Title Page
 Table of Contents
 Comparison of an Eniwetok coral...
 Plant community stability during...
 Standing crop and community survey...
 Fishery biology studies in Silver...
 Productivity theory
 Hypothesis regarding dependence...


PIlODUCTIVlTYOF FLORIDA SPRlIlGS NR163-106(IlONRS80-02) second AnnualReport to Biology BranchOffice oftlavalResearchProeress from J311uary 1to Dec_31,19S4b;r H.T. Odum andJ.L.YOlDlt ui th sectionsby Dalle Nate1BonDavid K. CaldwolJ.Fredorick H. Berr7r,,U-5llf,012'1\=GJloDepart.onentofBiologyUnive:roityofFloridaGainesville,Florida




INTllODUCTION 1... Preparedby,Howa.."nent ofBiology, Universit;r ofFlorida, Ga10asville (withBiology Bronch, OfficeofNaval Research)PRINCIPLF. INVESTIGATOR'(Thisprojectis administrativel;r listedunderW.C.AlleS, Head oftheBiology lJepsrUnent).Howard T.Odum (Fehl'llar;' 1 September I, 1954) JamesL.Yount(SeptemberI, 1954 Januar;r 31, 1955>Associate, DelleNatelson (September I, 1954 June IS,1955> Assistants,w.C. Sloan(Februar;y 1 June 30, 1954) F.H. Berr,y (June1 SeptemberI, 1954) D.K. Calch1ell (September1, 1954-January 31, 1955> TITLEOF PROJEXlT. PRODUCTIVITYOFFLORIDASPRINGSObjectives.A study of bo.sic factorsthatcontrol productivit;r andoftha effeeta of productivit;r on cOJImUnity structureand density byan analysisof tha unique conditionssupplied by selectedconstant temperaturaspr:l.ngs.ABSTRACT. a.During current reportperiodProduction at differenttimes ofthe yearindicate alinear relationship of 11ehtintensit:rand overellproduction at about 8%of thevisiblelight energy reachingplantlevel. r.t"""urementsofa coral reefat Eniwetok1nd1cate 6%. Furtherevidence ofbreedingat ell seasons butwithaquantitativepulseinthe seasons of max1Jllum lightindicatesthatthe seasonal fluctuationin primary productionisrouted through reproductionrather thanthrouehmajor chenges inpopulationsoTheBucaassionof planteand animals of theautwuchahas been shownwithglassslides andcountsf"rom Sagittariablades.Lossoroxygen bubble3 during the day and emergenceofaquaticinsectsat night have been messursd with funnels. Belljar measurements arereportedfor bacterial metabolismonmud surfaces. pH determined CO2uptake agrees withtitrationdeterminations.A fewrouehestimates of harbi1lDre production havebeenmade from cagedSIllllls,autwuehe succession,and fiahtagg:Ing. Nitrateuptakeatnight byauflo,uehecomunitieshas been confirmed ina c1rcul&t1ng microcosm eJ

2 ABSTRACT (Cont'd.)plants by Dr.Delle Natelsonindicateeessentialstabilityofaquatic plant comIlIIlJl1t1eaafter3yearsand about 10-20%reproducibilityin previous biaIlllSSest1mateeby Davis. Hork on anannualpictureof the fisherycbaracter1stics by Csldwll. Berry. and Odumis halfcallpleted.Thestudyofaquatic insectsin relationship to springgradientsby}1. C. Sloanhasbe...completed as H. S. Thesis. J.Yount hasbegun a studyofaffectof totalproductivityon _ty composition usingsut'wuchsorganisms on glass slidesplacedin different currentandlightoonditiona in Silver Springs. b.SinceStart of Project This contract >las begunJune1, 1.952. Inthe year anda halfpreceeding thepresentreport period,workof avery varied nature outlined the trophic structureandmetabolism of Silver Springswithcomparisons made with otherFlorida springs. The intGnsive study of Silver Springs is nownearlYc0m plete andquantitetivecomparisonsof productivity with other Springswill follow.Host of the teobn1ques andapproaches outl1nsdin the originsl proposalhave nowbeen applied.Thestudy of factors affectingqualitativecolllllUll1tystructure isthe maininCOtlpletephase ' ,,


" 3PLANSFORFUTUREImmediate:1.ByJ.L.YountDeterminetherelationshipofproductivitytothevarietyandthedominanceofspeciescomprisingthevariouscommunitiesof aspring.Ibis istobedoneasfollows:a)byobservationofthenumbersofspeciesperniche1nslnelehabitatswithinthespring,e.g.,Aufwuchs on Sagittaria blades andglass slidesindifferent reeions ofthespringwhereproductivitydiffersbutotherfactorsareconstant.Both hiBh and low trophiclevelswillbestudied.b)determinerelationshipsaeaboveunderexperimentalconditione1naquaria anqother vessels. Pl&lkton andAuf wuchs areexpectedtoserveastheprincipalexperimentalgroups,butother groupe mayalsobeexamined. 2. ByD.K.Caldwell,F.H.Berry, and H.T. OdumduringSpring and Summer.Completeanannualcycleof Fishery characteristics bagUft lastspring as follows;a)tagandrecapturemOrefishb)determinethe sienlfioance ofscaleannuli1n tllver Springfish c)furtherdeterminetheextentofwinterbreedingoffishes1nSilverlsconetanttemperaturewatersd)completefood assay ofdominatefishspeciese)determine growth rateofyougstumpknockersin cages f)determ1nethesignificance of length-frequencygraphsoffishcolleoted1nspringsthroughoutthe year 3. lly D.Natelsonduring Spring 1955ComparethecommunityoompositionofaquaticplantsWithcommunitiesnotinconstanttemoerature springs. 4.ByH.T.Odum during Bummer1955a)Relatetheoverallcommunityproduction of 20springsmeasuredbythedownstream flow methodto current velocitytotesthypothesisthattheoverallprimaryproductivityofcommunities1nsteadystate1safunctionofvelocityof water overplantsurfaces.b)CompletethepictureofmetabolisminSilverSpringsbyadditionaldata onorganic mutterlOBS downstream, effectsofsideboils,herbivore grovlth, repetltionaof measurements,spectrogramofwaterandplantaah.I" Range PlansByJ.L.Yount a) determinerelationshipofproductivitytovariety and speciesdominance1nother habitats,both inlandand marine,contrastinghabitatswithhighproductivity and1m. productiv1ty.


,I.,4b}def1ne tbe relat1onsh1pbetweenproduct1v1tyandcompetltion.2.ByH.T.Odum(DukeUn1vers1ty)a}applymethodofmeasur1ngcommun1tystructureandmetabolismtoothersteadystatesystemssuchasHotSpringsand tropical streams.b)constructmicrocosms1nthelaboratorytocontain amall statecommunities1nordertofurtherde11m1tpr1nciples. REPOF TS IINJ:! PUBLICATI ON SSupportedinpartbythisproject:Odum,H.T.1953.DissolvedphosphorusinFlor1daWaters. Invest1gat1ons#9,Fla.Geol.Survey,TallahasseeFla.Report40pp.Inpress:Odum,H.T.and B. C.Pinkerton Times SpeedRegulator: 'he optimumefficiencyformaximum power outputinphysicalandbiologicalsystemsAmericanScientistOdumH.T.andDav1d K.CaldwellFishrespiration1nthenaturaloxygengradientof an anaerobicspringin Florida. Cope1aCompletedThes1s 5181n, W.C.Someenvironmentalfactorsinfluencingthedistributionofaquaticinsects1ncertainFloridaSpringe.Manuscriptscompletedandsubmittedforpublication:Sloan,W.C. Acomparativeecologicalstudyoftheinsectsof t\'IO Florida Springs.(Submi ttedto Ecology) .Whitford,L.A.TheCommunities of Algaeinthe Springsand Spr1ngStreamsofFlor1da.(Subm1ttedto Ecol0l\ll:). Odum,H.T.andE.P.Odum.Troph1cStructureandProductivityof a Windward ReefatEniwetok,90pp.manuscr1pt. Manuscript inpreparationOdum,H.T.TrophicStructureandProductiVity otSilverSpr1nee Florida.


I5 PROroCTIVlfiOF SILVER SPRmGS Howard T. Od1llll ". Heaauromentfl:mddetailed studies continueon SUverSpringsin aneffort to determin.t.".detailedworkingeof afertile complex nquaticCOllllllunU,.1n .tend,-state. Ao SeasonnlTrml duction rr.,IIonth'-T--;-",, __ /\re.pirationand I downstreamlos8952 JProductionI'Figure 1.Annunl Cours.of Prlmar,Production,,


6 B. DeterminationofthoLightIntensUTReaching the Planta In ordertedetermineofficieneieeofpr1mary production,light intenGitioeroachingthoplant.woredetenoinodaofollowo: (1) Insolation roachingthe grotmd on agivendatafor a given cloudeovar 10determinedfromtab19s in Konned7 (1949.Bull. Amer.Iteteor.SOC.30:206..:21).). (2) IIali'of this istaken""infr...rod and half in thovisiblerange0)With a eulmarinephoteI:lctarcontaininga .... ton photroniccen senoiti... to visiblewawlength8Jthe percent penetration toplantlevelat6 it18determ1nado A wint0rCUI'V'Ql7'a8shO"fll onpage8or thelastprogreslIreparto A.UIlIDer curve i. nmr givan in Figure2bolcnr. (4)The percenttranoI:li.ciC!1for a givont1JDoof the ,..or10interpolatedbetlroanthotaoextrG:i1o curves representedb:rJan\1iU'7andlIal'oThed1tfeNOcebatmeenpercenttransndll1!51C1118ma..1nl;r duetoa d1ftorence:inangle ot incidence 0 (5)Th8 insolation reachingtiltspringcammitT10diIllin1ollodb7._from trecae ocspeclaU7wh.:nthesun i.ata lowrangleoDa. to theorlltat1aDoftlwspring anditstre. potWrDth1IIaUactisgreatestinthe",ming .. raflactedin the a;lc;rmtJtryofthediurnalcur.....Thop-aaaffect10 greatest1n the:'ftintor..Thl!!"fi 1. little efrect on cloudy' overcast days""an thalight 10difruso. To correct.torthe effectof treQ8, at leastduringthemorning, the daiq ineoIation C"J1'VOS abovo aNredrawn80that the morning 1s l)'DIMtriCalw1th the C\l.r'ft4S shann 1n Figure 2I> Theareabet_enthe;t,'Xloriginal cur ...represent.the light reDDvedbytoMtreesotocuch CotTGct.1on is madetorcloudtdI.1aoThocorrection onthe lf1ntorcurw1sgreaterproport1onate1T thanOIl tile8UD:IDGrcur'VOoLightintonsiti doterI:lined in this'ffa7areralated toproduction bolcnr. eurtacecorrectiontortree. "'"$00.Photometermcroampu Time ofFigura 2. Diurnalllarchof Light, IIa7 28, 1954 I' ,


7Co Production asaFunction atLightIntensity .for theWhol. Community" F""ooclldld1;r production figurogiven in Figuro 1,a dai1;r light Intenl!llt;rvaluawaadotorminedasdescribed inparagraphBabov80 Thggraph in Figure3show.totalcCilllllllll1t;r pr:lmar;rprcductirtt;rlUI a functi", oflight inten.it;r of rtsiblo....va lengthroaching plont levol. ThohorizontAlUna is th8entimatliofrespiratory and dOW'lI!1treamlossssfioom previous roports.Apparllnt1;rtho production ieprcporti",ol to tholiill>tintensity evonatth.so rolativo1;r high lightintensitie.. lIost :Ph;rsiologicalexperimentsan single plontsshowdecroasing.fficiencio.withincre ..inglight intcnoit.,'athiill> lightintensities.It "'"'alsobs notodthot thocOlIlIllonityrmlllbelowitsoverallcOOlp!!nsation pointon cloudl'w1nterdqIIoThisdoes not meilDthatindiVidualplzrtsdoth1a forthe communlt1' point includas not onl)'recpiratlonbutdOllllstream 1088&s. POBsibJTanuplmuation ofth. aWiJpir-'6 of haltthe1product1oncbwnstreo liesina requircmontthatstzc.dyatatotheamgleplantasstnevartbsleseneverbe&7.po<;Qdtoccclitions001011th3 individual compensation pointo Asseenbelowthecoralrea!do!:m notdo th1a and e1milarlydoss not have the low 11gbt of wintQroPerhQpstherei8 ageneralizationthat cOl!llfl'JIJ1t1eemust produceexcess organic matteras8011 or, peat or downstroam1068 tropicalconmuniti".ne.dnot d80Fromthedataa_lab)." itis notoloarwhether thore isabreak intho curveatthecompanaatlon pointas discussed inthe thaoreticalDfJct1ODoActuoll;rthobestfit to thopoints indicato. en increasing efficiooo;rwithincreasing light int0nsityo As diSCUBS.d belOW'thismightbeRecountedtOIfbyincrollBedrespira.tion of plantswhich lndi vidua.11 are bo1.o

".B 04SeaeanalPulJoeotEnargylareedinll and Pbotoper1od1l11 The _phs in Figures 1 and)indicate a .transpulaeofanarll7 intheeprinll andllUlIlIllar comparadto tholo..l1lll>tedmnths. Thie uaotua1l7 a greatord1rreronce in tha aquatic coJ!lDUl11 t,. than onland 'blIc&Uaoor the anglo etrectotthe treesand water reflection. One 18accustomed to assooiating big seuonaldit!sroncea in enera nux w1 th succeseionandblOOl:18 among theplanktonicorgwsma in lake5 andinthe oceano It iq'1ntcreat1ngtoCOD:sldl1rtho tat.of theenergeticpula. inSilver Springe whoro no large8tC8ss1onal changes have beanobservad even in too microscopic algae of theautwuCM 0In F1curee4and5 are shll1m amual p1ctureeofbraod1nll in tbeoppleona11 Poi:l2c.nlIhichaye ite eggeabon the water lin. and inPal.....meto8whichcarrieo itsegge. FrClllthoa.grapha it maybeconcl.ucWd thatin tlwsQ formsbreeding occursthroughout theyear inthis conetanttemperature envirorutelltbut at differtlntratesthatarelikely to bephotopar1odicall,y controlled.ThUllthe.. epoc1oe e ....adapted. to theenergetic pulseotthe wbohcommunit,. 0As forbes!on g agodoecrlbEild, thesurvivalof tInJ'particularcOlllDlunit;ycomplexrequires the components to oc.tnalthllr too much or toelittleoAs mentioned in the report anFieheryetud1esbelowthere18S)meevidencethats1m1lar roundthe :rearbrSGding occure1nthe tbheswith a l;rge:OP:;_pULe.. I Fill." Annual ReproductionofPomacea 2000 .. Eill Clumpeper125 IIot ehors 1Fig. 5Annual R.procluct1an ofPal',JDXletesPercent of thetamalellover 21 D:l.with.llgoPQ!ACEA1 F Jl\ lIonth0PAl.. ,fCJ:iETES<)110


,EoInitialAutwucbs !l1crocosm Eld. It hoBnatboon possible toget thoproduction upover tha colIi:lIUlI1.t,.compensation point which 121 vor,rhighbecause of too heterotrophic oslfQllascClDponente ..SilverSprings WaterAufwucbo COllllluni t:ran bladoUgh It1Ill.Ipo.o1hlo.lm;ev....toshowthatnitrato10rap1d17fixedb7 sano components intho o>=1t:r evanintho duIt during 1Illcll;rreopirato,.,.m.taboUsm. This o:xper1mont helpsto confil'a tho darkuptako ofn1tratofoundin blackbell jarsinthef10ld and thoalight deereRes of nltratai'rC!ll.the boil dcmnatream both in.thodayand at nlgh\.. In S replications.037ppm .decreaaed to.23ppmin1 hJ:o and nitratewas not detectablaafter 6bra. (jSUva'uwator .tandingin gIllS. bottle. los .. nitrate ata rate of about02 ppm. per montho Fig. 6. AutwuchsUicroco"",FlcnrS;rstolll I System."""ledboro11""011I0tal 1/8HP CirculatingPUI!IP ,,.


10F sare in etead7fltatethapercentloss atthetipof the clump 1s thepercent gro"Ii'tJI.. Acountof thelastsegments'When proratedoverthe"pringareagives anestiti'.ateorrate of autwuchagrowth and tbU8 ofthe component! of thatypicalbladetJPeorganlems.,Asan illustration orthesernethoda acurve of midges vorws time andthU8aleolengthisshawnin Figure 8.,Estime.tes ofmidge grOltt.hraterom thiscurveare discussedbelotr..SOOlO rough curVGe of wce.ssion on glases.idesareginn in Figure 9.The irregu1oriti in thll last Figura maybe lila tothe poa1tlomofthesUdebons in differentcurrentsOiIl1d depths .. Further work on this isbeingcarriedon byDr.Yount..The successionpattorn!l2r\l! much Ubplanktonic populations with bacteria first,8llIllll algal)next.thltn largeralgal,and finally herbivor ca.rn1voros3.8 somethingof a p3eudo-cl1m.axia attained. Thus one hascontinualf.succc891on inthe micro-environmentsotthe overall state..Similars:':louldbelooked forintropical oceanic plankton .. Fig.7.26 dqgrowth ot marked bladeaof Sag!.ttario AprilS. 19,4 ,,


,.Jadges bladoloogth 1n em Uean age in 10011 Fig"80SuccossionotHorblwrs. on Szg1ttarl&Bla.dss-------..,..---_._--------_..loo,ccovcrag& otCocconeit1P'dlatoDiatomgroups per slidsHerbivore. perslidoCarn!'Oresper elide '1&0 '0 Succes eion on elJ.de../Ul----IIl---@"ArClJlll.-v--/X"",/ ,, --rl.---nf----tjolTOar


, .12 0"Insect&oorgenceandO:cyS'l11)3ubblm 1l3as\lnUlfmtSThaY.rylargeamarg.anceormidges, caddis filesand other aqU3.tlcinsectajustaftersunsethasoosnraferredto in previous reportsoIt hasbaGlnnot!!dtobeintenseboth on fr0Qzingdays in 'fdntsr andhot summerd.aywoFol.1!'.funneldClvicQemtraplaced inthe spr1ns:asshawninFigura 100 SOIOOquantitlltiv. counts obtainod from thssl'1Belt ups asglvaninTabJ.g1.0&:oorgenc8 at dawnis not Signii'iC;llt in compn-ison to thQevsningoutburstsoRoughestimates of growthratea(net)are calculated below fromthis measurementofemM'genaorata. graduated centrifugetube....,' ',., '.'.. ,q", .', ......... '.:'.' &ilerlitni:pup.u'orollnen bubblo. are Figure10.. 'f.. '. ..:,"trapp.d'_.', Emerg,mc.'and I :',',: "'"'', n.-.nbubblo, ,".". '" j' ._ ApparatUli, i i::',11 'cmf'uim.l '. l;l '.. .', ." 'I',' ,," '.,"" '.'.''.' ":',';f';/ \////; '. //AutwuchoTha.....appantuo in Figure 10. 10 od during tho daytimo tocatchthebubblesof oxygantJh1cher2to form1n thaAutwuch.8 and thGnrise to thesurfaClIImthoutcO!Jf)lot8qdis601vingo This1088of oxygen18 a sourceof error in thaproductionmeasurements thAt causea and underestimationofthetooo.lproduction

13 Tablo 2. Bubblestil.Surface inSilvorSprlngs, 1954OatsBubble.Tim. CC/lir.inco. Lop ..in Hr80-NIC!lT:6:20-2:42 a.m. !lay13-14 06.306:00-11:oop.m. '107 26.6 .1.2.1 Mean .2 5.0.0Il 7:00-3:00a.llI. \lay 2J-24 0 0 0.1 mean .0256.0.00)1 IllY.4'45-9:30 a.m. lIay 23.12.7 oS .2 4.7mean.9 .192.20 p.II.-lp05p.II.lIay 6 .42 1.7 .25 9')0..... -).bS p lIay moan May 23.1 .93.1.1 mean HoRerb1wre ProductioniUl.tGo Fromdata1nprevious paragraphs80m rough estimates can nowbemad .. to thoproductionrate. of thedominantherbivor such Il8 Ar.11.tsdin previousprogressreportsoSeveralmethod. are ueGdas 1. FroIIl tho timate. of midges at tho tip.ot Sagittor1a blads.inFigure6 and tromtho averag. porcentSagittariagrowth ot trOll Figure1 CIllo compute.tho rats of midgegImlthnece.sar;y tokeepup wi.til tip1ina stoadT .tato.Thi.is anundsr timation a. it doe. ,not includeemergence and lossee to predation in the middlesoctionsot thllgrassblade:!.


