Title: Phytoplankton community structure and primary productivity in two Florida lakes
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 Material Information
Title: Phytoplankton community structure and primary productivity in two Florida lakes
Physical Description: vii, 83 l. : illus. ; 28 cm.
Language: English
Creator: Harper, Carol Lynn, 1942-
Publisher: University of Florida
Place of Publication: Gainesville, Fla.
Publication Date: 1971
Copyright Date: 1971
 Subjects
Subject: Phytoplankton -- Florida   ( lcsh )
Zoology thesis Ph. D
Dissertations, Academic -- Zoology -- UF
Genre: bibliography   ( marcgt )
non-fiction   ( marcgt )
 Notes
Bibliography: Bibliography: l. 78-82.
Additional Physical Form: Also available on World Wide Web
General Note: Manuscript copy.
General Note: Thesis--University of Florida, 1971.
General Note: Vita.
 Record Information
Bibliographic ID: UF00097671
Volume ID: VID00001
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: alephbibnum - 000549667
oclc - 13272416
notis - ACX3962

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oplakto CarouityStruct7ure alld Primary Productivity
illT rl~oridalas

























MS3ERTAION PFS`CTNiE TO THE GRADUATEG COUXCT L OF
THE UNIVERSITY l' UY 1RI'D IN PARTIAL
FULPTILDMTNT OF THE: REQUTRE. EN[ FOR '17EC DEGREE OF
DCOr' M 1 PHILOSOPHY








TPJUGU ilyfl intRIW














ACKNOC'.-.EDG E-iEITS

I would like to express my appreciation to my co.-mittce, Dr. Robert

M. DeWitt, Dr. Frank G. Nordlie and Dr. Hugh D. Putnam, for their advice

and guidance in the preparation of this manuscript. Dr. Patrick L.

Brezonik, Dr. William E. S. Carr, Dr. Carmine A. .anciani, and Dr.

E. Lowe Pierce also provided guidance during the research and writing

of chi.; project. Dr. Frank G. iUordlie .as especially helpful with

advice and equipment during the research.

The staff of the Depact-ment of Envirctniental Enginccrig provided

use oc, their facilities at Lake ;lI:Cloud, and I-ic. and Mrs. Jnmes i'ng

allo.-ed the use of their docking facilities on Bi..-en's Arm. Dr. Paul

Sy'.'oct, Dr. Thomas J. Krakauer andi Dr. Louise Racoy assi.std with the

use of thi ,c.incillationu counter. Dis. Paul E. anld iaroilyn R. M-aslin

.aisis'cd..with transportation.

:ly husband, Charles A. Harper, rrov'ided transportation, helped

,'ith collections iand illustrations, and coni:ribu'.ed valuable advice

and encouiragtinent.

I ..:is;1 to thank Lh'n Department of Zoology, the Departlcint of

Enlvirotnl2:\'al Engineoc.in:g, anrid the College of Education as '..:ll as

t'he Graduate School (Ceor .an U.DEA Title IV fellowship) for finr.incial

support ;th.t enabled we to carry out my graduate program.
en.7.?er! ~~ to carry~ out my~

















TABLE OF CONTENTS


Acknowl edzcements ....


,ist of Tables . .


List of ['igues . .


Abstract . . . .


Int-roduction . . .


The Study Area . . .


;-;ethods and materialss


POS Lsu11 L s . . . .


Prinmry Prcoductivity
(hloroplyll . .
Species Composition

Dis'u s sion . . . .


Lak..u McCloud . .
BiVrn'es Ann . .

'Srm ry . . .


nd x . . . .


Li iterat-.L e Cite.d . .


. . . . . . . ii


S 33


. . iv




S. vii





. 4

6



9
. . 9
. . 15
S 27

. . 42
S . 42


4 6

. . 51
- 51

. .. 53

. . 78














LIST OF TABLES

1. Chemical characteristics of the study areas . . . . 5

2. Primary productivity at two depths in Biven's Arm . .. 10

3. Relatibiships between chlorophyll and primary productivity 26

4. Relationship between number of algal units and primary
productivity .. . . . . . . . . . . . 33

5. Relacionship between algal diversity and primary
productivity .. .. .. ... ....... .... . 36

6. R-laLicnship between algal diversity and zooplankton
numbers . . . . . . . . .. ..... . 41

Al. Primary productivity, chlorophyll and algal unit count
data for unfiltered samples, surface waters, in Lake
McCloud and Biven's Arm . . . . . . ....... 54

A2. Primary productivity, chlorophyll and algal unit count
daLa for algae less than 158 yi, surface waters, in
Lake McCloud and Biven's Arm . . . . . . ... 58

A3. Pria..ry productiviLy, chlorophyll and algal unit count
data for algae less than 76 pu, surface waters, in
Lake McCloud and Biveu's Arm . . . . . . ... 61

A4. Priibary productivity, chlorophyll and algal unit count
data for algal less than 70 u, surface waters, in
L=ke !IcCloud and Biven's Arm . . . . . . . . 64

A5. Pr.unmry productivity and chlorophyll data for algae
greateLrc Lhan 8 it, surface waters, in Lake McCloud
and Biven's Ano. .......... ...... ...... .67

A6. Primary productivity and chlorophyll data for algae
grcdter than 5 p, surface waters, in Lake McCloud
end 3iven's Ann ....................... 69

A7. Primary prodict'vity, chlorophyll and total algal unit.count
dalc for unfiltered samples, deep waters, in Lake
;:cClodd and Biven's Ar. . . ... ...... ... .... 71

AS. Primary productivity, chlorophyll and algal unit count
data for algae less than 158 p, deep waters, in Lake
McCloud and Biven's Arm. . . . . . . . . . 72







A9. Primary productivity, chlorophyll and algal unit count
daca for algae less than 76 p, deep waters, in Lake
McCloud and Biven's Ann . . . ... . . . ... . 73

A10. Primary productivity, chlorophyll and algal unit count
data for algae less than 70 xi, deep waters, in Lake
McCloud and Biven's Arm . . . . . . . . ... 74

