ASSESSMENT OF REEF FISHES AT SOMBRERO KEY, FLORIDA
T.E. Thompson D. G. Lindqtast L E. Ckrvijo S. K. Bolden S. W. Burk N. C. Drayton Departmentof Biological Sciencesand Center for Marine Science Research Universityof North Carolina Wilniington,NORTH CAROLINA 28403-3297U.S.A.
Wequantitatively assessed the reeffish assemblageat SombreroKey, a bankreef off' the middle Florida Keys,during May 8-11,1988 using the stationarysurvey method. Wedivided the reef fish assemblageintothree depth zones < 3m,3'-6 m, and 6-10m!. Thesezones approrimated, respectively, the Millepora ridge zone 1!, the Acropora palmataarea zone2!, and the reeffront basewhich was characterizedby moresand and /esscoral cover zone3!. Eighty-eightspecies in 27 families, representedmainly by pomacentrids,labrids, and haemuiM were observedin all zones.The largest number of individualsand thehighest Shannon speciesdiversity occurred in zone1, however,species richness was higher in zones 2 and 3.
INTRODUCTION
Coral reef fish assemblagesare complexand there existsa lack of quantitativedata concerningabundance and distribution patterns of fishspecies, particularly for Caribbeanand Florida reefs Alevizon et al. 1985,Bohnsack et al. 1985!. This has resulted in a lack of understandingof the factorswhich govern the structureof coral reef fish asseinblages.Most studieshave dealt with smallpatch reefs, artificial reefs,or discreetportions of a largerreef systein Ogden 1982!. Only a fewstudies have dealt with reef fish abundanceand distribution by depth. Tilmant 984! surveyedthe literatureconcerning Florida coral reefsand found a characteristicspecies composition associated with differentreef typesand depthzones within a particularreef. Alevizon et al. 985! foundfish speciescomposition and abundanceto vary significantlywith depth at Deep Water Cay locatedof'f Grand BahamaIsland. However, variationin fish communitystructure was influenced by coral zonation rather thandepth itself. Bohnsacket al. 985! found that fish speciesoccurrence and abundancevaried with habitat associatedwith different depthzones on Looe Key Reef.
375 Divingfor Science-.19Ã
Coralreef nshery resources of Floridaand the Caribbean are facing increasing utiliza- tion andadverse environmental pressure by man. Usesinclude commerciai fishing, recrea- tional fishing,spearfishing, and collecting. Although someof Florida's coral reefs havebeen designatedas sanctuaries,all are facedwith an increasein diving and boatingactivities. 7i aditionallyviewed as nonconsiipptiv, diving activities may now be a significantdisturbance in someareas. In addition, Florida's coral reefs appear to be suffering from poor water quality associatedwith increased development of the Florida Keys and south Florida, in general Whrd 1990!.Proper manageinent of coralreef fisheriesrequires quantitative information concern- ingfish abundance, distribution, and structure of thefish assemblages.
Theobjective of thispaper is to providequantitative data regarding reef fish abundance anddistribution by depth zones within the Sombrero Key reef. Thisreef is currentlynot protectedunder any sanctuary designation and has received increased disturbance byway of divingand spearfishingactivities. Our null hypothesisis that there is no differencein Gsh speciesdiversity, incan fish species richness, and fish species abundance in three vertical depth zoneson the coral reef defined as < 3 m, 3-6 m, and 6-10 m.
METHODS
StadyArea
All visualsampling was conducted at SombreroKey located within the FloridaReef Tractapproximately 8 km south of Boot Key and 35 km northeast of LooeKey National Marine Sanctuary.Sombrero Key has well defined spur and groove forinations on the seawardside downto about 92 m. The tops of some spurs may be awash atlow tide. The shoreward edge of thereef is surrounded bysand interspersed withseagrass beds ?%dahlia testudiruurr!. Below10 m thespur and groove forinations give way to a combinationofsand mixed with low profilelimestone covered with octocorals andsponges. Wedivided the reef into three depth zones.Zone 1 beganatthe top of the reef and ended at a depthof 3 m.Zone 2 rangedfrom 3 mto 6 m.Zone 3 startedat6 mand ended at the bases of thespur formations in 10 m of water.
SamplingTechniiine Samplingwas conducted between 0900 and 1530 hours on May 8th through the 11th, 1988with the exception ofMay 9th when sea conditions precluded safe diving. Six SCUBA divers,working inpairs, used the stationary visual survey methods ofBohnsack and Bannerot 986!.Each pair of divers began a sample ina zonebyseparating from one another a distance ator just beyond the limit of visibility. In some cases, the pair were not able to see one another suchas when sainpling opposite sides of a spur.However, efforts were made to stay within easyswimming distance forsafety considerations. Eachdiver began by facing seaward and
376 Thompsonet aL ReefFishes at SombreroKey, Flonda listing all speciesseen within an imaginarycylinder stretching from the bottom to the surface. A radiusof 7.5 m was usedsince Bohnsack and Bannerot 9&6! determinedthat this radius maxinnzedthe number of speciesand individuals that a divercould observe including cryptic andshy species. The diverrotated clockwise and listed species for 5 minutes.This usually requiredseveral rotations. Except for speciesnot likely to remainin thearea such as carangids, individualsof a specieswere not countedduring the first 5 minutes.
Abbreviatedscientific names using the first threeletters of thegenus and first four of the speciesaided in efficientcoding of dataonto water proof datasheets. At the endof 5 minutes,numbers of eachspecies within the samplingarea were counted beginning at the bottomof thelist andworking up. Afterall specieshad been counted, the percentage of sand, limestone,gorgonia, and hard coral was estimated for eachsample. Each sample required 15-2Gminutes. The pair of diversthen swamto a differentsample site. The numberof swim kicksrequired to swimto thenew site was obtained from a randomnumbers table printed on waterproof paper. A completedescription and analysis of the samplingtechnique can be foundin Bohnsackand Bannerot 986!.
