grows around its den. In time feed on a number of coral species (Marsden 1960, 1962; the area resembles a small pit or cave. Domeciu acanttto- C;lynn 1962; Ebbs 1966; Antonius 1974b; Lizarna and phoru has rnodified second n~axillipedsthat allow the Blanq~iet1975. Glynn (1 973) reviewed western Atlantic crab to feed on suspended material. In this case. the coral predators. Same organisrns such as the gastropods relationship is both shelter- and food-related. Patton Corulliophilu ubbrcviuta. Culliostoma juvunicum, and (1976) reported that mobile crustaceans living in close Cyphotriu gibbosurn are mobile species that move to and association with corals feed on the sediment and detritus from the coral, 'They feed on the coral, but apparently from the water column and that which settles on the do not receive protective shelter. Fish within the families coral surface. Corals also release energy sustaining Scaridae, Ephippidae, and Pomacentridae feed on coral materials that other organisms may utilize. e.g, mucus, (Glynn 1973). They are not obligate to a particular coral fecal detritus, zooxanthellae, and ~lietabolic wastes. species; this major difference between Pacific and Competitive interactions are important in deter- Atlantic corallivores (lack of obligate species relation- mining comrriunity structure on a coral reef. ?'he major ships in the Atlantic) may reflect that the time of area of interaction is in the competition to acquire development for Atlantic reefs and coral species has spatial territory. Successful organisms can defend been relatively short. (Atlantic Scleractinia are only ca. their territory against intruders. They are also able to one ~r~illionyears old; Pacific reefs have a much older expand into the adjacent region at the expense of fauna.) Predation may directly cause mortality and organisxris that are less able to maintain their territory. morbidity. Injury caused by predation may lead to the An exarnple of local habitat manipulation is the effect of invasion of boring and rasping biota. When the coral is the threespot damselfish (Pornucrrltrus plirnifrons) on damaged, recovery may occur by regeneration of tissue; the corals Acropom cervicort~is and hlonfustracw un- but regeneration may be prevented by successful coloni- nularrs. Kaufinan (1977) and Potts (1977) reported that zation of the injured area by other organisms (filamen- the threespot damselfish was able to garden small areas tous algae, diatoms, sponges, Millcpora, and zoanthids). of reef by killing the corals and defending the area frorn This ahen growth is detrimental to the coral, which may l~erbivorousfish, thus allowing a crop of filamentous succurrib to the secondary invasion. algae to grow on the dead corals. These srnali fish are Rapidly growing species may grow over and territorial and defend the area against fish, invertebrates, shade out other organisms, denying them resources. and divers. As a result, the area has an abnormally high 'T'here is also evidence that some reef organisms are able algal cover. ?'his result is short term in A. ccrvlcortus to maintain and advance spatial resources through the habitats. Since this species grows rather rapidly, it can secretion of toxic chemicals (Jackson and Buss 1975). grow away from the damselfish territory and develop a Figures 24-27 show the cumulative effect of new area. fn the case of M. uanularis where growth is these interactions through time for a smd area on slow and propagation is mostly by larval recruitment, Elkhorn Reef. the damselfish may have a more significant negative impact on the affected coral colony. 7.7 PRODUCTIVITY A number of authors have reported on the effect of the black-spined sea urchin, Dudema antillarum, an The earliest study of overall productivity of a coral reefs. Sammarco et al. (1 974) reported that D. small coral reef was Odum and Odum (1955) at Eniwe- untillarum controlled coral community structure; Bak tok, a Pacific atoll. The comparability with Florida is and Van Eys (1975) reported that the urchin fed on poor; however, the concept of high productivity is valid. coral; and Sammarco (1980) reported that 17.antillarum Odurn and Odum (1955) reported that mean annual ing microhabitats for coral larvae production was 846 gm dry biomass/m2. On a 24-hour llarum is a common resident on basis there was a net gain of I gm elma. Sournia (1 977) most coral reefs; hence, its control is probably signifi- reviewed cord reef primary production and concluded cant. that gross production in coral reefs yielded the highest Another form of interaction that has been productivity of any ecosystem on earth. The magnitude mentioned earlier is the ability of the corals themselves of gross production ranged from 2-10 gm ~/m'/day; to maintairi or advance territorial superiority by extra- however, net production may, in some cases, be negli- 7 1 Flgwre 25. Time genes ab a 2x2 mcker area of Elkhorn Reef, 1979. Figrare 27. 