|
J
r]-]-L:'(... OF FLORID& SPRINGS
L '*W 580(02)
Firat Se~.=amnnu"a Report to
ZSIrJloCS Division
C.ica of NIval Resoarah
Pr'eore ...-..i rp:.i 1, 1952 to %January 31, 1953
:;.-.~.r T, Odum
.'. sections :'* ,'"..-i HD 3Davis, William Sloan,
David CAldwll, and Gordon Broadhead
Dopartmeon of Bioloa
College of Arts and Sciences
University of Florida
aainesville, Florida
TUNVERSITY OF FLORIDA
Gainesville
College of Arts and Sciencos
Department of Biology
31 January 1953
Chief of Naval Reaoarch
Attna Biology Branch (Code 446)
Department of the Navy
Washington 25, D3. C.
Dear Sirs
Herewith please find the first semi-annual report
of progress on a project, NOSR 580(02), concerning "Produc-
tivity of Florida Springs."
You will note that work during the first
has included much c.p o~- ; 5n of posaibilitles,
that Dr. Odum h",,. made real C:,oa.r'o and is now
to center down on a more intensive study of the
among the numerous prom iing opportunities that
covered.
six months
X believe
in a position
best possibilities
he has un-
Respectfully submitted,
W. 0. Allee, Read
Department of Biology
" *.(Aiv oi' lo-ra ", :'. ; (02)
,i. '.:' ., .,. .
ro '" 1.' : .... 1 ...-.
1. Chara. "i- i an d. abilityy of .n giving .nv c' 'it
,, t :,
.. 5lI 3 ca
; ard. x in mnacirobic ..r
: .. .ltiln expoarirent, by David :'.' r6w-''
'- 'ne i .ih ad. .- a
3 coposit O .'. o coammi tios (z t:. .: l)
1% '. of nram ai. n 'ber
Th 1 .1. and denuity of aquatic vegetation in four
spr.!t.t. hei'r riverI, by ,Jobn H Davii
Initial -.... .... in c.2s'f '.' .., ,..: of fish, by
40 Productivity
1. prod-rctiong
b, :eam drift rtt]'.,.
oC <.*vr-( Imc.ri"nt r3.'f .bol.d
ft.m :..%.tdat ivlty .*;
S L .... *. .states
Production pyramids in streams
Th'rmodyna~mic and food chain efficiencies
Efficiency, '-i.:..ty. and power in food ,_1inn
FL.e:, for 2nd '..::. ,' .
Productivity of Florida Springs NOFN 580(02)
Page 2
ABSTRACT
Productivity in the Springs of Florida
Work has began n on stdyin the factors responsible for productivity
in the Florida springs, t'hbiLc: are nearly constant temperature, constant
chemical, steady state giant laboratories, Progress has been made on five
aspects qualitn-i;.va description, quantitative description, completion
of knoTrledge of chu.i-iic.ari factors, measurement of productivity, development
of productivity theory.
Measurement of the .-i 1:.'.:; productivities in Silver Springs and Green
Cove Springs by two nen methodEs the raising of organisms in cages, andW the
measurement of night day di, '.. ,.' .'-i in oxygen downstream aree .*.'o,'l..,
Production in those springs is greater than previous production figures reported
for marine, fro.th i.fter, and lanrd lcn.m Instantaneous measures of production
show large variations w ith seoaon, time of day, and cloud cover. Production
estimatop rn'r..-c Lr., 11,,000 lbs. por acre per year to 70,000 lbs. glucose per
acre per year dwin.c.. day.Vi:. "- hours
Essential stability of the spriv,c environment has been shown with
respect to I.-r.-:,r."'t'.-w.j phoipho.rus, and plant cover. A correlation of species
number with lack of e'.,!;..'..A.;r has been shown with insects. uantitative
studies have shot' very large plan: t base to pyramids of mass. Correlation
of marine invasion 'iith Oh...v;iity has been shown. The essential aspects of
pH regulated ;phir.-iI.: 1,. geochemistry in Florida have been outlined. Some
theoretical ideas on productivity bave been evolved. Mapping of sessile
organisms in p:-.ti;- and. taxonoloi i.e ,tt.i'ication of dominant are half
completed. Plans for second six months include measurement of herbivore
and carnivore prod.netion ratos an:d completion of food chain efficiency
determinations in Silver ysr:J.ia,-.. as a preparation for subsequent comparisons
between springs.
rOi;.Oia'.c" rity of M ri.a Sp.ring. ... 580(o
PaWe 3
INTRODUCTION
By a rciiln,,. le circtumtanc.t o of nature there are many large springs in
the vicinity of Q'A-in,_z;,lll,, Florida,. Those springs have a relatively
constant tr.p,' -,rt.:.io of ,71 to 74 0;oro0i F. throughout the yer and. are
all at this tC:ri.p.:.'-".:.'o -.-i:. : are -,-:.n;y varied types, a a n. all contr,
aquatic corm..lAnitieo in their basins and their outflow channel. Each
spring d.ff'o:.; from the others ;,; a few factors. Tus there are seodium
springs, calcium -.p,:iv,; .. .-.'t.-, with high and other with low oc':.-n-
reduction potentials, saline springs, soft water springs, and other types.
Analyses of many of the '.:- cho~.1 mical elements in these waters have already
been published. *(2 .-!.. :, G. .X .L. ;.at, 0. W., Love, S. K., and Vernon,
R. 0., t'ri:;i of Flo ..'1 e .n ,:.'': l Li.tin No0 31, State of Florida
Department of *3c.. '".v:Mio-n, T'allaaszs, 1947) Oxygeno, T-. 3so.'.o .-.d
nitrogen had until .. .:-.1;r *:'o'nt boon : .eysi,.
Because of their pyeocal properties thbn;: npri :: are collectively
a ,::.;.-, constant ":...' *:"' 1 7*n. 1.'..:,,'r The flow of water of constant
quality in the varlioaus p'~ings bath~ a the spring comansition with a constant
medium in spite of the actions of the cowamnaity that rct ifjr the after Fior
the firat time it ia poasiblo to o':'c.*--r' whole communities 'i~':ch are W, Il..' ,"1a
under constant '-".tiona that .'' '." by on.y a few factors
The ifp:'.., '"'i.'-.c provide e. I.- ;;i of conditions in dhih tohe ,C;i::.;',
of the comunitaries rmy :' -'.'. ao.cause the rate of flow' of each rni
is relatively constant a7. te'oo ...:.. the run cor:-.po';n to a tie interval
following the first moe: '.. of sunlight, and water., 'tr.:.:, the ;-.':.'.I.. run can
be used to study rates of" ',' -, ;.::'..:. Several of the runs involve a tra--
ai'..c; :on from fr, '.t' water to ?Mea ,.,Yter of the ocean. '.", ., :.:...rI : '... On.,L,. r.':;n
of fresh water fish:. ,... ; .'- :.. slt water fish in .i:'..:h water spr:..'.,::
provides an qn T.':.. :: .ty-. to ",tiu.y tho differences between ccom i niwtios ..cc.
to predation of at watnr :. .'n; similar sprin' inland having only
fresh water fish. as ;--.. .. W spr.'. a:'. have oay2:.;r fl;t ow. W, Som0
have fish $.p'lT.'U t .. d soam ane too. -. from such IpoulatiS';.,
Thus there exists a n;.:'vloaus rc, ,.tr.ity to oti-'.,Iy proed.n,-'.!./ in tho
rcc'tt:-LA-'0.e natural laoba ,; in which. whole comamnities can be :*:.;r1 nuaier
controlled conditions. Tbo series of natural experiments that have been set
up seem ideal for -;..,.: ..; the role of the factors that control -c.-.Ityo
K Mi....- -*.' I. ,
The purpose of this reacsrch is to :"I.,"y the basic fadcors controlling
individual, population, and community prodad.tivity by an ~, ,.'..r of the
unit: conditions wre..i.0 'by oagrtflo .;,; selected constant tapecratrue
springs.
The general plan is to csM-Ar.Inh the qualitative and *;.Tr!(. tative
structttr of contrasting !-.:.'1,,.;, mneasne tha production rates, and determine
factors responsible for differences by .':.'it;,-; and. erperimient
.Productivity of :Florida :. :.:i 580(02)
Page 4
Funds from 0MR, University of Florida, and a small grant ($500) for
phosphate work by Florida Goological Survey have provided support of the
principle investigator full time in summer, one graduate student and one
umdergrsduate *unili-ntIn Lt o'uO the year, and other students and faculty
with field trip c.i:',n-ve,.
This project is a cooperative endeavor. Department head Dr. W. O. Allee
has made a significant contribution in planning. Dr. Howard T. Odum has been
coordinator and as principal investigator emphasized chemical conrposition,
production rate measurements, and productivity theory. Mr. William Sloan has
emphasized the qualitativo composition of inverte'rate fauma especially
insects, using species-:nuraber comsxnity analysis. This has been part of his
work towards the masters ..,,y0. Dr. John He Davis has investigated the
quantitative standing crop of aquatic plants in relation to salinity in coastal
spring runs, The Florida C.uolor;ic:l Survey has supported Mr. Richard Highton
during the summer, 1.,~o for analysis of pho~phorus in these springs and other
waters. Dr. Ao M. T. -..;-I.. hs develQped3 a her~'rium collection of ,*.itic
higher plants i:T.', the .ii'..];,.;1 '.:,:..1nti.l taxonomic aid has been provided
by 8 Dr. M. J. :.'.:;t..3., Dr. L. .:.r c, Mr. Ji9 Crenslha, Mr. Robert OCummins,
Dr. C. J. Gotoi, all of the Uii- ..*:; y of iorid; Dr. R J. ahmm, Oeanographic
Institute, Florida S.: -.1. Un.t;A'.;;.'. ~' and Dr. Horton Hobbs, University of
Virgin&ia Pr';.'r.!..h.i :'.r.y r.' ..' .i :t, in springs haIe been conducted by Mr.