Data Analysis
Data were analyzedusing Statistical Analysis Systems, Inc. software and the UNCW VAXcoinputer. Total number and mean number were calculated for eachfish species by zone. Additionalcomparisons by zoneincluded species richness mean number of species!and speciesdiversity. The Shannon-Weiner diversity index as presented in Smith 974! wasused to calculatespecies diversity.
RESULTS AND DISCUSSION
Threedays of samplingresulted in a totalof 47samples with 14,18, and 15 being taken in zones1,? and 3, respectively.The averagedepth sampled in eachzone was 2.7 tn in zone 1, 5.1 m iri zone? and 7.2 m in zone 3.
Habitat Description
Zone 1 wasdominated by hardcoral, followed by limestone, gorgonia, and sand. Coral found in this zone was primarily fire coral, MNeporacomp/anna. Limestoneand sand dominatedzone 2 followedby hardcoral and gorgonia. Elkhorn coral, Acmpora pzLmata, dominatedthe hard coralsin zone2 althoughsoine smaller heads of brain coral,Monazrtrea annularis,were present. Zone 3 wasdominated by sandfollowed by hard coral,limestone, andgorgonia. Both brain and elkhorn coral were present, however, brain coral donunated by formingmassive heads at thebases of thespurs. As depthincreased the relative percentages
377 Divingfor Science..J9N
ofgorgonia decreased andthat of sand increased consistently. Theincrease insand with depth wasa resultof thepresence of the wider sand grooves between the spur formations.
FishAssemblage Eighty-eightspecies representing 27families were observed during the study period ghble1!. Fainiliescontaining themost species were the Pornacentridae, Labridae, and Haemulidaefollowed by the Scaridae and Serranidae. Forty-one species 6.6%! were observedinall three zones. Five species .7%! were observed onlyin zone 1, 10 1.3%! wereobserved only in zone 2 andeight species 9.1%! were observed only in zone 3 Table2!. Theoccurrence ofa species inoiily one zone may be attributed tothe habitat present insome cases.For example, the large percentage of sand found in zone 3 mayhave resulted in the presenceofsuch species asthe yellow stingray and bridled gaby. The short study period may alsohave resulted inthe observation ofa speciesinonly one zone. A totalof 11,996individuals were observed during the study period Table 1!. The greatestnumber of species were observed in zones 2 and3. Meanspecies richness was also highestinzones 2 and 3 withvalues of19.9 and 19.2 respectively. Mean species richness for zone1 was16.5. Species diversity forzones 1,2, and 3 was4.5, 4.3, and 4.2 respectively. This indicatesthat although more species were present in zones 2 and3, thedistribution of individualsamong species was tnore even in zone1. Tenspecies comprised 78.1% of thetotal number observed for allthree reef zones Thble3!. These species included 3 labrids, 3 pomacentrids, 2 haeinulids, 1 carangid, and1 lutjanid.Bluehead wrasse,, sergeant major, smallmouth grunt, and the bicolor damsel numeri- callydominated theSombrero Keyfish assemblage. Although their relative ranking changed, theywere thefour most abundant fishspecies throughout allthree depth zones. The feeding requirementsofthree ofthese species xnaybe less restrictive than other similar species. This mayhave resulted intheir numerical domination ofthe reef fish community. Thebluehead wrassereadily takes plankton inthe water coluimL Other labrids consume macroinvertebrates associatedwiththe substratum. Thesergeant inajor isa rnidwaterfeeding planktivore. The bicolordamsel feeds on plankton asweil as algae. Barjackwas among the ten most dominant species in all three zones. Likewise, yellowtailsnapper andbluestriped gruntwere among theten most numerous species inall threezones. The striped grunt was included inthe top ten species inzone 1 aswas the French gruntin zone 3. Thegrunts are macroinvertebrate bottomfeeders and feed off the reef at nightusing the reef for shelter during the day. Theonly other poinacentrid fishamong theten most dominant wasthe dusky damsel inzone 1.One mullid «nd 2 labridsinzone 2 and2 labrids inzone 3 roundedoutthe ten most dominantspecies in each zone. Unlike the haemulids, thespecies from these last three familiesaredependent onthe reef for food aswell as shelter andbecause ofthis a pattern seemsapparent. A herbivorous bottom feeder, the dusky damsel, inzone 1 appearstobe
378 'Aompsoner aL ReefFishes at SombreroKey, Rorida
replacedby the macroinvertebratefeeders in zones2 and3. Thealgal food source required by the duskydamsel might not be asabundant in zones2 and3.
Family Accounts
Holoeen friday
Therelative abundance of all threespecies of squirrelfish was greatest in zone3 Pebble 1!. Nocturnallyactive, these fish hover in or closeto crevices during the day fresher 1984!. Thenumerous crevices and ledges at the base of eachlimestone spur provide optimum daytime habitatfor thesespecies. Bohnsack et al. 985! reportedthe holocentrids onLocc Key Reef tobe most abundant in theforereef and buttress zones. Both zones overlap in termsof depth andhabitat composition with the zone3 of this study.
Serranidae
The graysbyand harlequinbass were more abundantin zone2 liable1!. Bohnsacket al. 985! found thesespecies to be relativelyabundant in the forereefand buttresszones althoughthe highest abundance was recorded over deep live bottomhabitat.
Pomacentridae
The three most numerousdamselfishes were the sergeantmajor, bicolor damselfish, anddusky damselfish Table I!. Sergeantmajorswere most abundant in zones2and3. Bicolor damselfishwere most abundant in zones1 and3 whiledusky damselfish were most abundant in zone 1. The two chromisspecies were most abundant in zone3. YeHowtaildamse16sh were mostabundant in zones1 and2 whereasbeaugregory were moreabundant in zone2. Threespotdamselfish were about equally distributed throughout aU zones. Generally, the planktivores were more abundant in zones 2 and 3 while the strict herbivores were more abundantin zones1 and2. Theconsistent distribution of thethreespot damsel throughout all zonesmay have been a resultof itsrelatively more aggressive behavior. Bohnsack et aL 985! reportedsimilar distributions for the planktivorous species at LooeKey Reef.