'Ftnw ~erierof a 2x2 meter area of' Eikhirrxr et al. (1974), with some nlodification in terminology. v. Mesoepibenthic detritivores, 200-500 microns 1. Benthic Plants w. Macroepibenthic detritivores, >SO0 a. Nitrogen-fixing algae, nutrient recycling microns function b. Crustose coralline algae, framework or- 3. Benthos, Invertebrates ganism, cementing function a. Animal-plant symbionts, energetic sup- c. Benthic microalgae, primary producer port from autotrophs d. Turf algae, <2 cm high, autotroph b. Invertebrate "scrapers," remove sub- e. Carbonate producing macroalgae, stratum with food sediment production c. Invertebrate browsers, do not remove f. Boring algae (Ostreobium), filamentous, substratum within the carbonate frame d. Passive suspension feeders, collect nu- g. Detached macroalgae, Sargassum, trients from water column without floating autotrophs physical action on the part of the h. Marine grasses, blades, adjacent to organism the reef e. Active suspension feeders, actively col- i. Marine grasses, roots, alter sediment lect nutrients from the water column characteristics by creating currents or selective pre- dation 2. Plankton f. Microbrowsers (meiofauna), nutrition a. Heterotrophic phytoplankton, < 10 gained from material in sediments, microns <2 mm b. Autotrophic phytoplankton, < 10 g. Macro-deposit feeders, nutrition gained microns from larger (>2 mm) sedimentary c. Autotrophic phytoplankton, 10-100 fraction microns h. Sedentary micropredators, selectively d. Autotrophic phytoplankton, > 100 capture water column prey microns i. Small predators, motile organisms that e. MicroholopIanktonic omnivores, harvest invertebrates and vertebrates <200 microns j. Medium predators, mobile organisms f. Mesoholoplanktonic omnivores, that harvest inediumsized invertebrates 200-500 microns or vertebrates g. Macroholoplanktonic omnivores, k. Meiofauna predators, live in the sedi- >500 microns ment, harvest meiofauna h. Neuston omnivores, all sizes 1. Parasites and pathogens, very selective i. Microepibenthic omnivores, <200 predator that gains nutrients from a microns limited number of host organisms j. Mesoepibenthic omnivores, 200-500 m. Parasite cleaners, mobile organisms, microns specializing in removing parasites k. Macroepibenthic omnivores, >5 00 from other organisms microns n. Attached eggs 1. Mesoholoplanktonic carnivores, 200-500 microns 4. Nekton, Vertebrates m. Macroholoplanktonic carnivores, > 500 a. Grazers, also remove a portion of the microns substrate, i.e., parrotfish n. Neuston carnivores, all sizes b. Browsers, do not affect substrate o. Mesoepibenthic carnivores, 200-500 c. Bottom feeding planktivores, feed on microns epibenthc plankton p. Macroepibenthic carnivores, >500 d. Midwater feeding planktivores, feed on microns holoplankton and neuston q. Microholoplanktonic detritivores, e. Small predators, <50 mm standard <200 microns length (SL) r. Mesoholoplanktonic detritivores, f. Medium predators, 50-250 mm SL 200-500 microns g. Large predators, 250-500 mm SL s. Macroholoplanktonic detritivores, h. Top predators, > 500 mm >5 00 microns i. Parasite pickers, feed on vertebrate t. Neuston detritivores, all sizes ectoparasites u. Microepibenthic detritivores, <200 j. Detritus feeders microns k. Attached eggs 73 5. DerrirusiNutrtent The I)at.rl et dl rnodel cuntatned 104 cornpart- a. Sitrate, NOs dhsolved ments. A nlatrlx of the mterrelat~onrtupsimphed about b. Nitrite, NO2 difisolved 2,000 of the 1U.Kltl patentla1 mteractsons were active, c. Axnnlank, Ntfa dissolved and a materra1 and energy flow exists hetween thesc d. Organic carban, C dissolved categories. In terms of nzodehng coral reefs. Snlith ct al. e. Suspended detritus, < 10 microns (19781 reported that untal thc modehng techrurlue f, Suspended tletritus, 10-100 rnicrans advances to the po~ntthat it is reahstic, the budget g. Suapendcd detritus, > 100 microns analysis nlethod offcrs better snr~ghtinto ccosystenl h. Trapped detritus dynamics by fuc~ismgon an elernent wlthm tile ecosys- i, Orwn~cnitrogen, h' dissolved tern (e.g., carbon hudget. nitrogen budget). Smith et al. j, Inurgsnxc phosphate, PO4 (L978) reported an several budgets fro~na hypothetical k. Orsnic phosphorus, P djrisolved atoll in tiic 13aclf1c. I-igurc 15 provlded a quahtativc 1. Oxypn, Cjz graphic reisreserltat~crrl of a calc~urrt carbonate budget nr. Intzatitial ?do3,dissolved fronl Barbados. n, lilterstitlrtl NO2, dissolved o. Interstitid Nil4, dissc>lved 7.9 NATURAL IMPACTS p. lnterstitlal C,dissulved q, Intcwtitt~lparlicuhte wrgun~cC (dead) tiurncanes are the most scverc nattiral impact a. Interstlttsj organic N, dissc)lvcd faceci by coral reef communltres. Mega-r-hurncanes (those 8. Int~"rstitia1organic Ij*, di~olved with 200-mijhr or greater winds) have devast:ttcd reef 1, Interstitial I">,, d&sulved araas, and either recovery has heen slcrw or the coral reef u, fntcatilinl Oa, dissolved community has beex] replaced by another corttlnt~nity. Stotldart (1962, 1963, 1974) rcpurted titat ffurrrcane (I. C;coictg~, ifattie destroyed rcefs uff coast of Ur~tl~fiIIanduras ;i, Dlssolvad ~rrclrgsrrkcI: tC02,I1CO arid (Belize) and that recovery was nc&;liybte. Wcrudley et al. ~"0~1 (1981) reported that Ilurricane David tieelmated sfiallow h. SirspcrriXed tnorgaiilnic C' (fine C:a(J03J reefs near Usscovery Bay, Jasr~atca,and post-fiurricane c, t4ctPIitad ~nt,rgtkrricQ: f crjlarsc Ca(:Oj mortality of da113;igccl organwxris was signlfl~ant{Knowl- scdzxnensrr orx the ,yo& fhour) ton et at, tYHi ). d i.rtkrrtt! ~xtusn@~ticC: (r~offrarlre) fn corulclering tire effect of In~rr~carres,two c, Ifrkt;rnlsni 1n~>rgunir:I' (I';tC'L13 in lrvrxlg majar areas of negative ltrkpact arc rejrorteil l'f~yslc;tl nanwframe orgarrhnns) da~nageoecitrs as a result of tile huge seas penaratotl by f Hliltblz int~rgilrrtb:(i' (large talua) the t~urricanewinds. Shallow reef rones hear the brunt g. Xnc)!acatrlr (: in sand (C?aC-'03, (I 2 rltic- sf the wave lorces as they are expended