1420A tn cagesot PomaC32and '11v1parueSImlaDrsmaintainedlIithC1abundanceor food anCivolii'Ill! measurementsbefore and after a month growthperiod.ThsSf3:est.1Jnts8arow"ldereetimates since 10basnailsU!IDdwere alreadit of moderateBizetand past t!la morerapid juvan11.growthstage 30Fromthe 88CtiCl1 G abov;) on insectemergwnce the growth rate nBCesB&r1' tobalance theI!lQ8rgence in steadystate1'1'&8determinedusing.(D02gD'l13dryper..erging lndividual.Thi. figuro .hould be add.dto tn. Tip-lo figur. in method111abow.40Promtheostl'i'ateeor standingcrop biomaSllIofsmallinvertGbre.toherbivores in prevlouereports on. canobtainthe totalrespiration using orough figuro for .tr.einvertebrate. of.8 cc/wo/lrr. Thanif growth ofanimals1sabout 10%one maygeta rough figure for herbivore'roduction" Withtheso mathodsScm! herbivoreproduction' estimatesare given jnTable)",None ofthe 8"tirra tes areentire17 satisfactoryalthough theorder ofmagnitudei. indicated. Tabla 3. lI.thod Blade Tip10mothod(#l) &>o1la in Cagos m.thod (#2)SomeEstimat of Herb!vore Production Il.asured Quantiti.s plant 25.5plant .pring .26IIlant growtb/ll".pr1ni:!dq(1$bladegroWtb/dq)6%volume incro ..o/llonthViviparuo 1. volumeincreue/Month Vi viparul!I27%increase/montbPomace.lIoon'1l.6%/monthl 12 gmflil bioma.s Prod!!ct1ongm./Ji'/tr 219,P 17Inctem.rgonc.m.thod (#3)/P'; and.8cc/gm!hr(m.thod 64) 10 Carnivore Production 7.8individual./229 cm2/daygm./individual Total ofInsect" and fna:'1s 3204 gm./112 herbivorestandingcrop 196::;.:,2.'(03 In thofiohe17biology.tudT d cribedbelow a f

15 J. ot lbtGbo11sm.. Direct lleu;\Jr8mentotBacteriallletabollemEven thoughproduction rates otalltrophic levelsare not yet.satisfactorilydeterm1ntld. it isinstructive tocalculate metabolic rates b7 trophic leveleusing somo literaturevalues or metabolicrat. multiplied bY' estimates orstanding crop. Primary" production figures are takenfromprevious reporte. Th . m$jtaboll_ oftheherbivores1. taken as08 cc/iJDIhrwith a of32.4 f!jJI.s/1l'.Carnivore.aretok.n a 01 cc/wn/hrwithp./11biana... Top carnivor are t1=tedfrom.04cc/v;nlhrwith 4.0 VJMlfJrbiomae 1loU&b metabolism figure" come tremPr\D;Zoo Dir.ct Urnates or thebacterl.almotBboliem in th.algalgyttj&th.tmakes upthe COT1IJlunltybottcm'W(!re made withsmallbell jar.placedon themud aurface af'ter tho top zono lIhlchcontained algae waeekim..dotto A. withlarger belljar oxp>rimntsdescribed in jr.viouereport oxygenana.1yB war.madbefortJ endaftiU"experin:entalperlocb\meierblack cloth.Tbr.. replication. ga ..oxygendoere""e. in 95minute. or llr, .75. and1.610mg/l.with boll jar. 16 em in diameter2Ild a capacit" or1600cc.Thisturnsoutto be11S't'!.Thisestimate do.anot includetheconsiderablebacterial.nora of thw au.twucba whichwasmeasured 1n previously reported work. ItiB il:Jpartant tonote thatalthough much emallor in bi-.a the decomposerbacteriaa.re1:.01'"0il1i.portantmetabolica.l.qthan the regulArM!"blvorea. The variouarouiPe8t1m.....ormetabolismb;r trophic level are su:ena.r1zed I.in Fif,)ll'O II below:Thi. is thus afirststtempttoas.ignvalue.to themetabolism diagrampreviouslyprestmted......p1280 .H26 C, I


.,16I. Diurnal pH Curvob;rIloasuremont and D1e IIolllOgram rrClQCarbon D1oxl.doduplicate bottles4 IIII1II I::-,:I,-I,::-!I:-+'--,;-'-11,->-1..,JI,::-!I-<:I5 6789101112123456789101121234 S 6 m. noon p.m. midnight5 3 InFigura 12g oolo'n' 1s given another dqsdaterm1nat1onor production withox;ygenandc:aoon-dioxidecw-vea auch ashave beenginninp1"IIvioua progrereports. 'lb1otime.... tor SOllljlU8_rebroughtbade to tholaborat0!'7 III1dthepH determinedwithaBockman modela.The max1lIlwatimeot24 hours1.not110 eer10usinthe38100organic matteratera&II'ROUldbe tho casowithmost natural waters.Thecarbon dioxide value.war.thmconvertad into calculated pH vollieS tramtil.knownalkal1n1t7.Themeasured and c:urTecalculatedwithtileIqonomogram.... both shownbel"". 6I JlPlR at 43/4m1Ustetian 11 109876 Figura 12. DiurnalCurves tor 3/4m1U.tetienIIo;r23-24,1954


, 17 CCUPARISQI OFAN EllIWETO;(CORUREEFCOllllUNITY1'!l'nI SILVERSPRINGSI ,. In thoIlIllllIIlOI'of1954 H.T. Odllll andE.P. Odum(lhiv.ofGeorgia)""doa .tue!)"of thoprodueti Vityof" "llindmlrd roofon "'ilootok Atoll primarll;runder .pons.rahipot theAto:n:1c&lergyCom1ss1oDandthe Universit,. ofGeorgiab7means of aContr3CtextsnsiOD ofaproject directedby E.P. Odum.This endoavor .....aloo.upporwd inpart bTthoNa"7andindiroctl;rbTthotechnique.used in theSilv$!' Springe wal"k. On.main purpoSCi1'r.lSacoupa.rlson ofthe characteristics ofSilverSpring. andthoJaptanintar-1elond ro.f.Th.80pagareporton tho Eni"atok .tud)"hllSjustboon co:nploted and .ubllittad tothe AEC priorto.pUbUcation.A few ofthoconclusions which are algni'tlcant tothounderstandingot theSilvSI' Springe communit,' and stateaystelZl8in gen.ral mqho li.tedasfolio ... 1" Both commwtitiesarevery efficient,the coralreefbeing6%0 (of Vi.ible light ."orureachingaveragecOlllllUl1it,. depth.2.Bothare Imsnee17 fertile.Thecoral reatwithproductionratosabon7S,OOO Ibeo glucose p3r c..crl3peryear 1. amngthehigheet. on earth.Silver Springe18 JD:II"8productive in sumner month" but haa a lO'fflir annuAlpl"oductiono I 30 Both l!I;yatcms are aaasntiaUyeutotrophic 1n pr1m1U"1' trophic 1&ft10 (JohnTeal, HarvardUnivo, is rorl.;:L'Olg on productlvit,. ota smallB})rlng'Whoel)primar;r production aourco 1snotlight butallocthonoU8organicmatter.) sargent aUldAuatin'op1on13erstudies on roefproc!uct1v1t,..ere in generaleont1rmedo01l.S8olidz9dovm3 inbothSilver Springs and &n1lft1tok71eld coatingsof algQIJratherthanfoulingcomrnmltlea toundin inehoru oceania waterso4"Thep)Tamids ot standing cropor JJniwetokand SilverSprings ru-esurprisinglysimilar euggesting th2t in similarcurrentsituation a similar ratiootbig !llow produc$lra and 11 ttlefast producers maybenGcesGar,r to maintainoptimuml5uctureoNota thocomparative p;yramids inFigurQ13 Ol1d1n1lI'.v1oup IIro......Algae in calooraau ub.trata. \la timated 111 tila l"xtl'ac ... 1tmmthod alj:oa1nand donona of5 crabe. cma.,annelida at. oflarge h.ade)..'. her invert.brat ...,.::'fporif.c.a'it:::'0-:&.Lga&:..lf1:alp.1n,.: .::. "alga.:l.D .: 'algaeill' .,':. r'j.lin'..: : ..':'.caral.hGadfl :.'..: e!Wlll1e: 0-.' -. .. 5;ofwromida' 11 L'iJ2g,;.i.+-.harbivorea.. .. j.... : ..... Producera70). ... ....110'"P1rami.C1uedr (zoneat Figure1)"Pyrmn1daof live biomaa(noncalcareoue) 2on "'i ... tok--Japtaninter-i.landreaf;Figura.1ndr7_/II., .


18II. 50 Both comtnmd.ties s.rc cloSG to :Postaady'statowith production ballmclIdbyrespiration in thQcoral!'l')ctfwithinthel1m1tsotaccuraq ofthCl madoo In SilvorSpringstooreisal'wolYSSI. contributionof thelJpatracmatt6r to thedOiiiWtr2Qmcommuni.tieswhichthereufdoes not havwo (exceptto a vlJrysmallaxt0IlJG)Curve3showing production andrespirationar5givoninFiguro14.".longlrlth protous dIlta of Sargont andAu.tiniH.T.&E.P.Odum 19,4Hong.lapdataofSargent.&AU8tin..,.,"-r-+-"'...-'rr----------l.--1 3 gDlJ!I porII/I!r1o idnight 6a.m.n on Flgt1l"s:l.hoDiurnalCurv. atProductionMeasured withthoUpstrtl'lU!r'oo\'ffistream.flot... mothodon EhiwotokWindwardReelf I.60Both eystl11!lB3t'&lknown to hlOl.v\1 beenat leastroughly in stellcO' statUIbalance forlong po:-:1.odB of time andit1simportantthat contrastingsystemstend to 2.chi;vot.hooametrophicstructuroandcharacteristics inI. steaq, state 8uggGstingbasic!aweofbehavioraediscussed in tho theoreticalsection 70It 18further3"'.u-pris1ng that similar high lIfficlsncleoBra achiovod b,.Qvery high natri.ntnowinSilverondbT a vorylownutritlllt conditionin thecoralThusifthl!lprop8r adapted organlslll8are available thap2I't1cu1.llrdeficienciesofpossibleenviroI1ll1l)ntal limitingfactors ma.ybQcirCUl!lVontGdbyaconmunit,.byexpendingsoma partofits .nelU in mecllimis::ll3torcOBorvingthQlimiting factorsothatit iv no longer limiting oxcaptJGhntitmaintainfJasmallIInergetic taxontho overallsystGmo In thE c,)r:11reo!caBO this is theconservation of nutriElntsinthe calcarGou81sk&lotonso ,.1,


I.I 1,.19 PLAN'!'CCNuUNI'l'YLUR1NG 3 byH. 1'. OdumIn1951 maps "Ieremadeofthedistributionofplantsin 40sprlllfs. Oneof these,Farullng Spring.! 'WaB 1nthe first report.In the reportbyDr.Netelson,entirelyindependentmaps are reported, 3years later in6of thesameDprln[s without hav1nc seenbutoneofthe peevlousmapa. comparisonofthe earlierfTIo.powith thelater maps 18veryindicativeofthe debree ofstabilityin these chemoBtaticallyandthermostaticallyresulatednaturalcommunities.It1sapparent that in seneralserne speciesdominate after threeyears althoueh theexactpooltloDS or thevariouspatchesshifted.Itshouldberemembered :froID the work in511verthatthe higher plantshave atHGnover of one and 10timesayear60 therehasbeenampletimeformarkedchanges. ibis isfurtherevidence alongwith thepreviouslyreportedwork on a1eaeand insects thotcODaltionsare much morestablethan usuallyfoundinnature.This1snottoimplythatpulsesand chanees havenotoccurred1n 800e sprin5s; in somespringe moreaffectedbysur racewater,there 10 aleoa known variation orchem1calcondltlonso


,.I.1-II, 20StandlnE Crop Lind Community of 5ubmereed Vegetation Ibn Seven SpringeBy DelleNatelsonInthefirst report of thisproject1n January, 1953, Dr. JOhnH.Davis preeented onthe 8tandlIlt!crope otfour B9r1DEBand coastalrune and calledattentionto chanree indenaltyandcompositionoftheVSEetatlon, Bome ofwhich werecortelatedchanges 1nturbidityandchlorlnltyofthewater.fhls1e areporton workintendedtocontinueandextendtheabove 9tud1e:;.'l'he presentinvestigation, Hhlchbegan 1n 5eptemeer, 1954,hasforitssubjectboththe featureeofthe otandlnr cropof Bubmercedaquatic plants1nsome spr1nes andtheirrunsandthe tiualltatlve compositionoftheirprevalentcommunitlesoAn investigation ofthesubmersed veeetation inWisconsinlakesrevealedno discreterecurring communltieso Instead, there a patternofcontinual ofcommunitycompoeition along a gradientcOID91ex ofenVironmentalfactors Do, ThephytosoctoloEYor Gubmerged aquaticmacr09hytes1nrJ1ecooslnlakes.Ph.D. Thesis, Universityof Wisconsin,1954). Oneoftheprincipalaimsoftheoresentstudyistodetermineifsuch a situation exists intheaquatic of Florida, andif80, ",hqt 1sthepatternofvecetatlonhere. Aknowledge ofthepattern,referredtoas"vegetationalcontinuum",canbeusedinconstructing a classification system forthe commUnities andfacilitatescorrelationsamong veF.etation andenVironmentalfactors. rhe FloridastUdieshere reportedare baseduponmethodsusedintheWisconsininvestigation.Eachrun '\'la'Ssampledatseveral statioIBscatteredalongthelengthofthe river.Eome subjectiVitywas used inasmuch ascarewas takento sample characteristic,ratherthanatypicalordisturbedareas.butselectionof stations was objective,with two exceptions:1)insome instances,for comparisonpurposes,anattempt madetosample nt the Snme stat10nusedbyDr.DavisorMr. Sloan(bloan,'tIm., 'l'he dtstr1butionof a\luatlc insects1ntwo SprinGS,M.S.thesis, Univ.ofFla., 1954); 2)stations at Salt bprln,::.s runwerelocated rec:ularly at tllOmileintervalssincethe waterwas tooturbidfor rellabl selection.The samplingtransects madeacrosstheriverfrequently traverpedtwo ormoreobviouslydifferentcommunitiesandsuch stations weredividedintosubstations.Eachofthelatter were sampledindividually,sothatthedatacouldbeusedfor general communityanalysisas asfor standing cropand species compositionestimates fot' eachriver. For the latter purpose,the sev6ral subotatjons at each were weightedaccordins totheareaofthestation whichthey occup1ed,andthey weve thencombined. Thisteechlquewas alsousedforooois where amosaic ot communitiesoccurredo


II21SubstationsorstationswithhomogeneousvegetationwillbereferredtoBSstands.Eachstand waS sampledby5to25quadrats, according tothehomogeneityofthevegetation and/or thearea of' thestand.The quq,drat was a heavy wire frame onesquarefoot1narea.Withtheaidofaface..mask,therootedplantewithinthequadrate were uprootedandbroughttotheBUrface.Butsometimes, where thedepth and BubstrateweresUitable.arakewasusedtodenudeonesquarefootofthebottom(.etlmated),instead ot uprootingbyhando To determinethevolumeoftheplants,thedisplacementofwaterbythe plente removed fromthequadratswaSmeasuredusingthe method described by Dr.Davis(citedabove),andanaveragevolumepersquarefootwaacalculatedforeachstand. Since thespecificgravityofsubmergedplants1solosetoone,these were usedasanestimate of thewetweightoftheplants.Thepercentagecoverof the vegetationineachsectionofriverwasmultipliedbytheweightofthe thestandsampled1nthatsection,andtheresultswere toIbs./acre wetweight foreachsection.Wet weight ofthevegetationineachsectionwasweightedbytheestimatedrelativeareaofthesectionintheriverandtheresultingfigures were combinedtoproduceanestimateoftheaverage wet weight/acreintheriver. percent ale ofvolumecontributedbyeachspecieswasestimatedforeach quadrat, and an average was obtained eachstand. These flgureswerethenweightedbythe percentage coverand percentnee areaofeachstandintheriver,inthesame manneras describedaboveforthecombinedwetweights.Thusthepercentageofthe wetweight ofplantsintherivercontributed by eaohspecies was obtained.Thepercentageof water contentofeachspecieeaBpreviouslydeterminedbyDr.Davis\unpublisheddata) was ueedtoobtainthe dry weightcontributedbyeachspeciestothe averaeedry weight/acrefortheriver.Estimatesoftotalstanding orops, bothas wet anddryweight,andthepercentaeeofthetotaldry weight Which wascontributedbyeachspecies are presentedinTable1.Figures1and2illustratethespring-riversystems whose cropsweresummarizedin Table 1,andthe locationsor stationsandtheareaestimatedasrepresentativeofeachstationareshown. Acrosp a rlvar, fromshoreto ehore,. differentcommunitiesoftenoccurwithinsmallarees,evenatthesamedepth.Frequently,anenvironmentalcorrelationisobvious,e.g., substrates. However, inothersltuetionsnoreason for thedifferences1sapparentanditis thathistoricalfactorseuchasdisturbance,availabilityofpropagules,andconditions conduc.lve tocloneformation ,lere largelyreponslbleforthenon-uniformityofthevegetation., ,,


II.II r I ,22Incontrast, a regular'trend1nvegetation change downstreamoccursinsome rivers, superimposeduponthemore randomlocalizedvariability.Inthisinvestigation,suchtrendswerenotedinthespringewith conatal runs.InWeeklwacheeRiver,Charaoccurrednearthe head, ofteningreatdensity,butwasnotfound1nthemiddleorlowerparteoftheriver.Ceratophyllumdemersuml1kewiseoccurredingrea.testabundancenearthehead,butextendedmuchfurtherdownstreamthan lilhar\l. As Chara decreasedinimportance,Na.1asmadaluDens18, 'tlhlch wasabsent:fromtheupperpart of theriver,appearedandrapidlybecamethemostabundantspecies. Sag1ttar1a was moreprevalentintheupperpartwhilePotamogetonpeot1natus and Vall1sner1aneotrop1cal1swereapparentlyrestrictedtothelowerpart of theriver.Chassahow1tzkaRiverwassimilarto We.k1wachee Riverinsome of 1tevegetationalfeaturee. Sag1ttar1a was mostabundantheartheheadwhileVall1sner1aneotrop1cal1s, NaJasguadalupens1e, andPotamogetonpect1natusreachedtheirmax1mainthelowerpart of theriver.SuchtrendswerenotapparentInHomoeassaRiver,'exceptfortheoccurrence oflarge amounts of filamentousgreen algae 1nthemiddlesection of theriveralthoughitwas rare intheupper and lowerregions. However, adistinct change inthecharacterofthevegetationoccurredneartheGulfwheretidalwatersintroducesalinity. lhere Potamogetonoect1natusand algaeor marinetypeoccurred, While thecommonupstreamspeoieswererare.Thesethreer1vererunfromtheirhead springe totheGulfandthus a gradation 1nchlorln1ty,aewas shownbydatapresentedbyDr.H. T.Oduminthe January, 1953report of thisproJect. 2to5ehow,thequalitative changes invegetation Wh1ch occur1n someinstancesfromtheheadotarivertoitsmouth, and alsoshow how equallygreatoreven greater variationoftenoocurs among substations at the Sl.me generallocation.. Thus 1tseemsthatexcludingbrackishwaters, changes inchlor1n1tyare probably nota8muchthecause of community differences aearethe ohanges insubstrateandturbidity.ChangesincommunitycompositionintheSalt Springs run.whichdOBenotflo"',int0salt ''later butinta Lake G eore;sJ showednoconsistenttrends,exceptforthefactthatPotamogetonpectlnatus,wasthemostabundantspeciesinthepoolarea and the beginning run,and was absentorrareelsewhere.Hart Springs runhadessentiallythesameplantcomposition throughout itsshortlength.Sources ofError inthisworkfallintotwoprincipalcategories. The firstresultefromthemosaic arrangementof thecommunities Whichisrevealed by Tables2to5.Beoause of this,theerrorcanbeconsiderablewhenthestandingcropor