Labridae
Blueheadwrasse were most abundantin zone1 Table1!. The clownand rainbow wrassewere also most abundant in zone 1. YeHowheadwrasse were most abundant in zone 2 andpuddingwife were slightly more abundant in zones2 and3. Bohnsacket aL 985! reported that most of the labrids they observedwere more abundantin the lagoon rubble habitat. Althoughwe did not samplethis habitat,three species were most abundant in zone1 suggesting a preferencefor the shallowerhabitats similar to the lagoonrubble sampled by Bohnsacket al. 985!. The preferenceof the shallowerhabitat may result in a reductionin predation pressureon thesespecies.
379 Divingfor Science...1990
Scaridae Stripedparrotfish were slightly more abundant inzones 1 and2 TableI!. Therainbow andprincess pan otfishes were relatively more abundant inzone 2 TableI!. Theredtail and stoplightparrotfishes were slightly more abundant inzone 1. Redband and redfin parrotfishes weremost abundant in zones2 and3 respectively.
Haemalidae Smallmouthandbluestriped grunt were dominant inall zones Pable I!. Stripedgrunt wererelatively abundant inzone I but were notably absent inzones 2 and3. Frenchgrunt, torntate,and Spanish grunt more abundant inzone 3. Whitegrunt and caesar grunt were tnost abundantin zone2. Themost notable similarity between these results and those of Bohnsack etal. 985!is the relatively high abundance oftomtate they reported in thebuttress zone which is habitat most like our zone3.
Chaetodoatidaeand Pomacanthidae Thebutterflyfishes andangelfishes weremost abundant inzones 2 or3 TableI!. The completeabsence of thesetwo families from zone I mayhave been due to a lackof suitable forage.The butterflyfishes consume coral polyps and the angelfishes depend upon sponges. Bothfood items may have been more abundant inzones 2 and 3. Bohnsacketal. 985! reportedthese families tobe more abundant inthe buttress zone or deeper limestone habitat whichwas dominated by sponges and octocorals.
Gobiidae Theneon goby was most abundant inzone 3 TableI!. This was probably due to the largenumber ofbrain coral heads present inthat zone since this goby was observed onlyon thisspecies of coralat Sombrero Key.
Acanthuridae Doctorfishandblue tang were most abundant inzone 1 whileocean surgeon were most abundantinzone 3 Table1!.The greater abundances of2 species inzone 1 suggestsome sjtnjlaritywith those of Bohnsacket al. 985! since they found acanthurids to be more abundantinthe lagoon rubble and forereef habitats atLooe Key Reef, both relatively shallow habitats.Tilmant 984! also reported surgeon 6shes tobe numerous within the top zone of patch reefs.
380 Thompsonet aL.Reef Fishes at SombreroKey, Florida
ACKNOWLEDGEMENTS Thispaper is contribution No. 024, Center for Marine Science Research, University of North Carolinaat Wilmington.
LITERATURE CITED
Alevizon,W., R. Richardson,P. Pitts, and G. Serviss.1985. Coral zonation and patterns of community structure m Bahamianreef fishes. Bull. Mar. Sci. 36: 304-318.
Bohnsack,J. A., D. E. Harper,D. B. McClellan,D. L Sutherland,and M. W White. 1985.A resourcesurvey of fisheswithinthe Looe Key National Marine Sanctuary. NOAATech, Rep.OCRM SPDU.S. Dept. Cornrner.,NOAA,Office of Oceanand Coastal Resource Management.108 pp. Bohnsack,J. A., and S. P. Bannerot. 1986. A stationaryvisual census technique for quantita- tivelyassessing community structure of coralreef fishes. NOAA Tech. Rept. NMFS 41: 1-15.
Ogden,J. C. 1982. Fisheriesmanagement and the structureof coralreef fish communities. Pages147-159 Ig.. G.R. Huntsman,W.R. Nicholson and W.W. Fox,Jr. eds!,The BiologicalBasis for ReefFishery Management. NOAA TechnicalMemorandum NMFS-SEFC-80.
Smith,R. L 1974. Ecologyand Field Biology.Harper and Row,Pub. New York, N.Y. Thresher,R. E. 1984.Reproduction in Reef Fishes. TFH Publications, Inc. Ltd. Neptune City, NJ.
Tilmant,J. T. 1984.Reef fish. Pages 52-63. Ig. WC. Jaap ed.!,The Ecologyof the South FloridaCoral Reefs: a CommunityProfile. Fish and WildMe Service, U.S. Dept. of the Interior, Washington,D.C. FWS/OBS-82-08,MMS 84-0038. Ward,F. 1990.Florida's coral reefs are imperiled. Nat. Geogr. 178 !: 114-132.