on thr spur and rrfns.4 rrrrrr I groave and rcef flat. Smaller corals anti otl~e~att;~cbed h. Irrrtrgaful: C' rxt ~tzitd(c;:aC,'OJ, <<6,! ory,anxsm$ ,ire ofter; dislodged, fraglncntetl, ;lntl abraded ii11(LTr)l$b) by the sevcfc physlcizi poundmg. I hr otfier rtiqor 1111p;I~t t lrxtcrslrt~aidrs~~ivrd I' (C:OZ, I-lCOj, conccrns d~lut~urlot dnity ca~~sedby torrential rains. ~srtd('CfJ ittssofv~dfti. ~ntcrstirlalsetli- la reguns where rnuunta~nuus tcrrostnal larlci rriasses irt~rtt#iitcTI ,tbut the ~ctral-lmerfcoast, bcvcee rufloff may titxtt coastal wa tcrs hy posaline (Cureau 1'104) 1, roslun ut sod aild 7, j11f~tit! F 74 years after the passing of Betsy. Ball et al. (1967) and lae expulsion (Plate 24b) at Middle Sambo Reef (off I'erkins and E:nos (I 968) also discussed the effect Boca Chica Key) related to elevated water temperature Hurricanes Donna and Betsy had on the reefs from a that was created by synergistic meteorological and tidal geologcal standpoint. Jindricli ( 1973) postulated conditions. The overali impact was not significant in that that hurricanes were one of the controls on reef develop- recovery took approximately 6 weeks. Vaughan (191 l), ~iientat Dry Tortugas. Mayer (1918), Porter et al. (1982), and Hudson (in The nature of hurricane impact has several press) reported thermal stress damaged or killed reef interesting aspects. First, the hurricane causes consider- biota at Dry Tortugas. able physical alteration of habitat. Sessile biota are often Heat-related stress usually occurs in late summer dislodged and transported considerable distances from during solar noon to mid-afternoon. It is usually low their original growth positions. Fish are dislocated some tide, and the sea state is calm. Under these conditions, distance from the resident reefs (Springer and McErleari shallow reefs heat rather rapidly. Ambient temperatures 1962a). Smaller cryptic organisms are often left home- are already 30'-31' C. Since the upper lethal tempera- less, the resident coral or sponge being fragmented or ture for Acroporu palrnata is 35.8' C (Mayer 1914), it possibly destroyed. When the mobile organisnl is an does not take a great deal of heating to create stressful obligate commensal (having specialized niche require- temperature levels. Shinn (1966) reported that trans- ments), it is quite likely that it will be difficult for it to planted A. ccn!icornis expelled zooxanthellae at or near locate alternative housing. Following the initial impact, 33.8' C. In contrast to cold stress, most heat stress is turbidity may remain high for a week or more before localized and is not a transported water mass phenorne- water column transparency returns to normal condi- non. tions. After several more weeks, those corals that did Cold stress was discussed in Chapter 2 under survive will either exhibit growth over the scars and seawater temperature. 'The basic geographic configura- lesions and reattach to the bottom or they will die. tion lends itself to the creation of cold water masses in Several recent reports described in detail the process of sl~allowembayments, especially Florida Bay during the healing and secondary mortality . t Iighsmith et al. passage of winter cold fronts. The shallow bay allows the (1980), Woodley et al. (198 l), and Rodgers et al. (1982) water mass maximum surface exposure to the polar air reported on the effects of several Caribbean hurricanes. mass. After cooling, the water is moved to the Atlantic Acroporu cervicorrlis and A. pulrnuta displayed impres- by tidal pumping, density gradients, and winds. The sive recovery in Belize and the U.S. Virgin Islands water mass moves through the passes and across the shelf (I-iighsmith et al. 1980; Rodgers et al. 1982); however, and into the reef communities. Reef development these same species displayed considerable secondary requires no prolonged exposure to 18O C, however, mortality in Jamaica (Woodley et al. 198 1 ). In Florida individual species can tolerate significantly lower tem- rapid recovery was reported following IIurricanes Donna perature (Mayer 1914, 191:). Most reef corals are and Betsy (Shinn 1975). The differences relate to the severely stressed at or near 14 C. Shinn (1 975), Iludson magnitude of impact; those reefs exposed to major et al. (1976), and Roberts et al. (1982) reported that hurricanes that pass close to the reef will suffer severe cold water masses originating in Florida Bay caused coral impact, while hurricanes that are of lesser magnitude or mortality to patch reefs adjacent to the passes. Most a greater distance from the reef are a lesser threat. major offshore reefs do not suffer due to mixing of the Pearson (198 1) reviewed coral reef recovery; he reported cold water with resident water masses and the modera- that following severe impact, it requires several decades ting effect of the Florida Current. Dry Tortugas has for recovery. Pearson also reported that larval recruit- suffered significant mortality of staghorn coral popula- ment was much more significant than fragment propaga- tions as reported earlier due to cold stress. tion in the reef recovery process. This finding reflects Red tide (toxic phytoplankton blooms) are not recovery of Pacific reefs following the plaguelike preda- common off the east coast, but entrained water masses tion by Amnthaster plunci (crown-of-thorns starfish) occasionally carry a red tide into the Atlantic. Inore than recovery from natural physical