Gordon Broadhead, and Mr. O.'-,.ld A'> ,lz. !., UiAversity of Florida.
Sr:tcl.2I courtesies have been received from Mr, William Ray, :Ri..n;,;',
Silver Springs; Mr. Ross Allen and Mr, :ili.'( ,d Neil of Ross Allen Reptilo
Institute; Mr. P'I: :;..-:,' i. > ,Mnag of 'ir':n .:aSprings; Mr. Ray )tlni;.'d,
Manager of 'n :i'.--r',..- : r;; :.- ;,1-)i..':ed Dean, Chassahowitsla Springag
and Mr. T i',vv..7 'rC~a~a CoD '";l.r::rn
Investigations into the factors controlling productivity can be classified
under 5 h,: din prob:1-.y irn inverse order of importance
1, Characteristics ;', l stability of non-living environment
2, d.alitative composition of communities
3. quantitative composition of communities
4. Productivity
5o Productivity theory
During this first half year an effort has been made to obtain results
in each of the five divisions. Efforts to develop a methodology for measuring
standing state composition and productivity have been restricted to 7 conF-
trasting springs although systematic initial surveys have been made on 40
springs, Since the measurement of production rates is at the heart of the
objectives of the project, these studies have been began even though descriptive
qualitative and quantitative aspects of the varied springs are not yet completed.
Progress is summarized below .'.Iov 5 headings.
od cte I .Y of 'r T. '. -.: *n ."
1, CHAtRAJIf, L *TAM 'D ST. 0..1A7 OWF WI-IUTIVm; ; i':I'Ti
To date 1."rr0 1jrs *. .e1 rV'1i (-i*-An i and to.) booeen
made in 33 *:' z.i t to Cel ..2..' .. Bma .gni ,.n., <. o .iW..,1 and iT; of
plhoqt. l in those con,rt:'. ,, r I-;.r;..', l: flow systems. The a *- A:--:3 i
in LA '.*:'; .. haLu 1. .. .. .. with ..' p . o .. n.. 'n, V.'t i trn at ams,
01 C POO i ; ..,'.-estuavies :.'.1 with ." .i. ci..I support of the .' :.1''..
Result s so for hiwZfe C ."i : following '.or.:- of pho pbaruas
distribtOtiona
1. In tbhe '-, r '. flow the /pho':tern t content i
A- const Xt ,iuth distance .'.n :: rvn even W.t, ; -
.. '. .. T 3ta .'. Silver 1,:L- in table 1 inA.oalto
."," *propsrties,
2. The ..*, '::.:i ccr.onte : F thee clear waters in .:.s :-Ors 13 in.
^te :5 .1
3. The varlation of ,' .,.s conant with time is low, f-" c i ,
that. va? with ninsre ''e. *:.t! :.:r ..;r active olementus, sow n." *'.
sprL-; onviro'ents tat tIcat a;r; relatively constant,
4o The 6.!' *, 7.. ". i,.' ."' -".'. in p .WI. il. .- 'r, ': in conoces .trantionx
is available to o larnt in :;.ij. ?*-*'..r excess reserve amounts (Ite to
the contimwal "' *.* of vater. In some :.1,0 m.u; s ndh as ".1. O --'
(table 2) ,o.e ",.r.lu.'.tr evidence sh-ows definite A:.. ..o. in of
'Q h" A GcoantOM .o 1 7r var ialo W .'1-.- l?.-= ,.,... .. -,'v .: wore
o tb i.:13. O.ver the ride ..: .1.1 lower .:*...'i1 .,: of ., l;l,. '.,tkA
;Sp ',,-U' i t:o wr7..l' of water relative to y('A .... r.,v ia
5o The diss ol ,'. ,"'", ,1 ',.'.. in -* .', :. is rela.t.iv e) sm l ifn .i,.:>,; .. OI.-
to \ P! larga amounts .' 1 in the Mi. surface strea %m %'. .O
across :. ,. ., ric .. ..:. roo formations. A..'."'.. '.i '
prcr-orties of :7:'-i..? 5,' and ..T seem to be r1,i.ali-le. .v.T r..
nu.-.l:ne waters ...> .iL3.. ; into artesian Squifers l.YOcu.:. 't- ,-4 ad.
basic and lose the excessive phosphorus concentrations. A complete
discussion of this ,... .',"i with ..,.. to ;in:,'s:.: was subitted ;tx
,re)-po-'t. to the fl.orida G'rl*.':..- ': .*...... :, ?A .-I T,, i' .r with assio tance
of Richard A.., "'-n. ..j- :... : :."' ;t.,I. 'w;A i.n Flori'ada e waters, P.. .
to tho ..... ..- .- ., iT,'. ,.v :1 193 tontat ,v W:-
considered for. :.' o:;f.. of Fla, ~ Snrv:-y,, 70 pp ;,...':- .* :-.) o Th
3 include comparison pert ai t:.iE to phosphorus :,*..: n*.. :r*.,
od cte I .Y of 'r T. '. -.: *n ."
1, CHAtRAJIf, L *TAM 'D ST. 0..1A7 OWF WI-IUTIVm; ; i':I'Ti
To date 1."rr0 1jrs *. .e1 rV'1i (-i*-An i and to.) booeen
made in 33 *:' z.i t to Cel ..2..' .. Bma .gni ,.n., <. o .iW..,1 and iT; of
plhoqt. l in those con,rt:'. ,, r I-;.r;..', l: flow systems. The a *- A:--:3 i
in LA '.*:'; .. haLu 1. .. .. .. with ..' p . o .. n.. 'n, V.'t i trn at ams,
01 C POO i ; ..,'.-estuavies :.'.1 with ." .i. ci..I support of the .' :.1''..
Result s so for hiwZfe C ."i : following '.or.:- of pho pbaruas
distribtOtiona
1. In tbhe '-, r '. flow the /pho':tern t content i
A- const Xt ,iuth distance .'.n :: rvn even W.t, ; -
.. '. .. T 3ta .'. Silver 1,:L- in table 1 inA.oalto
."," *propsrties,
2. The ..*, '::.:i ccr.onte : F thee clear waters in .:.s :-Ors 13 in.
^te :5 .1
3. The varlation of ,' .,.s conant with time is low, f-" c i ,
that. va? with ninsre ''e. *:.t! :.:r ..;r active olementus, sow n." *'.
sprL-; onviro'ents tat tIcat a;r; relatively constant,
4o The 6.!' *, 7.. ". i,.' ."' -".'. in p .WI. il. .- 'r, ': in conoces .trantionx
is available to o larnt in :;.ij. ?*-*'..r excess reserve amounts (Ite to
the contimwal "' *.* of vater. In some :.1,0 m.u; s ndh as ".1. O --'
(table 2) ,o.e ",.r.lu.'.tr evidence sh-ows definite A:.. ..o. in of
'Q h" A GcoantOM .o 1 7r var ialo W .'1-.- l?.-= ,.,... .. -,'v .: wore
o tb i.:13. O.ver the ride ..: .1.1 lower .:*...'i1 .,: of ., l;l,. '.,tkA
;Sp ',,-U' i t:o wr7..l' of water relative to y('A .... r.,v ia
5o The diss ol ,'. ,"'", ,1 ',.'.. in -* .', :. is rela.t.iv e) sm l ifn .i,.:>,; .. OI.-
to \ P! larga amounts .' 1 in the Mi. surface strea %m %'. .O
across :. ,. ., ric .. ..:. roo formations. A..'."'.. '.i '
prcr-orties of :7:'-i..? 5,' and ..T seem to be r1,i.ali-le. .v.T r..
nu.-.l:ne waters ...> .iL3.. ; into artesian Squifers l.YOcu.:. 't- ,-4 ad.
basic and lose the excessive phosphorus concentrations. A complete
discussion of this ,... .',"i with ..,.. to ;in:,'s:.: was subitted ;tx
,re)-po-'t. to the fl.orida G'rl*.':..- ': .*...... :, ?A .-I T,, i' .r with assio tance
of Richard A.., "'-n. ..j- :... : :."' ;t.,I. 'w;A i.n Flori'ada e waters, P.. .
to tho ..... ..- .- ., iT,'. ,.v :1 193 tontat ,v W:-
considered for. :.' o:;f.. of Fla, ~ Snrv:-y,, 70 pp ;,...':- .* :-.) o Th
3 include comparison pert ai t:.iE to phosphorus :,*..: n*.. :r*.,
. .. :: i t *.
',-1. 6
T:. -' .. lpes in .' a.uer
Aug. 9, 1952; cloudy, 20 %w.,
.,.-, 'X.
1/8 Mile
&T I)
1 ", '*;
2 maile
r; milos
3 miles
5 nileo>
Mana
-. 1. '. "- '. ,atiO
'. with lo.a error' than
.'*., REVA
.041
o045
.040
.051
.043
S.(A
!: U'
*1,
(*rt*
( .'..'. .- '. 5 .'. of total .. ? -*. .. ..)
I
1 5
9 ".
Boil
t arlt 03 .loa rn T'
in ,ittoral I...1 .
0050
I ,
o047
.053
,046
'0.)
IF
Li
-P
'A.