381 Divingfor Science...I990
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382 Thompsonet al: ReefFishes at SombreroKey, Rorida
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Theswtrnmtnghabits of threespecies of carclurnbnidsharks, a tiger Galeocerdo cuvier!,a bull Carcharhinusleucas! and four brownsharks ~charhinus plumbeus!,held at TheLiving Seas Pavilion, were observed over a two yearpenod 988and1989!. All of thesharks are females with the exception of the bullshark. Alterationsin swimminghabits relative to frequently visited areas and depthsof the tank werenoted throughout a varietyof environmentalcPuutges. Changes includedthe intmhcction and removal of sharks,and the placement of a divider whichdecreased available space. Daily environmental stimuli consistedof diver anddolphin confrontation and artijicial feedingoperations. Vunughout thestudy, sharksshowed an avoidanceof the areasnear the dolphin. All the sharksswam next to the outer wail of the aquarium D&nlylit, open areas near the sharks' feedingstationwere frequented. Divers seemed to haveQtle effec on sharkhabits exceptingthe bull shark. Theintroduction of thedivider producecE no observable changesin the sharks'habits. INTRODUCTION Sharkslive in a veryconcealing environment and often have wide ranginghabits McKibbenand Nelson, 1986! and any study of theirbehavior is «iif6cult andpotentially expensive.It is often impossibleto determinethe stimuli that sharks encounter and hence their reactions to them. Weihs,et aL, 981! statedthat captive carcharhinid sharks in a largesimulated reef area,swam in establishedpatterns. This observation was verified by ~ow andHewitt 988! whodescribed selected swim patterns of sixcaptive shark species. The largeaquarium at The LivingSeas Pavilion places few restrictionson shark movementsand allows room for artificialreef structures and a variety ~f otherma .ne]j fe Thestudy of carcharhinids inthis extremely large controlled enviro~+nt maygeld a better understandingofspatial requirements ofsharks. 'His siinulatedenviron entenabled us to Divingfor Science...1990 documentthe sharks'swimming habits in responseto additionsand subtractionsof other carcharhinids.It was also possible to document the sharks' swimming habits in responseto dailystimuli such as diver and dolphin presence, a condition which may exist in theirnatural environment,buthas never been adequately observed. Myrberg and Gruber 974! reported aggressivebehaviors tooccur between resident bonnetheads anddiver-observers. According toThompson andSpringer 961! observations havebeen made of wild dolphins driving sharks awayfrom their young. However, dolphins and sharks may feed together without apparent conflict.Gilbert, et al., 972! observedthat "swimming habits of brownsharks were not significantlyaltered by the presence of porpoises". MATERIALS AND METHODS Thetracking of swimnung patterns spanned January 1988 through November 1989 and consistedof 492ten-minute observations 82 hourstotal!. The establishedcarcharhinid populationsince 19&6 consisted ofone female tiger shark, Gakocerdo cuvier, .4 m totallength TL!and 214 kg!, one male bull shark, Carcharhinus kucas, .4 m TLand 158 kg!, and one femalesandbar shark, Carchanbinus phcmbeas, .0 m TLand 57 kg!. Over a periodoftwo years,changes within the shark population occurred. A secondfemale sandbar shark, Brown 2, .0 mTL and 58 kg! was added in May1988. The tiger and Brown 2 wereremoved in Januaryof1989. Two fernale sandbar sharks, Browns 3 and 4 eachat2.2 m and44kg! were addedin February1989. Thestudy was conducted inThe Living Seas Pavilion which contains 21.5 million liters ofartificial saltwater, andis cylindrical inshape: 60m indiameter and8.2 m deep see Figure 1!.Three submerged guestviewing areas are the Inner Tank Module ITM!, Tunnel 1,and Tunnel2.Although these structures could be possible barriers, the sharks had been observed crossingover them. Man-made, fiberglass coralheads, ranging in height between 0.5 tn and 7.9m, are randomly dispersed throughout theaquarium. Allthe water parameters remained constantthroughout theduration ofthe study. Temperature andsalinity varied between 24.1 ~ 0.2C and31.0~ 1 ppt,respectively. Theyear round light:dark ratio is 17:7. Forobservations, theaquarium wasdivided into eight pie shaped sections numbered 1-8in Figure 2!. Eachsection was further divided into four areas lettered A-D!, to denote increasingdistance from the ITM. It shouldbe noted that although these sections varied in size,they were based onnatural divisions inthe aquarium, andwere the most accurate wayto pinpointthe sharks' positions. Observations weretaken between thehours of 0700and 2300 frominside the ITM or in the water using scuba gear. During each observation, a single shark wasobserved forten minutes. The subject's position was plotted at 15second intervals on handheld maps. The depth ofthe subject ateach point was noted ashigh, medium, orlow, correspondingto2.7 m increments.Thepaths between each point were plotted todenote the shark'sexact swim path. Environmental stimuhsuch asdivers, feeding, etc., were recorded to categorize observations. 386 Weber: Carclmrhinid Sharks at FPCOT Center Observationswere taken randomly,and coveredall the different conditionswhich existedin the aquarium,on a dailybasis. These conditions are the presence and absence of diversor dolphins,arid the occurrence of fish or shark feedings. The fish are hand fed a variety of seafood,from within the water. Divers perform these feedings twice a day,inorning aiid night.In additionto thesefeedings, the divers are present in theaquarium approxitnately 10 to 14hours a dayfor maintenanceand show purposes. The sharks are fed whole and cut fish, everyother day from the surface of theaquarium see shaded area of Figure1!. ThreeAtlantic Bottlenoseddolphins Tursiops truncatus! were present in theaquarium up untilFebruary 1989.Their holding pools extend off section4D asseen in Figure2. Thedolphins' play progressedto thepoint where it wasnecessary to separate thein physically froin therest of the animalsby theinstallation of a divider Figure 2! madeof PVCpipe and rope which allows smallerfish to swimthrough. Eighty-twohours of data were collected from January 1988 through November of 1989, whichwere spread over seven different time periods Cumulative 1 Cum1!: January 1988- May1988. 