impacts. It Natural events are a major factor controlling appears that following major storm damage Acropora coral reef development, community structure, and palrnata and A. cervicornis may benefit from fragment species diversity. Connell (1978) reported that high propagation. In Discovery Bay, Jamaica, however, the diversity in tropical rain forests and coral reefs was few fragments of A. cervicornis alive 5 months after the related to intermediate frequency and magnitude of storm still suffered heavy mortality from surviving coral interference by natural events. Without negative impact predators (Knowlton et al. 1981). Plate 10b shows by these agents, dominant organisms outcompete other fragments of an A. palmata colony that has been over- organisms for resources, and species diversity is reduced. turned and is now sprouting new branches. Highsmith The hurricane or other natural agent opens new spatial f 1982) reviewed propagation recruitment. habitat to pioneering organisms that can successfully The other major natural impact on coral reefs is exploit new territories. In a benign environment, one or thermal stress. This can occur on either end of the spec- a few species can potentially dominate, thus reducing trum; both heat and cold stress are reported to cause diversity and the overall ability of the community to negative impact. Jaap (1 979) reported on a zooxanthel- respond to outside stress. CHAPTER 8 8.1 HUMAN EhlPACTS northward. One kind results m spoil from deepening and maintenance dredging of navigational channels; it is Coral reef8 off southeast Florida are multi-user disposed of on land and at sea. When disposed at sea wsourses, experiencing rnc-rcased exploitallon that. there is potential for reef burial and increased turbidity. results in %me negative hurnan impact on the resource. (I)ccasionally, dredging operations wlLl cut through living Although natural events are far atore severe than sttan's reefs. Sewage outfail pipe burial has also cut through individual acts, human irnpact on the recfs rnust be reefs (Shinn et al. 1977). The other form of dredging mi~ltiplicdby the number and the frequency of occur- off the southeast coast is done for beach renourishment. rcnce, which in total ~nuynot allow the reef resources Sedimentary deposits are mined and brought ashore by sufficient time for recovery, Sorrie negative ~nlipactis pipes or barges and deposited on the beach. The nraterial focumd as ggatialfy small area anti rs chronic. In recent Is a slurry; the silty runoff leaches back to sea causing yoars a number of publicattons dealing with coral reef increased sedin~entloading of the water colurnn. During g~ullubxt>n,stress, and death have docunaented or par- the fall of 1981, a beach renourishnlent project on bsaycd tlxcse subjects in some detail. Loftas C1Y70), South Miarni Beach threatened a small Linear hardground McC:tosksy and Chesher 4 197 11, Sirxtrh et al. (1 973 ), cornnlunity conkposed of significant numbers of octo- Johanncs (1"575), kndeen (llk7@), and Weiss and corals and stony corals. [lade County Pollution Control, ik;radJard (1 977) dctsiled pirilution aad Erunl:tn lnlpact Datie County Marine Institute, U.S. Cieolugical Survey, rsn caral reof communities. Voss ((197.3) and Dustan and I:lorEda IINR personnel trattsplanted about 2010 il rd77b) rewicwed problcrns in Flvritla recfs, Ct~rrcntly stony corrtl colonies titat were threatened by burial. a rli~ntbex cif cctntrcjls arc in force to tnittgiite human A critical part of all dredging activittes should t~e itripact,, 111 wrn6 cases ccrnarc~leisave worked; in others, a coherent. field study to insure that reef comrnurtit~es ttre acgative impact hsconlinuccf, and increased cfarrrape arc not endangered by dr~dging.Occasionally tratieoffs kxds c~ccurrrd,Milb@trurr iu dlfficrrlt bccausc uctiv~ticsare mtrst be mlrde, but should be rninirni~ed.Straughan ~ar~tinucturaxid Itckrrran tnpact on coral rceh 1s cctntro- (1'372) condetnned tiredging for the demise of 1:lorida vmrrl. User groups arc lable 31 Keccrrt reef' slt~pwrecks. Si~e Vessel (ft) Ilatc 1,ocatton ltnpact fcc f+i~g;~r~tl 70 i 973 Molasses Reef I ug sarlk In deep water; bargc grounded un hdrgc. rtSel, both salvaged. 60 1073-1974 Middle Sarnbo ht~arrdonedon reef fiat. I,oiu 110 I976 tooc Key Aground on a spur for 18 days. (I'latc 2frh 2nd 27a) Nolttty Ilult' 00-7 5 1077 1.ooe Key Aground, spllied fuel illorutiru 1079 bort 1,audcrdate lfarnaged 5,600 ni2 of live botto~rr. 2 shrlrri p boats Nove~nber 1,ooo Key Heef f'xtcns~vcdan~age to a spur. i082 It'cl1i~'nod 397 August Molasses Kcef 1%.S acres impacted. The threat of a major shipwreck is quite real. more the impact will be unnoticeable (Pearson 1981). Much of the ship traffic into and out of the Gulf of Mexico is concentrated just seaward of the reef tract. Fishing Traffic into the gulf is between the coral reefs and the Gulf Stream; outbound vessels travel inside the Gulf Coral reefs concentrate marine protein in a Stream to take advantage of the current. Large tankers localized area, attracting both commercial and sport or freighters grounded on a reef because of mechanical fishing interests that use various techniques to harvest difficulties or navigational errors would cause greater fish and invertebrate stocks. Negative impacts occur as a damage than small vessels. 