23 speciescompositionestimateforariverisderivedfrom afairlyamallnumberor otatlonE.'he secondsourceoferrorarisestromthenecessity for estimatingtheplantcoverofeachsectionoftheriverandtheextentofthesectionrepretentedbyeachstandInaddition,therearesmallerrorswhichresultfromdifficulties1nsampling,1.e.,thecurrenteffectswhichoftenbendthevegetationhorizontallyanddonotpermitthequadrattobedroppedoverthetopoftheplantssothatonesquarefoot ot bottomcan be denuded. However,theseerrorsmayottencompensateforBIOhother.For example, theexcessweightcontributed by soilpart1c_les,whichueuallycomeup with theroots.istosomeextentcompensatedfor by thefactthatfrequently alarge partoftherootsystemisleftinthesubstrate.Anideaoftherelativelysmallsizeoferrororvariability1nresultsobtained1s byaoomparisonbetweenDr.DaVie'results(citedabove)andthoseofthisinvestigation.Lbs./acreLbs./acreDryWt ....!!qWtoU Error 39111 686 16% 4620 3667 26% 4000 3774 Average error-From Dr.Davls'data.#Fromthepresent work. convertedfromwetweights by oonversionfactors.Weekiwachee Springs andRiverChassahowitzka Sprin[s andRiverHomosassa Springs andRiver 1his comparisonofresultsofwork doneentirelyindependently bytwo personsleadstotheconoluslonthatstandingcropestimates are relativelygoodfirstapproximationsoftheproductiVityofspring-riverecosystems.However,itshouldbeemphasizedthatithasnotbeenpossibletodeterminefromthisstUdyWhatpartofthedifferencesbetweenthetwoeltimates fQi thesameriverisattributabletochanceerrorsinsamplingand Whichtopossibleactualdifferencesinthevegetation When theestimates were made. the poolsofsome bf the spring boils were mappedand areshown a8 diagramsinFigures 3 to7. ine areasofsimilarvegetation(stands)ineachweredelineatedandthecommunitiesanalyZedina mannersimilartothe oftheruns.Eachspecies1sI'epreeentedbya symbol ,.,hose freQuenoyineachstandonthemapindicatesitsrelativeimportanoeinthecommunityasdeterminedbyfrequencyoalculationsandvolumemeasurementsorestimates.I.,,


,. ..24 maps 111uct{uteieveral differentcommunitiesortenoccurwithin aWork oncommunitieswillcon Unue throuehthe or 1955andprobably w111 shed 11ght uponsomeofthecommunity mlu siterelationshipsNotes:1) Thespeciesmentionedinthisreportarelistedbelow with theirtaxonomicauthors:Ceratoohyllum L. LUdw151aoatan. Ell. NaJuguadaluneneij!(Spreng)!-lorong Potamogeton Morang P.pectlnatuBL.Rorlppa (Nutt.) A. HltoheVaillsnerlaneotroDicalisMarie-Viet.2) Thelocetlonadeacrlptlone orthespringsandtheirriversdlscussedhereare Giver. inFerguson,G.E C.W. s.K.Love,andR. O. Springs ofFlorida, BulletinNo.31, State of Department 01' Conservation, Tallahassee, 1947.


-------------------------,25Table1. Estimate: cropand percentase contri te.J"peelcs in5 springs andtheirrunts. Weeki. Cbassa.Homo.Salt HQrtFanning DateWetwt.1be./aereDrywt.1bs./aere11/8 35272 468610/2327495 343710/537926 377lf 10/12013710/31797710/76684 802 Percentage of the whichwascontributed by eechspecies:Algae(fIlamentous).820.1Anacharls .1ICabombaep.Ceratophyllum demeN':UI!l Charaspp.Hydroeoty1espp. Ludwls1a no.tanoNaJas BUeda1upenois Potamo8eton P.peet1nat\ls9.019.36.535.22.913.640.94.816.510. aess1flora Others3.3Valllsnerlnneotroplcalio1.811.06.1.629.1 Sas1ttarla spp.


Table2. HOiUosaw:aRtvo2'r:vegetation uttheComposItIonand standing cropofsubstationcommunity 26II. Importance*itSpeciel!! ABlB2B3CDlD2EF0 Algae ( file.ment0md 1320 64 6 32#35/1 Ceratophyllum demersl),!11 39183 29 3 33 19 3Najaa 56 3218283064 4840Potamogeton pectlno.tuB 822 142367 65 S"gittaria sp.12Val11enerla neotropicalis 37 3 42 3 21 331918Others2Ibs./acre wet\,le:1.ght 20'( 58558510717 604 604 elf 95 20*CommunitIes represent.ed bythe Bame letterand different SUbscriptsaresubstationsB.t tbe samelocatlou.*"Importance io by %F. x 2% V/3. where %F fre-quencyand %V = 1"Clat i \TEl volumeaemeasuredby dlsplv.ce'G:'G'nt of "rater.# Algaeof mlll'1netype 1Table3. \.ihe.sw'hmd Compositionand standingarop of vegetBt.!.oll thesubstationcommunitylevelImportanceSpeciesA BClC2DlD2ElE2F01G2 3 20710 1 10 35 2 '0 8 3 21 N. guadalupeilsis 1615 1 102644 52 82P.llUnoensI86 15 811P.pectlnatuB37 22 68 29 48RorlppaeeBslflora1 Sagittaria sp. 7" 100 804176 59 54 9 .. v.neotrop1calle '( 13 10 4 18lbs./acrewet "Jelc:ht 509 160 150 210 340 330 204 120 320 952 150I;


27 Table 4. Weeklwachecn1vcr: Compositionand standing crop or vegetation at substat10ncommun1tylevel.ImportanceSpecies A!! 0 !r!!-Alsae 7. Ceratophy11umdemcreum49.5 Chara sp.2449Ludw1g1a natans 9 baJas guedalupeno1.10059lOOt44PotamosetonpectlnatuB4225 Sag1ttar1a sp.2037533Vall1sner1aneotrop1cal1.372631Lbs./acrowet we1eht 6753201804401801180 Table 5.SaltR1ver:CO!lIpos1t1on and stand1ngcrop ot vegetationatthesubstationcommunitylevel.ImportanceSpec1es !1 A2 BC0102 --Algae4114 16 8Ceratophy11umdemeraum1821 41 2 4Charaap.2Najaeguadalupensls246023 497951821941Potamogetonpectlnatus 56 1149 2Valllenerlaneotroplcnl1s15182416 147659Lb/acrewet we1ght 200 200300828193 88323040 "


3 A.3.7HOMOSASSASPRINGSFigure 1. Locationsofstationsandsectionofriver represented byeach station. 1inch =.1 milstationI CHASSAHOWITZKASPRINGS GulfG=25jt; F =35 o A,B,C .5WEEKIWACHEE SPRINGS F .. Gulf AAlA2A3BB1cD E Fofriver8. "",Il, Substationsin WeeKiwaohe& Riverareestimationstorsectionsoftheriverwhichwereobviouslynotrepresented by thesampledstations.


Ii ... I ""il.Pli"lto"""''''"it,tfC .... dt".. t ...IAT,..u",,.'c,disoutIi., ...j",.... .-. .' z/?%sefeeT ..8.il It sVIta"SCI.ttCl'cd"".'tsO"nIo.tt.""Lo,.fi.,u of s"'''''l'li''JshTioltssTatio....' aTe;'"'l"'c.l..."'111<.-of,.j...,.


\ 0 '----:::0:--::-, 000o000o ooI, Y1ih.." ott.. 'r .. D',I .."UlfOTCl"rIec>tl"llriclli.s ... d.",n$w""9=AI&.."11.r. .. t.;'.','.&f. 0 0 0 00 0Qo ...s11'fiftJ'lofN",,"'n sp-, Sff'}Cent"rh,""".dc.,u\nl\.'rf'.CO'r.l,t._ .. ., ..


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.."J CrabCreekfoo I ."" ..,50 p/1,I5 F.. 'l'lt,.. I,' 2a"ic"C'Jli"p1/v.tr;$HovseNaj.sgjvadaJvp'ns;s spV ..lli'l"ne"ia.l1eoTroriGcllisre(tiY)...atChassahoklitzka SpT; ....dsOolobOT1'/'./ts't


R,verSI3iTTa:"'r'N"JuI,I"'rl:P1.aI' F luti'n.5"'?tt.tOf....d'wIth."JFeet so "'" Of Ch .... h.wild.A"uc.haTi..cOlT'lade".is -.'. .".40,..<,,..,,",,,,,


So Fut NSalt e. 7 F..Iand5epte."'ber .30,Upper I 5'tRun.:: PoV.llis"nui,N;j:a. rl.. 'a.t,

35FISHERYBIOLOGYSTUDIESINSILVERSPFINGSByDavidK. Celd9l'ell, H.Berry,and HowardT. Odum. A fish taggingprogrc:m inSilver Springs':&8 begunon March12, 1954,andhasbeencontinued byvurious,,'Orkexs todate.A total of19trips hLve been made\;ith the of8numberofvolunteerhelpers.Todate, 392 fish he-va been tagged inSilverSprings. T9ro typesoftagshavebeenused. PlasticPeUrson disc tegs ofvaryingcolors and sizes hbve beenusedonthe lcrger fish, and smallmetal clamp tags beenplacedonthe jaw oropercleofSMaller specinens. TheIbtter Ere numbered, the unnunberedplGstic havebeenusedin vLrious color combinations foreachfish,or beenusedcuttodifferentshapes with the sume color combinbtions. Mostofthefiah tugged hbve been Largemouthblackbass(Micropterus salmoides) and Stunlpknockers (Lepomis.E. punctatusl, with someotherCentrarchidsand otherspecies.The numbers ofeachspecies tsggsd andthetypeof tsgfore su..mmarized as Table 1. AsSUJnmiSrized inTable2,tenfishhave beenrecaptured. enough timeelapsed tagging and recaptureforsome growth tooccur, some fish h&ve not showany increase, and others show anapparentdecrease. Whether these'growthvalues Ere typicalor whether cnartifactor injury isinvolvedisnotyet cleer. The buss showthemost growth, and all ofthesespecimens are juveniles.Itisinterestingtonote that ellrecapturedfish wereretaken wherethey were initiallycaughtand thgged. Regular samples of 5tumpknockers (the domin&nt species) hove beentakeneachmonthbeginninginJune,andinMarch. Length frequencycurvesandlength-weightcurveshave beenconstructed with these.The length-weightretia doesnotvarymateriallyfrom monthtomonth, at leastforthesi2esmecsured. Atypical wetght lengthcurveisshownasFigure1.Enoughspecimensofotherspecies hbve notbeen taken fortheconstructionofsuchcurves.Monthlylength-freqnency gr<.iphs foI'theStumpknockel'donot show distinct age groups(Figure 2).A verylongbreedingperiod1sthusindicated.Thereis also evidencetosupportthisfrom observ(;tions of spawning beds ripe(ornearlyso)adultsduringmostofthewarmmonths. Though an entirewinterperiodhasnotyetbeen sampled, evidenceindicates thatspawning,althoughrare duringthisperiodinSilverSprings,doesoccur.Individuals vlith developed gonads r.ereteken onOctober 15 onDecember 15.Also,smallspecil!lens werewken duringthewinter(Figure 2).Samples ofscaleshave been tvken throughoutthestudyandaneffortis being mhde todetermineifthese can be used indetermininguge undrate ofgrowth. Thisstudyisprimarilybeing done ontheStunlpknocker, thoughsomeattempt bemadetostudythescalesoftheotherCentrerchids,parUcul&rlythebess.Preliminarystudies show thepresenceofrings, hutfurther studyis


, I,36necessvrytodetermineiftheseringsrepresenttrueannuli.Scales from thisconstant spring be compared withscalesfromthe some speciesfromother(nonconstant temperuture) Floridawaters,Bndifpossiblewithscalesfromnorthernwaters.Astraightlineratiohas beenshowntoexist betweenstandard length &nd totallengthfortheStumpknocker. This (S.L./T.L.-.79)existsthroughouttheentiresize forthisspecies as encounteredinSilverSprings, andwill behelpfulincomparingthe work doneinSilverSprings with thatofother workers on theseme speciesinotherareas.Coincident with thefishery work on lsrger fishes,ageneralcollectionofsmallfishes, algaehLS beenmadefor further studyof seasoml periodicityofreproductioninthis constant temperatureenv1r'onment.


37 NUMBERYiITH NUMBERWITHSPECIESlIETALTAGPLASTIC ThG TOTAL Lapamie R. punctatu583 147 230 I' Lapamismacrochirus 121224 Lapamie Burltus 3 0 3 Lapamie marginatus 1 0 1Lepamiemegalotis 25 934Lapamiemicrolophus1 1 2 Micropterus salmoides401959Cbaenobryttuscoronarlus1117 28amer!canus 1 12Pomoxls nigromaculatus0 1 1 ItepisosteusplatYrhincuB0 3 3 Anguillarostrata0 1 1 Erlmyzon sucetta0 1 1Ameiurus na taIls0 3 3 Grandtotals177 215 392 Table1. Summaryof'f'lsh taggedatSilverSprings,Florida,betweenMarch12,andDecember 29, 1954 I -i I ,


co'" SpeciesTagDate Date St.Len. (mm) St. LeD..GrowthType TaggedRecapturedout When tsggedat :Recap. (mm)Micropterussslmoldes MeW VI-11-54 IX-24-54 10512413410Micropterus se.lmoidesMeW VII-25-54 1-15-548211213119Micropterussslmoldes MeW IX-24-54 1-15-54 21116115-1Mlcropterus salmoldesMeW VII-25-54 II-24-54 12211012818 Mlcropterus salmoides Mets1IX-24-54 III-15-54821201244 1l1cropterus salmoides Metal IX-24-54 XII-29-54 961051072Chaenobryttus coronariu8PlesticVI-18-54 1-15-54 119116116 0Chsenobryttuscoronarius MeW VI-18-54 1-15-54 119112117 5 Lepom1a2punctatuaPlseti.VIII-31-54 XI-24-5485 142 139 -3 Leoomis mierolophusPlastic II-24-54 XII-15-5421 214214 0Table 2. Recaptures of taggedfishatSilverSprings,Florida.


I I' I,. 80-39 75706560555040mtl.-35-+'il.... m30-;: ..25- 20-0I,,, 15,10-0,' ,S.",..I,".L.I'I'"\III IIII l!?o1':301i.o O'10 201lO 40 50 60708090100110 Stondal'd Length(lllIII.)II" I" I.Figure1.Length-lfeight,Lepom.ls punctatu6 punctatus,SUverSprings,Florida,October15,1954


-1 Number <'or"Splcimens ,\I .... .... Ir-'....................'i"'! 'I"i'I'i"'19 9'(''i"!' 'i''f? ;j'''rP'f'.,. i 'i"!'9 I, ag-." c ':!'" 'l'i.g>.. t:'mf;'Jilr1mim:;" \I\-+ !O: II ------. ----..../, " -' 0 0" ,.. ,-. _...... .0" \o ,--\ ,, '." .............. o'o'r \, T 7 l?'f?q:?rd15t'lI':><'" II ,............,,\ ,I "" ,., '\,o i ... '. ,o' ," , o. \ ... ) ./:\ '"'Cl'"/'" 7,, !O,.o.;l ,I, I1 ,\ .,)", ,\ I,,.. .. /...',-0', (';.\ 1IIIII !O: I, .' o '. ,; J i, ,I, ,,\I, f,o rII 7 I II.. gCl;'ji!O: \, ,", (i \, : -;-/ .0,,I7 J I I, l IIII !O: lII,,, I), \!, ),, ) i)-, ,, \ < I,I III,\ I \, \, ,\0' , ,, ,'. J!0' ," ,III,IIIII III,II, !O:, ,I, > f,\I,,\.'! , II , , ) I, tlI 0-5_6-1011-15 16-2021-25-!g 36-40 El 41-45 f;j46-50Z551-5556-60i 61-6566-7071-75 76-8081-85 86-90 91-95-96-100101-105 106-110111-115 116-120 121-125126-130 131-135 136-140 141-145146-150151-155 156-160161-165166-170171-175176-180-


1-41PRODUCTIVITY THEORY HoT.. Odwa A. OptimumEfficionc;r--l/ax1m1lmP"""r PrinciploAppUodto fIl0t0';rntho.i. In provious reports. tbe principlehasbeen statedthatcpsn!Itead1'"tatesyoto""tond to ba adjustedatan optillllllll but 1""officiencythat corro.pcndatotho maDmumpowor output. (0dIm and Pinkerton,American inpro)Thatthieprincipleievalidfor phlltos;rnthoticoyotom.eeo"" indicatod inFigure1$bel",,:!beporJilpor IIr. -, ,, Figure1'0Photosynthesisversusefflcien07 indicating low but optimlmlefficiencyassociated1d. thmaximumoutput Thiagl"aph shows thathigh errlcisnclessuchR8havebeen achieved1nallthe workassociatedwithth.e quantum cantroverse}" haveall beenclawllght intensities sothatthoPO'l'l'l3r outputofgluCO!" hasbeenverysmall ..Ontheotherhandthenaturalpopulationsof algaeadjustedt.ohighnaturallightintenslties run atlowefficienciesand highlight intensitieDbat 80thatamuchgreateroutputof glUC080resulteo Ifplantsare evolutionarily adaptedto maximum output the,.must sacrifice efflc:;'enc,. for parrerby this hypothesis..This is il.way ofstatingthat attelll'to to increase'WOrld food b;r raisingchloreUaathighefficienciesJlIllst necessarily flop.Asecondpartofthis hypothesismaybe statedthat the optimum efficiencyfarmaximumpower output decreasesasthelightintensity inereasesoPlantsadaptedin nature to deepwaterachievetheoptimumadjustement ata higherefficiency thatatthe eurfaceoH01'Jover, aplant adjusted foronelight intensitycan notbe llX)vedimmediately to another light intensity lIIld achieve tho optimumodjustmentwithoutinternalmodification. Acarclimbinga hillinsecondgear at optiJl1lm.efficiencycalnot achievethe optimwnsrriclency for a straight anywithout changing gsarso Intheplantgears marbethoccncentrationgradiente.Th. efficienciinFigure 15 areofthe samemagnitude at optimum.adjustementas those foundin SilverSprings and Coral Reef DBo Organlamal Sizeversus MetabolicRatein Phototropha1nOptimum ItIs nowwellknOll' that mstabolism of heterotrophic organisms115 invere8 to bodYs100 ina 2/3pcwor function that10 preoumod to berelated to the l1m.ltsto diffusionprocesses. From the above section it wasconcluded that photosyntheticsystems in open steady"statetend toallbecane adjusted toa similar state of runningat ma:d.mumpO"I'fer output because orthesurvival valWI' 111 GColog1c.l competitionboth inanenvirolllD8ntalandevolutionarysenseo V:ith similar lightintensit.1es, similarstead1'statoplantsystems sh'ouldbe :il.djusted to similar efficiencies andthue81ml1ar totalphotosyntheticpOTier output orglucosoonan basil50