Aug. 16,
-.9 3t,
June 30,
rr '' ;
,061
.027
,,
~1 ~~ ,;?a-
S-'1 ,
' '1
. .a 2 ;d F ,t';-r. 1
" v r,:;. rties oa : Green W:'..-, ...'
* ~i:N '2 c
--A,,,
<'1 V2
* ,t.'
A :
,~ 11 ool
0 D
We yo, v I "AC)
S....r 16; clear ... '. .'o:
a'ci-' :..... : 2
*.*_ *,: .. 0 ,
i,'I ci. P, .*,.'-:
Aug. 10 .' .. ,
:..r ;.. ;.. P : ,
To"',. P, ppia
Silfur BacteriLa
Ye'O-.? .- .:.<.:... 'Lotl
;aa
53
,,0
,,022
'. w .i:.., 'i I A .
f '
"2
26.1
1,6
.4
.0.
-------------------------
Jan. 27, 1953; clear :- .:, A
(X2, ppm 45
*'U0 "<'i i'.-. -
Jan. 27, 1953,
O :y ;.:'.,, ppm
002 ppM
Nitrate N, .,
.-<..'C' BaWcterit
?3ti.a
r'in:,*., 10. p.m.
0.0
.4
0.00
I
- --------- re p t-- --- --------/
/ bare spots -------------- f
2
:I
i
- i 2
E
I?
t.... c
*-- r-- i~- -- -"nc -U .r~-rmr -rr\ r,-r~
25.9
4.6
.041
4.0
4,-0 j
.(.' !
5;,.9
4,8
.018
3.4
0004
,6 ,'-
;4.
5.5
.008
3.4
,005
.012
26,,7
6,1
3.3
94,
(2 .2-
1o6
W
2.4
169
24.6
2.6
1,6
1.4
1.4
asK
9,
12,3
2AO4k
2,2
1.4
0.00
2,1
1o6
.02
2.2
2.8
.09
2.6
2.2
.17
v -C W; A W -' -, *
~1~,J.; '
r. r-.L I..
; B -C i SP .9g Pa !.: :02)
..... *, .. in ,.a .* o Florida Waters
S(' of csos in parentheses)
". ".: .. district Other areas
:" 8 r) '".:^ (44)
S. ) .044 (21)
,.. (8) .6S (3:01
o0 (27
' .',t '.J.... .' 22 ', ,
MoanBuremontoI; :9r.'" ''o hT. weMO;P ..i i,,;.A1 to verW;.' the Oessental
o .. f n : r71 cn ::.centt h.;: W. ':- as one paf;aa Oiuro$a
the 5 mile 'SnA l., ., with an '. 1, .. ... I. of 12 ~, -... ,
the v.'- A.. -'.'. f in Silax r ar o i 's r ;' : \ 23.0, 22.5, R:O. .
*1 .. *. n i, % V 0,- tho variation is within a L*':r. even when the
air :, c.. 1.' low for: ave' winter er : .:. ..n o''.0 One Vigtoe '" t'heO Iwa'te
A:) ".. o :at boil ; .ar '... increased to 24r.0 5 miles r..:.'i th
river, La.rge: .- are. to be *., :.'...' in P ho lower reaches of more
S-. n. spr* w,, rru
The Wator '. tia at" a tito when th- air ..?r-r-,atn, is ,;!" o
lower tha, tha-t th' th '. t praure ca be '.,.. to locate zone of mixing
and onri ....' '.,.. w'. a To '. -'. Ta.. ;.'i iC- 'j;' of hori ontal
,y ""e 2
t;rkpe ran., r,.o, .IV ~. .a st LI r'itr. and ', "il *-', 1. j..
Portiona .. Silver o., l .a..- RA:a
18. 1952 .).> )) ,in 1.'-, .
:). ,O .0 tenths
l .
Productivity of Floridae Springs NONR 580(02)
Page 9
The current structure may be postulated to be relatively constant
since the variation in the rate of discharge of the springs is small.
Ferguson, Lingham, Love, and Vernon (cited above) show an aual variation
in the discharge from Silver Springs of about 20%.
The map in figure 3 was made with a midget current meter in order to
get some idea of the patterns of current velocity in horizontal aspect,
which may be important in controlling the distribution of organisms.
Figure 3
OCurent Velocities in Green Cove Springs
Feet per Second
Width Scale exaggerated 3 times
pool
CI s
Seep
6 ahes
Js
Productivity of Florida :~nri":i NONlR 580(02)
Page 10
Many of Florida s springs originate in strata which still contain
salt possibly left in the pore spaces by the last Pleistocene flooding of
lowland areas. These springs are slightly saline, some in the brackish
range. The arrangement and pattern of springs and runs are especially
adapted to testing effects of salinities on natural aquatic communities
in toto.
One group of 4 coastal springs of large size have parallel runs
710 miles to the Gulf. These spring complexes are, in order of increasing
salinity Weekiwachoe, Ohassahowitzka, Homosassa and Crystal River.
Initial analyses of the calinities in the runs of these springs are plotted
on the map in figure 4. 'Thiv,.Fr boils have salinities logarithmically spaced
but with the same cA.1cu1.1 concentrations The run, as they approach the
coast, increase in salinity first due to smaller tributary salty springs and
ground waters with grpa.l.11 increasing salinity, and finally due to tidal
waters in the last few milos. A strong distinct tidal wave moves up
these narrow runs many miles beyond any salt water of marine origin. The
wave affects the pulse of the spring run but does not reverse the flow except
at the mouth. The goochemical factors which are in operation have produced
gradual salinity gradients nich are of nmch more constant type than the
usual sharply fluctunting estery.
This remarkable natural salinity laboratory has already been the
basis of some investigations discussed below the invasion of marine
fish and crabs, the distribution of aquatic plants, the distribution of
insect fauna, and comparisons of fauna in stable and changing parts of the
same stream,
Further comment on the geochemistry of these springs is included in
a paper ready for publications The Geochemistry of sodium chloride as a
factor controlling the invasion of marine fauna into Florida fresh waters.
This was presented at American Society of Limnologists and Oceanographers
1952 session at Cornell in September. The essence of the biological aspect
follows below
Productivity of Florida Springs IOTR 5i80(02)
Page 11 \ _
Figure 4
Chlorinity Analyses in parts par
thousand
Crystal River
N ov. 29, 1952
S /" Homosassa, Springs
Sept. 28, 1952
QQ(
SChasahbowitIfka Springs
Aug. 27, 1952
Weekiwachee Springs
Nov. 8, 1952
0 VAu,
--~-~--
scales
Productivity of Florida Springs TONR 580(02)
Page 12
2. 4PUALITATIVN0 COMPOSITION OF COMMUNITIES
By far the most time has been spent on the initial mapping of the
boils of 40 springs using compass and tape. The dominant vegetation has
been plotted in position, roughly estimating the extent of plant beds.
A typical map is shown in figure 5.
If as discussed below many of the springs represent steady-state
environments, the conditions at any fixed place within the community
represents relatively constant conditions of current, temperature, nutrient,
and predator exposure. This horizontal spatial patterns are required in
order to compare factors and organisms. The methods for est.-i:!en i stand-
ing states discussed below require a map so that quadrat counts can be
multiplied by the area of similar habitat.
One result that has come from this work is an idea of the stability
of the vegetational parts of spring communities in Florida. Little Blue
Springs, Gilchrist County was mapped a year ago. Visited 5 times since
then, the pattern has been essentially the same except for a large bed of
floating JAFA which comes and goes primarily because of the dislodging 1y
summer swimmers. In gross aspects these communities seem indeed stable
systems.
The springs for which initial surveys and maps have been made areas
Ichatuclnee, Alexander, T;..hr^-,.., Blue (Gilchrist Co,), Blue (Jackson Coo),
Blue (Volusia Co.), Blueo (hrn1C.r'.'), Buckhorn, Crystal River, Crystal Springs,
Glen Julia, Fanning, Axniper, Lithia, Palma Ceia (dry), Poe, Ponce De Leon
(Volusia Co.), Rainbot., River Sink, Rock Spring, Salt, Sanlando, Silver Glen,
Sulphar, Wanm Salt, Wakulla, W1ekiva (Levy Co.), Wekiva (Orange Co.) Welaka,
Mud, Silver, Manatee, Orange, Homosassa, weekiiachee, Chaseahowitszkn, Bugg,
Morrison, Bonita, Green Cove, andQ Sn No Wa. Locations are given in Linghat
et el. cited above,
As a basis for all biological studies, the major taxonomic dominant
in the springs were collected at the time of the mapping. These cursory
sanples, representing only several hours collecting each, have been taxonomically
divided and the fractions are in pro~ese of being identified. The identifil
cation of the most abundant species of higher plants, insects, and fishes are
essentially completed.
Two striking results are emerging from this phase of the work
1. The spring communities seem to consist of a few species
abundantly represented.
2, Almost every spring differs as to the dominant species even in
cases where the chemical analyses of the major elements show
essential similarity.
Examples of the living components of spring communities are given in
the list in table 4.
Prod.u activity of F'Rarida '~1:~ Nr = R 580(02)
Page 13
Table 4
ENamples of the biota of some different types of Florida springs
(only a tentative classification)
...OT.~JLo y. ,.3..!J.n., Temp. 8?.0, bacteria, diurnal light turbidity
variation of peculiar chemical nature, gf s, -SJ G,
SS.%A,i frogs, blue crabs, requires Rideal-SteJswt modification
of Winkler method.
Examples Warm Salt Springs, Sarasota County,
Awn.robieM Teamip. 75, Sulphur and iron bacteria, blue green algae, midges,
'EMi '." .' ,., '11'Ag w, water clear and initially
C.l. t,'.Lhr..,u ...
Examrplea Orange, Groen Cove, Beecher.
SUM -.2.,1s-^ .-' ^ -,, ..102.1 ..- ... .'.nC' Temp. 75, :-^ .' Estan ajtan
a~e.JEf: ,.~'.n und froshwater fishes and blue uraus, heavy
green ;o n:'. t; .,1, .",''P.%t,,., amphipods, water greenish.
Example J.,lrt:.....' &alt, COhasahowiLtska, Silver Glon.
r Miat mih._ai anr- .cvacntigi, -. .. 74, clear, JAEte a, mullet,
":"'* "iv," r'-' S. ilver.
I ''- J'>.. _'L .' ,, ,' ';::xy ge 5 1 ..... "- '' na il i n. ; p r.' .: ,, i.,>n i
mixid. v tatcii; o n, t:i) .' H. :.- -. etc,,
c .. S .;; (rlbrint)xd, Ga, crayfish
h ,.e; ....J, coveredd~ at tieers by back water of tebid river)
vegetation almost completely ~CQgthI *
aEraple~ Morrison Spring.
F.te igc~ aggan West Florida, Temp. 69, EqaB oxygen
7. 0 ppm,
jPro iviv'. of r
Page 14
* .
-I-
- -~t
i ,'*
'
'
mlUm~N~a. I
,J/
o
a
IrB~s
\j%
o^.,
0.
Id L
CL.
Productivity of Florida .;, in3-. NOYA :"A 0(02)
Pace 15
SThe collections discussed above hae suggested the phenomenon observed
in pollution where communities have less varied composition but many
individuals of each species and where the particular species that is dominant
seems to vary radically. Dr. Ruth Patrick has suggested in communication
that a stable environment like these springs may possess fewer species than
a normal fluctuating enviroent because there are fewer transitory niches,
which in a normal fluctuating environment allow many species to at least
maintain a foothold. In both stability or in pollution one has a deviation
from the norMral fluctuating environment and possibly for this reason fewer
species.
As part of work toward a masters degree under the direction of Dr.
Lewis Berner, Mro il.'lli : Sloan is studying the distribution of insects in
relationship to contrasting facd;ors. The following section, written by himn
indicates pr~t o c;:> to d.ato.
Contsastin~ o' t':.er-' of Dnseot Distribution
by
Willia 0. :., .on
I":-.:n;r (."''' S :*..r.';,:. '.. that the .-....1t .Ic insect fauna of the upper regions
of tlhe Florida, prXing was scanty nand that the population increased a short
dist.anoa below thz boil, Thi' c;"l.*lrv'1...in tvas recently substantiated by
.e3itiUcll1:?:.;. of a series of collections obtratiin by Dr. H. T. Oditu from some
35 spri';',.:.' i,- .L;.ici.'.ve of the '-s:-uit study is to explain the oi:..city of
larvae in the boils and their relatively sudden increase below the boils. Xt
is also hIpr-i that some knowledge of the effect of small amounts of chloride
on the distribution of aquatic insect will be obtained.
J. -ri ;,'-'; the l'.' of 1952, the insect fauna of Silver, Weekiwachee and
Homosassa Spring (figure 4) was studied. Collecting stations were established
in the boil and rn of each of the three springs, and chlorides (figure 4)
and oxygen ,..*,loo:.ie were made at each station, Collecting apparatus included
Needhka s npe', dip net, ,;O1-t? wire screen buckets and strainern for larvae
and a butterfly net for adults, Adults vere taken whenever possible in order
to aid in identification of larvae,
Oxygen analyses indicate that, in general, dissolved oaxgen content
(DBo.) increases with distance down the run. The D.O. in the region of the
boils is fairly constant while that of the runs fluctuates from day to night,
Table 5 shows the range between day and night in each of the boils. Figure
6 indicates the total number of species collected at stations of varying D.O.
in Homosassa Springs. Table 6 compares the total number of species taken in
each of the three springs with that of Lake Wauxberg, a north Florida laki
located in Alachna County. This lake jwas studied by Mr. Richard Trogdon, then
at the University of Florida, in 1933-34 and the data on Lake Wauberg were
obtained from his unpublished masters thesis. The species list (table 6)
indicates the aquatic insects so far collected from the three springs. It is
almost certain that further collecting twill increase the list of all three
S springs, but it seems fairly well established that Homosassa has the most
abundant insect fauna, Silver the least, and that Weekiwachee occupies an
intermediate position,
Productivity of Flo.:.. ...-'-' MJR ,00(02)
Page 16
Several bypotheses may be advanced to explain the scarcity of aquati.
insects in the boils and their increase as the run progresses. As indicated
in figure 6, there is a possible correlation between D.O. and distribution
of insects, The boil of :-iorrs.as is the only one from which truly aquatic
insects have been taken and this has the highest DoO, of the three (see
table 5). The fact that even here they are scares may indicate that there is
insufficient food to maintain a larger population. Another factor may be
excessive predation. The wi-ater of the boil s is extremely clear and the
vegetation less dense than that hlco~r the boils and it seems possible that
fish predation in areas of L';.-,ufricient cowr may prevent the establishment
of large populations, In addition to DOo,, food supply and predation, the
nvwater of ocologi1al nicho available Lust be considered. The relatively
small number of species ;'- :.' t in Silver and Weekiwachee as compared with
Roiloos~i,,s may be o-:.-....ir .' the greater number of types of habitats found
in Homosassa. Silv-r aiod :.. :,..\..ih.r both have narrow runs, awift currentse
and 1w turbidity for the 1?.- ...h of their runs and practically no areas of
still water. The run of 'cr 7orc.-., on the other hand, broadens immediately
below tth run, the currexnt b~conmes lover and the tiurbidity greater, Along
the -'i.ti., in 'b --?'..:-r.- ares, the current becomes almost noticeablee. It
is in this type of habitat that the mayfly QaMRM &tfA3 an and the aquatic
Hemiptera EpI.m ;. and.. Q.?. c .; r : all predominately pond and lake forms
are found. That the nwmnbser of speciess of insects in Bomosassa compare
favorably with that of a lake mEV be seeo in table 6o
As indicated in fiIgre 4 the chloride content of Hoamoassa is 16".r
than that of either Silver or WIekiwadceeo E. n though chlorides are
present slightly in excess of 1,0/oo in many parts of omoasasa run and is
thas in the braclkish water .;-., it seems to h:vo little effect on the
insect population. Xt is likely, however, that further collecting will show
some forms to be ,;i'itiove to even these small amounts.
Methods of sampling quantitatively are being perfected. At the present
time, emergence er".:' are t.1i psal.-,.o c. at various points in the runs for the
purpose of ntJ..' ;.t..ir relative P i.-nd.in..e in different habitats.
Table 5
Oxygon range in spring boil areas
Silver Weekiwachee Homosassa
02 2. 6 p
Night 2.,9 20 4.3
Productivity f 1.c-clt Spri :.;' NNR I0(-02)
Page 17
Distribution of :9".Ti)cr of species in relation to disvolved ry:vn
in the spring boils
02 in ppm. in boils
1 2 3 4 5 6
7 8 9 10
02 in ipm r'''!. ;tUples. Each point represents one collection
at one station. .C!'0t:3.7 of species indicates the total number taken
in one c.lc.., period (1 -,2 hbr.~/tation)o Lettera indicate
separate or identical stations.
Ephemeroptera
Odonata
Anisoptera
7gopt era
Hemiptera
Ooleoptera
Lepidoptera
Neuroptera
Diptera
Tendipedidae
Heleidas
Stratioqyidae
TipUlidae
Tabanidae
Oalicidae
Total
Table 6
S.-.',,; of ;r0etoi0 recorded
S.i.' P Weekiwtache Homrosassa
1 4 6
A
0
0
0
fta
-ffrlU-M
Lake Wauberg
2
16
7
8
6
I
1
1
1
i
6
1
1
6
1
n
A indicates adult observed but no larvae taken
* the Tendipedidae have only been tentatively identified and
5ay consist of mrore species than are iadioated here.
rCU I -
emnBeahnwrM...c,...^.m
Productivity of .
Page 18
Ephomeooptera
Oa Uibaotie floridanu!A
Trichoptera
Kvdrootila op.
Hemiptera
Metrobate guomalu
MeGovelia a!"gantt
,Iagovolia chorouton.
Diptara
Tendipedidae o epeoiee A
IseAtarg'e op.
.;i Fi580(02)
Thble 7
Spooio List
Silver Springo
Oolooptora
Gogolagtua Qalagtgfilriv.
Q. chvrolsi
Gyrinua aMohsa 2as
G.o gokin anlas
Odonata
Zygoptora
Enallarma ape
-ialn rina hap a
Nohalannia sp* ,
Weekiwaohee Spring
Ephomeroptora
fTrico hodegge abiluco^
Stoi _____*-
Baotia op.
Triohoptera
gayoptoilj ap.
Leptoocridae apecles A
Homiptera
Motrobates heoerius
Trepobateo vgictua
Rhaovelia choreoutoa
Rydroumtra up.
D | Cy I ap
Dipteora
Tendipodidae apeocio A
Tendipe! pluMaosuo
Tanytarluo ap.
Odonata
Ani optora
Ashil MiSarcel
Hotaerina titia
Zygoptera
Arion gmaculatum
Argga species A
Argpi species B
NahMgg nis opeocies A
Nehalonnia speoles B
Lepidoptera
Elophilus ap* ?
Howmosaswa Springs
Ephemeroptera.