'She 400-ft vessel MV Well- result of gear deployment and harvesting. wood grounded on Molasses Reef on 5 August 1984, Lobster fishing is the largest commercial fishery causing severe damage to the reef. located in and adjacent to the coral reef habitat area. The coflisions of large vessels is like a bulldozer This fishery was worth over $10 million in 1980 (Table blade leveling the top of the reef. Other negative effects 1). The commercial fishery uses traps or pots to harvest of shipwrecks and groundings include fuel leakage and the spiny lobster Panulirus argus. Traps are set in waters cargo and other materials lost or thrown overboard. An inshore and offshore of the reefs, and on occasion close example of a vessel grounding incident was the wreck of to the reefs. Damage occurs when the traps are set on the i10-ft nlotor vessel Loh on Looe Key Reef on 5 corals, as well as during recovery. Trap recovery takes January 1976 (Plates 26b and 27a). The vessel was place while the boat is underway; the hydraulic puller loaded with construction steel when it began taking on retrieves the trap while the boat motors to the next water. 'She captain ran the ship aground to avoid sinking. trap. The trap is pulled along the bottom for a short The vessel ended up an a spur and remained aground on distance before it clears the bottom. When the lobster the reef for 18 days. Approximately 344 m2 of reef trap is on coral, the sliding trap breaks and damages ware impacted, The crew remained aboard ship while the coral and other sessile benthos. Large numbers of traps vessel was agound and disposed of garbage, sewage, are lost when buoy lines are cut by boat propellers. Lost damaged steel cargo, and other junk including batteries lobster traps decompose in time, but cement bottoms do and taols onto the reef. Several weeks after grounding, not rapidly degrade. To eliminate this negative impact, the ship was liyheened by the retnoval of cargo onto a fishermen should be required to keep their traps out of barge and towed to Key West for repairs. 'She Lola was the reef environment proper. foreign owned, and no legal action was taken, Sport divers who harvest lobster damage the reef Salvage operations pose a threat when they occur through their efforts to dislodge lobsters from ledges and around a coral reef. Techniques used to free grounded caves. The author has seen moderately large (1.5-rn or r;utzlccsn vessels are aften counter to reef conservation. diameter) brain coral heads (D. strzgosa) dislodged and Bxplosivcs, large anchors, and ground tackle pose a turned over by enthusiastic lobster divers. Smaller fragile threat to the reef. In 1976 the Robby Ilale carrying corals and other benthic organisms are damaged in the contraband clsugs was wrecked at Looe Key Reef. A process of lobster hunting. The lobster divers often sirlvagcr uscd explosives to remove the ship from the become so engrossed in their quest to obtain lobster that reef, despite warnings by the Bureau of Land Manage- they break off coral that is in their way to get at the ment that unwarranted reef damage would result in legal lobster. prosecution under the Outer Continental Shelf Lands Hook-and-line fishing methods are employed by Act. "i'hc salvage firm of Alexander was brought to commercial and sportfishermen in the coral reef habitat. court and found guilty on two accounts. However, 'There is considerable loss of line, leaders, hooks, sinkers, the Fifth Ckcttit Court of Appeals ruled that the Outer lures, and other paraphernalia on the reefs. When diving Cantincntal Shelf Lands Act .could only be invoked in on any reef, it is common to find line, hooks, etc., cases where rrlincrtl,l or exploration or produc- caught in the coral. The hooks and line impact when the tian act~vitieswere threatening coral, reefs. hook is dragged across the coral face, causing lesions or I11 cases where tughoats are used to tow the scars. Monofilament line wraps around corals and other groa~ndeciV~SFI off tit^ reef, dare should be exercised to organisms, often causing abnormal growth. Net fishing is keep towing cables off the bottom, During the Wetlwood not common around coral reef habitats; however the salvage, towing cables caused a great deal of damage to author found a net entangled in the coral at Eastern the reef (uorafr; were turi~edover, broken, and abraded Sambo reef in 1973. Shrimp vessels anchoring at Pulaski by the tawing cables). This could have been avoided. Shoal near Dry Tortugas have made a habit of disposing More stringent regulations should be enforced when junk on the reef. The area resembles a garbage dump, salvage operations are close tu cvral reef habitats. Coast with all manner of fishing-related gear on the bottom. Guard cooperation and reef scientists' advice would aid Fish traps are a controversial issue with ~olarized in moderating the severity of negative impacts. constituents expounding their viewpoint. Traps are not Numerous other incidents could be cited. The selective, but catch all manner of biota, much of which results have a general pattern that includes initial dam- is not consumed because of cultural biases or lack of a age; sinking or saivaging of vessel: and development of market. The trap is made of steel mesh and is marked successionaI cornn~uniliesin the disturbed area. If the with a buoy. if the buoy is lost, the trap continues to stmctural integrity of the reef platform is not adversely harvest fish, many of which die in the trap (Plate 29'0). affected, the connmunity will. recoveriafter 5-10 years or The fish trap. like the lobster