42I Nowif theplantsrunat thesamerateot output per are.andif sizeeffects holdfor plants ae wollasan1mala, th.en emall plantelikeOhlorellait growing in steadystateshould achieve thesame output per areabut lfithlees Thelarge climaxrain-torestwith big plants ontho otherhandifadju.ted to theIWIOlig/lt.ond optimum.tfic1.nc7willroqu1N amuch largerstanding crop biomaS8 of theslanrmetabolismper poundot tissues. InFigure16 hoJ.cty io ohcnm agraphof IDhotoo1!lthoUcrat.. of plonts ofvarious m:1.n1mumdiameter"under naturallightor max1DDtmphotosyntheticadjustmentsC)Thedat..;:!!'dfromVerdu1Do data from Verduin Max1mum .001 -3AnnualPhotos;ynthee18t>lumover Omglucoseper gilldrT .01 30...1ght.per Hr..1t 1. 3000 .001.01 5izG inC .Figure160 itf.ct ofor gan1smal sizeon Photosynthesisper1ftl'lght ..It 115apparent thatthere18 a sizo eftectovera wide rangejust aD 1n theheterotrophaoThus, knowingth& light lntenslt1'.the sizeoftheproducer"andtheroughefflcl.ncleafoundingiven environments, ane cancompute t.he.tsadTotatsb1omaso(carr"ingCllpac1t.7). C. P,yrlllll1dShape and Orgonlolllll1 51.e Ifasoutlin.dabow_11 .1z.d proaucoroput. out.the.....production perarea lfith a small biomass as do large producers with alarge biomass both working atsimilar erficienclaa, one canvisualizet'ftoextremetypee of P1t"amidoascalculatsd in Figuro 17.(Fer dataaho1d.cg olm1lar .rficienc1es andproduction formass8blorella cultureand graBSplote s (Burlew!Mass Cultureof Alga--ehapter$-Wassinkat. al ..,Camegieo1953) Inone the sizeofthe organismdecreasesasonegoes up the food chainasingraal-grasshopper-eplder. Inthe other the size oftheorganism increases as one goeaupthefood chain asin chloreUa,paramecium, and fish..It th.. eameenergrpaZ!ll'8a up through bothfoodchains withtheaama10%.ffici.nc)" for t.hohig/lsrtrophiclovelo,t...,ent.iral7difforont. .haped p"-,,,,1do of ateA.dT stateblomaal!lresult b8Cllua8 ofthe differentratesof turnover..Somemetabollc rate figures are used to computeFigure17 fromHeilbrunn's t.xto Photosyntheticvaluesaretaken from Figura 160I,


Thopyramid. in Figure 11helptovi.ualizothepo.sihlorolationshipof thetro.,lcal oceantothetropical rainforest.. The smalls1ze oftropicalplankters asWltllas thehigh temperatUNtend1;0..tt.reversed.pyramidcalculated for Chlorslla-paramoci\llll-fi.h. 43Spider GrasshoGraBS 162810. Figuro11. TlloExtro.... Pyramidawith tho samoEnorll7 Flux .hOllingthoIhpgndencG of biomasspyramidshape on organismalsize. Do Bnera ContributlC1'llS or Turbulence and Current.Plankton SizeTheefficienciesof pri.mary production 80tar estimated tar Silver Springe(8:')andforthoEni".tokroot($)areconsidorably higherthan man)' othernatural COlllll_ tio.or laborator)'_ori.ment."'" at high light inwns1 tio tarmax1J!Dlmproductlcm.oSinCG there 1a a largedif'l'erence in nutrientsbetnClthe r rand Silver Springs, it seemsthatoome other propertY'1a 1n partreeponrible tor thishigtl production.It1a reuonabl.topostulatet.b.athigher efficienciesarcproducCid by"thestrong currents thatserveas a comnunit;rcirculatorysystem,maintainingbetter nutrientconcentrations adjacent to 06118 and removing 'Wast. productsoThooffoctofthocirculation might bo.tatodin t1Io""70' AoThe comnunltyrecslveaenero fromboth the sun and trCID the currento BeTh d1rectlTfromthesun goesfurther because ofthe current condition.. If tho USWl1officionc)'atmaximumadjustllllint10 about n. about S:'mightbelpostulatedas dueto thecurr01lt system.. Thismight 1mpq that thee.noweyatemederive more of thoir enerafrom primary productionfrom thecurrentenC"gT than fromthe lightrece1 wd .. It current1s asiDportant5.8impliedasuggestion can bemada asto 1riIy largorand hoaviwph)'toplanktonlL!ually provaUovor .moller _cio. withfatdroplots sothat tho)'IllS7 noat with tho.lIlllO dansit)' as wateroRsavierplanktera are maintainedb,rthsturbulent eddie. 80 that thO)'arecontinually falling through thocurront. thotII11pport th ... oThi. mech.an.1smprovidesthe cen ti1 thalocal currentwhichthe organism atwater c.rurl.t,..lrould not have..1hsrelativel3'lowChloreUa .rticloncios Obtained1nhighlightinwn.it)'ma culture.in spits of highnutriont.U7 beaccountedfor mallsize of ChlorsUawhich decrealJ8athe .ffectiveness of .tirring""chani .....SoDecrease ot DaytimePlant Respiration Accounted. for by the fI1potheeie otPlant ReeplratoqS;r.stAms&8herbivores ofthe Autotrophicsysteme--4rctlc SignificanceThe data ot slIIIvGlral authors(10 Kok,sseRabinowitch)suggests that plant rospirationin da)'timo 10 mch10than whsn tholight i3below tho


1Ir I ... I 44 compel18atlon pointo The concept ofpyramidBofbiomass1Ih1chmay1 supportedduring steadystat,suggestsa reaSODo When$ver food Mara ia pas8Gd through ananctrg;rtranafonnationatep, alargepercent, perhaps90%,must gointo healasreqlliredbyth8 secondlawof thermodynamic!! for thlae typesof undttr optimUlll cfficlency-maJdmumpower adjustments".If apUnt inthe daycandrive80me ofits workeyateIZ18direct13 ratherthan making glueD!!. firstand then bumingthealucoBlilI, theplantcan saw astlPandthUD avoidtbt heat loS8fran theextraat.poThua ahigher.trieitmeyinthoheterotrophic system 1smaintain.d mth11;88 respirationoAtnight, bythisvi.w, itbecomes neceasaI'7 for the planttofallbackonglucose storei thuslengthening thQchain of transformationa and raquUi!lg increasedrespiratioDoIf tru.,thiBbecomesespecially significant 'When lengthof day andnight 1sconsidered. for & llttlo dirfDrence in daylengthmeans incre&sQd .frici.ncT aswellas additioral lighto There is lessglucose thatmustbestored for use atthe inefficientnightrateoBy this view Arctic plants duringcontinuoUB daylight of SW!IUsr should be muchmoreefficient thansimilartemperateplants0 On anannualbasishowever there wouldba nogainforthe communit,.would haveto storemanymonthsworth of organicmatte:r atthe'inefficientnight rateo F0Anaorobic Springe and thoSaproboSyet...Thrtaaprobesystem of classifying pollution communitieeassWIIJ!I theassociationof characteristic indicator spec1.t1fordifferentdegrees or sewagetypepol::t.utiono AconsiderationorFloridaanaerobicsprings shaws thatthe systemlfhileuSfilful lIhenproperl1' u8edhasafallacTthat leads tomisleading CQ'lcluBicns when usedon waters1ngeneraloBeecher.Orange.WarmSalt.and VolueiaCooBlueSpring are all e:Glllplos of lOl["gespringswithlow organic contentwator that1salso 101r in oxygeno The conmuniti_s that rGlsultare both ana.robioandautotrophic in natUl".o Incontrast sewage cOIJIDWlit1esan anasrob1cand heterotrophic. Inthespringsone getssulfurbacteriaand blue grssn algaebutnoc111atssoInshortone getssomeorthosl1probasystem biotain water that istheeoctremeoppodteofsewage polluted utero ThOUghtless Wle ofthe saprobeeyat.em leadstoa campleta mi!lclassificationofthe type of primary productionin thesecaselS. Go SomGDefinitionI'.SuggestadIDIthodolog In agriculturo,1IlllI1'. continuedlabor andsupervisionguiee a comple,. communit,. inadiroction heHOlmver inEcologicalEngineering the outcomeofproductionof a complexcommunity1s achievedb1' properaelectionofcomponentaat thestartwith!Iubsequsnthandsoffthuspermitting the communit,. toreach a unique steady'stateadju!ltment. Inthe usual eci.ntiric experiment. man controls onsvariable 80as to thebehaviorofanotherdepondantvariable 19hile holdingotherconditione canstant.Thustheprocess1e oneof analyzingcQllponentprocess8e0 In Uicrocoemuparimentation ontheother hand componentsareputtogether ana:the complox allowed to makeitsowntrendsun&lrobservat10Do The study' ofecological engineeringb;ymicrOCOSlllexperimentation 1s 2. practical methodologr for l!Itudyingthe!s1l1thGe1a ofecological l!IystamaoThisepringsproject 1a an of themicrocol!tnappro_cho


I .,."'. I ; I,45Ahypothesls dependenceof commbnlty structureanddensityonproductivitybyJ.L.Yount The hypothesis1sofferedthatundersimilar general conditionsthespeciesvariety1saninversefunctionofthecommunityproducttvlty.Theaalpeofaseriesofplanktonsamplesmade by PacificOceanicFisheryInvestigationsoftheUnitedStates Fish andWildlifeService1neplpelae1cwatersofthecentralPacificOceanwerestudleddurlng 1952-1954. Observatlonson themledtotheformulationofthehypothesispresentedbelow. The mostpertinentobservations were asfollows.In most ofthesamplesstudied,manyspeciesofsalpeweretakenWithlittlepredominanceofanyonespecies.Inonesample,however,there were bothafargreatertotalsalpquantityanda great predominanceof'onespeciesofsalp,onlya rewothers being takenandtheseininsignificantquantities.Allealpspeciesapparentlysimultaneouslyoccupysimilarniches(theconceptofthenicheusedhereisthatofElton,1927,AnimalEcology:63-4),andapparentlyalsoaresubjecttothesameenvironmentalconditions,thus apparen)ly areecologicalequivalents(inimpoverishedwaters;Beebelow).Observationsmade byotherinvestigatorsarealsopertinenthere.Studentsofmarineplanktonof high latitudeshavedescribeditae"monotonous",consistingpredominantlyofonespeciesoforganismineachnicheapparently,althoughthetermniche has notbeen applied inthesedescriptions. MOFt descriptionsoftheplanktonoflowlatitUdes,however.emphasizethe great varietyatspeciesWithlittleornopredominance anyonespecies(pernlche)(seeSteuer,1910,Planktonkunde:601-4;Russelland Yonge, 1936, Seas:123-6;DaklnandColefax,1940.ThePlanktonoftheAustralianCoastalWaters offNew South \'Jales, Un1v.Sydney,Dept.Zoology. 27-34).AnotherpertinentobservationdiscussedbySteuerandDakinis that productivityinthetropicsinwatersinfluencedbylanddrainuEeand1nregionsotupwellingmay equal orevenexceedthatof high latitUdes.Iftheseobservations are consideredtogether,it thatinepipelagic waters withrelativelygreatquantitiesofnutrientchemicals(theenrichedareas),productionoftheplank-tonisgreatinquantitybuttrendstowardfewspeciesoforgan1sms-probablyonlyonedominantspeciesoerniche--and1nepipelag1cwaters wlth relatlvelysmallquantltlesofnutrlentchemlcals(the1mpoverishedareas),theplanktonissmallinquantityandtrendstowardmanyspeciesoforganlsms--apparentlymanyspeciesperniche.


I -r I.I-. .I. L 45Ahypothesis dependenceofcommbn1tystructureanddensityonproductivitybyJ.L. Yount Thehypothesis1sofferedthatundersimilar general conditionsthespeciesvariety1saninversefunction of the cammun1ty productivityThesalpsofaseriesofplanktonsamplesmadebyPacificOceanicFisheryInvestigationsoftheUnitedStatesFishandWildlifeServicein ep1pe1ae1c watersofthecentralPacificOceanwerestudiedduring1952-1954.Observationsonthemledtotheformulationofthehypothesispresentedbelow.Themostpertinentobservations ae follows. Inmostofthesamplesstudied,manyspeciesofsalpaweretakenWithlittlepredominanceofanyonespecies.Inooesample.however,therewerebothafargreatertotalsalpquantityanda great predominanceof speciesofsalp,onlya few others being takenandtheseininsignificantquantities.Allsa1pspeciesapparentlysimultaneouslyoccupysimilarniches(theconceptofthenicheusedhereisthatofElton,1927,Animal Ecology; 63-4),andapparentlyaleo are subJecttothesameenvironmentalconditions,thus areecologicalequivalents(1nimpoverishedwaters;Beebelow).Observationsmadebyotherinvestigatorsarealsopertinenthere.Studentsofmarineplankton ofhigh !atitudeshavedescribeditasIImonotonouslt,consistingpredominantlyofonespeciesoforganism1neachnicheapparently,althoughthetermnichehasnotbeen ap911ed 1nthesedescriptions. Most descriptionsoftheplanktonof low latitUdes,however,emphasizethegreatvarietyotspecies wi th11ttIeornopredominance anyonespecies(perniche)(eeeSteuer,1910,Planktonkunde:601-4jRusselland Yonge, 1936,The Seas; 123-6;DakinandCo1efax.1940,ThePlanktonoftheAustralianCoastal '/aters off .New South v/ales, Univ.Sydney,Dept. Zoology. Publ.I:27-34).AnotherpertinentobservationdiscussedbySteuerandDakin1ethatproductivityinthetropicsinwatersinfluencedbyland draina8e and1nregionsafupwellingmay equal orevenexceedthatof latitudes. If theseobservationsareconsideredtogether,it that1neplpelagicwaterswithrelatively great quantitiesofnutrientchemicals(theenrichedareas),production of theplank-tonisgreatinquantitybuttrends toward fewspeciesoforganisms-probablyonlyonedominantspeciesoerniche--andineplpelagicwaters With relativelysmallquantitiesofnutrientchem1ca1e(theimpoverishedareas),theplankton1ssmall1nquantityandtrendstowardmanyspeciesofarganisms--apparentlymanyspeciesperniche.


1 I.I" .,.I, 46Ittheplanktonnowbecomparedtosessile oresnlsms ofthe Autwuchs andbenthos,itwillbeBeen that the two groupshave similar natures,1n that s6ss1leorganismsand planktera aremoderately passive 1nabilitytocapturefoodandtomove1ntheirmedium(mostsBseileorganisms,ofcouree,moveduringBornephase of theirllfecycle,andplankters have Borneabilitytomove,butdirectedmovement1elimitedforboth). For thisreason,atleasttheplanktonandsessileorganismsprobablyshouldbeconsideredtogetherinthehypothesis.It1spostulatedfromtheseobservationsthatifproductivity1S low andotherfactorsareconstant,speciesot eco1og1callypaRslve organismsatleast,mayoccurtogetherasecologicalequivalentswithlittle nopredominanoe ot onespeCies1neachniche;andconverselyifproductivityishighandotherfactorsareconstant,speciesofthese organisms shouldnotoccurtogetheras eoological equivalents,butratheronespeoiesshoulddominate1neachniche. 4ble leadstoafurther postulate. ifproductivity1slowandotherfactorsareconstant,competitionandothereoaationsshouldbereduced for these or8anlsma; whereasitproductiVity1s high andotherfactorsareconstant,competition and othercoactionsshouldbeincreasedforthem. 'be followingevidencestendtosupportthishypothesis.lnthetropic holoplankton,productivityis low andquantity19small,exce9t1ncertainregionsmentionedabove,andspeciesnumbersare great; 1ntheeplpelaglcholoplanktonofhighlatitudes,productivity18 hlBh and 4uantlty 1sgreat,butspeciesnumbersaresmall.Occasionally1ntropiowatere,swarmsotplanktersappear,consistingof few species oforganisms andarelativelygreatquant1ty--product1v1tyistherefore high andspeciesnumbers few, eveninthemidstofimpoverishedwaters,underenrichedoond1tlons.Inplanktontowsfromimpoverishedwaters,manyspeciesof organisms occurtogether that apparentlyareeoologlcalequivalents,andthisiseVidentlynot Drue of olankton towsfromenriched waters (evidencesforthisstatement are basedch1eflyonstudiesof salpst butit1spostulatedasbeingtrueofotherplankters 8B well).InFlorida Springs, ithasbeennoted(H.T.Odum,L.A.Whitford,W.C. Sloan) thatproductiVity18highandthenumber of speoiesorthevariousgroupsof orBanlsms islow. 'hiS hypothesisis being tested at present WithAufwuchsgrowth underthecontrolledconditioneoftheFloridaSpringsand,ifresultswarrant,is tobetestedunderotherfreshwaterandmarineconditione.Countsof Autwuchs speciesnumbersonslides,relativetocurrentcontrolledtotalproductiVity,havebeenstarted.Countsmade on the preliminaryfirstserieeareconsistent with thehypothesis.

Productivity of Florida springs
Full Citation
Permanent Link: http://ufdc.ufl.edu/UF00073979/00004
 Material Information
Title: Productivity of Florida springs NR 163-106 (NONR 580-02) Report to Biology Branch, Office of Naval Research; progress 1953-1957
Physical Description: 4 v. : illus. ; 28 cm.
Language: English
Creator: University of Florida -- Dept. of Biology
United States -- Office on Naval Research
Publisher: s.n.
Place of Publication: Gainesville
Subjects / Keywords: Freshwater biology -- Florida   ( lcsh )
Springs -- Florida   ( lcsh )
Genre: non-fiction   ( marcgt )
General Note: Frequency varies.
Funding: This collection includes items related to Florida's environments, ecosystems, and species. It includes the subcollections of Florida Cooperative Fish and Wildlife Research Unit project documents, the Sea Grant technical series, the Florida Geological Survey series, the Coastal Engineering Department series, the Howard T. Odum Center for Wetland technical reports, and other entities devoted to the study and preservation of Florida's natural resources.
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Holding Location: University of Florida
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oclc - 21432514
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Table of Contents
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    Table of Contents
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        Productivity of Silver Springs
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    Comparison of an Eniwetok coral reef community in Silver Springs
        Page 18
        Page 19
    Plant community stability during 3 years
        Page 20
    Standing crop and community survey of submerged vegetation in seven springs
        Page 21
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    Fishery biology studies in Silver Springs
        Page 36
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    Productivity theory
        Page 42
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    Hypothesis regarding dependence of community structure and density on productivity
        Page 46
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Full Text


NR 163.-106
(NONR 580-o02)

Second Annual Report

to Biology Branch
Office of iO'$,J- Research
S:i'.' ,es from JLanuar 1 to December 31, 1954

H. T., Odu4m rand Jo L. Yount

wtith sections by

D031o0 &Wn'-
M -id K Caldwo1ll

':-ck H l .ri;

opa nf ~of r .. ,..

.'.': of Florida

Gai O ," -, ,, : .

-477. .,


INTRODUCTION o o o e o e o o o o o 1


-7-)!oiVCTIVITY OF SILVER SPRINGS o o o o o o 5

WITH SILVER SPRINGS o o o o o .. o o 17







Prepared by:

Howard To, 01.'i aJd James L. Younx, with sections written by Delle
Natelson, Dr~'-.. K. C,!l't.Al and Frederick Ho Berry.