Oallibaetias flogidanu
oaesn1s diwminata
Caenia opq
Baseti pinosuas
Baotisa Bop
Tricory htodea olilinea&us
Trichoptera
heuis at pch ap
sgpoptggg .o p o
gonopgahq op-
Sdroutila up.
Hamiptora
Plgogr uap.o
Bolostomw luterium
Mesovelia mulsanti
Diptera
Tandipodidae s apoiesa A
O B
O
*1 D
*" E
Bazzia spo
Qdontogria ap.
poophelo ape
gulax apo
Odonata
Ani optera
Perith3mia seminol
Libellula pulchella.
Le auripenni.
Gomphus ape
Productivity ; .. :*i'j
b~r'>ic Yk'
Hariptprn (zo'tS)
o ... ,
rf ---, ... 1 1 v ,
I. :." '., (co:A.o
[ .- -.
Lii
SB
I.'..
~ >50010
doaM != h. acvn i ad on 'h i t1, d cHO' o V '
in tWhe catiocn o w .i : ly/au lne an Nuo c: '. .-'- 1 tho j.anoral x
for the z Won t : 0 j W : .' z down taa rua hnoe bin*
W 'l.o..,: !, LA cl : on d n3 diacuunod in t1w oC.Ation mi_
ducS' iv of t:: :' : : 'y
A* dica...ad i S Q-oti thn om n .: c0iw GU n
4d t. W. .edn o to: f n-:;-,i cyn a a masmro of voiat illity,
W in dir "O.W, Q 0 ny.n" n A 1 Q *p.aclow Ondor eao" 7 COUVNTio.s
* o C poM.i'^^n pr lw v n W:l 0 ; i ::: ro compOr !t;.;ons b:otr:;;:.;n b10il ''
u'rthor ch' aa an 2.H!X" O .M to va W1: ulai I toS. oA.
are Wto .. : .nvoc;A' W- :;i oemnivs in In,- kno vl or e m
4 1incnt 0 0 1 ::l by h ll :1V .M A MI ..Vwo
TO ot bli: OT Ala 2
O U00001aoc 101i Arcs uld
% CAleivly a VnQ
uniral:n c?'urmv WO ch"wa ;
crs'amon and l.;o l .:I -.i"o f
W... 14 .A .. 1. .
. botri end blu : i
U.S.Ha.ci':^ ni "Ta- ho 'hr0s o t hus a V:,v
ono qmoa downcatroy nad no theW iSn i
3 dcri.'? ono
pKo3 .c oo .' l va a p r n t f Zgri vnl=lt .i
in th*o ,on : .o aI4' 7 n ol:c vatsr. (on 0rook W: crop.; A c
' .' ) Qo i?:t' ialla y r' :. :1:' 1. n l o < t3 4o.: all7 is-e 000 W fo d yadi "A inQ.
knott -./ blu I s i 7ble that' thoe .. of tO Ai -
coito s ds i d f n3 Wg 1.a.:t in tha vm .2t' t=!.r U is ta
" cr'oolationC th ;, .. v A:: on the otia do but0 obu xit.AiK on toh, I 'r
tho rverre:o ." t K; ciNOW iltibn ta tMasoPU in 0.. r7W.'.;uQ,
I.!i /. r; '4! 'r
O r S .ca & .' -. C oM.'
L
u
..:-.~ ;
.s-- o-.- -~ 1711
Productivity C2 : .. (02)
Page 20
A4 iv"- Q e& t ,'rii for- r' .
S The presence of c-',i-uvnl of microorganisms in steady stat. in thooa
flow systems suggest ta z the idea of an ecological formula for a c,!:
is v'.'.-i. P."' i.;r this ol, tFtheorem is here practical, Perhaps onez cmn
list the oPr "itiona in : 'hi '. ya ijtl1 will grow pIu'i.,ln~ntlr;,t i.n ': ''* otate
in natuzra ,ondition : c"....::.". '-.t ., The clnic'l conditions Iknot:a for
anaeroble Gm:;..'. i Ocvo ::.r-: ar given. irn "-:,'L.e 2. ^:,',..-' d:a.: : et 1g.',;: o4"
t (citod .,r'. '.*.*- : ao ilol, 15 pmn; Iron, .03'; calci'u., *',.; .: i. 16
c..."., 2,..; "." .:iur ,, 1, *; ab..,-oa.. te, 100; sulfate, 49; chlorilon, 5.7;
? $1_..;'kc_.I, .A, ,'2 .
.- T -, ', it', l '. ; .-. ... *' I ., '.
f' G'"1 OF f tP 2 G AIR
. ., K6 C. i : .
1 4,.. :7{.'T '.. .:, c.:"X tclp'iL.no.V. ca. live rund.r anaorobic -ate:'
conditions &ueo t.'i, ., :b Tt of .. ul air. :Bcocher Sprins., .'..la.,, A.
anaerobie spir';, ;xotin m.'.ry of thpso fish 'hich are contiiTVlll; vo, :
A nimter .' eac.h p ci.s, s .. m. L.: h sp '. re
: in screen w o a c and 2 f' in '- ... : .. V "0
S .i-:--;''". so that t;. fi- ca.ld. ..?.2 air arei. the othorea Ci ./. at dif.: **..A
depths from ono 17.ch to t .oot boloew 'th ----. c so that tho firh coulA
not roach the e.'rf.'.c to ..:"~..
Those which .., .. .ir livdc without r'-r't di stras fo th
i;" t.ioz of th er' -. ;' "-'. hoUr. Tea, whieh coald not Croc ri.le. .
u'l -1. almost o.. o sin of diistross and. wore all deladat :. -;Otha, in
abou-t tw enty li.to .'
In ord'erI to ):u :; '- 1 di. not have'x-, o to ;.) air, as a .':.'. i'
life 7.;.'O r :. a. ,a,".n of :nrd.i-vida:ls were on...i below.the surfaceo in o r*;.: :.,-'.c.I
water r .': dlown the rvn, (1 1 ;.; 02). TheB ose lived .:. I1.7. without gulping
air until the oozeriment wa.s e.-.' ..::,Ui ", in thirty minutes.
''..i'.,. "- -~ :.. ., thA't the e t'..' -::.7,'.;ric, of fish can in r --i.litr
occur in waters with .l- o1.a o *;, t' :ly whicEh will not support other ,..:.' :1:
which do not gulp air. t i; clear that existence in Beechor Sprimn o..1:,:le
on gulping air. No other ;:-.r'. species wero observd.o
Peat y'0,. i-. -c:I
A preliminary experiment on d.eoay of wood and leaves was carried out in
anaerobic Beecher Springs. The floor of the springs is covZerd with brown
Fibrous peat about tt-o *r.t thick over which flows the very clear, anaerobic,
.,o c,.ct"i. .vit.y & Y_ .:,,'(02)
rP.-.. 21
sulfide i smellin water.' :' ~. rate of peat formation under this constant
temperature (Tl'.r'. 23.0 do., 0,), and constant chemical condition oomsd to
be '..a h c Ih.::.,;,, A h.ar.cl.a cloth n-;:: containing fresh elyat, ;. t-,.
w! ,'rs.... a ficks and. lsavoe a .*::*.X 2r:A on Juno 5, 1952. when rRozEcMned
on i-.... Iry 4, 19r -, :" w..r ood was ,fon',L to be essenti-llly ',, .....I
in yoss appearaavcs. The 'eo r.nl.ico.,l.d. so little modlficatioa that
no analysekC wero rk. JTho o:spo;'i2;nt continues. Control samples arVe torod
in the 1i ,i',.:..,;-, r.:..'. thoe ?'" leaves adbo decomposed. I, .:"7:
tr-i.'i.l fskoletoaB rCI': as ix;-.t have boon expected th.":t
*.:. rlo.''l,;'PJr-to :,,; ,*.l:.ion ixnor anaeirobio conlitiona is slow comp- ',* to
other fractions of the 3.,,..
The pheno.onon of t': ,:' invasion of 15 or more species of nmirio fishes
and the blue cr' ito ,l:.-..T .rulh waters and particularly springg lr oas been
studied as a nm-:. -":. r piroct; :ldoriceo maps for 1Florldaos freah .water have
been p;.: and t'e d.-iA ,t,.a of marine invasion correlated. ,. maaanu-
soript sumrxvrTIl. .'I; ts- :o a-' theo springs work bas been prepared for
publication (title cit:Oi :.-: .), The essence of this study is a series of
maps of cblori'i" cor~~Or: n i :A :-i..'.c.." esltuarine and .f:i~c waters, a i.i;.c.-.aion.i
of chloride geochao.i'..;: c :..ose'.pt...:. of the areas which reooiwv the toc<,.test
n.M?.-, of mari fii, ." ..: traplant .:,:,lc'l;',E,., of bluxe crabs in the
springs. 'T:i'l 7 sui-::wi o t he t i. on transplantation o..o T .... ., 'il
correlation of ':..).-: crat ith esenea of 25--1000 ppm roii..a cborido in
springs of constant i-.. *.:r ..1r the ability to survive vlo2i;. wbhn tros.a
planted in tho3 ep:;:il.".- U, :; t that inability to reab!::.,ib sodrliv clo.i-.'l.
from low con eontrations in the .:'i.':-..;..' is responsible for chloride ;noaitivity
as y 'Kr;, ": oa the ..u.rp.,.:b wool handling crab.