All. Zooplankton numbers for unfiltered samples, surface
waters, in Lake McCloud and Biven's Ar-m . . . . .. 75

A12. List of genera of phytoplankton found in unfiltered
samples, surface waters, in Lake McCloud and Biven's
Arm . . . . . . . . . . . . . . 77














LIST OF FIGURES


1. Variations in primary productivity for unfiltered fractions
fcom surface waters in Lake McCloud and Biven's Arm,
1970-1971 . . . . . . . ... . . . . . 12

2. Variations in primary productivity for nannoplankton
fractions from surface waters in Lake McCloud and
Biven's Arm, 1970-1971 . . . . . . . . . 14

3. Variations in primary productivity for unfiltered and
nannoplankton fractions from 1.5 m in Lake McCloud,
1970-1971 .... . . . . . . . . . . . 17

4. Variations in the ratio of primary productivity of
nannoplankton fractions to primary productivity of
unfiltered fractions from surface waters in Lake
McCloud and Biven's Arm, 1970-1971 . . . . . ... 19

5. Variations in the ratio of primary productivity of
nannoplankton fractions to primary productivity of
unfiltered fractions from surface waters and 1.5 m
in Lake McCloud, 1970-1971 . . . . . . . ... 21

6. Variations in total chlorophyll concentrations for
unfiltered samples from surface waters in Lake :IcCloud
and Biven's Arm, 1970-1971 . . . . . . . ... 23

7. Variations in total chlorophyll concentrations for
nannoplankton samples from surface water in Lake
McCloud and Biven's Arm, 1970-1971 . . . . . .. 25

S Seasonal fluctuations in phytoplankton in Biven's Ann,
1970-1971 . . . . . . . . . . . . . 30

0 Seasonal fluctuations in phytoplankton in Lake MicCloud,
1970-1971 . . . . . . . . . . . . ... 32

. Variations in phytoplankton diversity for unfiltered
sa;rle. from surface waters in Lake l;cCloud and Biven's
Ann, 1970-1971 . . . . . . . ... .. . . 35

11. Seasonal fluctuations in zooplankton in Biven's Arm,
1970-1971 . . . . . . . .. . . . . 38

12. Seasonal fluctuations in zooplankton in Lake MicCloud,
1970-1971 . . . . . . . . ... . . . . 40














Abstract of Dissertation Presented to the Craduate. Council
of th;i University of Florida in Pairia-l 'ul fillmcnpi of
the Requlrciients for the Degree of Doctor of Pliilosc.phy

PHYTOPLA 'rKfO; CO:-2-IUNITY S1P.RCTU'fUr A, Pf;~ L. Ri AR'Y PRODUCTIVE''
IN T-.'0 FLORIIDA LA;:LES

By

Carol Lymn Harper

August, 197]

Charinr.tn: Robcrt II. D'eW-itt
Co-Ch ai.i-nan: I'rank G. Nordlie
Major DIepanri-:cnnt: Zoology

The re]-tion.hips betucene Li pL i.n.ry pr i.- r icti. vity L a Cnd jga s;' c.iC.es

asseibla-iges, succession and diveisiLy were e:-;a ined. Pririary p.r-o-

ductivity was dcl-t-~nir.ed using Lte 14C technique. In an olig-ltro .hic

lake, a bimod;l pattern of primary ploJuctivity coincided '.. th l.hi-

appearanicc of diarc-ns, and blooz-s were follo:-.ed by increases in the

]oow-pliosphate- Lt(leraint gen.:s Dinobryon. N;annoplan!;ton :.ere fci.r.l to

dominate priminry produce ivi L in spring and vincer in tlh Eurf..-e

waters, aind throughout the :, ar at 1.5 meters. In a cutroplhic lake,

a persistent surunen blcom of heat-tolera.11 blue-g-een nlgae :.'as

superjim.posed on the bimodal pattern of sprinS and autu:-;n blooms.

Nanncrlan'kton were found to be of i;-porcance only in the spring

bl]ooi. No correlation. betti:e-'n .alal diversity or algal numnier::

and i primary productivity w. .:s f:'ud ji: either lake. Primary pro-

ductivity w;as signi."fic.-ntlv cor;--el.atd with chlorc.phyill 'a" con-

centr.c tons in the s:'rface. :n.aters o Lihe oligr-trop.ic :c .:c.














INTRODUCTION

Primary productivity and related aspects of plankton counmunity

structure have long been subjects of limnological interest, but most

research has .resulted in purely descriptive data for specific bodies

of w.ater. With the use of laboratory and field data from long-term

studies, attempts have recently been made to integrate these descrip-

tive data in order to explain the relationships between primary pro-

ductivity and phytoplankton composition, diversity aniid succession .

Iargalef (1960) emphasized the importance of this approach in opening

new avenues of investigation and analysis.

Studies of algal succession in lakes over long peiicds of time

have led to the formulation of hypotheses about the species or

species groups expected under certain biological and chemical con-

ditions (ilMrgalef, 1958, Olive et al., 1969). lbe physical, chIc-Al;al

and biological causes of succession have been directly correl-tced

:Jith the rcquireenn-.s of several species groups (Rcund, 1958, Brook,

]965, H;:itchinson, i967, Holland, 1963) and have resulted in the

recognition of c certain indicator species for various after r types.

Thie jircect relationships betowon a particular aIlgtl cpecici or

speciess assemblage and primary productivity hv.c been dcterLinild by

use of trA.-ce studies (Olive at al., 1968), studie-s of diversity

(P.tton, 19(3. :l.irgaLte, 1965, ;'icherson et al., 1.970) and chlorophyll

conlte'Lt (Prrsons and Strickl.nr.d, 1963, l.orenze-n, 1970), and direct

oh. ,:v-atLun. One result of the iieLcgration of Iihese d. ta has hben







the. dia3overy of the relatively larse contribution of nannoplaitkton

(those aL:-ca and bacteria that are nor retained t :,y- a #25 mesh plankton

net) to pr:i-:ry productivity in lakes. Rodhe (1935), Pavoni (1963),

Lund (1964) and Anderson (1965) indicated that nannoplankters are

often nu-rerically dominant, even when not dominant in biomass. Rodhe

(1962) attributed the L-nportance of the nannoplanlfton to the possi-

bility of their utilization of organic material in the water and to

their apparent tolerance of low light condLtions. These conditions

would allow nannoplankton to survive and reproduce .hen larger

ph.yto-pll.a ers could not. The numerical domiinance results in the

estabiishmlenit of an important food source for zooplar~nton (Rodhe,

1955, Lindar.an, 194.2, Gli..icz, 1963, 1969a, 1969b, 1969c).