NONR 580(02)

ANNUAL RATE: $5,C0 (h -,.:-3)

CONTRACTOR: Departeont of Biology, University of Florida, Gainesville (w-ith
Biology r.:~~r'c.h Office of Naval Research)'

PRINCIPLE INVESTIGATOR: (This project is administratively listed under W. C. Allee,
Head of the Biology Department),
Howard To Odum (February 1 September 1, 1956)
James L. Yo'-~ (September 1, 195h January 31, 1955)
Associate: Delle Natelson (September 1, 1954 June 15, 1955)
Assistants: W. C. Sloan (Y:7'.-: -y 1 June 30, 1954)
F, H. ro!:.. (June I September 1, 1954)
D, K. Caldvell ( .1. 'I,.,c. 1i 1954 January 31. 1955)



A study of basic factors that control productivity and of the effects
of productivity on coiniunity structure and density by an analysis of
the .- si'. *: conditions supplied by selected constant temperature
3pi k':.'tVl.,:


a. During current ..'. :;; ..'.iod

Production measurorents at different times of the year indicate a
linear relationship of I2 .' v'-,iitj and overall production at about 8% of the
visible light energy i :'-a ~.. level, clbtri.'eien s of a coral reef at Eniwetok
indicate 6%. Further evidence of breeding at all seasons but with a quantitative
pulse in the seasons of maximum light indicates that the seasonal fluctuation in
primary production is routed through reproduction rather than through major changes
in populations, The succession of plants and animals of the aufwuchs has been shown
with glass slides and counts from Sagt iai blades. Loss of oxygen bubbles during
the day and emergence of aquatic insects at night have been measured with funnels.
Bell jar measurements are reported for bacterial metabolism on mud surfaces, pH
determined C02 uptake ,.,j:.'.:': with titration determinations. A few rough estimates of
herbivore production have been made from caged snails, aufwuchs succession, and fish
tagging. Nitrate uptake at I!':.,-i by aufwachs communities has been c.nf,inod in a
circulating microcosm experimuet as well as in bell jars in the springs. Distri-
butions of oxygen and or-;_. -:r" have been used to criticize the saprobe stream
classification system. Theoretical consideration of maximum photosynthetic rates
in literature data indicates lCirthi3ic rate variation inversely with organismal
size just as for respiratory metabolism. Extreme pyramid shapes are thus shown for
communities in which or-:ni:-.-'l size decreases up the food chain and for other
communities with the same ~.... ~. influx but with organismal size increasing up the
food chain. Literature data is used to further demonstrate the validity of the
optimum efficiency--mBaxim0 i po:er principle for Photosynthesis. Work on aquatic

ABSTRACT (Cont'd.)

plants by Dr. Delle Uatl,:l ;i..-ates essential stability of aquatic plant comma
munities after 3 years and about O10*-20% reproducibility in previous bioaess
estimates by Davis Wo rk on an annual picture of the fishery characteristics by
Caldwell, Berry, and Odu is half completed. The study of aquatic ic':; in
relationship to spring g:.'T *:.:~s by W, Co Sloan has been completed as M14 S.
Thesis. J. Yount has begun a study of effect of total productivity on community
composition using aufnuche .',: i;.irs on glass slides placed in different current
and light conditions in '.-l-. Springs,

be Since Start of Project

This contract was begun June 1, 1952. In the year and a half pre-
ceeding the present report period, work of a very varied nature outlined the
trophic structure and metabolin of Silver Springs with comparisons made with
other Florida springs. The intensivee study of Silver Springs is now nearly com-
plete and quantitative comparisons of productivity with other Springs will fol-
low. Most of the techniques and approaches outlined in the original proposal
have now been p.Pi.. Tl. study of ..':tors. affecting qualitative community
structure is the main S'u,... '.,.: ::.;.Co


1. By J. L. Yount
Determine the relationship of productivity to the variety
and the dominance of species comprising the various commun-
ities of a spring. This is to be done as follows:
a) by observation of the numbers of species per niche in
single habitats within the spring, e.g., Aufwuchs on
S~.iLtJ-; "._ blades and glass slides in different regions of
the q;. w.ng where A l'!i.otivity differs but other factors
are constant. Both high and low trophic levels will be
b) det, .>.' *' relati',on.~ ps as above under experimental
conditions in ; uaria another vessels. Plankton and Auf-
wuchs are expct .. to serve as the principal experia.:..1tl
groups, but other groups may also be examined.

2. By D. K. Caldwell, F. H. Berry, and H. T. Odum duW:.w. Spring
and Summer.
Complete an annual cycle of 5 3hro' characteristics
'Wb.?:_'. last s.~ ing as follows:
a) tag and ... pture more fish
b) determine the significance of scale annuli in silver
M'IA. ; :g. lA.
c) further determine the extent of winter b~:-:.: Tg. of
fishes in Silver's constant temperature waters
d) cow.,t.. ;: f:,;] assay of dominate fish species
e) 1 .termine Cr't-- h rate of young stua.pl1,hn'e..-er in o.ses
f) determine the sigrni- chancee of .;m ,,h -.ij: ..... y -.C.-.- .:'.. o
fi." fish collected in sprl,.a' thr:.. --out the year

3. By D. Natelson oa'.'i..a. Sp,~0~3i; 1955
Co.mp,.'e o .1 community composition of aquatic plants with
Comi!!'.: ties not in constant inr.rn-ature 5-.M 1 r..;s,

4, By H, T. Odum ';. -in;, summer 1955
a) Relate the overall community p.ola.ction of 20 springs
measursl- .' the downstream flow method to current
veloci' to test hypothesis that the overall primary
productivity of communities in steady state is a function
of veloci.,- o' water over plant surfaces,
b) Co;,.pl..te the picture of metabolism in Silver Springs
by additional data on organic matter loss downstream,
effects of side boils, herbivore growth, repetitions
of (u ..; ,t.t ...-.i-urements, spectrogram of water and plant

Long Range Plans
1. By J. L. Yount
a) determine relationship of productivity to variety and
species d-'l,Jr."se in other aquatic habitats, both inland
and marine, contrasting habitats with high productivity
and low productivity.

b) define the relationship between productivity and

2. By H. T. Odum (Dulke University)
a) apply method of measuring community structure and
metabolism to other steady state systems such as Hot
Springs -. tropical streams.
b) construct microcosms in the laboratory to contain
small stee- state communities in order to further
S-limit ...Inciples.


Supported in part by this project:
Odum, H. T. 1953. Dissolved phosphorus in Florida Waters. Report
oC Investi.l tons #9, Fla. Geol. Survey, Tallahassee Fla. 40 pp.

In press:
Odum, H. To ... RT C. Pinkerton
Times Spr-..- Regulator: The optimum efficiency for maximum
power outp.'-. in *...olcal and biological systems.
American Scientist
Odum H. T., : David _. G,:3:,9:Cll
Fish respiration in the natural oxygen gradient of an anaerobic
spring in Florida.

Completed Th. -:., j
Slain, W. Co Scmse onvirr .-.,.i al factors influencing the dis-
tribution of aquatic insects in certain Florida S.ji-...:;3.

Manuscripts cc.,bl..ted ", submitted for publication:
Sloan, W. C. A c. -:-:::.tlve ecological study of the insects of
two Florida Spr'c.. (Submitted to Ejo~ y).

Whitford, L. A. The Communities of Algae in the Springs and
Spring Streams of Florida. (Submitted to EcO1U.)).

Odum, H, T, and Eo P. Odum. Trophic Structure and P.. .-,,;ictivity
of a Windward R::.' at Eniwetok, 90 pp. manuscript.

Manuscript in pre.a..r'ation
Odum, H. T.o 7,'.. i- Structure and Productivity of Silver Springs


'o -:" -' To Odum

i:. "...v.'.... ':, and d-'. "~.-. studies continue n Silver SpIt.rA in an
effort to determine the da- :- working oC a fertile e-.-.*n.1-,: aquatic
.... ?-.. ,'.A in s ..

A o ..: ., ..

" i 'n ',...N in Chanostatic Conditions

The upstreamsLdownstrean method of measuring productivity as reported
in :.., pr- .. :rr--t" has now been carried out in Silver !.K..-*
in various saseons of the y.r and with different loud covers From these
data an annual cwr. f- of total- j r -../ primary production is dramn in
Fi..ure 1l The .:,ro, *arundier, -3s curve is th total'primary production.
'.', horizn'-tal '' .. the respiration and downstream losses as
prevo..7-. dsto.rra~W.ao tth ..-i. is in itood,;r state the area above
the horicntal lian! shoud c the area below. These areas do not
"12. -carr- !; .; in ..me asurement of downstream lo sse
in the "-'. K c"I::t'in., .d in the :...a of aide boils will be
:required to ac.co..t i t .I excsss production
From, the
A cuirve ". '.. :.l: iij p .c '.; experiments is
also showa. in '.' .. f2erence -.. .: '. this ,;. i <,1 and the
total p-.',: .. r.oproGuK ; tho ". -p.:-X production which turns out to be 7:,
of the total Tven .' '. 1ho "'.. :'. biomass is ,r<...e.i.o That most
of the produce., ; community is algae and Sagittaria is evident
from plant maps of silver: Springs referred to in previous



*" :" total ,' ..'...vr production

S/ \ \ -12

-** '---- . r :)
,. -.," ,f

r3 ,, lXo 'n.ual ..*8 of .:'... Production

Fr ',"T' Ci>


'o -:" -' To Odum

i:. "...v.'.... ':, and d-'. "~.-. studies continue n Silver SpIt.rA in an
effort to determine the da- :- working oC a fertile e-.-.*n.1-,: aquatic
.... ?-.. ,'.A in s ..

A o ..: ., ..

" i 'n ',...N in Chanostatic Conditions

The upstreamsLdownstrean method of measuring productivity as reported
in :.., pr- .. :rr--t" has now been carried out in Silver !.K..-*
in various saseons of the y.r and with different loud covers From these
data an annual cwr. f- of total- j r -../ primary production is dramn in
Fi..ure 1l The .:,ro, *arundier, -3s curve is th total'primary production.
'.', horizn'-tal '' .. the respiration and downstream losses as
prevo..7-. dsto.rra~W.ao tth ..-i. is in itood,;r state the area above
the horicntal lian! shoud c the area below. These areas do not
"12. -carr- !; .; in ..me asurement of downstream lo sse
in the "-'. K c"I::t'in., .d in the :...a of aide boils will be
:required to ac.co..t i t .I excsss production
From, the
A cuirve ". '.. :.l: iij p .c '.; experiments is
also showa. in '.' .. f2erence -.. .: '. this ,;. i <,1 and the
total p-.',: .. r.oproGuK ; tho ". -p.:-X production which turns out to be 7:,
of the total Tven .' '. 1ho "'.. :'. biomass is ,r<...e.i.o That most
of the produce., ; community is algae and Sagittaria is evident
from plant maps of silver: Springs referred to in previous



*" :" total ,' ..'...vr production

S/ \ \ -12

-** '---- . r :)
,. -.," ,f

r3 ,, lXo 'n.ual ..*8 of .:'... Production

Fr ',"T' Ci>

B0 Determination of t- ".. 2 :..on i.-v Reaching the Plants

In order to deterine -,.:. "'...-;. ..-;r of ;...lianry production, light
i n::..it ; --;:~:.".ti thoe -'.-to were determined as follows:
(1) ntr.:, ..' ,- .1 :' -"' th.o .n." on a given date for a givz.un cloud cover
ie d ..inw from '.:' .:- in -...V (199. Bullo mero Meteor. Soce 30:208-213.)o
(2) 1.. C of this is talre, ,, ",'.. rA :.d- and half in the visible rango
(3) With a tubmarina .. '.:: "- containing a veston photronic cell sensitive
to visible %wave '. .. the p',r.oz,t penetration to plant level at 8 ft is
determined A winter curv'oe mas shaon on page 8 of the last progress ::.,i -'cl.
A sammaer curve is now g:,'" in Figure b,,. ,.
() .:h, :.':. - trans.fidcion :;.r' a given tirea of the year is interpolated
between th two xtriOmo curvm. ;:-p.*:.~.:-:.. t'lod by January and MaQo
TI- 'i;-: *.. t batean percent transmission in mainly due to a diffarenco
in angle of incidnasco
(g) 1". i.Asolat io : Ith '*.n".; comanmity is di; ti;i&..' .'. 2:
frm tres : '. wn. the sun is at a low angloo Due to the orienta.tin
of the -sd its. tra ,.* ..v t ;t is greatest in the ..
as ref.loCted in .' of the diurnal curveSo The ree effect :is
greatest in the anter. i. li OVeA on cloudy o-.:*. '2 .7 1 .
wen the 1 1'". in c :.7 .
To correct for Lc .- of trees, at least during the ? -.'r -: the
llily insolation erb 'suI s :tioned above are redrawn so that the n .t.j .;
is symatrical ih the r-a 7 curve as shTa in Figure 2o T,..
area betyv en the rexa'r. ..L. 0 '. curve *.'k-..- -Q- .L the light removed
by the treaso No Iuch ct':on'tion is made for cui.,; day:. The correction
on the winter ournvS is -proportionately th than on the summer curve

S!U.. intesnsitis do~ ruined in this ray are related to production I-- tr-..

':T' .: 'on A'. trees


Time o f f~ ,
..arch of V..t., May 286 "1'.;"

Co r.iu.-:., as as a Fiuactin of I.L:,% Intensity for the Whole Community

For each -.07 production iSu:. .' given in rugye 1 a WdO.iN1. :. .
intensity v..r'. as 0:- .-.. :.as described in p:,-!.'yvph B above The
.'ph in :.-a 3 shows total community ir.'.i productivity as a 2 o'.' -
of light intense si_ ..1' .', k','. wave lr,;th reaching plant le.-, The
horizontal a3.: is the eatAmate of respiratory and dconrstream Dow rmrn estimated
from I .:..ious .2 -'.! ,

?' <..:, 0..'. the '. ";ion is : .. 'Jcyct.cri- to the light intensity even
at these relaSt i'. '.." '. ',. 1'. intnsitieso A.'-t physiological u pr -.-; .s
on :Irn: ;-. :' how' efficiencies with increasing light inteonity
at high ; ':.' intensitisol It can r'.? '. be noted that the conmtunity
runs below its overall :../:'.;.n point on cX'.rA y winter dayso ThiQ does
not mean that individuvl. :i do this for the mmni common compensation
point i.Ot. .: not : r ton butaton b downstream losses. Possibly
an o; -.- .'. of the '. .. ..j th .c '.n. r 'wnstream lies in a
requir-emet that at st;'; sA.nte the .:.. plants must neveBrthelss
never be e oa d ;.;:.d'Wions bl' ow the individual compensation point,
As seen below th (o.ral ro o do% not do this and similarly dogs not have
the low '. :,int.o '. o there is a generalization that
temp ratee coarmnitKies :mut producec excess L:::n.i. matter as soil or
peat or downstrsana Ova wtich, tropical communities need not d*o
Pro, the data *' .". : it is not : '.:.? whether .h'vy is a break in the
curve at @ ': as -0'::. .:-A in the theoretical oectiono
A,'h.rN :;.7 the :.. .. '.. .. .. .' t ....'':-r an i oxn an .og.- i..: *..a. .. r;.r
with .;.-," intrnit'i.'o As discuss ..W:'-rw this i., :...t be
accounted W'-;* by incvr's .'..bation of -.' which indil.iA l1;
t're ;.1 the ir .. j; 'i. n I:s..- da:ys in more '..-, phacw,

S.A :.'*.-^. yday ,. -
S4(504 W ..TW :mr:7 line
Production ., -

50 ---. s--------- cr't.mir compensation

i% .i............. ..........-.
-| ,.i"r lin. ,-

'- ;'.? ";'.' visible light

S'.ty p ..,'-0- product on v ., .; : : '.. 7

TO..9 i '

Do Seasonal Pulse of ..;' Pr..?,dingj and Photoperiodism

The m.:apl~ in Figu 's. 1 .: 3 indicate a strong pulse of energy
in the spring and summ P ~'.- .: --,d to the less lighted months This is
actually a :r "'-' difference in the aquatic community than on land because
of the angle ....aglo ot the trees and water reflection

One is accustomed to r. '.,I.-...aAng big seasonal differences in energy
flux with succession and blooms :..W-.on;. the planktonic organisms in 3.-'- s
and in the ocean It .. : i" '. l-.. to consider the fate of the .-::. ..u;..
pulse ir Silver -.01M; 1:. n. o v.:r.: sucoessional changes have boon
ab.-.'. ven in the mic1-oscopic n-.lj. of the aufmchso

In i'. :-es h and 5 are shoirn r.':i.w.l pi'tu.res of br.andlI'z in
the zp-.pl; snail Pon ~ .3Mca which .. '.y its eggs above the water line and in
Pa&",.onstes ehich carries its :g2s,. From these graphs it -:- b
cr....*. that in the so .br.-e. .,i; occurs .'"..~:; hoo '.: -. the -.r in this
constant temperatureo en-viron;menat T7.t at 'i',:-rent rates that are
b..y to be ... % ... ...;:'o.L.fl. Thus these species seem
adapted to the ': pulse of the whole community As i;: 'i.-,
ago doscr::.b. : .. th survival of -o"T p'.ivl,. community complex r'-..p '.
the components to eat neither too much or too little As mentioned
in the r:.p. on :. .. ..' .. .3 below there is some evidence that
similar round theo '. "' : occurs in the fishes with a l g..,o

Figo ) Annual,


P.-*- !' .. ..... \

per 125 .
of shore l

-. ,- 4 ;. h S j. 3f .. ^
"1'. Month off the .V '
Fig.o 5 Ar.v-:
Reproduction of
Pal oneteS PAI.. T.',

the females
over 21 gi
with eggs /

Ho Initial r,-u:' -n Yicrocosm Eolperiments Nitrate uptake in the Daak

As mentioned in the theoretical section below microcosm exparimants
are a useful type of .'; .im ent for studying ecological open steady state
systems A ...' -- -: -i. .. blade of Sagittaria with its encrusting mat
of aufTv-', was placed i th. e tVo., > G .'j ? *.-- tube of the apparatus pic -..
in Figure 6 below '..- .';:.'ings water was circulated thus -I .j.ul.,T.
the natural ~ ?. ;'iono i. a time interval some of the 650 cc of
water vi'..i was t: a nd tested for changes in oxygen, carbon dioxide
and -n it .- .."

The .".;. co: .clusion ws .: in spite of all the light sources in
thp e I-::' ;.;- ,. h could i.. 1 brought to bears the aufwuchs micrc-.-. .:.:~ix-ti
normally adjusted t) -Utd or ..... intensities rapidly used up the
y P.-.,n :p-:Lx and dicL ".- as not yet been possiba to get the
production up ove '..:. ooam:Iunity compensation point which is very 1.'i.h
because of t he tcrotrophc a ,s .A1, as autotrophie camponentso

It was.g ':. "'' ,' .. to show that nitrate is rspi.l.,7 fixed
by same coiaponents in te unity even in the dark d'ri- ;.];.
:,pir :;<:...i metabolismso '.:" experiment helps to confirm the ..
,iptar @f nitrate fou.d in ''1 I :-1. Jars in the .j.1 and the
slight decrease of .itrat, .'. tihe boil ,.,:, :'--" both in.the day and
at night In 5$ .- .. .':':...'.. o037 ppm
decreased to o23 ppm in 1 I- and nitrate
was not deti -. i-. lafti .,.'.-. "Silver'
water standing in :..L-:a bot' tlo l.s:c
nitrate at a rate of 2. pm
per month, S l ve
Fi Springs
Flow "- ..w....a\

.MoneL Metal 1/8 HP
Circulating pump
l'h-. re :

Fo AufNucho Succession

The rates of CL-':.' and succession of autwuchs have been measured on
slides submerged in the .;-i- aMd by a count of distribution of pl-.n. and
animals along thae I.'. i' : -': i bladeso Since Sa ttaria blades grow
from the bottom, it has ': pF'ible to measure their gro h rate as a
means of th.t.e .;i::p'o, th age of the attached aufwuehs at any place on the
blade The distance fe..-. the base of the plant indicates the ti=e
since the succession hb.., .

As sha~n in 1'.,f 'u re the rates of growth of single blades is far from
equal Small wires were inserted in a young blade and an older blade in
the same clr:.'p.. After 26 days one blade hd shown a rapid 182% growth
pushing the attached wres tith the tip whereas the other older blade
had hardly ,r":. :, Appar?,,arn, a blade shoots out ard then as the aufwachs
covers it growth ceases and goes into new blades. Thus one gets old and
,;~.-. g4r ~16. d::: of near.., the same length next to each other, one bi5:j
clean, the other -rv ".r covered with the periphyton community

Figo 7, 26 day
growth of
marked blades ->w
of iria '
April 5, -'

j.-4 1416 inches

For a whole ,. of ,.I: -, however there is an average r.?...:which
tend to average tout the spurts, Th:r,,f.',- by cutting 50 blades into
segments of 2 inches a,., '., wd placing all the first shen~t '.:..'.h:.
all the ...a etop one may relate the attached
average aufwuchs to the average rate of clump gr otho Knowing thL
area of the blades scraped in each ~I- group and knowing previously the
percent growth of F-h'.taria '..,ra the planting experiments, one converts
length into time and ",on counts into area estinateso If '.'
are in '-,,.:;-" state tho parent loss at the tip of the clump is the
percent growth A ounft '.' the 1 : segments when fraJl over the
spring area gives an estimate ofatf rate of auwuchs growth and thus of
the components of the ty "..'"i blade type organisms As an illustration
of these methods a curve of A;i-es versus time and thus also length is
shown in Figure 8o Easti;s:es of midge growth rate from this curve are
discussed below Sqe .: ,.' curves of succession on glass slides are given
in Figure 90 The l.r .lariti'e in this last Figure may be due to the
positiorsof the slide boxhe in different currents and depths Further work
on this is being carried on by Dr. Younto
The succession i.*;:-. :'n ar much like planktonic populations with
bacteria first, small sn.., n~ x t then larger algae, and finally herbivores
and carnivores as :..:.-:: ". of a pseudo-climax is attained. Thus one has
continuasucession in the ia.cro-environments of the overall .i.:r, state
Similar microcomns should be 1. 4 :.; for in tropical oceanic plankton


blade 1 in yc>m

on a B L do e100

of Co :..
type J."'. "




Herbivores -
per slide

per slide

Fig. 9o Succs=
sion an slides

. ---- ------H


.. ------- P-

"0 /
H .' Craspedacusta






anJi m.o

of bdS
106 -

day s

"'.-* ..'r,

'. ., 8 ,,.


o Insect En rj;,-.o anzd ::... Measurement

The Very rgo ; caddis flies and other eq-i. .,; insects
just ~.f'Zt, sunset has bo-n *. orr d to in previous reports. It has been noted
to be intense both en fr. -.. days in -r.'. :oi aid hot summer days Foie
funnel devices waer placed the s aprai.: as shown in Figure ,.. Soz .'.2:.'-:.:-.ive
counts were ofL;..L, from thse sot '.,:i as given in Table o 1 E ncS-r;;.-..:
at dawn is not -i.ln..;. t in comparison to the evening outbursts Pc-'. ..
estimates of h"r.-: r.:' ". r.oates(net) are calculated below from this
measurement of ....*. -' rato.
_,..raduated centrifuge tube
.. .. ..! '
... ;'- merging" b'.. 2o oxygen bubbles are
Figure' .. ... 'hera ." .
2 -, a -
Oxygen bubble .
/ ra d. funnell @-

.' coated with
/ ,I/ / ""

The a is :. during the daytime to catch
the o ,i to 's' in the Aufwuchs and then
rise to the surface ithot co, ;L ( 1 This loss of s'.:;. ciO
is a source of er o in t:~l p.ouc:tlion inmasurements that causes and
underestimat-in of th VtttL .don: ? 'c.r-a g the tubes exactly
at the water t7 r' + the '"': .n..n can be converted into a ::' -uh
'.-t, i since t h bubble arS at ~attosphl r ic pressitre It seemj s ';-
that these 1 .'- a .... ."' c.r... a s Vhour measurements ibow a
maximum effect at time of ,xia ,,' ....'. ':..... See Table 2o

.:'i 1 Insect Di,-r,!g.:.-': In Silver :',..,,.. '
in ap'~:.).:tu3 of Figo 10o
Place Caddis
Distance from shore Time i l :.in

1 fto 1, ; 20 7:30--12:00 pM 9 5
S28 6:00--1:00 p.m. 1 1
25 ft 2 f 20 9;390-12:00 p.m. 8 2
S: 28 -':00=-3..,00 p.m. 2 1
.; 13 8:20--2:42 p.m. 15 2
40 fto Station 1 fto 20 7:30--12:00 p.mo 5 2
Station 1 Fay 28 6:00--11:00 p.,m 3 5
Station 2 .:-... .r 28, 6:00-:100 p.m. 0 2


5.o 2,4

Table 2. ]r','.:c' ";hing the Surface in Silver Springs, 1954

4. h,,.

,''blo Time
in cco Laps.
in irso

(t~l~ilI= = --~IPE-~-PI~ nnmr~~lg~P Pnsz~if~B~~i

NIGHT: 8:20--2:42 a.m, May 133l,-

6:00--11:00 p.m. May 28


7:00--3:00 a.m, *;; '-?Th 0

2:20 p.m.-4:05%
9*30 a.m.-3:s4

p.m.May 6
p.m. May 23



4:00-=7:00 p.m. **.- 23 ,1
moa I

Ho Herbivore Production '-..'-.,-,

From data in p.,:.'..:,C:.
as to the production rates
previous progress reports

:. 9." ,;-..:;:n some rough estimates can now be mad
of the dominant herbivores such as were listed in
Soavral methods are used as follows:

lo From the estimates of t, ... at the tips of Sagittaria blades in Figure
8 and from the ae~rago parcont C:..*.ttaria growth of 1%/day from Figure 1
One computs the rtes th e of midge growth o;,ary to keep up with tip loss
in a steady state. This is an u.derestimation as it does not include early
emergence and losses to pred-s,.aton in the middle sections of the grass blades






~i~p~L~~-z mn ~ I .?__ ___ __.cDI~.-^.






DAY:s !$--9:30 a4,m, 9:.0: 23



2o A few cages of Pomac "i a ~tJ ivigg ra snails were maintained with
an abundance of food -rnc v.:, meas-urments before and after a month growth
period. These estimatQs aro underestimates sines the snails used wore already
of moderate size and past the mor i rapid juvenile growth stages.

30 From the section G above on insect emergence the growth rate necessary
to balance the emergence in steady state was determined using 8002 gms dry per
emerging individual 'I,." fig~ r should probably be added to the Tip-loss
figure in method #1 above.

4, From the estirates of '.t-i..l g crop biomass of small invertebrato
herbivores in previous reports one can obtain the total respiration -Ji;.:.-;
a rough figure for stream invesrtbrates of o8 cc/gs/hro Then if '...;;.h
of animals is about 10 one ;.:a-' get a rough figure for herbivore froductioon

With theso rmthods s~mi herbivore production estimates are given n Table 30
None of the estimates are en tily satisfactory although the order of magnitude
is indicated.

TJ.'.? 30 Ste Estnates of Herbivore Production


Blads Tip loss rmthodla o .i

Snails in emaenc method i.':)

Insect emergence method (.')

Measured Quantities

Pro idtion

~4 An idge/M2 plant suIface
2$5 M plant surface/A spring
So26 M plant growth/i spring/day
(1% blade growth/day)

6% volume increase/Month Viviparus
i 7% volume increase/Month Vivipar=us
:I'-- increase/month Pomacea
1Man: 11.6%/month; 12 gm/Mr biomass

7.8 individuals/229 cm2/day
S002 ngs/individual




Total of Insects and ,;ila

Assumedeafficiency and
.8 cc/gm/hr(method #4)

32 ot. gm/M2 herbivore standing crop

Io Carnivore Production iY.-..

In the fishery b:,1.:Yr study described below a few recaptures gi.v
some minimal estimates of growth rates of fishes. The average? growth rate
is about 25%/yr. If tho :,;, .'rinj biomass is about 7.3 gns/ia fish the
production is around 2 .: :, Lr o This figure does not include the
main stumpknocker populations Satisfactory biomass estimates of these
fish have not been corplts. ,:, These estimates based on large tagged fish
are probably much too low, If 7.3 gas/Me fish had a metabolic rate
of about .0 cc/gm/hr and xa i: .ricirncy of 10% the Rish growth rate would
be .6 g~m/M /yr. Considei.-- :.- ,ore work is r,;qi:red on the higher t.:,":.c
levels to establish the o,.,.-', ,::aa production and efficiency

l y 03

Jo Pyramid of t '..:'.- :, 'r:ct. Measurement of Bacterial Metabolism

Even though production rates of all trophic levels are not yet
satisfactorily determined, i i s instructive to calculate metabolic rates
by trophic levels using some literature values of metabolic rate :.- v-1;i:7.!.6
by estimates of standing cropo

Primary production '- .*os are taken from previous reports. The ma abolism
of the herbivorea is taken as 08 cc/gm/hr with a biomass of 32.4 cws/a o
Carnivores are taken as oO7 co/,;'..r with 7o3 ms/2 biomass. Top carnivores
are estimated from e04 co/ .: with 4oO gns/ biomasse Rough metabolism
figures come .TCav Psaeto"
Direct estimates of the baetC' s.:.: n ; .^i;blitbi in the algal gyttja that
nakes up the comnnuni' botto.-a werQ made with small bell jars placed on the
mud surface after the top zono .which contained algae was skimmed offo
As with larger -. ..11.. -..nts .. ri i-d in previous reports, oxygen
analyses were made bforse : d af tor -y. Ain.rt'l. periods under black cloth.
Three replication C"`- oxyg no decreases in 95 minutes of I1, o75$ and lo84
mg/i0 with bell jari.. .'' cm in.. .etar and a capacity of 1800 cco
This turns out to be 775 ., This estimate does not include the
considerable bacterial flora t -?:s o"'.;*o.'bo, which was measured in p.r-....'c.!.:.ly
reported work, It is "' ',an:t to note that .' l;:a.iri much smaller in biomass the
decomposer bacteria are oo -..o me4. h'iWiAUy than the regular herbivoreso
The various r:- a sti mats of rme..e'1:I. by cbroplioc level are suanarized
in Figure 1, b lo:. ia t"s a first o ;'.mp-; to assign values to the
metabolism cdit u' .'o *v Eszntsd -
: .-rxfl)

Figure 3J
tPowr' c-rty aolc.lc aitt.noa
toward a tabolic diagram
NCai yro

matter lost-

t? 6"?i",~

Ko Diurnal pH Curv y : .-ir~'.ornt and Dye Nomogram from Carbon Diodxda

In Figure 12, l -.-' is ..von another days determination of production
with oxygen and carton-:* S:.' : curves such as have been given in previous
progress reports Thi.r time water samples were brought back to the laboratory
and the pH determined with a ,.:.ln model Go The maximum time of 24 hours
is not so serious in thesa .-:" o-.;::-. ic matter waters as would be the case with
moat natural waters h-:. carbon dioxide values were then converted into
calculated pH values from thE known alkalinity The measured d and curv
calculated with the DyS naorogriya are both shown below:


duplicate bottles

'0 12 1" 2 3 5 6 7 89 10 11212 3 4 5 6
; y Y*.'.:0 a.113 1 2 3 4 $ 6 7 8 9 10 2 11 2 3 45 6
no noon p.m midnight


CO 9
ppm 8

6 /

saattered cumulus
direct sun
s*" 7. ,,,.. \ 1 O noylight

7 7 )( measured
A calculated from
O 706 s CO2 =&ans of above

Figure 12o !:'. .'.L. Curves for 3/4 mile station May 23-24, 1954


In the summer of *1. H.T- A, and EP-n.' ,s*,iv. of Georgia) made a
In the aummor of "1-1'.-"H, T l"voi and E.P.
study of the productive .I of a windward reef on Eniwetok Atoll primarily
under s;-'" !.C.o-; '": of the Atomic .'-:ir Comission and the University of Georgia
by eman of a otract extension of a pr'-:u directed by EoPe Odum. This
endeavor was .:' .' .." : I' *:.In "" Y. by the ..;.7 and indirectly by the
tecl:.t s used in the Sl .. work.

One main purpose 'was a *.. .-;.', of the characteristics of S;'.vr,
Springs and the J.- :% *-iland T.So The 80 page report on the
Eniwetok study has b.' ( bei .. '.. too: and submitted to the AEC prior to
publication A f.e .7 theo -, '. ion, which are signiibant to the
understa.:L ~ of the "-'.-r ;..- community and steady state systems in
general may be :'..': I v .

1. Both com~aniuioJ s a&eo very :i..r.'1. .', r~. the coral reef :.i :-C~i r .
(of visible "',' r '.' -~:-i. average commauity .ap thi,

2o Both are "T fr ., *' .. coral reef with production rates
above 75.,000 .. .. .- .. .. :.re per .: r, is among the highest on earth
Silver Springs in Moe produ-ctiv in sui er mtoths but has a lower annual

3o Both .,.. are .; atotr. .'' in primary trophic leveo
(John Te:,:1., H.arvard '.vb is .." on ..a.: -.... ry of a small :-i..' .;,
whose primary I'... --ton o 1ci : not light but alloothonous organic mAttero)
Sargent and Austi.n9 '*.. ..:tude. on -' .:." productivity were hi general
confirmedo Gass ." .don 3 "eoskss n both Silver Springs and .;.i.-.1.. :
yield coatings of rather t.an .''" ,; com cities found in inshore
oceaniO waterso

o0 The .
surprisingly similar
ratio of 1.': t7.' produc~ns a:
maintain optimn a s3aiGrcur-o
'and' in .- ;.i..' .-- -
ith a t^Ji'f.'.-.-1" :L ..... '"

Pyramid on one of 5
(zone of large hEada)
other i. :.2.:.r' .b..

f.ll -. *.. :. 7. .
:^~ M 1 ..- .. _* .

-i., .- nodv:


a- end Silver Springs are
in similar, arrestt situaticm a aim xer
.1 ji- ma,,y be t.~*'y*1o
-.' In caloa-rf;os oubstratas vm estim-ted

*Fisjh crabs, cones, annelids

it;- --
;' tjiJ'l" :.Ln "~un :L .jJh;'~7,I? *. i. +-:Y;- in,

U.. u' nivores
of 5 pyrAids
-I ,me *7*~ -- -
_________ L"-;s..-* .1 .

on~g 1><.

*. hi v.Pse I. :- caic~
ftpsland~ .~ PFi gwes In dry go

SBoth coam:initi. ; ':; a' close to a stUJr state with production balanced
by respiration in the coral.! within the limits of accuracy of the measuraments
madoo In Silver Springs there is always a contribution of the upstream
organic matter to the do1us 1 c:m coamrnities which the 7. .:c does not have
(except to a v..': small xtont) Curves 'ho.in, production and respiration
are given in Fig ure3, o0 .o : i:'. preYious data of Sargent and Austin


t 6 a,.m.

E.Po Odum

- *~,,


6 pmor

*' -' '. :ial C.r-. of Production I.e5ai'c-.d with tVh
. a".:: ., :,' ::ahsod on Eniwetok Windward P. 7

60 r .- *
state balance for' 7-
systems tend to .':
he7ore tati suggs!
theoretical ''

: ":-' :,o to have been at least r-;mnhl!y in '..r
~: 10.Q.. timne and it is important that coutr. .. ,
vo O;h. ..to taophic structure and characteristic. a in
:\27:,r basic laws of behavior as discusesd in tho

To It is "' that to .r-.'. eafficiencles aire
achieved by a v. .nuint "-7 in K MV:C... 0 .. and by a vary low
nutrient condition n tIh: cornl ."-- 'o '.,n if the -. ,:, -' .: .,. "E', orn..ina
are available the par.' ."... .. Y.. environmental I. .,'.. .t
factors may ba circnM-., td "' a community by -.:o. :'n som part of its
energy in mechanisma '- co oar-' th .: ...e .. : ', -.. so that it is no
?i-;r MI.':..;". -* -. tha it mainta"ns a r1s energetic tax on the
overall i-; t'.:.-,., In t' : case this i$ the conservation of nutrients
in the oalicrou .' -

:o.'L ... ..-,
per :, li.'

aa (


: A ,



by H. To Odum

In 1"'' X.-,. were made of the distribution of plants in
40 sorinfs. One of tho' e, VanninG Sprli.:r was j~:pcf;jced in the
first process report.

In the foT.':.'.. tr by Dr. NEtelson, entirely inde-
pendent maps are repori:od, ;'.'.: 3 years later in 6 of the sane
springs without having soen but one of the previous ;,i.r.o The
comparison of the earlier maos with the later maps is very
indicative of the de' of stability in these chemostatlcally
and thermostaticaly .regulated natural communities. It is
apparent V.'...t :la gaircl s.me e ecies dominate after three
years alth.'.. h the e..act positions of the various patches -.-.1 ,,
It should o.. ...' *ered !r:nom the work in Lilver that the higher
plants have a r between one and 10 times a year so that
there has been ample time for marked changeSo This is further
evidence along with the pr'sviously reported work on g .1 : and
insects that .. tonss are much more stable than usually found
in nature. This is not to imply that pulses and chlin.. have
not occurred in sone sprsings; n some ..'.- y'le more affected by
surface water, there .i, .io0 a known variation of chemical con-

Standing. CGrop and Community Lurvey of
Submerged Vegetation In Seven Springs

By alle atelson

In the first semi.-annual re a.ct of this project in January,
1953, Dr. John HI Davis presented figures on the standing crops
of four springs and their coastal runs and called attention to
changes in density and comp.-,osition of the vegetation, some of
which were correlated w vth changes in turbidity and chlorinity
of the water, i. Is a report on work intended to continue and
extend the above stu.dic.:, .-e present Investioation, which l.y.e.n
in leptL.ic 19'.7, has. for its subject both the quantitative
features of the standing crop of large submer .:ed aquatic plants in
some spr.: and their : uns and the qualitative composition of
their prevalent coryrmm unit.e.

An invent -tion of the submerged vegetation in Wisconsin
lakes revealed no discrete recurr.r, communities, Instead, there
occeMd a pattern of continual c- .. e of community composition
alone a gradient complex .' environmental factors (Natelson, Do,
The phytosociot.., of c:bme -.d aquatic macroohytes in Wisconsin
lakes. Ph.oD Ths"ol.s, Univ;ersity of Wisconsin, 1954). One of the
-principal aims .. the oresent study is to determine if such a
situation exitts in the aquatic vegetation of Florida, and if so,
wh4t is the pattern vegetation here. A knowle .3 of the pattern,
referred to as "veg.eta.tlcona continuum", can be used in constructing
a classification ;-:te, for the communities and facilitates correl-
ations among ve.oetation and environmental factors. i'he Florida
studies here reported are based .i' ~ methods used in the Wisconsin

Each run was 4 sa ipled at several stations scattered al.;!,
the length of the river. Lome subjectivity was used inasmuch as
care was taken to ... characteristic, rather than atypical or
disturbed areas, but selection of stations was otherwise objective,
with two exceptions: 1) in some instances, for comparison pur-
poses, an attempt v.as made to sample at the same station used by
Dr. Davis or Mr. Sc.an, (bloans, Wm., The dstribution of ..--tic
insects in two Florida Springs, M.oB Thesis, Univo of Fla,, 1954);
2) stations at Salt ', s run were located re ulo.rly at two-
mile intervals since the water was too turbid for reliable selec-

The s..i l tranucts made across the river frequently
traversed two or more o:-.. ously different communities and such
stations were divided into substations. Each of the latter were
sampled individually, so that the data could be used for generall
community analysis as well as for st':din,; crop and species
composition estimates for each river. For the latter purpose,
the several substations at each station were weighted according
to the area of the station which th, ... occupied, and they were
then combined. This teechique was also used for pools where a
mosaic of communities occurred

Substations or stations with homogeneous vegetation will
be referred to as stands. Each stand was sampled by 5 to 25
quadrats, according to the homogeneity of the vegetation and/or
the area of the stand, The quadrat was a heavy wire frame one
square foot in area. With the aid of a face oi mask, the rooted
plants within the quadr.ate were uprooted and brought to the sur-
face. But sometimes, where the depth and substrate were suitable,
a rake was used to den:de one square foot of the bottom (estimated),
instead of uprooting 1 -I.o

To determine the volume of the plants, the displac;er.'.+Lt
of water by the plants removed from the quadrats was measured
using the method described by Dr. Davis (cited above), and an
average volume per squ.iure foot was calculated for each stand,
Since the specific l r ".ity of submerged plants is close to one,
these p figures were u. -:. as an estimate of the wet weight of the
plants. The percent'.'.: cover of the vegetation in each section of
river was multiplied by the weight of the vegetation+ of the stand
sampled in that section, and the results were converd to lbSo/acre
wet weight for each section,, Wet weight of the vegetation in
each section was ;'..:hted by the estimated relative area of the
section in the river and the resulting figures were combined to
produce an estimate of the average wet weight/acre in the river.

'he percentag- of volume contributed by each species was
estimated for each quadrat, and an average was obtained for each
stand. These figures were then weighted by the percentage cover
and percentage area of each stand in the river, in the same manner
as described above .. the combined wet weights. Thus the per-
centage of the wet '-...:" t of plants in the river contributed by
each species was obtairedo The percentage of water content of
each species as previously determined by Dr. Davis unpublished
data) was used to oU:'. the dry weight contributed by each species
to the average dry r .' ;ht/acre for the river

Estimates of total s' .'n'fig crops, both as wet and dry
weight, and the percent., ". .C the total dry weight which was con-
tributed by each species are presented in Table 1.

Figures 1 and 2 illustrate the spring-river systems whose
standing crops were a..i..-.:.:'-Ied in Table 1, and the locations of
stations and the area estimated as representative of each station
are shown.

Across a river, shore to shore,' different communities
often occur within small areas, even at the same depth. Frequently,
an environmental correl..tion is obvious, e.g,, different substrates.
However, in other situ.t.io.ns no reason for the differences is
apparent and it is prol. '... that historical factors such as dis-
turbance, availability of propagules, and conditions conducive
to clone formation were largely responsible for the non-uniformity
of the vegetation.

In contrast, a regular trend in vegetation change downstream
occurs in some rivers, superimposed upon the more random localized
variability. In this investigation, such trends were noted in
the springs with coastal runs. In Weekiwachee River, Chara occurred
near the head, often in great density, but was not found in the
middle or lower parts of the river, Ceratophyllum demersum
likewise occurred in greatest abundance near the head, but extended
much further downstream than @har. As GharA decreased in impor-
tance, Ngjgs ~eadalusenis which was absent from the upper part
of the river, appeared and rapidly became the most abundant species,
Sagittaria was more prevalent in the upper part while Potamogetn
peotinatus and Vallisneria neotropicalis were apparently restricted
to the lower part of the river,

Chassahowitzka River was similar to Weekiwachee River in
some of its vegetational features. Sagittaria was most abundant
hear the head while Vallianeria neotropicalis, aae guadalugensi s,
and Potamogeton gecSinatus reached their maxima in the lower part of
the river.
Such trends were not apparent in Homosassa River, except for
the occurrence of large amounts of filamentous green algae in the
middle section of the river although it was rare in the upper and
lower regions. However, a distinct change in the character of the
vegetation occurred near the Gulf where tidal waters introduce
salinity. There Po~fSo3Sgon pectinatus and algae of marine type
occurred, while the common upstream species were rare.
These three rivers run from their head springs to the Gulf
and thus contain a gradation in chlorinity, as was shown by data
presented by Dr. H. T. Odum in the January, 1953 report of this
project. Tables 2 to 5 show the qualitative changes in vegetation
which occur in some instances from the head of a river to its mouth,
and also show how equally great or even greater variation often
occurs among substations at the s.me general location. Thus it
seems that excluding brackish waters, changes in chlorinity are
probably not as much the cause of community differences as are the
changes in substrate and turbidity.
Changes in community composition in the Salt Springs run,
which does not flow into salt water but into Lake George, showed
no consistent trends, except for the fact that Potamogeton pectin-
atu was the most abundant species in the pool area and the beginning
of the run, and was absent or rare elsewhere.
Hart Springs run had essentially the same plant composition
throughout its short length.
Sources of error in this work fall into two principal
categories. The first results from the mosaic arrangement of
the communities which is revealed by Tables 2 to 5. Because of
this, the error can be considerable when the standing crop or

species composition estimate for a river is derived from a fairly
small number of stations. The second source of error arises from
the necessity for estimating the plant cover of each section of
the river and the extent of the section represented by each stand.

In addition, there are small errors which result from
difficulties in sampling, i.e., the current effects which often
bend the vegetation horizontally and do not permit the quadrat
to be dropped over the top of the plants so that one square foot
of bottom can be denuded. However, these errors may often com-
pensate for e ch other. For example, the excess weight contri-
buted by soil particles, which usually come up with the roots,
is to some extent compensated for by the fact that frequently a
large part of the root system is left in the substrate.
An idea of the relatively small size of error or variability
in results obtained is shown by a comparison between Dr. Davis'
results (cited above) and those of this investigation.

Lbs./acre Lbs,/acre
Dry Wt.* Dry Wt Error
Weekiwachee Springs and River 3941 4686

Chassahowitzka Sprin:s and River 4620 3667 26%

Homosassa Sprines and River 4000 3774
Average error =16
*From Dr. Davis' data.
#From the present work, converted from wet weights by
conversion factors,

rhis comparison of results of work done entirely independently
by two persons leads to the conclusion that standing crop estimates
are relatively good first approximations of the productivity of
spring-river ecosystems. However, it should be emphasized that
it has not been possible to determine from this study what part
of the differences between the two estimates fS the same river is
attributable to chance errors in sampling and which to possible
actual differences in the vegetation when the estimates were made.
the pools of some Of the spring boils were mapped and are
shown as diagrams in Figures 3 to 7. The areas of similar veg-
etation (stands) in each were delineated and the communities analyzed
in a manner similar to the substations of the runs. Each species
is represented by a symbol whose frequency in each stand on the
map indicates its relative importance in the community as determined
by frequency calculations and volume measurements or estimates.

These maps i.,. .i ,,e ,J .d veral different comiauniti..es
often occur withinn a r,. 11 ;rea. Work on communities will con-
time fth. the epri, 1:'55 probably will shed .li.h~ upon
some of the cormmuni.:ty a .::il ite relationships,

1) !'. aP !S mentio.od .: r
their tax -. aut .
f. ,- A.i"r* .

n ;V ,';. .. ) A.D Hito
fl-v 1.'1- 4 qI Marie-ViC

report are listed below with.


2) -.. loc tic. A-,: riotions of the spr&D s.3 and ther rive
discussed here r. ,. n F.. .. --on, G.E., C. W. Linuch.am,, F.
Love, and R 0.O V ....:, p.ngs of Floril- Geoloical Bu.lleln
No. 31, State of -.. cart ment .' o Cons0-ca:.ctlon, Tallahnorea,:

Table 1. Esti ..- 1 '::'.. c rop and percent .:. con-
tribute d. b: v, : eceis in 5 springs and their rv-.: .


Wet wt. lb!4/acr:>,

Dry wt. lbs.J/areb

4 D

''f" :Y flfl




H ono.








H rt


Percent- > thr tot:.

Algae (fiilamentous)

Ano ch n. is carnSre u

Cabomba op.

Cerator'Illui r,, .zA,

Chara : ...

Hydrocotylo -, ,

Ludwigia n atia;a

Naji. n e ..-.. is
Pot ,_r cton illi. oi n)o '

P, ... ,"'i" natus t

Rorippa s4-s5.fo ra

Sagittaria Oppa

Vallisne t


il'- t which was contributed by each



7' 33











16.5 o0. 1


66.4 ;2




9 1



22 4



6 .1






Table 2. H-..... .'a .i-i.v: Composition and standing crop of
... action n at t .he :,*. station community level.*


1 B i B2 B3

S .'10 .

C B1 D2


Sc i ttaria s
VvI11s.. -ria rrot-ic.: ... 3Y

lbs /acre wet UI.:tl b W

z 20

S 18

53 19
48 A'.


67 65

42 3 21 33 19 .1

"' .5 10 717 604 6'. C4 95 20

* Communte iit n:..n :
are subte tat:on t
* Imaportanc- e V. .l.V. 1
quency and V
# Al' f moY iwe i

17. the same letter .::.. diff:ery
no. ':;a Stiow
1 ; "- Vx .. d. i -l
1.: o 4 as .* &.u*. d by dis" .

r ':; eripts

117 C.i fre-
cut of water

Tc.l.n 3.

veg t ...a

'f:1'er 0*...;:,-: 'ition and standing; ::.Co of
" ...u. t.ton community levels


A ?_ ... o f

,i "S l "; '...
N, g- .'; '" 1 7 ,
Pa. 11100t'u V1
P. pectinatusa
Rorippa aesa i.lo-.ra
Saglttaria sp,
Vo neotrop an -l., :

lbs acree wet y, .

A B C G2 D 1 D2
' 7 10 1
S8 3
.: 15 1
6 15
37 22

El E2
10 "'

; 100 ", 41 76 59 54 9
;' 13 10

. 1 ,.'i 210 340 330 204 120 320 952 150

.-1 G2

52 82

Weekiwache. n .v:r Composition and standinR crop of
v-,:_ .., tion at ti.e substation community level.

Ceratophyllum .2"sum
Uhara sp.
Ludwigia natana
Lajas guadalupenasi
Potamogeton pectinatus
Sagittaria sp.
Vallisneria neotroplcalis
Lbs./acre wet wei'ht


24 49

100 59


20 37

675 320 180 440 180

1 180

Table 5.

Salt vor: C opposition .An.-
at the subat.ation community

1 standing crop

of 'c.L station


Ceratophyllum demersum
Ghara sp.
Najas ar: .t:, i'ensil
Pot.ear:, [,eton y.v. tinatus
Vallisneria neotropical s

Lbs /acre wet weight

41 A2 A3 B
4 11 4 16
24 60 23 49
11 49 2
15 71 24 16

C Dl D2 El E2
21 41 2 4
79 51 82 19 41

14 76 59

A0 200 300 828 193

Table 4.

8 83 230 40

Figure 1. Locations of stations and section of
river represented by each station.
1 inch 1 mile
c station



Giulf I




of river
A1 25
Al 2.5
A2 7o Substations in Weesiwac-e
A3 8-6 River are estimations for
B 3o7 sections of the river which
B1 25-0 were obviously not represented
e 18o8 by the sampled stations.
D 6,2
E 16.2
F 2.5

A,B,G C m

sa 7% C. P57 -33%3%
oj river

L o0c atis fsa ?'o sTations

StatoC-S are in reJ.

Hart Sprias

So Feet

o-nj, small
scatTered 4j1l a ats
on bo0tte-m

) This side o
i\ has:;
t57. Nasit evedalumtn u
S Ceraltepogilw

s57. Char
5%1 4se- e
Scirpvs conryoidCe f0

Pflar -o-mmynit recordC- i-n TblCe .
A-rea 'asp ed is ou'iinae in red.

5alt Springs d oe trn
I ch a Imile

Fi 4re 2.



\c Feo 'To

31V .ifS V
N~ D R"''-s /.- --"


Nephalr Sa, H ~droof /
I ,' .:
Lem je- tl. ?ede a c I
C~ ~~~~~~i &'< .'* e-*s^^t!'v "*
^ ** i ., ; ^'
^em*-.* s!'1 c.~aa~a ~iyis~a.^;

i -

if, "~:



I -' ...... ---:f .
-. f i- ^ .'- /


-if-u-~c )



,;\ .. s" --,,, ^-,.,, \..

i''-.,.< N",F \ "
-' N .-

-'y .: ^;i i^ \- v

-|if|% 3


C~Lvd;3aa vnta-hs

A:_O'fp SB~ij~


: I II -

i.\ \

e 5 11 f we 'a

cot yle sp.

L aaw dt a r$ s T

Fdia -nTi ll s sp.


NadJs dv uada fvpen rsi

MA rl '-i-llum
1 '

Bethkir Area

*" .) o-, ^ A **.
*i c '''^ ^ e: ^

o, :o
so 0e4t

/ 1

0 e
/ .3 s

*o:;0 tha~tuckte Headspri^Q Poo!

);, ~SeTevmber 21 '5

'SO -e t.
Afc-io~f't o^ea o-ac ? S~aty r
fcse y o _iTeatv T f jen h

Figure 5

Gr a Creek oof

at. Chassahowi tz ka Spr s

October 17, ?S1t

so Feet

iva Njas udlpns

L~q~iri&r Sp*

2Aksln!2 eer-i -ctr,? i ca

A a3 e

pIvi CC~'toph/Ilaw dCperustis

2ikn;cheli~a paItstris

a -
N- -' d?1
-Ar'- .. .1
-t .--,'g ~
5' -)


- Chassa


40.^ I .f<
u^^ Bwt ---Y -- ----- --~
Goat fat
d Revials

F% ure b

fleaJsprno' ok Ctassahowdzka
P4o ata-n eIanoensi5

o~t CeratoehlIntl c~ erxPsuvaj


/)To c Ier f 16 t

2'o Alqte
FIcan l i sa of alg ae w-th
ius saa'TTaTia Pistia s raeles aind Eichanor-a cvtssipes
NadJas 5uabeIElanisl

p-N 'I ';

r~; ----
e't~3 .0? i'e -0
--. -'i~ "-- -.4r
-- ''r- JI I) beGfl

: .Ci-, 't- i
'SJ ri-" t~~, "
1 9 r (C
-t on r :-

; '4: ~ I;`


S.. I ^n.

-.-.---4-.- -- -j -

P --
0 -'


j \ ^ ^, -- -'"-l~li^ '/^

V -

5,pT tmer 3a 0 19S1'

*;, Po ost os eaon- ,: ;.'ates

V a l SW b V i0 i e'l 0 S
t<.a' sjas ,~ ad p si

pIus a l a

C50 F-e
I^_r__--P9^L~II-- \

J y


By David K. r ''' .i'j -.ck H, Berry, and Howard T. Odum

A fish '. ': program in Silver Springs was begu n on '.i, ch 12,
1954, and has been continued by various workersto date. A total of
19 trips have been made Jwith the assistance of a number of volunteer
helpers. To date, I?.'. fish have been '.,.:-nO in Silver Springs. Two
types of tags have been used. Plastic Peterson disc tags of varying
colors and sizes have been used on the larger fish, and small metal
clamp tags have been placed on the jaw or opercle of smaller specimens.
The latter are .: I 1 while the unnumbered plastic tags have been
used in various color combinations for each fish, or have been used
cut to different shapes w;.ith the same color combinations. Most of
the fish 1.- have been Largemouth black bass ([J-ot-,i- salmoides)
and Stumpknockers (T puinctatusy, with some other ..;-;;. i.d-.:.lds
and a few other nuibbers of each species -'. .-:d and the
type of tag are summarized as T. :.. 1. As summarized in T ,b)., 2, ten
fish have been reca:ptired. although enough time elapsed between
tagging and recapture for soEe growth to occur, some fish have not
shown any increase, and others show an apparent decrease, Whether
these growth values a-re ty 1 or whether an artifact or injury
is involved is not '.clear. bass show the most growth, and
all of these specimens are jui'.enil.es. It is interesting to note
that all recaptured fish were retaken where they were initially
caught and t. .'

p,-- r .. 7 .. Stulpknockers (the dominant r',-...:p ) have
been taken each month s : 'in "i.,- : and in '=.:-:.ah, Length-
frequency curves and lengit-weight curves have been constructed
with these. The length-eight ratio does not vary materially from
month to ... :t at least for the sizes measured. A typical r.'1.. -
length curve is shown as 'e 1. Enough specimens of other
species have not beae taken :. the construction of such curves,
Monthly length.-: ,y gr 1.. for the Stumpknocker do not show
an distinct age ,; .. ('- .P 2). A -::. long breeding period
is thus indicated. '-. -' is also evidence to .;',r:, this from
observations of :..-. .' .beds and r,.. (or nearly so) adults dur:.
most of the warm months. 1'0 I an entire winter period has not
yet been .: !'i .", evidence indicates that spawning, although rare
during this period in Silver Springs, does occur. Individuals with
developed gonads v:ere taken on October 15 and on ':...:1.r 15. Also,
small specimens were taken ".;& : the winter (Figure 2).

Samples of scales have been taken throughout the study and an
effort is now being made to determine if these can be used in
determining .,. and rate of ...:-. This study is primarily being
done on the St-;-.-- -, K. some attempt will be made to study
the scales of the other Centrarchids, particularly the bass.
Preliminary studies show the presence of r;w-.i., but fuj ;h::'.! study is


necessary to determine if these rings represent true annuli. Scales
from this constant t'...:-, ,i:..; spring will be compared with scales
from the same species from other (non constant temperature) Florida
waters, and if pon.. i.?~l with scales from northern waters.

A straight line ratio has been shon to exist between standard
length and total 1..;! :..1 the Stumpknocker. This ratio (S.L./TL.*.79)
exists throughout the entire size range for this species as encountered
in Silver E ::..i.;s, and will be helpful in comparing the work done in
Silver Springs with that of other workers on the same species in other

Coincident with the fishery work on larger fishes, a general
collection of small fr' :, invertebrates, and algae has been made
for further study of seasonal periodicity of reproduction in this
constant temperature environment.


Lepomis j. punctatus 83 147 230

Lepomis macrochirus 12 12 24

Legomis auritus 3 0 3

Lepomis marginatus 1 0 1

Lepomis megalois 25 9 34

Lepomis microlophus 1 1 2

Micropterus salmoides 40 19 59

Chaenobryttus coronarius 11 17 28

Esox americanus 1 1 2

Pomoxis niromaculatus 0 1 1

Leiosteus platyrhincus 0 3 3

Anguilla rostrata 0 1 1

Erigzon sucetta 0 1 1

Ameiurus natalis O 3 3

Grand totals 177 215 392

Table 1. Summary of fish tagged at Silver Springs, Florida, between
March 12, and December 29, 1954.


Micropterus salmoides

Micropterus salmoides

Miropterus salmoides

.icropterus it, c.l.:

_.r---. to salmoides

Chaenobz ttus coronarius
w:w-: _:- .. ..

Lenomis n _nr___*ct_
T... I .- "L i .; '



























X I -54















St. Len. (mm)
When tagged











Table 2. Recaptures of tagged fish at Silver Springs, Florida.

St. Len.
at Eecap.





































10 20 20 40 50 60 70 80 90 100 110

Standard Length (mm.)

Figure 1. Length-weight, Lepomis gnctatus punctatus, Silver
Springs, Florida, October 15, 1954.

120 130







Figure 2. Length-frequency graphs for bE mis aunctatus, Silver


Springs, Florida.

I \

, / / I I
I' 1
*ft i


1954 -

j? D
.-.. 4
ft 8, '


,- ,' JUNE 1954
/ a --. '" ..


0 '4


N /\
0 %"

10- --
0 - ---- ---*---- -.1- <-- --

10- / SEPTEMBER 1954 -
5- \ ... -*
.- o--^ e------.a -

10- OCTOBER 1954 -

--" ^ x-^---^-- ----- :-`-`---

10.- /- NOVEMBER 1954
5-A $ DEE\ R/ 19541

__--a-B ~t>~~I-----D--ECE-MBER- ~ 1954
"-' DECEMBER 1954
'i 0-__

I I I I I I I I I I I I I I I I I I I it I I I Ir- r-I I I
", A r-1 r A 0- A -- -4 H- H -H r-H H r-1 r-4

' 25-








/ N 0 /


q .



- -~-- ------ -~----CIII-~-*IUII-~ --L---ZIWIII-I.I.IC~LI .-LlltlWIIIII-YI- I__

HTT.rviTV T55o-

HoT,:, 0''."

A., p .'iTi D tlu t.. :,.. --.. .'. ", ;.r Principle Applied to itasyNiTh.;*,!L

SIn --prcr.....s :;.,*., ::i;,. tha principle has been stated that open steady state
systems tend to be adjusted at an optimum but low .[f. cl :.acy that corresponds
to the maximum power -....;.. (" ., a.nd Pinkertonp American Scienti f.., in press)
S That this pr:inY.. is ,.,L .' -*:,: ~A~.vi.. systems seems indicated in
Figure 1$ below:

Snatural or high light intensities
(kW -Ao -.'I photosynthesis

per fir quantum controversy
'.. ;;.!.1 ..;.t ./ -.

low but,.;.. i-, -""e. associated with maximum output

This graph shows that '.,-, ..,,.i. .'. such as have been achieved in
all the work associated P .0, the quantum controvercey have all been r.-'-v.!
light intensities so that the .:.: output of glucose has been .'.n.
On the other hand the .";."': populations of algae d.3ui.:d to h~gi .natural
light intensities run at .., ....t..a.s and h... .light intensities but so
that a much greater :i ,.". of .'-.-.S rrn.s ',-.. If pl nt. are evolutionarily
adapted to maximum output A%"- must sacrifice efficiency for power by
this hypothesis This is a .- of ett ..,': that attempts to increase world
food by raising chlorella at ..,;. .,A"Jci...., must necessarily flopo A
second part of this ..,i;. .."..j may be stated that the optimum efficiency
for maximum power output :.'; -- .. decreases as the light intensity increases
Plants adapted in nature to deep water achieve the optimum adjustment
at a higher effic:-:.m-vr that at the surface "-iv.,v a plant adjusted for
one light intensity can not be moved ir,~,O. *'it.ly to another light intensity
and achieve the ct-n., i', '.... n without internal modification. A car
climbing a hill in .*;:..:'' ,.. *at optimum ..':..irj .nr.cy cannot achieve the
optimum efficiency for a -1,; .t away without -'.n..r Q a-. In the
plant gears may be the concentrate tion gradientso The efficiencies in
Figure 15 are of the same .d a, at optimum adjustment as those
found in Silver Springs and the Coral Raeo?,

Bo Organional Size versus .l .~".: TA.L: in Phototrophe in Optimum Adjustment

It is now well know that amtabolism of heterotrophic organisms is inverse
to body size roughly in a 2/3 o'..- '.... function that is presumed to be related
to the surface/volume 7514:. to V02ll' n processes. Ma the above section
it was concluded that 1 .... .' y, .'; in open steady state tend to
all become adjusted to a similar state of running at maximum power output
because of the survival 4 '''. in : l. ":7*. .":. n'....'..n both in an environmental
and evolutionary sense Xith .r,.- intensitiesA similar steady
state plant ~stfli ..' ...& be ;". .-..-i to similar efficiencies and thus
similar total -.ho.-. .,.:. .. power output of pl-I ....co an onAREA basis

Now if the plants run at the same rate of output per area and if size
effects hold for plants as v:.:.'l as animals, then small plants like 6hlorella
if growing in steady state ~:.I-.'il. achieve the same output per area but with
less biomasso The large climax rain-forest with big plants on the other hand
if adjusted to the sams light and optiImum efficiency will require a much larger
standing crop biomass because of the slower metabolism per pound of tissues

In Figure 16 -~lhcr is sih~ n a graph of photosynthetic rate of plants of
various minimum diameterP under natural light or maximum photosynthetic
adjustments, The V-;-;.. a j. '1.'Ci V :..c'l 3i-,

data from
mum o001l- 0- 3
Photosynthesis a Surnover
Gm glucose
per gm dry o.01 -3
weight per
Hro ,
r 01 *'3 / ""300

1o .,. 3000

,o01.01 .1
Size in Cm

Figure 16o Effect of organismal size on Photosynthesis per weight

It is apparent that there is a size effect over a wide range just as in the
heterotrophso Thus, 0.-i1' ,o; the light intensity, the size of the producer,
and the rough offL'.-icJ-'.?.,! f*-.:.'.. in given environments, one can compute
the steady state b.:'l.,C car.. rying .-p.ci. o

Co Pyramid '. '... nd r':-:; .:. ..'z. Size

If as o.tl ,.d above siall sized producers put out the same production
per area with a . r.1.J, biomass as do large producers with a large biomass
both working at a.L.:.e.~,: offici'rl:,. one can visualize two extreme types
of pyramid as c gap .:i.'.; in C..r 17 (For data showing similar
efficiencies and o.'..;.:..i ?.:, mass Shlorella culture and grass plots
see (Burlews Mass Culture of 1~.:...b --.[.t-e ---Wi.e.,r-b..,k et. aloCarnegie@ 1953)
In one the size of the organism decreases as one goes up the food chain as in
grass-grasshopper--o.idA lr.'o In the other the size of the organism increases
as one goes up the food chan as in .L.0i' ulla, paramecium and fish, If the
same energy passes up tbr',^ .':cr:t food chains with the same 10% efficiency
for the higher trophic .?;:'- l, two ~I tir'l different shaped pyramids of
.at~Judy state biomass result because of the different rates of turnover
Some metabolic rate fV. i;::-.. are used to compute Figure I? from H.il';nn s text
Photosynthatic values are taken .:.i..:.i Figure 16o

The pyramids in Figure 17 I'.>ip to visualize the possible relationship of
the tropical ocean to the tropical rain forest. The small size of tropical
plankters as well as the h.;I -..: .'..r: tend to cas o the reversed
pyramid calculated for Chloralla--paramecium-fisho

Spidr 22 Fish
Grasshopper 62 Paramecium
Cas* '810
810 _.6 Chlorella

Figure 17. 'i *... !,.:;-:. : with the Same Energy Flux sho' rinytr the
,p.,.;..c, of A.., ,. : .,d h '. on ,rE:.niaimsl size.

Do Energy Contributions of T'dr C::. '.:' and Curr~nl-, Plankton Size

The efficiencies of primary production so .?--ir estimated for Silver Springs
(8%) and for the ,;.-i.,';,'.. .-E ', are 'o:;.!? .;"i2 nbi.;- higher than many other
natural communities or 1.:.- V-'..:v experiments ran at high light intensities
for maximum production 2:icj there is a ila.s difference in nutrients
between the roof and Silver Springs it seems that some other property
is in part responsible for this high production. It is reasonable to
postulate that hi, e-:.. efficient ies are produced by the strong currents that
serve as a community cir .. :' y.:d, rnintai,'i)g better nutrient
concentrations a ..1: .t to alls and r: p.ovini waste products

The effect of the circulation might be stated in two ways:
Ao The community receives .-;*'.*; ," :'i both the sun and from the current
Bo The energy directly from the sun goes further because of the current condition.
If the usual efficiency at maximma adjustment is about 2%, about 5% might
be postulated as due to the current system This might imply that these
flow systems dlerive wre of their r ':,vrgy f..'<::- primary production from the
current energy than *:..,n tho 1~-.ht received directly

If current is as important as ipi,-i' above~ a suggestion can be made
as to why larger and heavier : r...rkton usually prevail over ~:-..-.~I'r
species with f?: (cr"%:.'s so that theyy may float with the same density as
water Heavier 7: 'e.,"'t.rs are ,;.i;.i 'i::d by the turbulent eddies so that
they are continually iall.l-:,. through the currents that support them This
mechanism provides the ce ll vth a local current which the organism at
water density would not have.o Th relatively low Chlorella efficiencies
obtained in high light ,nt.. 'i.-.': ;.7 mass cultures in spite of high nutrients
may be accounted &.'."r small size of 'ic:a.'l" which decreases the effectiveness
of stirring mechanismso

BE Decrease of Daytime Plant !epi:ation Accounted for by the iypc '~.;i.is of
Plant Respiratory Sy.'~-.rv' as herbivores of the Autotrophic systemai- Arctic

The data of several ainT.: ,o (ioeo Kok, see Rabinowitch) suggests that
plant respiration in the 2 .~'me is much less than when the light is rii n ; the

compensation point The :;o;,: ;, of pyramids of biomass which may be
supported during - :'..-'- state suggests a reason, Whenever food i'rg,
is passed through an energy '.. ;.::li.l:. :-.s' step a lirg-c percent, perhaps
90, must go into heat as ri.- ..-:'r A, by the second law of th..y--. .:y;ir.ii:': for
these types of reactions under optimu.n officiencyrmaximum power adicij:.;..,:,t.E.;-,

If a plant in the day can drive some of its work systems dir-c.,-,
rather than making glucose first and then b.'niiC the glucose, the
plant can save a step and thus avoid the heat lose from the extra step,
Thus a higher efficiency in the b.,:.'o'"phic system is maintained with
lss respiration, At l1'-.ht, by this view, it becomes necessary for the
plant to fall back on .'. ,,;-. stored thus lengthening the chain of
transformations and :. ,'..'-.".? increased respiration

If true,this becomes : ,-i '. all.y significant when length of day and night
is considered, for a .-.;,: .:.'.? 'ence in day Ien, means increased efficiency
as well as additional .i.,:1'., i!l; .- is less glucose that must be stored for
use at the inefficient ,'*-i-,. rate, By this view Arctic plants during
continuous cla5.,'<. ,, of suaenr should be much more efficient than similar
temperate pla'r: On an aual basis however there would be no gain for
the community would have to store ::..:-i.;y months worth of organic matter
at the 'inefficient ::..(. rate

Fo Anaerobic .i,:,- o .. nd tbh -:. .;r-., ?::. t,-.I

The saprobs system of classifying pollution communities assumes
the association of characteristic indicator species for different degrees
of sewage type pollution A c:.;:-'i4o.t.!t,-n of Florida anaerobic springs
shows that the e'_.' while ": r'1 when properly used has a fallacy that
leads to misl:,'..u.,; cc,.'-. .:..J when :.ei D- on waters in general

Beecher, Orange, Warm 2 11.., and Volusia Co. Blue ?pr)m: are all examples
of large springs with low or: .: content water that is also low in oxygen
The communities that result are both anaerobic and autotrophic in nature
In contrast sewafg communities are anaerobic and heterotrophico In the
springs one gets sulfur bacteria and blue green algae but no ciliateso
In short one gets some of tho saproba system biota in water that is
the extreme opp.s'!it;1 of *.. .:.- polluted water, Thoughtless use of the
saprobe system -:.i to a : .:,*' :.-' misclassification of the
type of primary production in these cases.

Go Some Definitions, S, ~;-! methodology

In agriculture, mans r:.i h..-di labor and supervision guise a complex
community in a direction he desires o However in Ecological Engineering
the outcome of production of a complex community is achieved by proper
selection of components at the start with subsequent hands off thus
permitting the n:.:_nity to reach a unique steady state adjustment.

In the usual scientific experiment, man controls one variable so as to study
the behavior of another dir:.d-irjt variable while holding other conditions constant.
Thus the process is one of analyzing component processes In Microcosm
experimentation on the other hand components are put together and the complex
allowed to make its own trends under observation

The study of ecological ,. :. I: .-I.ng by microcosm *;r,.-:rr.,.ian ,.To,: is
a practical met.ho:_:..1o,-,r for h.-..., the ";[:.l 0.,'i ,, .--.. ,i .l .--,.:.t.',.1.
This ;.'.. n i'.. ; .*..*. ". is ma : .., % ..: of the icrocoom :,' .:.-.,

A hypothesis rei;.eirding dependence of community
structure and density on productivity

by J. L. Yount

The hypothesis is offered that under similar general conditions
the species variety is an inverse function of the community pro-

The salps of a series of plankton samples made by Pacific
Oceanic Fishery Investigations of the United States Fish and
Wildlife Service in epipelagic waters of the central Pacific
Ocean were studied dur:i., 1952-19549 Observations on them led
to the formulation of the hypothesis presented below. The most
pertinent observations were as follows. In most of the samples
studied, many species of salps were taken with little predom-
inance of any one species. In one sample, however, there were
both a far greater total salp quantity and a great predominance
of one species of salp, only a few others being taken and these
in insignificant quantities. All salp species apparently simul-
taneously' occupy L~Ial.l!.r niches (the concept of the niche used
here is that of Elton, 1927, Animal Ecology: 63-4), and appar-
ently also are subject to the same environmental conditions, thus
apparently are ecological equivalents (in impoverished waters;
see below).

Observations ii':r: by other investigators are also pertinent
here. Students of marine plankton of hiLn latitudes have des-
cribed it as "monotonous", consisting predominantly of one species
of organism in each niche .-ppr~arently, although the term niche has
not been applied in these descriptions. Mo-t descriptions of the
plankton of low latitudes, however, emphasize the great variety
of species with little or no predominance by any one species (per
niche) (see Steuer, 1910, Planktonkunde: 601-4; Russell and Yonge,
1936, The c71 ; 123-6; Dakin and Colefax, 1940, The Plankton of
the Australian Coastal Waters off i'ew South Wales, UnvJ _Sdney,
Dept2 ZoolJg Publ. I: 27-34). Another pertinent observation
discussed by Steuer and Dakin is that productivity in the tropics
in waters influenced by land drainage and in regions of upwelling
may equal or even exceed that of high latitudes.

If these observations are considered together, it appears
that in epipelagic waters with relatively great quantities of
nutrient chemicals (the enriched areas), production of the plank-
ton is great in quantity but trends toward few species of organisms--
probably only one dominant species oer niche--and in epipelagic
waters with relatively small quantities of nutrient chemicals (the
impoverished areas), the plankton is small in quantity and trends
toward many species :.' organisms--apparently many species per

A hypothesis r;,.arding dependence of community
structure and density on productivity

by J. L Yount

The hypothesis is (-'..:.red that under similar general conditions
the species variety is an inverse function of the community pro-

The salps of a series of plankton samples made by Pacific
Oceanic Fishery Investigations of the United States Fisi and
Wildlife Su-.-'ice in epipelagli waters of the central Pacific
Ocean were studied during 1952-1.951., Observations on them led
to the formulation c"., the hypothesis presented below, The most
pertinent observations were as follows,. In most of the samples
studied, many species of salps were taken with little predom-
inance of any one species. In one sample, however, there were
both a far greater total salp quantity and a great predominance
of one species of salp, only a few others being taken and these
in insignificant quantities. All salp species apparently simul-
taneously occupy lmiinlr.r niches (the concept of the niche used
here is that of Elton, 1927, Animal Ecoo: 63-4), and appar-
ently also are subject to the same environmental conditions, thus
apparently are ecological equivalents (in impoverished waters;
see below).

Observations made by other investigators are also pertinent
here. Students of marine plankton of high latitudes have des-
cribed it as "monoonon.i:-", consisting predominantly of one species
of organism in each niche .ppar-r.ly, although the term niche has
not been applied in these descriptions. Most descriptions of the
plankton of low latitudes, however, emphasize the great variety
of species with little or no predominance by any one species (per
niche) (see Steuer, 1910, Planktonkunde: 601-4; Russell and Yonge,
1936, The Seas: 123-6; Dakin and Colefax, 1940, The Plankton of
the Australian Coastal Waters off New South Wales, Unlv Sydney,
Dept. Zoo2001 Pub.l: 27-34). Another pertinent observation
discussed by Steuer and Dakin is that productivity in the tropics
in waters influenced by land drainage and in regions of upwelling
may equal or even exceed that of high latitudes,

If these observations are considered together, it appears
that in epipelagic waters with relatively great quantities of
nutrient chemicals (the enriched areas), production of the plank-
ton is great in quantity but trends toward few species of organisms--
probably only one dominant species oer niche--and in epipelagic
waters with relatively !':,jl.l quantities of nutrient chemicals (the
impoverished areas), the plankton is small in quantity and trends
toward many species of organisms--apparently many species per

If the plankton now be compared to sessile organisms of the
Aufwuchs and benthos, it will be seen that the two groups have
similar natures, in that sessile organisms and planktero are
moderately passive in ability to capture foo., and to move in
their medium (most sessile organisms, of course, move during
some phase of their life cycle, and plankters have some ability
to move, but directed movement is limited for both). NFor this
reason, at least the plankton and sessile organisms probably
should be nonsideri'. together in the hypothesis.

It is ',vi.,.,.l:' .ted from these observations that if productivity
is low and other factors are constant, species of ecologically
passive ori...,al:n at le -"1, may occur together as ecological
equivalents with little of no predominance of one species In each
niche; and conversely if productivity is high and other factors
are constant, species of these organism should not occur together
as ecological equivalents, but rather one species should dominate
in each niche~ X'hia :,!.s': to a further postulate: if productiv-
ity is low and other factors are constant, competition and other
coactions should be reduced for these organisms; whereas if pro-
ductivity is high and other factors are constant, competition and
other coactions should be Increased for them.

the followli.,. evidences tend to support this hypothesis. In
the tropic epipelr., .c holoplankton, productivity is low and quan-
tity is small, except in certain regions mentioned above, and
species numbers are great; in the epiul..'ti.c holoplankton of high
latitudes, product. ity is high .:.nr. quantity is great, but species
numbers are small. Occasionally in tropic waters, swarms of
plankters appear, con.~Mt'_:u, of few species of orgo.icon and a
relatively great quantity--productivity is therefore high and
species numbers few, even in the midst of impoverished waters,
under enriched cond lt:ons In plankton tows from impoverished
waters, many species of u. .in1..l:.ik occur together that apparently
are ecological equivalent, and this is evidently not true of
plankton tows i:' , enrich... waters (evidences for this statement
are based bchefly on studies 0i salpB, but it is postulated as
being true of other pl.ankters : well). In Florida Springs, it
has been noted (H. T. O(.::;, L, A. Whitford, W. C. sloan) that
productivity is high h*-.. the number of species of the various
groups o o organisms is Low,

his byothesls is being tested at present with Aufwuchs
growth under the controlled conditions of the Florida Springs
and, if results warrant, is expected to be tested under other
fresh water and marine conditions. Counts of Aufwuchs species
numbers on slides, relative to current controlled total pro-
ductivity, have been started Counts made on the preliminary
first series are consistent with the hypothesis

U5-7- 1 92 9
46 PF 3/,
If the plankton now be compared to sessile organisms of the /150*f
Aufwuchs and benthos, it will be seen that the two groups have
similar natures, in that sessile organisms and plankters are
moderately passive in ability to capture food and to move in
their medium (most sessile organisms, of course, move during
some phase of their life cycle, and plankters have some ability
to move, but directed mcnc~rtnl is limited for both). f'or this
reason, at leW.:,n the plankton and sessile organisms ,.. .'l-bly
should be considered together in the hypothesis.

It is postulated j't L.I theeose Observations that if productivity
is low and other factors are constant, so:pci.es of ecologically
passive orclr:n..r at least, may occur together as ecological
equivalents with little of no predominance of one species in each
niche; and conversely if productivity is high and other factors
are constant, species of these organisms should not occur together
as ecological equivalents, but rather one species should dominate
in each niche. 'his leads to a further postulate: if productiv-
ity is low and other factors are constant, competition and other
coactions should be reduced for these organisms; whereas if pro-
ductivity is high and other factors are constant, competition and
other coactions should be increased for them.

t'he followvrin evidences tend to support this hypothesis. In
the tropic epipe:.!ic holoplj.nhl; ton, productivity is low and quan-
tity is small, except in certain regions mentioned above, and
species numbers are great; in the epipelagic holoplankton of high
latituderJ, productivity 1j. hi1,h and quantity is great, but species
numbers are small. Occasionally in tropic waters, swarms of
plankters appear, consistin5 of few species of organisms and a
relatively great quantity--productivity is therefore hlgh and
species numbers few, even in the midst of impoverished waters,
under enriched cou.'llUons. In plankton tows from impoverished
waters, many species of .., 0;:if.l",- occur together that apparently
are ecological equivalent;, and this is evidently not true of
plankton tows ,'ru. enriched waters (evidences for this statement
are based chiefly on studies of salps, but it is postulated as
being true of other plankters as well). In Florida Springs, it
has been noted (H. T. Odum, L. A. Whitford, Wo C. sloan) that
productivity is hI.'. and the number of species of the various
groups of organisms is lowo

'his hypothesis is being tested at present with Aufwuchs
growth under the c ,,. tro) c.' condi tions of the Florida Springs
and, if results warrant, is expected to be tested under other
fresh water and marine conditions. Counts of Aufwuchs species
numbers on slides, relative to current controlled total pro-
ductivity, have been started Counts made on the preliminary
first series are consistent with, the hypothesis.

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