o 7
Transplantation. of :Blue Crabs AiA, in M i Florida. Springs
Crab Source, (Date (Date MC No. of Spring of Ca T l1 Days of
1952)) 1952 ?IP crabs Expt PPM ;.;,. PPM Survival
F,
Bayport estuary Feb. 4 3,300. 7 Homosassa 50 75 570 >14
Bayport estuary Feb. 4 3,300. 5 Chassahovjitzka 49 74 53 7-18
Salt Springs Aug. 7 2,800. 8 Silver 70 73 8 5
Bayport estuary Feb. 4 3,300. 4 weekiwachee 49 74 5 1-
Barport estuary Feb. 5 3,300. 4 WeekiLachee 49 74 1
Chassahowittka run Aug. 22 730. 7 Rainbow 21 74 4 2
----- mmm-lra .tw~,,*rna'"n crr mm i~if ln;I r an~*1~
Productivity of Iluida ::. .:-,.?: NOifm 580(02)
Page 22
30 QUKANTITATIVE COMPOSITION OF COMtMUITIES (STANDING CROP)
The quantitative doscri. ,lon of a natural aquatic community is
difficult because each typo and size of organism requires a separate
means of sampling. It is only the essential stability of the spring
that permits one to do one thing one time and come back to finish the
job after dov-lP:,l't: neo tw.-.O.:X.n'to.e
To date in .i"i- :1 nativeive :~:r li;L : has 'caF carried out in the
largest spring (:.!l-.':) oWe of tbh sialler springs (Green Cove)
Whereas the job i. the. tc .. even in these two are incomplot.x',, onov',;h
has been done to otli:no t*:: ..'. l of p'.:z.ui. of mass and :;.-'
as in figure 7, Th nt3Fo of the pyramid is evident. Thi Is suggestive
of Dr. Archie Carr"s thoori;, .;t.i-. food chains as applied to marine
high plant Veir:".ti.io:. Co7n,:<::t on *.;. ,'I., is discussed belov in the
theory section.
From these ?'.., :. i..:. it sem~8ii O.,i.o..0L that a valid picture of
standing states can be oln1ained these ec..cJc. .rou,' spri, The
clarity of the waters p:rits one to census the fishes by eye. AI'I -., 2
rough, it is ..:'.1. of Tvgnit. --- with which one is here at first considering.
Start of ;:. '~ on ;.. is ,. x..-....' ",-1.co by Gordon J.rco,' h.- .,
Figure ?
P''i;' '..i of Mass in .it ).'.-' Springs
`1 a o at ycEst Y
S. ..- . ...
.......... ;....... ....... o ,
~i~~ . ./ .... .-- .--::, .. .
Productivity of IF'lorida. S2p' ;i. 5380(02)
page 23
Methods of quntitative sampling include planimeter measurement of
the areas of eaeb type of association and counts and weights of samples from
each association. Sampling procedures under way are as follows Algao-
scrapings from known plant weight; higher plants--by sq. ft. visual cropping
under water with face mask; microscopic organisms--scrapings from known i.l-'a
weight and pouring through plankton net; invertebrato animal---sampl ing with
sq. foot grabs with box samplor; attached microorganisim-counts on 2Afmor.vl
glass slides which become coated in 3 weeks.
Dry weight o''qiv:l-.~ ,' aree being obtained for field wet weights.
Special attention hacs boon paid to the aquatic higher plants, their
weights, and total tonages in the spring runs. In addition as a b .so to the
food chains eand. pyram.is, thiI information constitutes interest in itself in
relation :o -.: ..e. i .pr:portCie of the springs. Dr. John ., DIavis hkars
studied -~ i;i qiant.i'ak tivre vi:.-tion of the plants in relatioY o0 the chalsn
ing proportios Cdown the csal; rims i2nr.ied. in figure 4. *,... report follows.
i'.;N' U." A,";' of Aquatic Vegetatio n in our Springs
',.nd thoir 1Runa of Florida
by
Joh1n H. D) '.. 1, Jr.
This is a 1.-:.':.: .r-:'r reOport of p~'t of the investigation of the
biolo.ic.-.l produ dAiv2,:;: of plant communities in four Florida spring :.: ,s.
These areas G.;, 1, HoAoasa, i ''t'-ts, and pictured
with chlorinites in fi., [4. The data in the present report include the
weight per unit aroa of subma:-. ;.d and floating aquatic plants. The waters
of the four :prl ; systems flow into the Gulf of Mexico, all of thom entering
the salt water marsh and littoral zone within three to six miles down stream
from their spring heads, Their waters therefore vary from almost no salinity
to salinity values thl.t are over half the concentration in the open sea.
They all contain hard waters with greater than 144 ppm i..nLL.'h.,o and thr'b,.out
their cotuse the mean water temperatures during the season so far inve;~.t:. tod.
did not vary over 5 ... : .n ..ti.do (20.0--25,0).
The chief objective of this part of the investigation was to determine
the density of growth of the different plants or groups of plants in terms
of weight per unit area and tlas arrive at some basis for comparing total
plant growth in terms of salinity, turbidity, hardness, and other factors,
and some basis for computing productivities when growth rates are established.
The main rtTlsd emilored was directed 'at first toward estimating the
extent of each characteristic type of area of plant growth, and second
obtaining representative samples from most of the distinctly different types
of areas. The first objective wnas partly accomplished by inspection of the
runs for apparent differences in both density and species composition.
Sampling was then made in areas that were most nearly representative. This
very selective type of sampling was supplemented by some random sampling in
certain parts of these systems so that the sampling was about 70 percent
Productivity of ',,.i -. '. ;, D" (.
selective and 30 percent random. The latter method is recommended for further
investigation. The sampling was accomplished by collecting the attached or
floating plants within a onoe-aquare-foot iron frame. From these samples
both wet (green) condition volumes and oven dry weights were obtained. Th~
greenEwet plants collected after draining 1 to 5 minutes were immersed in a
large can that had ,a ;n.rlike spout and which was filled with water to the
spout. The overflow water resulting from the immersion of the sample was
measured in cubic centimeters and this volume figure used as the rou4gh wet
weight since the specific gravity of water and the plants are similar. Percent
water was computed from the wot and dry weights.
The oven dry ,:.l '." were obtained by first air drying with caution taken
to prevent molding and for;aentation. Then the samples were oven dried to nearly
constant weight at a t; 'u.- '.stue 100-105 dog. C. The weight val-ies in grazm
wore then used to estrimate tlh pound-s per acre in aroan for which the a*ir....
were representativ-. A fin- enatimate of the approximate total poindsa r.g
acre in the :-pr'],:.-: .'**' rivs;zo was calculated on t'he basis of ynrc':.. ,, 3 of
cover of the yl:vi :,,i aa eoutiatoe in the field visually tb .'M~nt the clear
spring waters. The data nre .. s...- a. summary in table 8, Similar:, but
incomplete data not J. :.'" -11 in the table .11.e,....to that the lower rea.has of
these same runs poaesa 1.-'.. t.elr'J.:ties, diofite marine salt, o3tu.arine
fluctuatioans, : a. aicsh caller C--: ty of +l->:-'G-e. In the clear offshore
marine taters rooted .. :.:-. n..- stationn again b' co,~na dense in the shallovw
flats off the '~..."' cast.
Plant species constit...' ... the -.t' c vegetation were tr. .,. *:, in
nearly all cases .''i;.i ,; for 1 :'.i-LiX '.: .l-,:v.-. of a few algae still *P..',,
Some characteristics of percent water and growth density are summarized in
table 9. This table cen be used as a basis for wet;dry conversions in
further work.
So far, data on chlorinity, oxygen, and phosphorus have been obtained
for these runs. Definite correlations between chlorinity and. specioe have
been observed as in the correlation of ?tya:.~;L!, tSAAt.e with chloride
values between .100 and 1,0 parts per thousand. Observation of/oculiar Soroe
of depleted chloride (o000 ppt) ::.'. heavy 7',rc beds in Weeklwaches springs
is being checked.
The tendency for SA tp&.L to dominate the low. chlorinity waters andfor
D-11UgI 1 to cover the bottoms in the slightly higher chlorinity ranges
h3b been observed. Zones of eimu2ltaneous growth of the two species occurred
nalivay down these runs. Thc;x- tvwo species are very similar in general growth
form. An ideal situation for the study of competition is indicated here.
*1'.-bTl, 8
.u..:.-.'v of a"-.-'"*.; crops of plante in upper
part of 4 coastal spring runs
o I of sq. ft.
vweifheod
Range of
% coverage
Average Total
lbs/acre
dry jo :/'lhi
Weekiwachee Spriv.je
(strift current)
Crystal River
(wide, sluggish, 0--:p:.,
turbid)
GhaBsahowitska. '-.i::.' ,-;
(sla1llow, clear, ms.&dium
current)
(clear, wide, -;':.-.!.
15f-60%
30%-70%
25%-70%
15%-70o
3,9413
2,561
4,000
.007--.o025
.3:) 4.39
.0o0,' .840
Donsity, porcC.t ater, .:.-:.'.: i':.' f.ch-ts of come aquatio
pA.T:.. : in 4 coastal spring unms
Species
localities
sampled
Sq.
Ft.o
Oms. dry wt.
Me.=an ;-ic
%te
iater
85.3
84.8
86.0
77.4
90.5
91.5
113
73
1141
105
59
65
86-135
43--102
74--177
104--~06
56--62
55--115
Range of
01
Ai'
rmr ~,rsrararwrt~~------- -------l~~tn*lruranr~mlrqrL~zr~mm~ri
C-aLlllrdII~"""*"*2~li~~Y11\-I1UI-m
I~-iL_~ICli~~m~(~~~-PI7~N~l~ll~pl~~
rrrm*l^-r~lc^-~l~----nUI-r*lrrror r~nrcbanrr.l~cul-r~;r~~
I" g
~
;;r: 29
ia$ ::~
nonasesatrfino
Spring
I12404 (Macatf?,A)
EnhamthSiliaim ABQiL
SUCIssad, UasQ
7Sj]SO&9x. 2,A1'
USalma& ffi
The ...- of Color '*r-,' in the Stuwl; of 'ish
i''l -tions in ITomaouneoa Springs
by
Gorcdon C. Broadhead
Fmr..q.nri S..-.:'1:;:. provides an excellent site for the study oi' marine
fish prcl.at.: o1u. (): c ''. ;~3elo of marine fish enter t',: spring" area
where tkr, '. o.,-vv':.,t ,''"i' been erected for tourists, and boacuse
the clarity of the water an'. the narrow limits of lI.o .,pli..ig area mike
obserwtiot of t.-il- moveniat s ;r.,*!., Casual ob:,t'vti. : It:. ..:e Uit the
large popaClationa of fish in ':"t. springs chr n-i' from day to day both in species
and in the numbers of each spreies present. By marking with cotloj..-(d P1-j't:j
fish t,~ t-1.j moveoeonts of the fish can bo recorded, the drni~ly.; !*,.:,'.t : .'1
of fish estimated and& ;.'.:* turnover of the fish in the sprinLs: also est ;,.. ,.
ITIp,'L:. will also permit "'. '* of the behavior of the ta;:,.z.d fish .1i.l estAimation
of their growth rate if r .-..;,V~:.'i at aa later data.
;, i' o:,,ter 6, .".'". a :.. i-.a,- .n.,.y experiment was carried out to test the
feasibility i uising tlh co,..,fdi t 3., Nine ir:.llot ::..a, c:.r';hi in the boil of
the .;'pi:.-",; by n, c: of a eia,-nt. The fish were .::;(e. with lar&'I) .nro tags
and r.l :.ci ... ',. into tho .,.. a In a.'::,:...c c rporilmnt a rlf:ff'rfL;.', color combi
nation ,..l 'o cl b for each flaisb The f',lo~.'.n;: morning five of the iino
mullet 2 ,'. be eeen I: ,.-'1,..i: in the .;;.in:,:, and at one time feor of them were
observed t.o;:'-:'.., in a mal.l school., *"':o::o fish tmere bore.-'d by the .ip:.c.'n
.tterndrd for two moresa .1' .a i:d .after that they disappeared and were not seen
again.
AlilA.,jL it iis -cil:, to draw any definite o ncilL.:.Eri,,;. as yet,
indications are that the irullet do not romain in the springs .q ny great pe.ricrl
of time and that the 1;-..:' o. 11y population is maintained by fish movinE to
and aout of the l:'.'"ri into the much wider run area. The faot that four of
the tagged i...,h were o :':.:,l. in the same mall school could i~.9.1.S:-t that the
t;gg,:,g fish vere not &, .'... ;,;r distributed throughout the ovpuct.m... Since
they were .tuoxi: t. one at a timee, i s t nt likely hat they were in the same
school ibufor' o.)..in;..
IV PRODUCTIVITY
The study of the production rates in a community involves the estimation
of production rates of each class of organisms in the food chain separately.
By trophic levels, there are the primary producers, herbivores, carnivores,
secondary carnivores, decomposers. Satisfactory measurement of the production
of any of these tEder natural conditions is difficult and has rarely been
accomplished.
To date, att.sliptf to measwue production have been directed at measurement
of the primary production in situ of the plants in the springs. Most of the
effort has been directed at Silver Springs, After some trIal and error. some
Productivity of Voi: a pi.g ... ......*2)
Page 27
success has been attained by three direct and indirect methods. Silver
Springs is almost completely covered rith a thick bed of A~E LwhoZ4ao
growth form is that of eel grass. These heavy plant beds are coated with
algae which in places forms heavy mate. Some other plants are found around the
edges in relatively insignif ica~nt quantities. Measurement of the growth rate
of the n,. ..r-'.!, l~ s boon accomplished by the following methods
a<. Growth of wet t,,;.beilt, transplants in cages,
b. Measurement of dom.fstream drift of plant fragments,
C, ZIR:aao'on t of difforonces between oxygen content of
downstream stations in the day and at night.
Measurement of growth of transplants probably produces mininamu values
since cages shut out some 1l.. 't, exclude turtle herbivores, and in the case
of rooted aquationl inrvolvo roo a.ili..l.i-t of root systems. The diroot Loeaszu'e,
meant of plant :<,th involved! two different procedures for two different
purposes. in o.- ': to ia~ra-ure the production of a community where a study is
being made, the species aused rai~t be the dominant in the community, Thus in
Silver Springs I':. *-I.: ,is :3 l1.A1! up, weighed in a cage with a two minute
drain, and replanted. in the soft ri.'nl3 bottom muds. within the 4x6x4 cages
in about 3-4 feet of tateo ..*'-.';".!,i:,' typical bottom conditions, Howeasr,
a comparison of .e,':d.1'! fertility of the spring water also is needed in order
directly to comp c :' cheomical. potentialities of different communities So a
* single species should to be ed in all the ,priL'~.o For this purpose a floating
aquatic such as hyacinth or z"'.l'., should be used so that the type of mra 'ci'c,,i
will have no offoot. -.,u. I'.T.v-~ .., a minor natural plant in Silver Zivl',,y:n ia
being grow~ in 'r-a,.
The data from (:direct meast'irment to date are meaer aand preliminary but
mey show that the method will work. Plants of ,A&g^MUI with 1 foot blades
had grown to 3 feet length aand developed blossoms. The growth of the r-ij-?. .
9er.id to be much less than t~at of the gis in the summer, possibly partly
due to light differences. The values of productivity obtained by this method
in table 10 seem reasonable,
The second method for nes:,Curing productivity involves stretching a gill
net across the run 1 mile downstream from the boil. The water by the time it
reaches this spot has filtered across a great volume of waving :QCf.?,;8.i..
Now suppose the ,prinsTi is in a, true steady state, and all our evidence suggests
that it may be ,certainly with respect to 3 i:J._~ ,i,;. since it remains clogged
with it all surfaces being covered except the sides of some of the limestone
out rops. Then the rate of production should be balanced by the rate of plant
loss to the food chain and to drift downstream,. wg I, wheno it breakE
loose, floats to the surface. Thua the dou';:trcam drift which is large, obvious
;snd continuous is caught in the .Ill net and later wet and dry weighed., This
Is done early in the ,,:,'.:'i so that effects of boats during the day will have
least influence on eeu th uts h this estimate will be minimum since more
would break off during the day. As seen in the data on standing state, the
amount of SACUr plants is large in proportion to the animals that possibly
draw much of their nutrition from ;il-y,, If further observations are conaittent,
it will be concluded that only a small part of the Wa- ~st production is
drifting downstream, The initial test of the drift measurement gave the figure
Proda'tl-i;vity Of Floid. 0: ;.'U(02)
Page 28
for Silver Spring in table 10. This drift catching procedure is further
discussed in relation 1o ordinary streams in the theory aeotion,
c. aemsga gr1iAi;Z-Cit.whod.
Xf the a :.rf-; are a flow system in i-hich the constant flow of ;v.tritLve
clear water a rixed at the t1oil with sunshine to start a biological chain of
reaction, then the distance "C.u the run is proportional to the rate of reaction
in some chemical kiLnotic apparent s. Thas this property is the basis for a third
measure of plant production in the springs. As water comes out of the springm
it possesses a nearly constant oxygen value diurnally and annually as seen in
figure 8. 0:<;:..'r is :l'e i.l as the lEarge riSvr of water poiur dowastr-eam dvu. to
two nctionr. diffauion fr:om the air, and photosynthesis .slSocD.:t:., tlth
production. At the constant ICp:ri'.:..tuv day and night the rate of dii:ff'; o:
will be constant. Tihds t:he diffrence between the oxygen valwu ioi thi r'n
in the day and that at 5;.. is the different~ between photosynthesis ead
respiration, Thus one h1as :-:: '.:.'el. the metabolism of the whole comemnity,
the size of a small lak, l d.ic-':.E 7 aln instantaneon.,-ly. Xndeed one can measure
inrtantaneously the '-',. ":;..lo.a rate
etc.
he data in tables 2 and 10 for Silver and r.on Cove, and figures 8 aand
9, abundantly illustrate that this l:jilr indeed is practical AiI- a;.
incomplete, the .. at h'.dz so-iW a large difference between 0C1.CP\f i-.'l. suB r
days and wbtp'y.o' winter and smrmer. This io exciting especially as it ,
rapid compare oi:.-:. n of :.v ..n. r and prol.:ii.ty can be adapted to stream andc rivers
and even estuaries if a !.li..- of current and. siotltaneous observations can
be made. Certainly, the critliismi that communities are too big to work with
directly seems ci'X':'r.v-i':c hore. Some initial data on other springs c'e,ur
similar orders of magiitud. These springs are giant respirometers.
Green Cor Springs as one of the small springs showed (in table 2) an
especially striking contrast beti'eeng ,we-fjor afternoon where trees are sush
that sun reaches -'..ho spring run, ourmer night where oxygen actually decreases
down the run, and winter n.-i't and day where the sun does not reach the plants
:ind where because of a removal of plants the production and also the night
respiration w.'as do'reased it.'U.!.i.U.in': the difference between diiTfr ion and
respiratioinin yT~oc1cl,, the D.O.
If these oxygen data are correct it should be possible to develop check
techniques with carlbon-dioxide or pH shift of constant alkalinity water. The
constancy of the dowstream alkalinity has to be proved.