trap, can damage the coral 78 by physical impact during setting and recovery. National indiscrinlinately or in strong concentration (Plate 25b). Marine Fisheries Service laboratory studies show that 'The most frequently employed chemical is Quinaldine some reef fish species can enter and escape fro111 fish (Jaap and Wheaton 1975). The State of Florida requires traps at will. a permit for its use. When used in a restrained manner, it Spearfishing is the major means divers ernploy to poses a minimal threat to corals. In strong concentration harvest finfish. Spearpoints damage coral if they are it may cause zooxanthellae expulsion and stun cryptic fired into the reef. Some divers use a "bangstick" (explo- ~r~icrofaunawhich are rapidly consumed by certain sive head on the end of a spear or a stick) to harvest fish (e.g., the bluehead wrasse) that are i~nmuneto the large fish or for confidence. When exploded on the chemical. The use of Quinaldine is minimal; hence, it surface of a stony coral, ~t creates a concave crater does not pose a great threat. Other chemicals cited as about 0.5 ft in diameter and of equal depth. It is ~Ilegal major threats to coral reef co~nrnunitiesinclude a great to use spearguns in several of the marine parks, and list of materials (Johannes 1975). The use of commercial within much of Monroe County. laundry bleach (sodium hypochlorite) is a potential threat. It has been cited as a fish and invertebrate Diving Activities collecting chemical, but it also does severe damage (mortality) to sessile benthic invertebrates (Johannes Diving as a sport and hobby has increased and 1975). developed into a major industry in the Keys area. As reported earlier, it is a major economic contributor in Pollution some parts of Monroe County. With the numbers of persons involved, it is not surprising that negative im- General reviews of the effects of pollution on pacts occur. Unskilled divers grab coral for stabilization. coral reefs are Johannes (1975) and Endean (1976). The dive guides try to enforce and instill an axiom of Weiss and Goddard (1977) reviewed a case history of don't touch or collect anything. The efforts are for the coastal pollution and its effect on coral reefs off the most part successful; however, there are so many people coast of Venezuela. Bright et al. ( 1981 ) added additional to monitor that it is difficult to police everyone. Those references on the subject. Loftas (1 970), McCloskey and persons who visit the reefs in private boats are not Chesher (1971) and Smith et al. (1973) documented the governed by similar restraint. A small number of coral effects of pollution on coral reefs. Many of the previous collection violations occur within the parks and sanctu- studies were either unrealistic field and laboratory aries. Enforcement is spread very thinly; hence many experiments or from Kaneeohe Bay, Hawaii. Certainly persons probably do collect coral as a souvenir. there is cause for concern in Florida coastal waters as the Marine collecting as a hobby includes both live population continues to grow and municipal sanitary and dead material for aquariums, Some individuals sewage systems use ocean outfalls as an expedient means harvest mollusk shells and other curio items for collec- of disposal of sewage effluent. Kaneeohe Bay, Hawaii, is tions. For the most part, these specimens are dead; a classic case of what can go wrong when untreated however, some live material is taken. Collection pressure sewage and siltation from poor land management impact is quite heavy on colorful and distinctive species such a semi-closed ecosystem. Eutrophication and siltation of as helmet shells (Cassis spp.), thorny oyster (Spondylus the sea bottom extirpated a large number of the reefs spp.), and the flamingo tongue (Cyphoma gibbosum). (Smith et al. 1973). Florida coral reefs represent an open Queen conch (Strombus gigas) is harvested for food ecosystem where eutrophication is less likely to occur. and for its shell. Much of urban Monroe County uses septic tank Live marine specimen collecting has developed as sewage disposal; therefore, input into the marine envi- a minor industry and popular hobby in southeast Flor- ronment is likely. Bright et al. (1981) reported that the ida. The collection of attractive fish and invertebrates is porous limestone strata in the Florida Keys did not focused in the coral reef areas primarily between Palm retain the sewage effluent for sufficient time to allow for Beach and Key West. Brightly colored fish, crustaceans, decompositon. The aforementioned article also reported mollusks, and other species are collected and sold on the massive amount of liquid wastes pumped into the to pet shops and aquarium stores. ocean by southeast Florida coastal communities. As Specimens are collected by divers using various growth in coastal southeast Florida continues, potential techniques and transported in tanks aboard their boats. for pollution impact on coral reef communities also Professional collectors use nets, traps, and in some cases increases. Dilution in the ocean is not the solution for narcotizing chemicals to collect target species. Damage is sewage disposal. inevitable; in the quest to harvest illusive fish and Manker (1975) reported on heavy metal accumu- invertebrates, certain amounts of physical damage are lations in the sediments and corals off southeast Florida; bound to occur. Frequency and magnitude of damage he noted higher concentrations of mercury, zinc, lead, are correlated with the experience and conservation and cobalt adjacent to population centers. ethic of the collector. Part-time amateur and hobby Disposal of wastes from existing lighthouse collectors with limited experience and time pose a navigational aids may be a minor problem, yet is clearly greater threat than a professional collector with several a cause for concern. Plate 30b shows batteries and other years of experience. refuse disposed of on the reef flat at Carysfort light- Chemical collecting agents threaten corals if used house. Coast Guard maintenance crews have, over a period *rS yas, xdhposd of fapcne batteries by Bhsa>wkrng panhrr9itg.y rl:\e.