Rodhe et al. (1960) showed thatr nannoplankton may daninate pri-

mary productivity even when they lack numerical dominance. Fogg, (1965),

Findenegg (1965) and Olive et al. (1963) stated that nannuplankton

.a.inmilarcion of carbon is greater than chat of larger algal cells

due to the larger surface-to-volume ratio of the smaller- cells. The

'.annojplapkton dom-inrance of primary productivity .varies with the trophic

staLte of the lake, the time of year and depth of the lake. Nannoplankton

,contri.bu.ut the major portion of primary productivity in ol igotrophic

lakes in '.inter and spring at all depths, and throughout the year in

deeper waters (Gold:,an, 1961, Goldian and Uetzel, 1963, Pavoni, 1963,

Ecerly, 1964, Gcen and Hargrave, 1966, Gliwicz, 1967 an.- Frey, 1969).

Thi.; study was undertaken to determine the relaticnslhip between

the plhytoplankt'on com-unity structure and primary productivity, with

p.:t i.cular emphasis on th, relative contributions of different si.ze

grouping,. '-Tvo sc-mi-tropical Florida lakes at ei.tlIrr end of the




3


oligotrophic-cutrophic continuum were selected in order to present

clear-cut corrparisons between lakes of different trophic states.

Determinations of productivity, chlorophyll concentration, algal

composition, succession and diversity, and zooplankton composition

urire made for each size grouping.














THE SIUDY AREA

Lake IlcCloud, an oiigotrophic sand-hill lake located 40 kilometers

(25 miles) east of Gainesville, Florida, has a surface area of roughly

8 hectares and an average depth of 5 to 7 meters. The lake thermally

stratifies only rarely in summer, and light penetrates co a depth of

about 3.3 meters. Chemical characteristics of Lake icCloud may be

seen in Table 1. Other studies on various aspects of the chemistry

and biology of Lake McCloud may be found in Colson (1969), Mlaslin

(1969), Miaslin (1T70) and Brezonik et al. (1969).

Biven's Arm is a shallow: eutrophic lake south of Gainesville,

and has a surface area of 60 hectares and an average depth of 1.5

Eotors (i;irdlic, 1967). The lake never then-.ally stratifies, and

light penetr;tion is limited year-round by dense algal bloorns.

Chemical parameters are outlined in Table 1. Detailed studies of

the che-i.uistry and biology of Biven's Arm may be found in Nordlic

(19S7).



















TABLE 1

Chemical characteristics of the study, areas


Constituent Lake McCloud BLven's Arm

pH* 5.2 7.9 7.8 9.8

acidity, ppm CaCO3 3.5 2.0 4.0

alkalinity, ppm- 3.0 99.3 176.0

dissolved organic 0.012 0.72 1.84
F04, iig P04/1

nitrate, mg [03/1 0.041 0.0 0.3

fromn this study (1970-1971);
all other Lake McCloud data Ci ii
Brezonik et al. (1969), and all
ocher Bivcn's Arn data; Crom
1;ordlie (1967).













METHODS AND MATERIALS

Sampling occurred at two-week intervals from February, 1970,

through February, 1971. Surface samples were collected directly

by immersion of a five-gallon plastic carboy, while samples from

deeper levels were collected with a Van Dorn sampler and transferred

to a carboy. The water was then returned to the laboratory for

analysis and treatment.

Algal size groups were separated by filtering the lake water

through bolting cloth of three mesh sizes. Each sample was divided

into four groups unfiltered, less than 158 p (#10 silk), less than

76 p (#20 silk), and less than 70 p (#25 silk). The latter group,

for purposes of this study, has been designated as nannoplankton,

in accordance with those definitions reported by Dussart (1965),

and Williams and Murdoch (1966). Filtered and unfiltered water

was then placed in BOD bottles for further processing.
14
Primary productivity was determined by the 1Carbon method as

described by Strickland and Parsons (1968), modified for fresh water

as suggested by Arthur and Rigler (1967) and for liquid scintillation

counting as reported by Lind and Campbell (1969). Light and dark

bottle pairs were inoculated with 2.5 pCuries of Na 4CO3. Samples

were incubated for 4 to 8 hours in a shaker at 200C and a light

range of 500 to 700 lux rather than at varying conditions, or in

situ, to facilitate comparison of the two lakes. Fifty-milliliter







aliquots of each size sample were fixed with 40 percent fonraldehyde,

filtered through 0.8 p millipore filters at low pressure, dried over

silica gel for a minimum of 24 hours, and counted in a liquid

scincillation counter. Some samples were filtered through 0.47 p

millipore filters, and samples of filtered water from both size

filters were counted. No apparent difference was noted between the

cwo size filters, with each size filtering out all algae found in

che samples. Equal portions of the unfiltered size grouping were

further subdivided by filtration through 5 p and 8 1 millipore

filters.

Carbon fixed as mg C/m3 hr was determined from the formula:

(RI Rd) x 1I x 1.05 x: 1000 I/m3
h x A

where: R = counts per minute (cpm) in the lig;t bottle,

Rd= cpm in che dark bottle,

W = weight in mg/l of carbonate present in water,

h = incubation time in hours, and

A = activity added in cpm.

The figure 1.05 is a factor that adjusts for possible isocope effects

caused by the diflfrence in size between 12C and 14C.

A replicate BOD bottle for each size group ans analyzed prior

i:o incubation for water temperature, pH and alkalinicy (Standard

Methods, 1960). These three factors were used to determine the

.jnount of carbonate in the sample according to the method outlined

by S..au:idrs, Tramn and BaclLmann (1962). Portions were also preserved

with 40 percent formaldehyde and set aside for the counting,

identification. and measurement of phyto- and zooplankton. Identi-

fication to genus was :nade when possible for algae, and to class for

zoopl.d;'ton (PLrscott, 1954, Edmondson, 1966).







Additional aliquots were filtered through millipore filters for

Sol.c-ccion of and analysis of chlorophyll according to the methods

o-tlined by Pichards (1952), Richards and Thompson (1952) and

Creitz and Richards (1955), and as modified by Parsons and Strickland

(1963). Size groups w.erc the same as those established by filtration

for primn-ry productivity. An additional replicate BOD bottle was

incubated uith the radioactive samples, and analyzed for chlorophyll,

algae and zooplankton in a similar manner after incubation.

Statistical analyses of correlation were used as described in

Snedecor and Cochran (1967). Calculations were made with the aid

of programs for the Olivetti Progranmia 101 and the ;onroe Epic

calculators.















Priasry Productivity

Primary productivity (ft carbon fixed/m3 hr) of unfiltered

samples from surface waters is shown in Figure 1, and that of

nannoplankton samples from surface waters is shown in Figure 2.

In Lake McCloud, peaks in productivity occurred in May-June, 1970,

and in October-November, 1970. At these times, productivity of

Lake McCloud exceeded that of Biven's Arm even though Biven's Arm

had a persistent algal bloom throughout the year. Similar trends

in all data were exhibited for all size groups and are shown in the

Appendix (Tables Al A6).

Productivity for deeper waters in Biven's Arm was measured in

March, September and December, 1970, and in February, 1971. Due to

the shallow nature of the lake, wind mixing of the waters and low

light penetration due to algal blooms, the euphotic zone was limited

to one meter or less. Sample, were taken at half the depth of the

euphotic zone (0.3 m), and no differences in trends from surface

were noted in these samples. Table 2 shows that productivity at

0.3 m exceeded that of the surface with the exception of the

December, 1970, sample.

Primary productivity at 1.5 m below the surface in Lake McCloud

was measured at more frequent intervals than at the 0.3 m depth in

Biven's Arm. The depth of 1.5 m was selected as being half the

depth of the euphotic zone:iii.n order to compare data with those taken

from Biven's Armi Productivity for the unfiltered and nannoplankton























TABLE 2

Primary productivity,' at tw:o depths


in Biven's Ann


Surface 0.3 m
Date unfilt. nanno. unfilt. narno.

M3Lach 17, 1970 1.44 0.00 4.40 4.07

Sept.ciber 24, 1970 6.49 3.96 7.73 5.90

December 6, 1970 1.37 1.01 0.33 0.72

February 19, 1971 1.78 1.78 4.19 2.35


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samples from 1.5 m are shown in Figure 3; and Tables A7 through A-10

show these values for the other size groupings. Trends are similar

to those shown in Figures 1 and 2, although the magnitude of pro-

ductivity and the fluctuations were less in the deeper water.

Ratios of the productivity of the nannoplankton fraction to the

productivity of the unfiltered fraction are shown in Figures 4 and 5.

The ratios often exceed 1.0, and this phenomenon has been attributed

by Gliwica (1967) to the reduction of inter- and intraspcecific com-

petition. Filtration results in the renoval of algae and a subsequent

increase in available nutrients, light and space for the remaining

cells. In addition, the reduction in numbers of algae could result

in the reduction of possible inhibicive activities of some species,

and filtration reduces grazing pressure by the removal of zooplankton.

The ratios for Lake McCloud exceed chose of Biven's Arm in April-May,

]970, and in October, 1970 larch, 1971. The ratios for deeper

waters in Lake I'cCloud ece:ed those for surface waters for the same

lake from June, 1970, through ::oveber, 1970.

Chlorophyll

Total chlorophyll in mg/ml in unfiltered and nannoplankton

fractions is. shown in Figures 6 and 7 respectively. In Biven's

Arm, chlorophyll content roughly corresponds with the curve for

prLi-ary productivity, with peaks in chlorophyll concentration

appearing to lag behind productivity as much as two eeks. Total

chlorophyll concentrations in Lake McCloud appear co be much nore

stable. Primary productivity showed no coicelation Aith total

chlorophyll or with chlorophyll "a" with rh., exception of a positive

correlation between productivity and chlorophyll "a" in surface waters

in Lake IcCloud (Table 3).


































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0








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ro 0



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TABLE 3

Relationships between chlorophyll and prir.iary productivity

Chlorophyll "a" Total Chlorophyll
Lake-depth r 95' bounds on 4 r 95', bounds on

Biven's Arm surf. 0.28 -.216 -L .664 0.11 -.330 .551

licCloud surf. 0.55*-- -.020 P- .804 0.28 .2.81 .706

MicCloud 1.5 m 0.58 -.438 1- .947 0.59 -.422 !.948


: significant at .05 level







Chlorophylls "a" and "c" were the major types present in both

lakes throughout the year. These types are indicative of the presence

of diatoms, dinoflQgellates or blue-green algae (Delevoryas, 1966,

Hutchinson, 1967).

Chlorophyll concent-ations from incubated sarpleq showed some

differences in content from those measured prior to incubation, such

as those variations reported by Geen and Hargrave (1966). In some

cases, chlorophyll content increased during incubation while in others,

chlorophyll concentration decreased. .:o consistent trend for loss or

gain w.as noted for any size group in either lake, nor could losses or

gains be correlated with changes in plhyto- or zooplankton numbers.

These variations were possibly due to random variability in chlorophyll

in natural populations as noted by Yentsch and Rythcr (1957) rather

than to the influence of the process of incubation.

Species C-omposition

It became evident during ex:rumination of samples that certain

algae, corronly occurring in multicellular colonies, had cells that

were either too small or too numerous to count. Each colony was

therefore counted as a "unit" equivalent to a cell of a solitary

species. Figures 8 and 9 show the percent of the total algal count

for the algae found in the two lakes. Total unit counts (Tables

Al A10) and species (Table A12) were similar for both depths in

eich lake. In nainoplankton samples, species were similar to those

found in unfiltered samples in both lakes. This latter phenomenon

w:as reported by Pavoni (1963).

Algal assemblages show a dominant blue-green algal bloom

(Anabaena sp., licrocyscis aeruginosa) in the summer months in




23


Biveni's ArmL, and a winter and spring assemblage of diatoms (Nelosira

granulata) and small colonial green algae (PediastrLon spp., Schaerocvstis

sp,, Sccnedesius sop.). In Lake ilcCloud, algal species were pre-

doniinantly small dinoflagellates, chrysophytes (Dinobryon sertularia,

Mallomonas sp.), and desmiids (Staurastrum spp.). Total numbers of

algal units did not correlate with primary productivity (Table 4)

and did not change with respect to numbers or species composition

diiring incubation.

Algal diversity (Figure 10) for unfiltered waters was calculated

using the Shannon index of diversity (Hutchinson, 1967):

D = Pi(log2Pi)

where pi is the probability of occurrence of the ith species, i .oing

fro.n i to n. Diversity showed no correlation with primary productivity

(Table 5). Diversity remained relatively constant throughout the year

in Lake :cClcud while it decreased in the sr.mmer in Biven's Ann.

Zcoplanrkton composition in percent of total count for each

la're is siho.-n in Figures 11 aind 32. Biven's .rm wJas dominated by

.all r'jt.i.fers anid a colon.il cL'.[L atl i ..-ile a shelled amoeba Was Lihe

do;"iina::t species in Ln..k. TiCr; loud i-''. iln numbers of socplankLon

occurred *.i .June, Sepctcbbe-r and Decmc.ber, 1970, in Biven's Arm, ar.d

in July and October, 1970, in Lake IlcCloud as well as January, 1371

(Ta-bI l. ). Comparisons betWcen algal diversity anrd r oplaiidton

.-h0o:..:d .o correlation (Table 6). Numbers of zooplankton were simiilar

at the surFace and at 1.5 m in Lake IlcCloud.


























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TABLE 4

Relationship between number of algal units and primary
productivity

Lake-depth r 95,'. bounds on

Biven's iArm surf. 0.12 -.291 ._ 9 .493

McCloud surf. 0.11 -.345 i .523

McCloud 1.5 m 0.13 -.635 A .766







































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TABLE 5

Relationship between algal diversity and primary
productivity


Lake-depth r 95A bounds on \

Biven's Ann surf. 0.24 -.159 < .578

McCloud surf. 0.20 -.264 1.592

McCloud 1.5 m 0.55 -.254 .905































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TABLE 6

Relationship between algal diversity and zooplankton
numbers

Lake-depth r 95% bounds on

Biven's Arm surf. 0.01 -.454 .470

IIcCloud surf. 0.25 -.216 43 .438

IicCloud 1.5 m 0.66 -.080 s .932














DISCUSSION

Lake McCloud

hle spring primary productivity pulse in Lake McCloud coincided

with the appearance of diatoms (Figures 1, 2, and 9) and an increase

in productivity by nannoplankton (Figure 4). However, total algal

biomass, as indicated by numbers (Tables Al A10) and by chlorophyll

(Figures 6 and 7) appeared to remain at a low level. Peaks in pro-

ductivity in Lake McCloud might therefore be attributed to an increase

in the efficiency of nutrient utilization reflected by the increase in

nannoplankton productivity (Findenegg, 1965, Olive et al., 1963). Low

numbers of nannoplankton during the primary productivity peaks (Table

A4) and the relatively stable numbers of nannoplankton thlcughout the

year reflect the efficiency of these small cells ii productivity.

The fall pulse also corresponded with the appearance of diat-ons

:ut there was no corresponding increase in the productivity of

nannoplankton. The gradual increase in the summer population of

dLnoflagellLaes seems to be more closely related to this autumn

productivity pulse.

Dinrobr.-von species are often found following nutrient depletion

folioirng an algal bloom (Lund, 1963, Hutchinson, 1967). The decrease

in prii1mary productivity and the parallel incr-ase of Dinobryon

sc tularia after the spring and fall pulses could be attributed to

a decrease in t::;sential n,;.:riecrts, specifically phosphates (Guseva,

1947, as reported by Lund, 1965).








The decrease in the magnitude of the biomodal pattern at 1.5 m

migir. be attributed to lower light penetration at that level, not

species composition, since species were similar in number and kind

at both depths. Since the samples from 1.5 m were incubated at the

sane light intensities as the surface samples, some form of light

inhibition could be postulated. Stratification is rare and never

uarsistent in Lake McCloud, so the algae could become shade-adapted

only on a short-term basis. Such major physiological adaptations

as enzyme shifts which were proposed by Yentsch and Lee (1966) probably

are not responsible for the lowered productivity.

The importance of nannoplankton in the productivity of Lake

NMcCloud, as sho.n in Figure 5, fits the pattern predicted by Goldman

and Wetzel (1963) in that nannoplankton appeared to be most important

in winter and spring at the surface, and throughout the year at 1.5 m.

'lie increase of nannoplankters at the time of the spring bloom has

bcen attributed to their efficiency in the utilization of nutrients

and to the inability of the larger species to capitalize as quickly

or: the niuirienCt pool due to their slower reproductive rates, growth

rates and germination time from overwintering stages (Lund, 1965).

The lack of nannoplankton dominance in the fall pulse indicates that

resources were tied up in the biomass of the larger algae. Once the

larger species died, the stored nutrients would be released, and the

naunnoplani:ton could increase in winter. 1he nannoplankton species

appear to be able to utilize organic nutrients and to be tolerant of

lo; light conditions (Rodie, 1962), which permits reproductive

success in winter and in deep waters.







As expected from studies of other lakes by Pieczynska and

Szczepanska (1966), chlorophyll "a" concentration was a good indicator

of primary productivity in Lake McCloud (Table 3). The lack of

correlation at 1.5 m may be due to the small sample size (n = 8).

An increase in the n:-nbers of samples analyzed would probably show

that chlorophyll "a" concentration correlated with the level of

primary productivity throughout the lake, at all depths. Chlorophyll

is a better indicator of productivity than numbers (Table 4) since

numbers can include dead individuals, individuals with lowered or

raised activity, or individuals of varying sizes and shapes. Fogg

(1965) suggests that surface area is a more accurate indicator of

productivity than numbers as it reflects biomass more accurately.

The species present in Lake McCloud, as indicated by chlorophyll

content and microscopic investigation, were predominantly diatoms and

dinoflagellates. Species such as Peridinium sp., Sphaerocvstis sp.

and Dinobrvon sp. are indicative of low nutrient conditions (Hutchinson,

1967) while desmids such as Staurastrum spp. (Brook, 1965) are indicative

of both L-v nutrient conditions and acid waters. Colonies of Dinobryon,

as meiutionad previously, are especially numerous in lakes recently

depleted of nutrientby algal blooms.

The pattern of succession followed that described by Glivicz

(1967) and Margalef (1958). According to Clivicz and Tiargalef, a

spring bloom of algae that are rapid reproducers and efficient

producers (such as diatoms, nannoplankton) is followed by an increase

in importance of larger, moce sluggish spe-ies that are more tolerant

of low nutriic::t conditions (such as Dinobrvon sp.). These species,

being lon3-lived, slow nutrient turnover, leading to lowered pro-

ductiviLy. In the fall, die-off of these larger species releases








stored nutrients to the water, allowing an autLumn bloomn, followed by

another pulse of species tolerant of low.nutrient conditions. The

winter population of efficient, low-light tolerant species (such as

nannoplankton) forrmsthe "seed" population for the coming spring bloom.

In Lake licCloud, algal diversity was not indicative of primary

productivity. Productivity appeared to be more closely related to

[he actual assemblage of species present rather than to the number

of algae, che total biomnass or the numbers of species present. Ilargalef

(1965) proposed that algal diversity was directly correlated with

primary productivity but this was not the case in Lake IlcCloud.

Zooplankton species (Figure 12) did not appear to change

chason.lly but those animals found in my samples were fewer and less

diverse than those found in the samec lake by other researchers (Maslin,

1969, !la.lin, 1970). This was due to my failure to adjust collecting

Lc.chiniques for zooplankton, the more agile species being able to avoid

the i'.krrow mouth of the collecting devices. A subsequent underestimation

of certain groups resulted. -iaslin (1969) showed that some groups,

notably copepods, appeared to be correlated with changes in productivity,

:l.id the peaks in zooplankton number that he found closely follow the

inc rE.as s in I:.-inn:oplarkton productivity found in this study. The

!:ir.noplanLtoun ace thought to be a principal source of food for

herb iv.nrC~s plankton (NauLwerk., 1963, as reported by Lund, 1967,

Cliwic::, 1969a, 1969b, 1969c).

In .; lr.- ary, Lake !.cCloud, defined as an acid, nutrient- poor lake,

can be described using primary proSductivity, algal species and succession.

In .-uch a lake algal species and size groups typical of the nannoplankton

dcni.inate pci:iriry productivity.








Si'ven's Arm

The s.-asonal periodicity in Liven's Arm did not completely re-

semble that in Lake McCloud (Figures 1, 2). A persistent swum~er

bloom of blue-green algae, sho\.n in Figure 8, coincided with a summer

increase in primary productivity. Such blue-green alg3l blooms are

not uncon-inon in shallow, wind-mixed lakes where nutrients can be kept

in circulation throughout the year (Lund, 1965, 1967, !.hitford and

SchuLmacher. 1968). This apparently was the case in Biven's Arm.

Spring and fall pulses in productivity appeared to coincide with

the appearance of the diatoa., Melosira granulata. The spring bloom

was also dominated by nannoplankton productivity (Figure 4) although

:.o such dominance was apparent in the fall. This pattern is similar

to that described for Lake IIcCloud, although the actual magnitude of

productivity and the numbers of algae in Biven's Arm were generally

greater chan in Lake McCloud (Figures 1 4, Tables Al A6).

lhe tendency for productivity at 0.3 m to be greater than pro-

ductivity at the surface (Table 2) was not due to an increase in

nuL-b,--r in the deeper waters (Tables Al, A7). Since 0.3 m is very

near the bottom of Liven's Arm, it is possible that the increased pro-

,1uctivity. is due to the proximity of the algae to the nutrient-laden

bottoma sediments. In addition, fe.er numbers of algae permit a

rn'duct:i.on in competition bet.jeen cells for available nutrients,

*-v n H.h.1,gh light penetrLation is limited.

.'*.r. i-cc' :se in nanpcjplankton numbers in the spring bloom might

be expec ted if an increase in nutrient content in the water accompanied

a '-t~in :a.ii-ing tra'nd, ;ordlie (1967) indicated that both nitrate

and .'.osphatc- concentrations increased in May, following a slight








decrease in !larch and April, and coincided with a temperature rise

beginning in March. Such a rise in available nutrients would allow

the more efficient nannoplankton to reproduce rapidly until nutrients

were exhausted and until the summer species began to appear.

During the remr.ainder of the year, nannoplankton contributions

were equal to or less than those of the nannoplarnkon in the summer

in Lake !icCloud. From this pattern, it appears that nannoplankton

are not as important in the productivity of a cutrophic lake, except

during a period of rapid increase in numbers such as in the spring

bloom.

Neither chlorophyll "a" nor total chlorophyll content showed

any significant relationship with primary productivity (Table 3).

Pieczynska and Szc;epanska (1966), and Winner (1969) indicated that

the use of chlorophyll as an indicator of total viable biomass or of

primary productivity became limited when chlorophyll concentrations

increased above a certain level. Much of the chlorophyll then measured

would be from green but inactive cells and detritus, and more chlorophyll

would be found in cells operating below maximum efficiency due to

trading or self-inhibition. Chlorophyll levels in Biven's Arm were

rarely below 40 mg/ml, far above the level of 800 pg/ml set by

Pieczynska and Szczepanska. Similarly, in Lake McCloud, the

chlorophyll level usually fell within the range of zero to one ng/ml

and a positive correlation between primary productivity and chlorophyll

"a" lcas noted in Lake McCloud.

SpecLes present in Biven's Arm, as indicated by chlorophyll

corccntraticns and microscopic examination, were predcsninantly

diatoms a:id blue-green algae. Species such as Melosira sp.,

Pediastrum spp., Scenedesmus spp., .'nabaena sp. and .Microcvsctis sp.







are typical of eutrophic systems (Brook, 1965, '-.-,itford and SrhuTacher,

1968, Hutchinson, 1967, Holland, 1968). Many species of blue-green

algae, such as Anabacna and ilicrocystis, are also tolerant of high

temperature ranges and bright light conditions. This results in

blooms in heated surface waters in the summer w~hen other algae are

inhibited by light and temperature (Lund, 1969). In addition, several

species of blue-green algae are capable of nitrogen fixation, notably

Anabaena sp. (Lund, 1965). Blooms of Anabaena such as in June and

July in 3iven's Arm could then occur in nutrient-depleted waters when

not even other blue-green algae could survive. Such blooms Could also

add to the nitrate "pool" in the system, permitting a rapid regrowth

of formerly limited species. The frequent reappearance of the diatom,

I'elosira granulata, during the sLmmenr may be due to changes in the

turbulence of the water. This species, although still viable, may

sirn to the bottom of the lake where it survives. Increased circu-

lation in the water due to winds lifts portions of the bottom of the

lake in'o t:hre water column, and such reappearance does not necessarily

indicate a change in nutrient status of the system (Lund, 1969).

The pnl:tern of succession in BLven's Arm an association of

de:nmids, diatoms and colonial green algae followed by blue-green

algae also fits the pattern described by Margalef (1953) and

Gliwic- (1967), and that found in Lake McCloud. i.ith the exception

of the smmnmer bloom that resulted fron wind-iixing that did not

occur at deeper Lake McCloud, identical cycles of succession were

followed.

lhe clgal diversity of Biven's Arm decreased in summer during

the bloon, cf blue-green algae (Figure 9). Often these blooms were








mor.ospecific, or nearly so. It has been theorized (HarL-tan, 1960,

as reported in Lund, 1965) that blue-green algae e::ude growth-

inhibiting substances toxic to other species, and that these sub-

stances are rrost effective when the population is nearly a mono-

culture. Under such conditions, reproduction and growth of new

speciesare curtailed until a die-off of the blue-green algae occurs.

As a result, inefficient producers such as blue-green algae can main-

tain high population levels and a relatively high productivity level

on the basis of sheer nur.bers. Because of flis summer phenomenon,

algal diversity did not correlate with primary productivity (Table 5).

In fact, even higher productivity might be obtained from such mono-

specific cultures if total numbers of individuals were reduced. A

higher productivity during the simmner bloom occurred in the nanno-

pl.inton sample of Biven's Arm than in the unfiltered sample (Figures 1,

2). Ho.ev.eir, the species associations were the same. Filtration

r.sIltcJd in a reduction of numbers only, allowing more light penetration

pcrv c ll, and nore nutrients per cell. In such a situation, the blue-

graen alg.c c'ixhibited an ability for increased productivity that is

pcesur:.abii hLl.d down by intraspecific competition in nature.

The ::oop'.arkton samples from Biven's Arm suffered from the

same undc.restimation as th)se from Lake McCloud. However, a marked

scaronility ..-as apparent. (Figure 11). Decreases in the importance

of t'-e rotifer population during an Anabaena bloom in June, 1970,

and an absence. o)f cladocera throughout the sumuLcr bloom of blue-green

a1..ae s.),-cies .,are observed. This decrease in sone zooplankton groups

...y be due to a lack of tolerance to varmer watcrs, higher light

inten;itiesli o the algan themse.lvs. Ilumburs of zooplankton did







not correlate with algal diversity (Table 6) since som-e species of

small rotifers increased in numbers during the sumLmer months. These

species may have been feeding on bacteria associated with the sheath

of Micrccysris aeruginosa (Lund, 1967), a food source not accounted

for in the algal counts.

In summary, Biven's Arm exhibits many of the patterns shown

by Lake McCloud with respect to primary productivity patterns and

nutrient cycling. These patterns are superimposed on patterns of

productivity, algal succession and species composition common in

eutroph'ic systems '.here "net" plankton assume the dominant role in

productivity.














S U.EIARY

In this study, the relationships between phytoplankton community

sc.-'.:ture and primary productivity in two Florida lakes of different

tror-.ic status were examined. The most important conclusions were:

1. Nannoplankton (algal species with a diameter less than 70 p)

dn:inaced productivity in surface waters in winter and spring in

oligaotrophic Lake McCloud. Lannoplanklton also contributed the major

po,-tic'n of primary productivity throughout the year at 1.5 m. In the

iu.riophic la!e, Eiven's Arm, nannoplankton were of major importance

in the spring bloom only.

2. Spring and fall primary productivity pulses coincided with

che appearance of diatoms in both lakes.

3. Spring and fall blooms were followed by species capable of

rcproducti.ve success in nutrient-depleted waters in both lakes. In

L:-ke2 :cCloud, this species was Dinobrvon sertularia, a species

rolcr.-at of low phosphate concentrations, while in Biven's Ann,

:i.Ln-'.:a sp., a nitrogen-fixing blue-green alga, followed the pulse.

4. in Miven's Arm, a siinmer primary productivity pulse coincided

with a blocirn of blue-green algae.

5. Chlorophyll "a" concentrations .were positively correlated

%..'Lt!. primacy productivity in the surface waters of Lake McCloud only.

C!.loioply'1 co..centr:ations in BivAn's Arm were too high to be useful

aj an i'd.iicator of productivity.








6. ;;o correlation was found between primary productivity and

algal diversity or algal numbers.

7. Increases in uannoplankton productivity were followed by

increases in numbers of zooplankton, and by the reappearance of

certain zooplankton species. Zooplankton species and numbers de-

creased during blooms of blue-green algae.







































APPlETDIX




















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    Zooplan!:con numbers (it/liter) for unfiltered samples,
    surface waters in Lake McCloud and Biven's Arm.

    Date Lake licCloud Biven's Arm

    1II/70 ---------- 1.0 x 103

    12/111/70 1.3 x 103

    17/111/70 ---------- 3.3 x 103

    26/III/70 0.7 x 103

    4/IV/70 ---------- 1.7 x 103

    13/IV/70 2.3 x 103 ----

    17/IV/70 ---------- 0.0

    23/IV'/70 ---------- 1.0 x 103

    30/IV/70 0.3 x 103 ----

    4/V/70 ----------- 1.0 x 103

    11/V/70 1.7 x 103

    22/v/70 --------- 0.3 x 103

    29/V/70 0.0 ----------

    6/I/70 ---------- 0.3 x 103

    16/VI/70 ---------- 29.0 x 103

    26/VI/70 1.7 x 103

    8/VlI/70 ---------- 5.0 x 103

    17/VII/70 --------- 0.0

    21/VLI/70 3.3 x 103 -----

    29/VLI/70 ------.--- 0.3 x 103

    6/VIII/70 2.7 x 103

    21/VIII/70 ---------- 1.3 x 103

    11/1X/70 ---------- 8.0 x 103

    15/I:/70 1.3 x .03 -----









    TABLE All Continued


    Lake ;icCloud


    Dace


    2'/IX/70

    23/IX:/70

    523/X/70

    12/1/70

    23/'X/70

    7/ XI/70

    17/XI/70

    24/XI/70



    6/X11/70
    6 1'::11/70

    13/X11/70

    23/XII/70

    29/XII/70

    9/1/71

    15/1/71

    22/1/71

    31/1/71.

    5/11/71

    12/T1/71

    19/II/71

    1/III/7 /


    ----------

    3.3 x 103



    3.3 x 103
    3. 3 x 103


    ----------

    2.0 x 103

    ----------

    1.0 x 103

    ----------

    1.0 x 103

    ----------

    1.3 x 103

    ----------

    1.3 :; 103

    ----------

    3.0 x 10

    ----------

    1.0 x 103



    1.7 103
    1.7 x 103


    Biven's Arm


    3.3 x 103




    0.3 x 103




    2.6 x 103

    0.7 x 103




    0.3 :x: 103




    6.0 x 103




    2.3 x 103




    0.7 x 103




    0.0




    1.3 x 103




    0.0


    -~---








    TABLE A12

    List of phytoplankton species found in unfiltered samples,
    surface waters, in Lak2 McCloud and in Biven's Arm.


    Division*
    Cyanophyta


    Eul enophyc
    Cllorophyta


    Chrysophyta


    Prroph,; ta


    Lake McCloud
    Agmenellum sp.


    colonial green algae

    filamentous green alga

    PediastrtCln sp.



    Scenedesmus sp.

    Closterium sp

    StaurastrumL spp (5)

    Staurastrum cornutum
    Malllo.nonas sp.

    Dinobrvon sertularia

    Ceratium sp

    unidentified dinoflagellate
    (2)


    Even's Arm
    Microcvstis aeruginosa

    Anabaena circinalis?
    unidentified flagellates

    Sphacrocvscis sp

    Coelastrun sp

    Pediastrum spp (2)"

    Ankistrodesmus sp

    Scenedesmnus spp (2)

    Tetraedon sp.

    Staurastrum sp.

    colonial green algae

    Botrvcoccus sp.





    s


    Bacill.rioophyceae Ielosira granulata

    Melosira sp. Cyclotclla sp.

    Cyclocella sp. unidentified diatoms

    Asterionella Eormcsa

    unidentified diatcins


    SDivisions according to Edmondson,
    1966.
    SFi.gures in parentheses refer to the
    nu-mber of d-stincc species found in
    the genus.













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    BIOGRAPHICAL SKETCH

    Carol Lynn Harper was born September 28, 1942, ir. Camden,

    New Jersey. She attended Wellston High School, Ohio. In September,

    1960, she entered Bowling Green State University, Ohio, where she

    received the Bachelor of Science degree in 1964. She attended the

    Ohio State University field station in the summer of 1963. In

    September, 1.964, she began graduate school at the University of

    Southern California, where she received a masterr of Science degree

    in 1967. In September, 1967, she began graduate work at the

    University of Florida and has since pursued work toward the degree

    of Doctor of Philosophy. She received support from the department

    of Zoology (1967-1970), Environi-ental Engineering (sumLers, 1968,

    1969), the College of Education (surmnner, 1970) and t1ie Department

    of Health, Education and UielCare in the form of an idea Title IV

    fellowship through the Graduate School of the University of Florida.

    M'rs. Harper is married to Charles Alan Harper. She is a

    member of Phi Sigma, and the .Anerican Society of L.-Lriologisls and

    '3c -ea.'.ogra -i e rs.









    I certify that I have read this study and that in my opinion it
    confomns to acceptable standards of scholarly presentation and is fully
    adequate, in scope and quality, as a dissertation for the degree of
    Doctor of Philosophy.



    R. !M. DeWitt, Chairman
    Associate Professor of Zoology



    I certify that I have read this study and that in my opinion it
    conforms to acceptable standards of scholarly presentation and is fully
    adequate, in scope and quality, as a dissertation for the degree of
    Doctor of Philosophy.





    F. G. Nordlie, Co-ChairmTan
    Associate Professor of Zoology



    I certify that I have read this study and that in my opinion it
    confonlis to acceptable standards of scholarly prc.::cntatio-jn a] is fully
    adequate, in scope and quality, as a diisertLLi'on for the degree of
    Doctor of Philosophy.





    i. D. Putnam
    Professor of Environmental Engineering


    This dissertation was submitted to the Dean of the College of Arts and
    Sciences and to the Graduate Council, and was accepted as partial fulfill-
    r.ient of the requirements for the degree of Doctor of Philoscphy.

    Augusti.- 1971


    Dean, Graduate School




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