1 hourscollected on three sharks!, Transition I: May 1988. Brown 2 added. hourscollected on four sharks!, Cumulative 2 Cum2!: June1988-October 1988. 8 hours collectedonfour sharks!, January 1989: First three weeks inJanuary 1989. Removal ofTiger andBrown 2. hourscollected on two sharks!, Transitiori II: January1989-February 1989. Browns3 8r.4 wereadded. hourscollected on foursharks!,' '[Before %hll, March 1989-May1989! After Wall May 1989-Noveinber1989!. Dividing wall installedbetween sections4 and5 see Figure2!. 5 hourscollected on four sharks!.After one month adjustmentperiod. 9 hourscollected on four sharks!j. Chi-square tests were performed to determinerandomness of svtimming. RESULTS Thesharks' swimming habits were not random throughout the duration of the study x = 7.13-152.2df= 3P >0.001!. Table 1 showseach shark's preferred sections throughout theseven time periods. A preferredsection isone in wbich the subject spent greater than 10% of itstime. Random swinMiiag patterns would be reflected in 3%time spent in eachsection. Theaverage amount of timespent in eachnon-preferred section was 1.6% ~ 0.5% 387 Dnkg for Science...le% F ~ ~ or gag}yski~ QvcFtbc cMIFsc ortlpo ~. rtlhcil @+~ t t ~ Clm I ~ ~ Wall! to represcateutque eavlreamcolal coa@tt DEPIHS: Depthdata were analyzed usingthe three depth divisions ofHIGH, MEDIUM, tuel H!W,corresponding to2.7 m depthincrements. Comparative relative depth proflles are showninFigures 3-5.Percent timespent atdepth isgraphed asa functionofdepth foreach subjectthroughout theseven timeperiods. Theaverage amount oftime expected tobespent atone particular depth division would be 33%. Chi square tests were performed totest randomnessineach sharks depth distribution . Results arepresented inFigures 44. DISCUSSION InCum 1,it is believed thatBrown 1 was ableto occupy section SCdue to her size wassmaller andmore rnaneuverab]e whichwould enable herto easily pass through thela'Ã coralheadoccupyingthat section. Thismaneuverability isalso reflected inher preference for MEDIUMe OJMdepth!. depthdivision during thistime as the opening inthe coralhead isat « DuringTransition I,both thetiger andthe bull changed swimming habitsslightly > ~ moretimewas spent inthe HIGH depth divisioiL Brown 1'shabits immediately changd duringTransition I,whea both preferred tanltsections anddepth levelchanged tocorresp withthe preferences ofthe nay added ~nsp~dfi~ Br~2. This immediate ch~eg ' g ' 'J~.eflect anitlterspecific relationshipof schooling, whichwould supp" thefindings ' gsof Gilbert ' aI 967!eg aI andCastro 983! orit cou!d beattributed todo Gilbertet al 967! statedat th individualsofthe same sizehave been known tosc"~ Weber:C roc/garhinidSheets gr Epcor Center in nature.~ II Castro 983! statedthat sarrdbar sshark ar "t ten d tot segregateby sex...oftenforming scnools . nngWm 4 thetiger still did not reflect any changes inswimming habits but maintaineda preference forthe HIGH depth division. This small change inhabits could have beendue to lack of space atLOW depth division. Ingeneral thetiger displayed theleast amountof changein swimming habits. Also during Curn 2, the bull shark was observed to changehispreferred sections byavoiding theone that all the other sharks occupied. A possibleexplanation could be due to the lack of space. Throughout Cum 2, Brown 1 main- taineda preference forsections anddepth divisions withor near Brown 2,again possibly due to either schoolingor dominance. In January1989, the bull did not migrate back into sectiori 8D, but remained in the sametwo sections and displayed nodepth preference. Brown 1 stillretained a preferred sectionnear ones previously occupied byBrown 2 mostofthe time, however, shedid migrate overto 7D which was occupied bythe bull. Similar tothe bull, Brown 1 alsodisplayed nodepth preference. DuringTransition II,the bull migrated back into 8D, possibly tohave more space since noother sharks occupied it. He alsopreferred the same sections as in Curn1. Thebull also developeda depthpreference for the LOWsection which was the sameas all the othersharks. Brown1 changedhabits irrunediately and preferred sections and depth divisions which were near,or occupiedby, both the newsandbars and the bull. Again it is believedthat this could be attributed to the schoolingtendency of browns,or to establishdominance. Both of the new brownstended to prefer sectionswhich were near Brown 1 andthe bulL Thosesections were alsoclear of anylarge coralheads. They also both preferred the LOW depth division. During both time framesBefore Wall and After Wall, the bull maintainedthe same three sectionsnear the backwall, andcontinued to preferthe LOW depthdivision. Brown 1 alteredher preferredsections during Before Wall, by occupying two preferredsections which were along the outer wall. One of the preferred sectionswas occupied by the bull and the other was occupiedby the newbrown. However,through both time framesBefore Wall and After Wall, Brown 1 maintainedthe preferreddepth LOW. The newbrowns both decreased the numbersof sectionsthey preferred from four to three.It is possiblethat during Transition II, they were more spreadout in order to testdifferent areas, then afterone monththey each settledinto fewer preferredsections. After the installation of the wall, the bull still occupied the samethree sectionsmost of thetime, as weII as the samedepth preference for LOW.Brown 1 alsopreferred the same three sectionsas the bull, andshe also maintained a preferencefor LOW. Brown3 preferred onlytwo sections, both near the outer wall and with the bull and Brown 1. Shealso maintained theLOW preference. Brown 4 preferredallthree of thesections preferred by the others and shealso preferred one additional section near the others but unoccupied bythem. This could Po»bly be attributedto lackof space.And again she also