+lt'~$iitfSi" UI rit) ildandge to ;lac ~c~f~$2 tlacm mrar the $a&;acnbsequentiy. acadb am released snto ~c?ara!s k.tgitotr S,r,a~ti~~z~ tlr,tr ha% high strEua~~B~, the area, poses grcv?(-fi{l~~*~~t S~?u$heabrernk"Surnria is a majar aruck faran are:* Kink$-vo~ia'gild l-r>).r ( lQr171 rcpc>rhra! aidril,agc rrr kavr vegetab!~?~itnd fm~t,Ijw of a&crtltur~l ~lr~a~rical\ grrn:ii!al tisarc\ ~niff~ao ictr.ii rrcsitibrtcrxt In sitdii~surec; (fartI.lar,cn, Ir~chlctdca,pe6tr~xdcs, cfc.) ip nnlcnsc Prlrl~us r.sc;ts off k tiat, X~r~ei,tl~it sere. 'hrorrlbaily j>r.rilutctiii) c;rrrh, CBII~\ylat~fiali ICIJI~Z$t~ the 11~~5,and rap%&rurrratf UIC' Other fnu!il nru?r~rwdh?i,lzsrcpc)rtt:tJ tl~t. lt r.ijrct~u: of surf;fce ~gntcrtalsfq-~liuwurp rams and ~mg~i.lllr,niarc qq7, pnirszsactl ~r pn:dler rrftxntty frar oil tf~~n,l scirri ,s: cxatrscs far can~~rtr,Mc( loskcy rnrrd ('tlc$kr.r l lc3 11 1 1 (I,cwzs 1'1 7 1 ) i)if fn~xcd witit sedirnc.ntr aim cdiu\crl rcg>~rt6d titat c-~rgataocBrlorr~P(:~01~1pk)t~11d6 xedticzd xrooshadity in rordll (8aL anti % igershuren 1"37>) Oil ~xntdaactawity(~*rhslnrsyntfrrsrk.~esyrrratrsira ratlo$) $18 rlivt~, iln.%per$,int in cran~ccnlrrt?tons (31 IOU-500 pprn %a jrrrra ~'t~~ri~'frp;ttisf he IIIC~~Q~~~LO~U~Y ,IIXPL$ CO~(CC~~~S~BI~O~S f~arrntul tca &orals {Lctbrs lW1r Od 2nd it5 effcct c~tritlreef& ia frikr%p.fy txard~nr~tt~ciKltwr~we surrrnranlarrg tv!Eil' pc>Ilubrut~rs J~nrrnfrtlfu coral reef so~lltxrun~ttesI tit. the ti~fd~rintllt~tl~IICIUB~: d~itaitfk~~ tlV?lj, i,clyi( dtld rtrrrcltnpic rr,kbuxe' i)l mitny reef tf~cXLa~~gUX~~~~ISX~IS, and Huihcvlictr (IY63111, Hay i IUXX ), and Bnplkt c.1 it1 (1 ")HI I tixc Isact that rn:nny arc SV~P~IC,Ieacts 11%) thi" ZPP~~LIP~~~I)~P r~rrrekulraplrl ci~httxtentr(i WB) svvlrwarl rsrl jltrltutaarrl rn tll,:t whzlc ruort ctl tltr' resrarcti rn cca'tl rech txr IYili fit+? ;ttar#tt~~it~tf~)ttd~te~tf g3<~il~tlt~r1Itii~ d~:ilt witti CIIT~~~?,chr~rtlt~ i9r n>iaxaise A tirvsrgity of mcstkorjh ftrr~sbcrn osctl tr~evulnaitc cr~ncr;'iltr;~trc~rrsue f'li6' wcrulil ;~lrolrsrrn the c~tirrr lknc etfeCtlr4.11 RtC ufr cur&i\, rndkrry rr't aha gmroieduresi tlf~diflklllb Ii~rptltatlundyna~riicr treprtrd~rc tion, rlicvclcrft~ uaed xs*pr~"~e~lfe~p~"rtt~~% u~b: UOXBC~~-Z~T~~LUIIX ttmt PIC rrtcrrt, Larval rccrurrntcrrt, scttlenrcnt, :mrf juvcnde drflli'xrlt lo rriats* rt, fteXrS cclritliliocrr. Sutnr: ukuattu,at:, ura griwtir) Jgrycltrs tra trc the b~rlopicdl prore%$ rnurt P c rzzdl accf acrltl it) irkagtnlfy @x~&(rbitre,wfkile otl~c*rs d.rlfccfa.tL hy Itif( pr~ilntion M~YU~L~Iv'rbrti afltgtdct, i~brlrtt1t)rytc&f"lfte8t ~rrvur~rc,$!rb- k24otc*ntidl (3rl piflfuticm &orilcra tn~lttdc ts1rkc.r it6 ~u!#tpa$ed~1tt8 tila ectrra! ex~vixunalratrt btuiluu rref clrnnkng and cdrgv tfrsctrargr*, vachirc! sinkings. irxtd JGrl- fleb, Iggoot~a,drrd otk~kjr wc1 c~li~~r~~t~~lertt~watt) /501>1 dcnt~,dnd d~~rcicana,iitirritrurgui irurmt ~tc.trr~lu~:~stjlrtx!isc- h8rculallt)ti cq~vw~)cgrng%s~xh cn Irrd%emcrrt~cvtitr;rfjotrr ttrm and t~~r~hpt~rfdti~t~tJC t~vit~~h, l he 'Lllncrrb M:rradgr- Ril&al fRPPrl oC."CIEi ~t>ildOll(.fttgl.~~ialk4nl B~CII. 111d ~COOV~ Itxcrrl Srrvrcv IMhiSi (i)cp,irtnicnE of the inte~lnr) kt8rratalukt1t wtltr lwtt turbtxieeret: probably tciiiu~e ccrrrlrul&Iertslrrg .knd uprtr:sttoniil ,Ispe~i$01 olt\ttcrrc r,aI sxsrurx~rc E xper rnnentl* txavc iio~tkn~cxrlctPrcsg+car~sg srt erpXx~r~trr,nand d~velopnxrntIfr tealcr,~f water\ r~tr91B8htuv 10 C 8.2 CORAL RLEk KLSUURG'E MANAGEMENf Xl~nrocZ ounty prtrfatt,rt\ crsc 01 spsar&uns for ,i unique. rtlan:agerlrvnt for subsea rc>c)\trLc:, ttordc~to I orrg, Key. f'srks, Sanctuaries, and Moi~~trner~rs State of Florldu IZlsca 5 ric ,Y~ztmi~trisii'l~ilr6 IIE.YC'/ i 'lO.822 Ild , ile+f>ar:rr~t~tif<~f !ktzfuml Rtwt~ri~e~(t!>~Vl<). DSR I ikrurc. f I ocdti'd oft wutRe,~'rt I ~UIIL~I~,~IILIU~~II~: is rc.;prrnslble for r~iar~agcnicrtt01 all r~ldrttuefrsherlcs ;and nortrecf ~rtvs111 f%nc,ryncLLiy, Rect ,ire,ts ;trr ht"j\i(.iril t>t rest?trrc t.s IJI St~tewatrfs In tliir dren 01 consldcrattort. i lliott ,ittd the KdgecZ Keys to 'i 60-f t ( 18.3-rrt t tiepbh thrs ~ncludcs lobstrr, stkook, nndpper, &roug>r*r, cilltcs I ci the. southwest thrr park ~ttfjurni Jolifi I9cel~nrkdtrrp co~xr~iiercral,,ind spcrrt spectra, nrr,irtgr;ruuc, se:tgrrcss, ~ctd C crr'il Kret St,rtc Pdrk JII~hey 1,dxpcr Ndr~i,fl~iliM,~ranc ~vr,tlteet ~oni!rltts~~f~t.$f)N 13 tias spc.ctfl~*police (?owur\ b6inctu.ir> Io the r~cjrtltthe. I-INI' dtt~oir~sk+ii;c,ryrrz l3;rq. tflrouph the L*lorrd,t Mirnrte I",ttrtd 10 erttosic Stdtr i <* if7 4 f,fijif ttidlt%%Gi idlvdpc cjt fri\t Adey, W. 1975. The algal ridges and coral reefs of St. Aildredge, A.L., and J.M. King. 1977. Distribution, Croix, their structure and Holocene develop- abundance, and substrate preferences of demer- ment. Atoll Res. Bull. 187. 67 pp. sal reef zooplankton at Lizard Island Lagoon, Great Barrier Reef. Mar. Biol. 41.317-333. Adey, W. 1977. Shallow water Holocene biohergns of the Caribbean Sea and West Indies. Pages Aller. R.C., and R.E. Dodge. 1974. Animal sediment xxi-xxiv, Vol. 2 in Proceedings: third iriter- relations in a tropical lagoon, Discovery Bay, national coral reef sympc~sium. University of Jamaica. J. Mar. Res. 32(2)'209-232. Miami. Miami. Fla. Alt, D., and H. Brooks. 1965. Age of the Florida Adey, W. 1978. Coral reef morphogenesis: multidi- marine terraces. J. Ceol. 73(2):406-411. mensional model. Science 202:831-837. Antonius, A. 1974a. Final report of the coral reef Adey, W., and J. Vassar. 1975. Colonization, succes- group of the Florida Keys Project for the pro- sion, and growth rates of tropical crustose coral- ject year 1973. Harbor Branch Foundation, Ft. line algae. Phycologia 14:55-69. Pierce. Fla. 193 pp. Agassiz, A. 1885. The Tortugas and Florida reefs. Antonius, A. 197411. New observations on coral de- Mem. Am. Acad. Arts Sci. ll:107-134. struction in reefs. Association of Island Marine Laboratories of the Caribbean, 10th meeting Agassiz, A. 1888. Three cruises of the U.S. Coast and 4-7 September 1973, Mayaguez, Puerto Rico. Geodetic Survey steamer Blake in the Gulf of (Abstr.) Mexico, Caribbean, and along the Atlantic coast of the U.S. from 1877-1880. 2 vols.. Boston, rl~~to~~ius,A. 1976. Kranke Korallen: Riffzerstorung. Mass., and in Bull. Mus. Comp. Zool. 14 anti 15. Unlschail 76:493-494. Agassiz, A. 1890. On the rate of growth of corals. Antonius, A. 1977. Coral mortality in reefs: a prob- Bull. Mus. Comp. Zool. 20(2):61-64. lem for science and management. Pages 617-623, Vol. 1 In Proceedings. third international coral Agassiz, L. 1852. Florida reefs, keys and coast. reef symposium. 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I9 78 t rtv~ri~rxrncrtxr1mud pact $trrtzmetrt rxad frvhery rnlttt Erc~u~,$,)it , bf, Hrrlao, and J 5ttunder.i 187X R~cvfb an3 sr.!errrtul crorbunatc%,ec011ig) asrd xcdrimaafnln- g) Studre* *stn gectlogy 4, pZmcriratl Awsc~~tarott of Iktr~le~~rti(ieoil~grsfh, fuhr, Ijlile 42 R pp Ery, W C; IP17&X ike IPkc~lopj raf tire Porrfcia 6.or~l %~BCL-tlntt 25 Acodzrrxrc Pte~!~NK*% \t;oek 59 2 Crerrx)alstte. F , and A,C3 &lrclartp t 970. Se.ularrrvttr 1x8 8;titfnptftgg Pirle: d8f ?at%' KT<&P~CL~CptiRPII" f X;larolrf. J , wad P karar\ 19h2 Suro~Pr f*lc$rrJsapractu reef%rrt C~dtf~oft~te~PO~UCIIOII (.rf I)&@ CUIPE lh- ~dlurrra ~~rb~~f#nrPst~rr.bFtri~~Mrur Geed 4%3b 1 296. (;;tlatr.i, P lVal Ilre pullshaeoe Bslurtu ol (trc $*rltrnautr: fslsrrcib Bu-1). Be Mu9 @~d? fit+%1 X~.oii 2 LtS I 99-2t l jtnqt~tkrg, K.3 i9SR k~r~xrca~rclinttualziktp!, of gr~litta~e and eloa$rrtmrnt prrtrcles 88% sox%@%oath FLurrd~ crsbitrartc wdxxrtcz~ktr Buik Am A%uc Petrol Ccok QO(i(lbk.2 $84-2427 Ginsburg, R.N. ed. 1972a. 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CIX~~P~XO~JPICIbiltorm effatl~'19 an ~Xtd Bfitidl E#QX~~~L~CB$r9?6ft6 mii cay%. NIL~WI~ Tailir~zt,F.H. 19BS. A de*riprion of the curd ~rlnrc- IP6[48s4): 511aSX 5, (urr: arP 'Ekiisr We& (Tang~rayika "fcrritury, E;I,+~. Al&r) and the leh faunla Prm. Zaut. *e. Stdd~rlrf,D, R. t86.3. Efiwts of Narricrrnt: tfsttis obi Londi, $45 831470. the itrfrhk Hortduxm tacfe and cay& Wt . 36.5 1, 196X. Atcxlf Rw, 1Xlull, 95: 1.842. r*arlba€, F,I.t., B.C. Rawell, annll G,R,'k". Anlttmtl, 15378. Card tmf Ik& ccrtnmunitscs. urntabis, St&drrrl+ B.& PPB9, k~lsgy merpfrat@g~uf lW4gvedtw ~~8tatrmahs@X. M~nogr~48:42%- kvreent cord xcet, Biuf, Rev. Caantb. Pbties* SCX, 640. 443433-491. Faylor, D.L. 197 3. Aigrl symbiuals of iarrertcbwtea. &an@,Re** M$erubIaI. 27. t 7 1-Xki;7. $34 Taylor, D.L., ed. f 977. Proceedings; third inrema- Ttlrmel. R.J., and R.G. Swanson, t9"f. TXle develop- tional coral reef symposium. Miami, Fla. 2 vols.: nwnt af Roltriguer Bank, a recent carbonate 1 Biology. 7, Geology. 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Saitmplinga clcmerbal rooplank- eiomcb, fUnlv. eI tcm: a ca~vtgruribc~nof field calfectirrns using three diffcrertt ctttcqenerr traps. 3. Exp. Mar, Bid. Erol. 6f:111-124. Zerlicxs, W. 1974. Tropical rtrnrlrtu Irrvertrhrates x,! arxufi8rrxr FIurida arid the U~~BOI~PICXIIIII~~~*, John Wrley and Suns, New Yrtrk 132 pp, Zrrrrr~tx, J 1982, l%e eet~logy of the seagrasses crf murh i-lrrrrdr. a cnn~trsunity proflie, I2.S" I. xsh Wildl, Scru, EtioX, Serv, fbrogrrtn FWS/C)BS.82/7 5. I "i OPp, the gcart~#$id@~#srrr"cl Tortugas Lab. (452) -- -..- 4. Title and Subtitla THE ECOLOGY OF THE SOUTH FLOR . Jaap ," ------.- '." "- s Affiliation: Department of Natural Resources Research Laboratory --.-.-"%.--",--,.-.-"*"-,- rsburg, FL 33701 Minerals Managernent Service Gulf of Mexico OCS Region P. 0. Box 7944 Metairie, LA 70010 "...." .. ~.- " - 15. Supplementary Notes ( * U. S. Fish and Wildlife Service Report Nunrber, FklS/OBS-02/08 I ------I ---."..-.-..." - -l^-ell-p-XI -*--^- --X17^i- 18. Absbrac An overview of coral reef research in southern Florida i\ provided as cl prclude to a genui ne descript.ion of the coral reef ecorys PI TI i rr the Florida Kcyc arid rurrnuriding environments. Coral reef comnluni ty types, rr.ef henthus, planktor\ drid reef fish are given specific treatrnerat ' Coral reef ecol agy and mar~dqerneritdre dcscri bcd. "---" ------17. 00cumsmt Analys8r 0 Oexdpron Coral reefs b. IdentlCinlDosn Ended Tarms Southern Florida, subtrop.ica1, marine, ecosy'it;ems, ecology, biology