With valid methods of ;..::,iu'ini' the large dominant aqaetic hiT;i.-r plants
in most of the spring, attempts are being made now to develop -,1t2'U,.- of
measuring production of algae, animals, etc. Cage measurements can be mado on
the medium and larger animals, bottle respiration can be resorted to for algae,
fish tagging has begun,
Productivity oa .'loA pi::
'`;, '- 29
Firsro 8
'* .J S -
C. '' to' ,
~^-. -.
.. .. '
'4 *1 f* '
(2; :o,
Fi&gre 9
;-,i:*-. at station 1 mile down Silver River
/ ./)
Coy
01,
d y W/sdlr-
tnd0 er V
/a Arei Utinder caub.th
4.
S 1 0 V .
SDay
.'i .,,,,02)
E'
c ill.. ......
-", i". "
i '
3' a
*ILS-~ IIU~-~-. I~_~
~
''"
iI~.i-1_^~-- --
.~. 3:t~
lit I # e
-rod.uc tivity oif .'.b n, ...: *. 1."(."
P:P'"a 30
V ....'. i,' .VIT THEOR?
In the course of a studyr such as this which involves gropirng4 for new
procedures, some questions arises that seem to bear on general :r;. -:t;I of the
science, such as the :Aol.io, li
1, Can an ecologicJal an.i:..:': community xis~t in a stable steady state in
which matter and energ-y flo? t:'.2ow.., the system but standing staat;e lantitaftively
and qualitatively remain .-v. :.r ... The observations so far s -:. ; .t that this
is possible in a flow <.i::.-. in nature J'st as in an algal puro culture -:o.'or.;!
madhino. It is to be :.'."- -c. however if a steady state aqjatifc C..,'..ical
system can. be etablived u nadcr xmtter closed conditions The reason for feeling
that it can not bo st-'.: i to :o folds first, most relatively cle..r.a-tri't~ e
systems, like bottle cultures, pondi and lakes oscillate and flu.ctvuate. Second,
in a closed system a liW:'. ,, lc:riont miut always give rise IBV suaccosionra
and relative ~.pi'l.tion to '~ae other lir'iting nutrient. The stability pri.'Jri'lr.!
says that all r -: -' -, .'..1 by a nItural selection pro css toward a self.
reigulated systea. n: **. i. 0 .: does a ij .th;Ar.iccall,' :.:c.,
solution exist to :.':-.:..- ci.rcular transfer of material corqponents through
oramisms with each component "..'; at a different yet constant rate. Even
more simply and op' .: k ....,--'.". if placed in sealed bottles and placed in the
light how ~7:.1V J kind .-. of simple communities will stabilize and how many
will o.3c.cill,-t ,
2. In .foi.lnt; streams of uv.su-al type a striIind4 feature is the small standing
Crop of plants in cCoparison to imall animals and especially fishes. T'iL.
seems to be an important interpretation other than that there is an autochthonius8
source of primary food matter. In constructing pyramids of production (not
standing crop) for a strean ona is intoer-n.i.-,. in production per time such as a
year. in this time the fish production is the growth of fish located In one
place since they have not left the stream. But for plants and for iaall animals
the production is wtht has drifted downstream for a whole year as well as that
which has passed up the food chain. Thus the standing crop of diatoms drifting
down stream from ourcro in bayous is an infinitely small part of the annual
crop which involves the whole years downstream drift. It is probable that such
pyramids when constructed will show a more usual wide bottom. In fa~t, production
in rivers may be the easiest of all to measure by catching what drifts down as
in springs experiment described alove. In the springs, of course the volume
flow is such that initially there is no true plankton.
3. In discussion of food chains, the concept of efficiency of food chain
transfer has been a useful operational measure that has helped understand
natural ecosystems. However, there is one fuzzy aspect of this concept that needs
clarification. It is often pointed out that photosynthetic efficiency is of the
order of magnitude of 1% or less but that energy transfers further up the chain
are of higher order of magnitudes of 10-20% and under some artificial feeding
experiments with fish of 50%. Those interested in increasing natural food
resources naturally ask what basic differences there are and whether photo-
synthesis can't be made to possess a higher efficiency.
Productivity of ;'. p i:;-: : ...(' :
Page 31
Xt seems that one thing being confused here is the diffr_--.'.." bsetuon
a true efficiency of energy transfer and a food transfer fo'ci. -.-e. in which
not all of the nn-,ry transferred hdanees state, When a fish eata protein
food it may diiGoao and separate amino acids of the proteins and i;;;.-v.tc.lJ
stick them back to.:c;:ot.r :. to form new tissue. Etch of the energy in ',h: imino
acids ti such a case never cr;-'..o form. In the case of photo3yn.'t..-.io all
of the energy ch.e. ._. form,, As discussed below it is eurgested that there
is a definite thermodyzname reason why efficiencies must be low and vAIy
increasing efficiency ,would I _- .pi, total production. When one refers to
a 50% food chain transfer efficionacone is referring to the sum of energotio
efficiency. for the energy actually changing form in the metabolism and the
energy transferred in unmodified chemical molecules.
4, The total ... ::..!... oof c living or nor.living mahin is a .s .
of the rate of inflow of .., ond the -.l.L -o;i';f; or u tiiiaJio. ,i.
is iu.rrtc.d. here th5t o-'.,-: a ro two extremes in living and r a:-.1'.ing ac i.ne
that both produce a zser enoary~tic output per time. One extreme ia a ras.chin
with a 100 efficiency ns ia a ..'v: rvli'le cannot cycle but an infinitely slow
utilization so that the o'..'L: iso sero The other extreme is an infinitely
rapid energy intake and tr': *.-, r vhich. is so fast that the thermodyrnaiio
efficiency is zero and so the totca output of this high speod system is again
zero for all the -:r.',.,. -r-. into heat. Tlhse two extremes seem to be a
necessary result of t2~ ~conda. L of Tharmodyna.micse If this r...-: ,.*I~i ia
correct (r,'d a actual calculaoviion lhs been made on a physical ,.,-, the
Atwoods i: :-hi -'. -iK) 2: tharo Ea3t be an optimum efficiency for the maxim.BMa
power. 1f thia is I' obc;eo tihena one sees a reason for Icf*. ceccr.s of
photosynt~ei, ..e7L, 1., hat ~ they ar, Tho machines are set to go at ctie to i'
speed to et the best ;:. ...i.lc.,L of both efficiency and 'pc.. At t".-:.,
under some z3:,loli.,'. c.'y.:.iton.;~u organism that laei. faster and. less efficiently
may have the odge over one thvt is slow but efficient, Hoavews, if correct,
these notions sugest no hope for h~is.hcr h,~o: food production w lihiO% repeal
of the second Law.
Second Six Monthsg
Having made bo.',nvi::;a in the 5 divisions of the study, the imi~mdiate
objectives seem ,.r-. ly 1 n't-l. Except for completing dcnildn ,iciroi,:,
community u3ocLi..:-:.r-.i.' ;';,. s of coastal runs as part of '-'o '-ion'
master s program, and the chemical survey of nitrogen motazioirm in the fre~h-
water springs with the help of an undergraduate assistant, Mr. Hampton, all
attention is to be directed. to completing the production measurements of all
trophic levels in Silver Springs. To complete this and to insure adequate
attention to t~e all important algae Dr. L. A. Whitford, of North Carolina
State, has been engaged to work with us this summer on identification and
production. Tentatively, objec~ti ws following these are the comparison of
productitivtles between spring using methods aich worrk best in the intensive
Silver Springs study.
Measurement of the amount of light diurnally and seasonally that reaches
the organisms in these clear water aquatic communities has not yet ~omrmenced
pending the procuremBnt of a suitable instrument.. So far physical and chemical
measurements have been made solely for their implications for productivity. A
number of splendid problems in physical and chemical limnology and oceanogr-aphy
await future investigation,
Productivity of i -., : ,( 2)
Page 52
Table 10
Pri a~'y Production Rates in Springs .
vy~5 rasCa
Method, Place, Date, Plant
Operationally determined
figure
Usisim
P6S3p
jet, /I -
Pounds per ore .
dry weight (or
000. O ,e.
On e ag Enol osuro e. falrg rs
Silver Springrs
Sagittaria
Aug. 16-cot. 9
Oct, 99--aNov. 15
Pistia
Aug. 181-Oct. 9
.. ., '
% wet weight' increase
per days
046%
..Q$-
9C,
1.9%.
2. Downstream(rirft of Plant
FrMents (- production lost
from the system)
Silver Sprin:g
8:o00 a.m. Jan. 14, 19"5
lbs et weight
caught per half
hours
8.6
(289 gma dry)
5. Oxygon Gradient M'suroe t
'(day minus n.{ht)
Silver Springs
Area undor curve of fipre
9 (winter day, clouify)
Clear day in cumer,
June 50
Green Cove Sprir,:s -sun
Suammer day, July 16,. olear1
Winter day, -Jan. 27, ceolea
shaded by treqs in a nt"er
PPM difference
between day and
night' Oxygens
0--1 o5
0-2.8
2.2
*2
15,000.
11,600.
7546
27,700.
77g000.
62,6o00
4,900.
For comparisons productivities reported for literature in marine, fresh-
water and land ranges 1,200 ---53,900.
-,.~.~L~t'';~d~a -~~--~ ---~~ --~I-~-I-- L ---._- ------- ~I Ir --
Oa 4P~- ~ F~V-r ~rl2I~IYC 4i EWC~~R7bl~ .t ;bnL~DLLI I~pIl. r M. Imam--Dvna -~sm ~ rn -Y~I~ IICLLI rI~d
J-~.nJ;
/~3~
I-'
|
RSS
HIDE