preferred LOW. 389 Diving for Science...1990 Thefindings of this studysupport those of Crow andHewitt 988! in that tiger sharks were found to prefer the upper one-thirdof the water column. Crow and Hewitt also found thatbull sharksoccupied the upper one-third of thewater column. The first year of thisstudy confirmedthose findings, however, upon the reinovalof the tiger shark,the bull preferredthe lower one-third of the water column. Crow and Hewitt also found that sandbar sharks occupiedthe lower two-thirds of thewater column. Again, those findings are confirmed by thisstudy, although more specifically, the sandbarswere observed to prefer the lower one-third of the water colunm. Changesin swinuninghabits due to thestimuli of divers,dolphins and feedings, were recorded,however, statistical analyses have not been performed. The following are notations on the effects of those stimuli. Whendivers non-feeding divers on conventional open-circuit SCUBA! were present in theenvironment, the tiger and the brown were generally unaffected. They seeined unaware of thedivers and it wasnot uncommon to bumpinto the sharksif the diverwas not looking. Thebull shark,however, would veer off suddenlywhen confronted by a diverin hispath. The dolphinsin the aquariumwere extremely playful andwould vocalizeat the sharks and"shadow" them followalong beside, behind or on top!. This occurredrarely with the tigerand the bull. During these few instances the shark would react by increasing it's swimming speed.Most of tbetime the dolphins chose the sandbars for theobject of their'play'. They wouldnip at the shark'sdorsal or caudalfins. Theywould poke the sharksin the sidesand front of headwith their rostrum.Often times this would increase to the point whentbe dolphinswould end up in pinningthe shark to thebottom so that it wasmotionless. Whenever possiblethe sandbars would react by increasing their swimnung speed. The findings from this studydo not coincide with observations byThompson 961! of dolphins driving sharks away fromtheir young. In thissituation, the dolphins had no young to protectand therefore it is believedthat the dolphinswere simply 'playing'. Due to the fact that the sandbarsharks did changetheir swimming habits in thepresence of dolphins, the findings of Gilbert,Irvine and Martini 972! in whichsandbar sharks were not affected by dolphins, are not supported. Thefish feedings in theaquarium seemed to haveno effect on thetiger or thebull. Thebrowns however, were attracted to theareas of fish feeding with varying intensity. Often, they seemedattracted to diverswith herring. On someoccasions the sharkswould retrieve anyherring that other animals dropped. It waspossible for thedivers to wardoff the sharks but the effectswere not longlasting. Thesharks' swimmiiig habits seemed tobe unaffected bythe installation ofthe dividing wall. Althoughthe dolphins could still see the sharks and vocalize at theinfrom the other side of thedivider, this did not seem to have an effect on the sharks swiinming habits ie. they continuedto swim nearthe divider!. Thisstudy confirmed thatcaptive sharks generally move indefinite patterns and occupy specificzones Weihs et al, 1981; Crow and Hewitt, 1988!. All ofthe sharks seemed toprefer 390 Weber: Carcharhinid Sharks at SCOT Center sectionswhich were away from the dolphin holding areas, near the shark feeding station, and closeto theouter wall. Thesesections also tended to bemore dimly lit andclear of coralheads compared to other sections. Documentationofhow various stimuli affect sharks' swimming habits in captivity, may yieldan insightof spaceutilization of captivesharks under a varietyof conditions.With additionalstudies, a betterunderstanding of the behavior of captivesharks may be attained. ACKNOWLEDGEMENTS I wouldlike to thankSherri Seligson forher contributions tothe first year of the study. I wouldalso like to thankA. Wangof theUniversity of CentralFlorida, M. Carroll,staff and managementof The Living Seas Pavilion for theirsupport, as well asM. Xitco andJ. Crow for their comments and criticisms. LITERATURE CITED Castro,J. I. 1983.The sharks of NorthAmerican Waters. Texas AkM UniversityPress, CollegeStation, Texas. Crow,G. L, andJ. D. Hewitt.1988. Longevity records for tiger sharks Gakocerdo cuvier! with noteson behaviourand management.Int. Z00.Yb. 27: 237-240. Gilbert,P. W.,et al.,eds. 1967. Sharks, skates, and rays. Johns Hopkins Press, Baltimore, Maryland. Gilbert,P. W., B. Irvine,and F. H. Martini.1972. Shark porpoise behavioral interactions. MoteMarine Laboratory, 9501 Blind Pass Rd. Sarasota, Florida. 33581. 3 pp. McKibben,J.N., and D. R. Nelson. 1986. Patterns ofmovement and grouping ofgray reef sharks,Carcharhinas amblyrhynchos, at Enewetak,Marshall Islands. Bull. Mar. Sci. 38 !: 89-110. MyrberqA. A.,Jr., and S. H. Gruber.1974. The behavior of thebonnethead shark, Sphyrna ttbum. Copeia. 1974:358-374. Thompson,J.R., and S. Springer. 1961. Sharks, skates, rays, and chimaeras. Publ. Dept., U.S. Fish.Wildl. Serv., Washington, D.C. circular 119, 19 pp. Weihs,D., R. S. Keyes, and D. M. Stalls. 1981. Voluntary swimming speeds oftwo species of largecarcharhinid sharks. Copeia, 1981 !: 219-222. 391 Diving for Science...1990 ah~ rl reading tet.ion ooealae 'l.i ~ his'h dolph heidi pool~ StIIII ooeala> ~. 4a hleph sharkfaediaq station Figurel. Overheadview of 'fbe Uvtng Seas Pavilion. Shaded coruls represent those whichextend into the 'highs depth division seemain text!. Diameter= 60 m. Figure2. Overbeadview of thcaquarium depicting labelled sections used for observations, Walls' ~ betweea sections 4 aud 5 aud aLsobetween sections 2 and 3 above 1hnnel 2! werc iustalkd May L989. Radius A-D! = 21.0 ~43 m. 392 Weber: Carcharhinid Sluvks at SCOT Center IDD II I m I I1oI I DD IG fhEu ~ 2 'II 1 I CD E h E h DEI'TH DETTII ~hhhh2 DD DD TI h CD CD E Il DEDTh Figurc3. Comparative relative depth profiles foethc tiger, bull, Browa t, aud Brown 2 during Cum1, %mssition I, ond Cum 2. PercenttiTnc is graphedssa functionofdepth. DD DD ED T I 1I h DD CQ 2D D E h h h DDTTh DEFT!I Figuvc4.ComparITtfvc relativedepthprofiles forthc bai ond Broun >IdmTh2g Joannes'f pep. Percenttime is graphed asa faadionofdepth 393 Divingfor Science...1999 T00t DIIOvrlI i LDDv 00 l0 I 1 I I 00 0 00 L L ZD 0 0 DEPTH DEPTH 08 00 T I T I 00 0 K E ZD 0 L FigureS. Comparativerebtiwe depth pro6les for thebalt, Brown1, Brown3, aud Brown4, during the time periodsof fiansltiou II, BeforeWail anElAtter Wall. 394 BUBBLE MODEL IMPLICATIONS FOR MULTI-DIVING B. R. Wienke Applied Theoretical PhysicsDivisions Los Alamos National Laboratory LosAlarnos, NEW MEXICO 87545 U.S.A. jvfulti-diving multi-level,repetitive, mLdti-day, deeper spike!, using present models, witnessesa greatershare of problemsthan both bounce and saturation diving. Part of the troublearises from incompatibletreatment of bubblesand gas nuclei Within bubblemode&, ilh areredressed through: I! reducedno-stop time limits,based on varying-permeabilitybubble skins; ! safetystops or shallowswimming ascents! in the 10-20ft zone; ! ascentrates not exceeding60 ftlmin; ! restrictedrepetitive exposures particularly beyond 100 ft, basedon reductionin pernu'ssiblebubble excesses; ! restrictedspike shallow-to-deep!exposures based on excitation of additional micro nuclei; ! restrictedmulti-day activity based on regeneration ofgas nuclei; ! smoothcoalescence of bounce and saturation limit points, consistent with bubble experiments; 8! consistenttreatment of altitudediving; Discussionofthese pointsis the focus, with implications fordiving practice. A bnef descriptionof the reduced gradient bubble model RGBM! is given. INTRODUCTION Validationiscentral to diving, and much testing of non-stop and saturation schedules BoycottetaL, 1908; Buhlmann, 1984;Workman, 1965;Spencer, 1976;Weathersby, 1984; YountetaL, 1976; Kunkle etal., 1983; Thalmann, 1984;Thalmann, 1986;Farm etal., 1986; Langetal., 1989; Lang etaL, 1990; Vann etal., 1989; Walder, 1968; Pilrnanis, 1976;Hills, 1977; «mplemann,1957!has transpired. Inbetween, repetitive, multi-level, deeper-spike, and multi-daydiving cannot claim the same benefits, though some programs Thalmann, 1984; Thalmann,1986!are breaking newground. Application ofthe Haldane algorithm inthe latter caseshaswitnessed higherbends statistics thanin the former one, asreported byVann VarUi «al,1989! inDAN newsletters, anddiscussed atworkshops Latigand Hamilton, 1989;Lang »dEgstrom, 1990!and technical forums. Reasons canbeconjectured, someofwhich directly impactdecompression algorithms. 395 Lh'vingfor Science...19% TheHaldane approach Boycott etaL, 1908! isbased on a dissolvedgasmodel, and h f l th b 1kof tissue gas Ee~ inthe di~lved state, the more corre~md thereforeusefulwillso prove long such ast ean approach. u o issuButas increasing proportions offree phases Yount et~ 1976-Hennessy andHemplem~ 1977 Wie~e. 1989' Yount and Hoffm~ 1986; Yount, 1982;Yount, 1979; Yount etaL, 1979; Wienke, 1990! grow, by direct excitation ofcritical micronucleiorgradual bubble coalescing transitions, theclassical algorithm loses predictive basis.Such conditions might attend diving activity extrapolated outside model and test range~ maybeasa surprise.Thefact that some! divers push Haldane meter algorithms toiijn,t, beyondtested Haldane tables Farm etal., 1986; Lang er aL, 1989; Lang et al., 1990; Vann er aL,1989! underscores theneed for more globally applicable schemes, possibly with greater focuson free phase buildup. In lockstep, procedures such as shorter no-stop time tuni Spencer,1976! safety stops, Lang etaL, 1990; Pilmanis, 1976! and slow ascent rates Lang et al.,1989! are consistent with bubble dynamics. Though not proven, these conservative protocolshopefully lower bends incidence statistics. The problems associated with multi divingmight also be addressed through reduced repetitive gradients or equivalently,tissue tensions.While reduced gradients are very diKcult to codifyin tableframeworks, they are relativelysimple to implementin digitalmeters. BUBBLK DYNAMICS Bubbles,which are unstable,might grow from stable,micron size, gas nuclei which resistcollapse due to elasticskins of surface-activatedmolecules surfactants!, or possibly reductionin surfacetension at tissueinterfaces. If familiesof thesernicronuclei persist, they vary in sizeand surfactantcontent. Large pressures somewhere near 10 atm! are necessary to crushthem. Micronuclei are small enough to passthrough the pulmonaryBters, yet dense enoughnot to float to the surfacesof their environments,with which they are in both hydrostatic pressure! and diffusion gas flow! equilibrium.Compression-decompressiort is thoughtto excitethem into growth.Ordinarily, bubble skinsare permeableto gas,but can becomeirnperrneable when subjected to heftycompressions again 10 atm!, outsidenominal activity.Such a modelof skinbehavior, called the varying-permeability model VPM!,was proposedby Yount 979! andStrauss, Yount et al., 1976!and extendedby Kunkleand Beckman 983! andother co-workers Yount er al., 1986;Yount, 1982;Yount 1979;Yount @ al., 1979!.Rudiments of nucleationmodels can be tracedto early observationsof Waider 968!. By trackingchanges in nuclearradius that arecaused by increasingor decreasing pressure,the VpM hascorrelated quantitative descriptions of bubble-countingexperun carriedout in supersaturatedgel Yount,1982; Yount er aL, 1979!.The modelhas also been usedto tracelevels of incidenceof DCSin animalspecies such as shrimp, salmon rats ~ humans.Microscopic evidence has also been obtained which indicates the spherical gas nuc " thosepersistent microbubbles, actually do exist and possess physical properties consistent ~+ earlierassignments. For example, nuclear radii are on theorder of 1micron or less,and th + numberdensity in bio-mediadecreases exponentially withincreasing radius, characteristi« a systemVPM nuclei in equilibrium with their surroundings atthe same temperature. 396 +enke:Bubb& Model Implications for Multi-Diving A VPMcritica} radius, rp, at fixedpressure, Pp, represents a cutoff for growth upon decompressionto lesser pressure. Nuclei larger than rp will all grow upon decompression. Additionally, following initial compression,hP =P Pp,a smal}erc}ass of micronucleiof criticalradius, r, canbe excitedinto growthwith decompression. Pib}e 1 listscritical radii, r, excitedby sealevel compressions Pp -33fsw!,assuming rp=.8 microns.Deeper decornpres- sionsexcite smaller, more stable, nuclei. Apparent}y the bodyis able to supporta certain numberof safe micronuclei,and a certainexcess for varying periods of time,decreasing with cuinulativeexposure time. Short deep dives excite inany small nuclei, while longer shallow divesexcite fewer larger nuclei. Since tissue deformation and impairment of circulationshould dependupon both the sizeand numberof bubbles,it seemsplausible that the total volume, V,nt, of evolvedgas would serveas an effective criteria in anymodel. For shorterdecompres- sion times, bubble nuclei havelittle tiine to inflate.The permissiblecritical radiusis then sma}}er,and the allowedsupersaturation larger, resulting in manysmall bubbles. Conversely, during long decompressions,bubbles may grow very large, so that only a feware permitted. 'Ihbk l. Kxdtatlou Radii CRITICAL GRADIENTS AND NUCLEI Anyset of non-stoptime linuts can be plugged into model equations, andmaximum tensionsacross allcompartments anddepths assigned asthe M-values. Corresponding critical supersaturationgradients, 6, are obtained bysubtracting offainbient pressures, P.Using a set ofreduced time limits, listed in Table 2,we can construct a conservative setof gradients for purposesofillustration anddiscussion. Thebounce gradient, G,is computed foreach compartment,r, across the tissue spectrum, lsr~720 minutes. Non-stop exposures, with surfaceascent, thus al}ows Gpfor that compartment. BothGp and dG are tabulated inTable 3,with hG taken from Buhlrnann 984! and representing thechange incritical gradient with depth,d. Because thetime limits are conservative, thesupersaturation gradientsare a}so coriservative.Maxitnum tensions occur at threshold depths, did, for no-stop limits, ted. Whatisreflected here isthe body's ability tosupport increased degrees ofsupersatura- tioriwith increased pressure. Bubbleand micronuclei tendtoboth shrink andstabilize under pressure,permitting increased levelsof supersaturation becauseofgreater surface tension pressure.Under decompression smallerbubbles andnuclei also grow more slowly forthe 397 Divingfor Science...1990 samereason. Surface tension pressures, varying inversely asthe spherical rad;us h l l ' thepocket bysqueezing andbuilding up a diffusiongradrent across Q boundary.expe gasin Unless e pocnuclei are stabilized sothat the net surface tension iszero gl n l; eventuallycollapse upon themselves because ofthis squeeze. When nucle; ~e + increasingpressure, experiments established thatthey stabilize atnew smalle growingback to earlier size unless pressure isreduced. I%hie2. Bounce%me Limits. depth time limit depth . 'time limit d ifsw ! I t~ mtnutc s ! d fsw! l~ ttullQh'S! 10 1 tt40. 130 10 4tLA. 140 30 230. 1.50 7. 40 10tt. 160 50 65. 170 5.5 60 40. Ett0 5. 70 30. 190 4.5 tt0 >j 200 90 1 tt. 210 3,5 100 15. 220 3. 13. 230 2.6 L": 11. 240 ZD Ihbte 3. Bounce Gradients. ld depth surface gf adient gradient change tft > 192.4 819 150.0 $19 35 '93,9 $19 5 Althoughh thetheactual size distribution ofgas nuclei in humans isunknown, experiments Yount, er aL 1979! ! in vvivum suggest thata decayingexponential isreasonable. Undersuch circumstances,a largernumber ofsmaller nuclei areexcited intogrowth bydeepe«ecornp sions,and a smallernumber umberoflarger nuclei excited byshallower decomp«ssio~- 398 %ienke:Bubble Model Implications for Multi-Diving SEPARATEDPHASE HYPOTHESIS AND MULTI-DIVING FRACTIONS Therate at which gas inflates in tissuedepends upon both permissible bubble number, ~, andsupersaturation gradient, G. Thephase volume hypothesis requires that the sum of theproduct of the twoover time must always remain less than some limit point,a Vent,with a a proportionalityconstant and V Extending Wienke, 1990!the phasehypothesis to multi-diving,it wasshown that any conservative!set of bouncegradients, G, suchas Table 3, canbe employedfor repetitive diving, provided they are reducedat successiveexposures. Denoting the reducedset, G, we take, G=gG, with 4 aset of multi-divingfractions bounded in relativeterms, that is, 0 sf<1. ! As repetitivetime intervals decrease, appropriate 4 should get smaller and staging approachsaturation linuts. As repetitive time intervals increase,4should getlarger, and staging approachbounce limits. In between, total elapsed time, total surface interval, tissue compart- ment,and profile determine g.Considering interpolating behavior, a checklist ofproperties of 4, correlatingwith divingpractice, is desirable: ! 4 equalsone for a bouncedive, but remains less than one for repetitive dives within some characteristic interval; ! 0 decreasernonotonically with increasing exposure time; ! b irlcreasemonotonically with increasing surface interval time; ! 4 scalefaster tissue compartments the most; ! 4 decreasewith depthof dive segment; ! k scaledeeper-than-previous dives the most; ! 4change withevery dive segment, butonly within anydive segment whena greater depth 8!isthe reached;time mmt t controllingg isrelated tothe regeneration timeto mcronuclei, ir,an the permissiblebubble excess, M. Consistentwith the above set, a multi-divingfraction can be constructed fromthe inverseof nuclei regeneration times,4, theexcitation raradii, ii, r, anand the e characteristiccara time constantfor permissible bubbleool excess intissues issue, M. These f are written in simplest form, 399 Divingfor Science...1990 [1-exp ~ r~r! j exp Am imp!, ! withtgggr interval surfaCe!time, t~p tOtal elapsed!repetitive time and tpmln the SmalleSt ratiO of permissiblebubble excesses on consecutivedives nevergreater than one!. Permissible excessesdepend oa 1 -r/rg3, with r givenin Table1 forvarious pressures. Every factor in Eq. ! isbounded lFzero and one. Corresponding timescales inthe exponentials arenear a minutes!and 2 r a days!.In repetitiveapplication, these 4 possesssome general properties, consistent with the above checklist: