PT'i ..TIVITY OF FLORIDA SPRINGS
NR 163.-106
(NONR 580-o02)
Second Annual Report
to Biology Branch
Office of iO'$,J- Research
S:i'.' ,es from JLanuar 1 to December 31, 1954
H. T., Odu4m rand Jo L. Yount
wtith sections by
D031o0 &Wn'-
M -id K Caldwo1ll
':-ck H l .ri;
opa nf ~of r .. ,..
.'.': of Florida
Gai O ," -, ,, : .
-477. .,
TABLE OF CO'ii.U(;
INTRODUCTION o o o e o e o o o o o 1
PROGRESS
-7-)!oiVCTIVITY OF SILVER SPRINGS o o o o o o 5
COMPARISON OF AN ENIWETOK CORAL iREF j r,'::. .ITY
WITH SILVER SPRINGS o o o o o .. o o 17
PL.iAi'T. COMMUNITY STABILITY DURING 3 F:',., o o o o. 19
STANDING CROP AND COMMUNITY SURVEY OF .'.
VEGETATION IN SEVEN SPRINGS .. o. 2 o o o 20
FISHERY BIOLOGY STUDIES IN SILVER Si-... 3 . 35
PRODUCTIVITY THEORY o o . o o o 41
A HYPOTHESIS REGARDING DEPENDENCE OF C( '.' TY STRUCTURE
AND DENSITY ON PRODUCTIVITY ... . 45
INTRODUCTION
Prepared by:
Howard To, 01.'i aJd James L. Younx, with sections written by Delle
Natelson, Dr~'-.. K. C,!l't.Al and Frederick Ho Berry.
NR:
CONTRACT:
NONR 580(02)
ANNUAL RATE: $5,C0 (h -,.:-3)
CONTRACTOR: Departeont of Biology, University of Florida, Gainesville (w-ith
Biology r.:~~r'c.h Office of Naval Research)'
PRINCIPLE INVESTIGATOR: (This project is administratively listed under W. C. Allee,
Head of the Biology Department),
Howard To Odum (February 1 September 1, 1956)
James L. Yo'-~ (September 1, 195h January 31, 1955)
Associate: Delle Natelson (September 1, 1954 June 15, 1955)
Assistants: W. C. Sloan (Y:7'.-: -y 1 June 30, 1954)
F, H. ro!:.. (June I September 1, 1954)
D, K. Caldvell ( .1. 'I,.,c. 1i 1954 January 31. 1955)
TITLE OF PROJECTS PROTri .i- .., ,.i OF FLORIDA SPRINGS
Objectives:
A study of basic factors that control productivity and of the effects
of productivity on coiniunity structure and density by an analysis of
the .- si'. *: conditions supplied by selected constant temperature
3pi k':.'tVl.,:
ABSTRACT:
a. During current ..'. :;; ..'.iod
Production measurorents at different times of the year indicate a
linear relationship of I2 .' v'-,iitj and overall production at about 8% of the
visible light energy i :'-a ~.. level, clbtri.'eien s of a coral reef at Eniwetok
indicate 6%. Further evidence of breeding at all seasons but with a quantitative
pulse in the seasons of maximum light indicates that the seasonal fluctuation in
primary production is routed through reproduction rather than through major changes
in populations, The succession of plants and animals of the aufwuchs has been shown
with glass slides and counts from Sagt iai blades. Loss of oxygen bubbles during
the day and emergence of aquatic insects at night have been measured with funnels.
Bell jar measurements are reported for bacterial metabolism on mud surfaces, pH
determined C02 uptake ,.,j:.'.:': with titration determinations. A few rough estimates of
herbivore production have been made from caged snails, aufwuchs succession, and fish
tagging. Nitrate uptake at I!':.,-i by aufwachs communities has been c.nf,inod in a
circulating microcosm experimuet as well as in bell jars in the springs. Distri-
butions of oxygen and or-;_. -:r" have been used to criticize the saprobe stream
classification system. Theoretical consideration of maximum photosynthetic rates
in literature data indicates lCirthi3ic rate variation inversely with organismal
size just as for respiratory metabolism. Extreme pyramid shapes are thus shown for
communities in which or-:ni:-.-'l size decreases up the food chain and for other
communities with the same ~.... ~. influx but with organismal size increasing up the
food chain. Literature data is used to further demonstrate the validity of the
optimum efficiency--mBaxim0 i po:er principle for Photosynthesis. Work on aquatic
ABSTRACT (Cont'd.)
plants by Dr. Delle Uatl,:l ;i..-ates essential stability of aquatic plant comma
munities after 3 years and about O10*-20% reproducibility in previous bioaess
estimates by Davis Wo rk on an annual picture of the fishery characteristics by
Caldwell, Berry, and Odu is half completed. The study of aquatic ic':; in
relationship to spring g:.'T *:.:~s by W, Co Sloan has been completed as M14 S.
Thesis. J. Yount has begun a study of effect of total productivity on community
composition using aufnuche .',: i;.irs on glass slides placed in different current
and light conditions in '.-l-. Springs,
be Since Start of Project
This contract was begun June 1, 1952. In the year and a half pre-
ceeding the present report period, work of a very varied nature outlined the
trophic structure and metabolin of Silver Springs with comparisons made with
other Florida springs. The intensivee study of Silver Springs is now nearly com-
plete and quantitative comparisons of productivity with other Springs will fol-
low. Most of the techniques and approaches outlined in the original proposal
have now been p.Pi.. Tl. study of ..':tors. affecting qualitative community
structure is the main S'u,... '.,.: ::.;.Co
PLANS FOR FUTURE
Immediate:
1. By J. L. Yount
Determine the relationship of productivity to the variety
and the dominance of species comprising the various commun-
ities of a spring. This is to be done as follows:
a) by observation of the numbers of species per niche in
single habitats within the spring, e.g., Aufwuchs on
S~.iLtJ-; "._ blades and glass slides in different regions of
the q;. w.ng where A l'!i.otivity differs but other factors
are constant. Both high and low trophic levels will be
studied.
b) det, .>.' *' relati',on.~ ps as above under experimental
conditions in ; uaria another vessels. Plankton and Auf-
wuchs are expct .. to serve as the principal experia.:..1tl
groups, but other groups may also be examined.
2. By D. K. Caldwell, F. H. Berry, and H. T. Odum duW:.w. Spring
and Summer.
Complete an annual cycle of 5 3hro' characteristics
'Wb.?:_'. last s.~ ing as follows:
a) tag and ... pture more fish
b) determine the significance of scale annuli in silver
M'IA. ; :g. lA.
c) further determine the extent of winter b~:-:.: Tg. of
fishes in Silver's constant temperature waters
d) cow.,t.. ;: f:,;] assay of dominate fish species
e) 1 .termine Cr't-- h rate of young stua.pl1,hn'e..-er in o.ses
f) determine the sigrni- chancee of .;m ,,h -.ij: ..... y -.C.-.- .:'.. o
fi." fish collected in sprl,.a' thr:.. --out the year
3. By D. Natelson oa'.'i..a. Sp,~0~3i; 1955
Co.mp,.'e o .1 community composition of aquatic plants with
Comi!!'.: ties not in constant inr.rn-ature 5-.M 1 r..;s,
4, By H, T. Odum ';. -in;, summer 1955
a) Relate the overall community p.ola.ction of 20 springs
measursl- .' the downstream flow method to current
veloci' to test hypothesis that the overall primary
productivity of communities in steady state is a function
of veloci.,- o' water over plant surfaces,
b) Co;,.pl..te the picture of metabolism in Silver Springs
by additional data on organic matter loss downstream,
effects of side boils, herbivore growth, repetitions
of (u ..; ,t.t ...-.i-urements, spectrogram of water and plant
ash.
Long Range Plans
1. By J. L. Yount
a) determine relationship of productivity to variety and
species d-'l,Jr."se in other aquatic habitats, both inland
and marine, contrasting habitats with high productivity
and low productivity.
b) define the relationship between productivity and
competition.
2. By H. T. Odum (Dulke University)
a) apply method of measuring community structure and
metabolism to other steady state systems such as Hot
Springs -. tropical streams.
b) construct microcosms in the laboratory to contain
small stee- state communities in order to further
S-limit ...Inciples.
REPOF.TS ANIB PUBLICATI,. '
Supported in part by this project:
Odum, H. T. 1953. Dissolved phosphorus in Florida Waters. Report
oC Investi.l tons #9, Fla. Geol. Survey, Tallahassee Fla. 40 pp.
In press:
Odum, H. To ... RT C. Pinkerton
Times Spr-..- Regulator: The optimum efficiency for maximum
power outp.'-. in *...olcal and biological systems.
American Scientist
Odum H. T., : David _. G,:3:,9:Cll
Fish respiration in the natural oxygen gradient of an anaerobic
spring in Florida.
Copeia
Completed Th. -:., j
Slain, W. Co Scmse onvirr .-.,.i al factors influencing the dis-
tribution of aquatic insects in certain Florida S.ji-...:;3.
Manuscripts cc.,bl..ted ", submitted for publication:
Sloan, W. C. A c. -:-:::.tlve ecological study of the insects of
two Florida Spr'c.. (Submitted to Ejo~ y).
Whitford, L. A. The Communities of Algae in the Springs and
Spring Streams of Florida. (Submitted to EcO1U.)).
Odum, H, T, and Eo P. Odum. Trophic Structure and P.. .-,,;ictivity
of a Windward R::.' at Eniwetok, 90 pp. manuscript.
Manuscript in pre.a..r'ation
Odum, H. T.o 7,'.. i- Structure and Productivity of Silver Springs
Florida.
k; NO 'J.V'.' C1 .. SILVER SPRINGS
'o -:" -' To Odum
i:. "...v.'.... ':, and d-'. "~.-. studies continue n Silver SpIt.rA in an
effort to determine the da- :- working oC a fertile e-.-.*n.1-,: aquatic
.... ?-.. ,'.A in s ..
A o ..: ., ..
" i 'n ',...N in Chanostatic Conditions
The upstreamsLdownstrean method of measuring productivity as reported
in :.., pr- .. :rr--t" has now been carried out in Silver !.K..-*
in various saseons of the y.r and with different loud covers From these
data an annual cwr. f- of total- j r -../ primary production is dramn in
Fi..ure 1l The .:,ro, *arundier, -3s curve is th total'primary production.
'.', horizn'-tal '' .. the respiration and downstream losses as
prevo..7-. dsto.rra~W.ao tth ..-i. is in itood,;r state the area above
the horicntal lian! shoud c the area below. These areas do not
"12. -carr- !; .; in ..me asurement of downstream lo sse
in the "-'. K c"I::t'in., .d in the :...a of aide boils will be
:required to ac.co..t i t .I excsss production
From, the
A cuirve ". '.. :.l: iij p .c '.; experiments is
also showa. in '.' .. f2erence -.. .: '. this ,;. i <,1 and the
total p-.',: .. r.oproGuK ; tho ". -p.:-X production which turns out to be 7:,
of the total Tven .' '. 1ho "'.. :'. biomass is ,r<...e.i.o That most
of the produce., ; community is algae and Sagittaria is evident
from plant maps of silver: Springs referred to in previous
reports,
.I
"C~
*" :" total ,' ..'...vr production
S/ \ \ -12
-** '---- . r :)
,. -.," ,f
r3 ,, lXo 'n.ual ..*8 of .:'... Production
Fr ',"T' Ci>
k; NO 'J.V'.' C1 .. SILVER SPRINGS
'o -:" -' To Odum
i:. "...v.'.... ':, and d-'. "~.-. studies continue n Silver SpIt.rA in an
effort to determine the da- :- working oC a fertile e-.-.*n.1-,: aquatic
.... ?-.. ,'.A in s ..
A o ..: ., ..
" i 'n ',...N in Chanostatic Conditions
The upstreamsLdownstrean method of measuring productivity as reported
in :.., pr- .. :rr--t" has now been carried out in Silver !.K..-*
in various saseons of the y.r and with different loud covers From these
data an annual cwr. f- of total- j r -../ primary production is dramn in
Fi..ure 1l The .:,ro, *arundier, -3s curve is th total'primary production.
'.', horizn'-tal '' .. the respiration and downstream losses as
prevo..7-. dsto.rra~W.ao tth ..-i. is in itood,;r state the area above
the horicntal lian! shoud c the area below. These areas do not
"12. -carr- !; .; in ..me asurement of downstream lo sse
in the "-'. K c"I::t'in., .d in the :...a of aide boils will be
:required to ac.co..t i t .I excsss production
From, the
A cuirve ". '.. :.l: iij p .c '.; experiments is
also showa. in '.' .. f2erence -.. .: '. this ,;. i <,1 and the
total p-.',: .. r.oproGuK ; tho ". -p.:-X production which turns out to be 7:,
of the total Tven .' '. 1ho "'.. :'. biomass is ,r<...e.i.o That most
of the produce., ; community is algae and Sagittaria is evident
from plant maps of silver: Springs referred to in previous
reports,
.I
"C~
*" :" total ,' ..'...vr production
S/ \ \ -12
-** '---- . r :)
,. -.," ,f
r3 ,, lXo 'n.ual ..*8 of .:'... Production
Fr ',"T' Ci>
B0 Determination of t- ".. 2 :..on i.-v Reaching the Plants
In order to deterine -,.:. "'...-;. ..-;r of ;...lianry production, light
i n::..it ; --;:~:.".ti thoe -'.-to were determined as follows:
(1) ntr.:, ..' ,- .1 :' -"' th.o .n." on a given date for a givz.un cloud cover
ie d ..inw from '.:' .:- in -...V (199. Bullo mero Meteor. Soce 30:208-213.)o
(2) 1.. C of this is talre, ,, ",'.. rA :.d- and half in the visible rango
(3) With a tubmarina .. '.:: "- containing a veston photronic cell sensitive
to visible %wave '. .. the p',r.oz,t penetration to plant level at 8 ft is
determined A winter curv'oe mas shaon on page 8 of the last progress ::.,i -'cl.
A sammaer curve is now g:,'" in Figure b,,. ,.
() .:h, :.':. - trans.fidcion :;.r' a given tirea of the year is interpolated
between th two xtriOmo curvm. ;:-p.*:.~.:-:.. t'lod by January and MaQo
TI- 'i;-: *.. t batean percent transmission in mainly due to a diffarenco
in angle of incidnasco
(g) 1". i.Asolat io : Ith '*.n".; comanmity is di; ti;i&..' .'. 2:
frm tres : '. wn. the sun is at a low angloo Due to the orienta.tin
of the -sd its. tra ,.* ..v t ;t is greatest in the ..
as ref.loCted in .' of the diurnal curveSo The ree effect :is
greatest in the anter. i. li OVeA on cloudy o-.:*. '2 .7 1 .
wen the 1 1'". in c :.7 .
To correct for Lc .- of trees, at least during the ? -.'r -: the
llily insolation erb 'suI s :tioned above are redrawn so that the n .t.j .;
is symatrical ih the r-a 7 curve as shTa in Figure 2o T,..
area betyv en the rexa'r. ..L. 0 '. curve *.'k-..- -Q- .L the light removed
by the treaso No Iuch ct':on'tion is made for cui.,; day:. The correction
on the winter ournvS is -proportionately th than on the summer curve
S!U.. intesnsitis do~ ruined in this ray are related to production I-- tr-..
':T' .: 'on A'. trees
microampn
Time o f f~ ,
..arch of V..t., May 286 "1'.;"
Co r.iu.-:., as as a Fiuactin of I.L:,% Intensity for the Whole Community
For each -.07 production iSu:. .' given in rugye 1 a WdO.iN1. :. .
intensity v..r'. as 0:- .-.. :.as described in p:,-!.'yvph B above The
.'ph in :.-a 3 shows total community ir.'.i productivity as a 2 o'.' -
of light intense si_ ..1' .', k','. wave lr,;th reaching plant le.-, The
horizontal a3.: is the eatAmate of respiratory and dconrstream Dow rmrn estimated
from I .:..ious .2 -'.! ,
?' <..:, 0..'. the '. ";ion is : .. 'Jcyct.cri- to the light intensity even
at these relaSt i'. '.." '. ',. 1'. intnsitieso A.'-t physiological u pr -.-; .s
on :Irn: ;-. :' how' efficiencies with increasing light inteonity
at high ; ':.' intensitisol It can r'.? '. be noted that the conmtunity
runs below its overall :../:'.;.n point on cX'.rA y winter dayso ThiQ does
not mean that individuvl. :i do this for the mmni common compensation
point i.Ot. .: not : r ton butaton b downstream losses. Possibly
an o; -.- .'. of the '. .. ..j th .c '.n. r 'wnstream lies in a
requir-emet that at st;'; sA.nte the .:.. plants must neveBrthelss
never be e oa d ;.;:.d'Wions bl' ow the individual compensation point,
As seen below th (o.ral ro o do% not do this and similarly dogs not have
the low '. :,int.o '. o there is a generalization that
temp ratee coarmnitKies :mut producec excess L:::n.i. matter as soil or
peat or downstrsana Ova wtich, tropical communities need not d*o
Pro, the data *' .". : it is not : '.:.? whether .h'vy is a break in the
curve at @ ': as -0'::. .:-A in the theoretical oectiono
A,'h.rN :;.7 the :.. .. '.. .. .. .' t ....'':-r an i oxn an .og.- i..: *..a. .. r;.r
with .;.-," intrnit'i.'o As discuss ..W:'-rw this i., :...t be
accounted W'-;* by incvr's .'..bation of -.' which indil.iA l1;
t're ;.1 the ir .. j; 'i. n I:s..- da:ys in more '..-, phacw,
S.A :.'*.-^. yday ,. -
S4(504 W ..TW :mr:7 line
Primary
Production ., -
50 ---. s--------- cr't.mir compensation
i% .i............. ..........-.
-| ,.i"r lin. ,-
'- ;'.? ";'.' visible light
S'.ty p ..,'-0- product on v ., .; : : '.. 7
TO..9 i '
Do Seasonal Pulse of ..;' Pr..?,dingj and Photoperiodism
The m.:apl~ in Figu 's. 1 .: 3 indicate a strong pulse of energy
in the spring and summ P ~'.- .: --,d to the less lighted months This is
actually a :r "'-' difference in the aquatic community than on land because
of the angle ....aglo ot the trees and water reflection
One is accustomed to r. '.,I.-...aAng big seasonal differences in energy
flux with succession and blooms :..W-.on;. the planktonic organisms in 3.-'- s
and in the ocean It .. : i" '. l-.. to consider the fate of the .-::. ..u;..
pulse ir Silver -.01M; 1:. n. o v.:r.: sucoessional changes have boon
ab.-.'. ven in the mic1-oscopic n-.lj. of the aufmchso
In i'. :-es h and 5 are shoirn r.':i.w.l pi'tu.res of br.andlI'z in
the zp-.pl; snail Pon ~ .3Mca which .. '.y its eggs above the water line and in
Pa&",.onstes ehich carries its :g2s,. From these graphs it -:- b
cr....*. that in the so .br.-e. .,i; occurs .'"..~:; hoo '.: -. the -.r in this
constant temperatureo en-viron;menat T7.t at 'i',:-rent rates that are
b..y to be ... % ... ...;:'o.L.fl. Thus these species seem
adapted to the ': pulse of the whole community As i;: 'i.-,
ago doscr::.b. : .. th survival of -o"T p'.ivl,. community complex r'-..p '.
the components to eat neither too much or too little As mentioned
in the r:.p. on :. .. ..' .. .3 below there is some evidence that
similar round theo '. "' : occurs in the fishes with a l g..,o
Figo ) Annual,
Pomaoea
P.-*- !' .. ..... \
per 125 .
of shore l
/
-. ,- 4 ;. h S j. 3f .. ^
"1'. Month off the .V '
Fig.o 5 Ar.v-:
Reproduction of
Pal oneteS PAI.. T.',
the females
over 21 gi
with eggs /
Ho Initial r,-u:' -n Yicrocosm Eolperiments Nitrate uptake in the Daak
As mentioned in the theoretical section below microcosm exparimants
are a useful type of .'; .im ent for studying ecological open steady state
systems A ...' -- -: -i. .. blade of Sagittaria with its encrusting mat
of aufTv-', was placed i th. e tVo., > G .'j ? *.-- tube of the apparatus pic -..
in Figure 6 below '..- .';:.'ings water was circulated thus -I .j.ul.,T.
the natural ~ ?. ;'iono i. a time interval some of the 650 cc of
water vi'..i was t: a nd tested for changes in oxygen, carbon dioxide
and -n it .- .."
The .".;. co: .clusion ws .: in spite of all the light sources in
thp e I-::' ;.;- ,. h could i.. 1 brought to bears the aufwuchs micrc-.-. .:.:~ix-ti
normally adjusted t) -Utd or ..... intensities rapidly used up the
y P.-.,n :p-:Lx and dicL ".- as not yet been possiba to get the
production up ove '..:. ooam:Iunity compensation point which is very 1.'i.h
because of t he tcrotrophc a ,s .A1, as autotrophie camponentso
It was.g ':. "'' ,' .. to show that nitrate is rspi.l.,7 fixed
by same coiaponents in te unity even in the dark d'ri- ;.];.
:,pir :;<:...i metabolismso '.:" experiment helps to confirm the ..
,iptar @f nitrate fou.d in ''1 I :-1. Jars in the .j.1 and the
slight decrease of .itrat, .'. tihe boil ,.,:, :'--" both in.the day and
at night In 5$ .- .. .':':...'.. o037 ppm
decreased to o23 ppm in 1 I- and nitrate
was not deti -. i-. lafti .,.'.-. "Silver'
water standing in :..L-:a bot' tlo l.s:c
nitrate at a rate of 2. pm
per month, S l ve
SSilver
Fi Springs
Flow "- ..w....a\
.MoneL Metal 1/8 HP
Circulating pump
l'h-. re :
hero
Fo AufNucho Succession
The rates of CL-':.' and succession of autwuchs have been measured on
slides submerged in the .;-i- aMd by a count of distribution of pl-.n. and
animals along thae I.'. i' : -': i bladeso Since Sa ttaria blades grow
from the bottom, it has ': pF'ible to measure their gro h rate as a
means of th.t.e .;i::p'o, th age of the attached aufwuehs at any place on the
blade The distance fe..-. the base of the plant indicates the ti=e
since the succession hb.., .
As sha~n in 1'.,f 'u re the rates of growth of single blades is far from
equal Small wires were inserted in a young blade and an older blade in
the same clr:.'p.. After 26 days one blade hd shown a rapid 182% growth
pushing the attached wres tith the tip whereas the other older blade
had hardly ,r":. :, Appar?,,arn, a blade shoots out ard then as the aufwachs
covers it growth ceases and goes into new blades. Thus one gets old and
,;~.-. g4r ~16. d::: of near.., the same length next to each other, one bi5:j
clean, the other -rv ".r covered with the periphyton community
Figo 7, 26 day
growth of
marked blades ->w
of iria '
April 5, -'
j.-4 1416 inches
For a whole ,. of ,.I: -, however there is an average r.?...:which
tend to average tout the spurts, Th:r,,f.',- by cutting 50 blades into
segments of 2 inches a,., '., wd placing all the first shen~t '.:..'.h:.
all the ...a etop one may relate the attached
average aufwuchs to the average rate of clump gr otho Knowing thL
area of the blades scraped in each ~I- group and knowing previously the
percent growth of F-h'.taria '..,ra the planting experiments, one converts
length into time and ",on counts into area estinateso If '.'
are in '-,,.:;-" state tho parent loss at the tip of the clump is the
percent growth A ounft '.' the 1 : segments when fraJl over the
spring area gives an estimate ofatf rate of auwuchs growth and thus of
the components of the ty "..'"i blade type organisms As an illustration
of these methods a curve of A;i-es versus time and thus also length is
shown in Figure 8o Easti;s:es of midge growth rate from this curve are
discussed below Sqe .: ,.' curves of succession on glass slides are given
in Figure 90 The l.r .lariti'e in this last Figure may be due to the
positiorsof the slide boxhe in different currents and depths Further work
on this is being carried on by Dr. Younto
The succession i.*;:-. :'n ar much like planktonic populations with
bacteria first, small sn.., n~ x t then larger algae, and finally herbivores
and carnivores as :..:.-:: ". of a pseudo-climax is attained. Thus one has
continuasucession in the ia.cro-environments of the overall .i.:r, state
Similar microcomns should be 1. 4 :.; for in tropical oceanic plankton
Flies
blade 1 in yc>m
on a B L do e100
of Co :..
type J."'. "
Diatom
groups
per
slide
I1,,
50
Herbivores -
per slide
Carnivores
per slide
Fig. 9o Succs=
sion an slides
. ---- ------H
7
r-
.. ------- P-
"0 /
H .' Craspedacusta
-t,
st.'
Ii,,
Syear
Dry
Gins
4)
anJi m.o
per
of bdS
106 -
day s
"'.-* ..'r,
'. ., 8 ,,.
Arcel".
X'
o Insect En rj;,-.o anzd ::... Measurement
The Very rgo ; caddis flies and other eq-i. .,; insects
just ~.f'Zt, sunset has bo-n *. orr d to in previous reports. It has been noted
to be intense both en fr. -.. days in -r.'. :oi aid hot summer days Foie
funnel devices waer placed the s aprai.: as shown in Figure ,.. Soz .'.2:.'-:.:-.ive
counts were ofL;..L, from thse sot '.,:i as given in Table o 1 E ncS-r;;.-..:
at dawn is not -i.ln..;. t in comparison to the evening outbursts Pc-'. ..
estimates of h"r.-: r.:' ". r.oates(net) are calculated below from this
measurement of ....*. -' rato.
_,..raduated centrifuge tube
.. .. ..! '
... ;'- merging" b'.. 2o oxygen bubbles are
Figure' .. ... 'hera ." .
2 -, a -
Oxygen bubble .
/ ra d. funnell @-
Saggitteri
.' coated with
/ ,I/ / ""
Aufwuchs
The a is :. during the daytime to catch
the o ,i to 's' in the Aufwuchs and then
rise to the surface ithot co, ;L ( 1 This loss of s'.:;. ciO
is a source of er o in t:~l p.ouc:tlion inmasurements that causes and
underestimat-in of th VtttL .don: ? 'c.r-a g the tubes exactly
at the water t7 r' + the '"': .n..n can be converted into a ::' -uh
'.-t, i since t h bubble arS at ~attosphl r ic pressitre It seemj s ';-
that these 1 .'- a .... ."' c.r... a s Vhour measurements ibow a
maximum effect at time of ,xia ,,' ....'. ':..... See Table 2o
.:'i 1 Insect Di,-r,!g.:.-': In Silver :',..,,.. '
in ap'~:.).:tu3 of Figo 10o
Place Caddis
Distance from shore Time i l :.in
1 fto 1, ; 20 7:30--12:00 pM 9 5
S28 6:00--1:00 p.m. 1 1
25 ft 2 f 20 9;390-12:00 p.m. 8 2
S: 28 -':00=-3..,00 p.m. 2 1
.; 13 8:20--2:42 p.m. 15 2
40 fto Station 1 fto 20 7:30--12:00 p.mo 5 2
Station 1 Fay 28 6:00--11:00 p.,m 3 5
Station 2 .:-... .r 28, 6:00-:100 p.m. 0 2
-LB~e~rPCnBrreuyl~a~urmaoP-~r~~r;
5.o 2,4
Table 2. ]r','.:c' ";hing the Surface in Silver Springs, 1954
4. h,,.
,''blo Time
in cco Laps.
in irso
(t~l~ilI= = --~IPE-~-PI~ nnmr~~lg~P Pnsz~if~B~~i
NIGHT: 8:20--2:42 a.m, May 133l,-
6:00--11:00 p.m. May 28
02
.1
Mean=iT
7:00--3:00 a.m, *;; '-?Th 0
0
0
.1
meana"rm^5
2:20 p.m.-4:05%
9*30 a.m.-3:s4
mean
p.m.May 6
p.m. May 23
.21
2.7
.5
.2
.42
2.6
3.1
12o0
3.2
"'-g7
4:00-=7:00 p.m. **.- 23 ,1
.9
3.1
.1
moa I
Ho Herbivore Production '-..'-.,-,
From data in p.,:.'..:,C:.
as to the production rates
previous progress reports
:. 9." ,;-..:;:n some rough estimates can now be mad
of the dominant herbivores such as were listed in
Soavral methods are used as follows:
lo From the estimates of t, ... at the tips of Sagittaria blades in Figure
8 and from the ae~rago parcont C:..*.ttaria growth of 1%/day from Figure 1
One computs the rtes th e of midge growth o;,ary to keep up with tip loss
in a steady state. This is an u.derestimation as it does not include early
emergence and losses to pred-s,.aton in the middle sections of the grass blades
Time
CC/Hr.
6,3
8.0
.0031
~i~p~L~~-z mn ~ I .?__ ___ __.cDI~.-^.
1,7
6.2
o19
.25
.83
3o0
DAY:s !$--9:30 a4,m, 9:.0: 23
.nl~~~~ll~~PUI1WII~~L~~P-~L~Y~~.Y~
mean
2o A few cages of Pomac "i a ~tJ ivigg ra snails were maintained with
an abundance of food -rnc v.:, meas-urments before and after a month growth
period. These estimatQs aro underestimates sines the snails used wore already
of moderate size and past the mor i rapid juvenile growth stages.
30 From the section G above on insect emergence the growth rate necessary
to balance the emergence in steady state was determined using 8002 gms dry per
emerging individual 'I,." fig~ r should probably be added to the Tip-loss
figure in method #1 above.
4, From the estirates of '.t-i..l g crop biomass of small invertebrato
herbivores in previous reports one can obtain the total respiration -Ji;.:.-;
a rough figure for stream invesrtbrates of o8 cc/gs/hro Then if '...;;.h
of animals is about 10 one ;.:a-' get a rough figure for herbivore froductioon
With theso rmthods s~mi herbivore production estimates are given n Table 30
None of the estimates are en tily satisfactory although the order of magnitude
is indicated.
TJ.'.? 30 Ste Estnates of Herbivore Production
Method
Blads Tip loss rmthodla o .i
Snails in emaenc method i.':)
Insect emergence method (.')
Measured Quantities
Pro idtion
ms/M^/r
~4 An idge/M2 plant suIface
2$5 M plant surface/A spring
So26 M plant growth/i spring/day
(1% blade growth/day)
6% volume increase/Month Viviparus
i 7% volume increase/Month Vivipar=us
:I'-- increase/month Pomacea
1Man: 11.6%/month; 12 gm/Mr biomass
7.8 individuals/229 cm2/day
S002 ngs/individual
2195
17
196
Total of Insects and ,;ila
Assumedeafficiency and
.8 cc/gm/hr(method #4)
32 ot. gm/M2 herbivore standing crop
Io Carnivore Production iY.-..
In the fishery b:,1.:Yr study described below a few recaptures gi.v
some minimal estimates of growth rates of fishes. The average? growth rate
is about 25%/yr. If tho :,;, .'rinj biomass is about 7.3 gns/ia fish the
production is around 2 .: :, Lr o This figure does not include the
main stumpknocker populations Satisfactory biomass estimates of these
fish have not been corplts. ,:, These estimates based on large tagged fish
are probably much too low, If 7.3 gas/Me fish had a metabolic rate
of about .0 cc/gm/hr and xa i: .ricirncy of 10% the Rish growth rate would
be .6 g~m/M /yr. Considei.-- :.- ,ore work is r,;qi:red on the higher t.:,":.c
levels to establish the o,.,.-', ,::aa production and efficiency
l y 03
Jo Pyramid of t '..:'.- :, 'r:ct. Measurement of Bacterial Metabolism
Even though production rates of all trophic levels are not yet
satisfactorily determined, i i s instructive to calculate metabolic rates
by trophic levels using some literature values of metabolic rate :.- v-1;i:7.!.6
by estimates of standing cropo
Primary production '- .*os are taken from previous reports. The ma abolism
of the herbivorea is taken as 08 cc/gm/hr with a biomass of 32.4 cws/a o
Carnivores are taken as oO7 co/,;'..r with 7o3 ms/2 biomass. Top carnivores
are estimated from e04 co/ .: with 4oO gns/ biomasse Rough metabolism
figures come .TCav Psaeto"
Direct estimates of the baetC' s.:.: n ; .^i;blitbi in the algal gyttja that
nakes up the comnnuni' botto.-a werQ made with small bell jars placed on the
mud surface after the top zono .which contained algae was skimmed offo
As with larger -. ..11.. -..nts .. ri i-d in previous reports, oxygen
analyses were made bforse : d af tor -y. Ain.rt'l. periods under black cloth.
Three replication C"`- oxyg no decreases in 95 minutes of I1, o75$ and lo84
mg/i0 with bell jari.. .'' cm in.. .etar and a capacity of 1800 cco
This turns out to be 775 ., This estimate does not include the
considerable bacterial flora t -?:s o"'.;*o.'bo, which was measured in p.r-....'c.!.:.ly
reported work, It is "' ',an:t to note that .' l;:a.iri much smaller in biomass the
decomposer bacteria are oo -..o me4. h'iWiAUy than the regular herbivoreso
The various r:- a sti mats of rme..e'1:I. by cbroplioc level are suanarized
in Figure 1, b lo:. ia t"s a first o ;'.mp-; to assign values to the
metabolism cdit u' .'o *v Eszntsd -
: .-rxfl)
Figure 3J
tPowr' c-rty aolc.lc aitt.noa
toward a tabolic diagram
NCai yro
organic
matter lost-
t? 6"?i",~
16
Ko Diurnal pH Curv y : .-ir~'.ornt and Dye Nomogram from Carbon Diodxda
In Figure 12, l -.-' is ..von another days determination of production
with oxygen and carton-:* S:.' : curves such as have been given in previous
progress reports Thi.r time water samples were brought back to the laboratory
and the pH determined with a ,.:.ln model Go The maximum time of 24 hours
is not so serious in thesa .-:" o-.;::-. ic matter waters as would be the case with
moat natural waters h-:. carbon dioxide values were then converted into
calculated pH values from thE known alkalinity The measured d and curv
calculated with the DyS naorogriya are both shown below:
6
duplicate bottles
O
'0 12 1" 2 3 5 6 7 89 10 11212 3 4 5 6
; y Y*.'.:0 a.113 1 2 3 4 $ 6 7 8 9 10 2 11 2 3 45 6
no noon p.m midnight
10
CO 9
ppm 8
6 /
7-
saattered cumulus
direct sun
s*" 7. ,,,.. \ 1 O noylight
7 7 )( measured
A calculated from
O 706 s CO2 =&ans of above
graph
Figure 12o !:'. .'.L. Curves for 3/4 mile station May 23-24, 1954
COMPARISON OF AN FrT"F:7 : ...- Tr F COMMUNITY WITH SILVER SPRINGS
In the summer of *1. H.T- A, and EP-n.' ,s*,iv. of Georgia) made a
In the aummor of "1-1'.-"H, T l"voi and E.P.
study of the productive .I of a windward reef on Eniwetok Atoll primarily
under s;-'" !.C.o-; '": of the Atomic .'-:ir Comission and the University of Georgia
by eman of a otract extension of a pr'-:u directed by EoPe Odum. This
endeavor was .:' .' .." : I' *:.In "" Y. by the ..;.7 and indirectly by the
tecl:.t s used in the Sl .. work.
One main purpose 'was a *.. .-;.', of the characteristics of S;'.vr,
Springs and the J.- :% *-iland T.So The 80 page report on the
Eniwetok study has b.' ( bei .. '.. too: and submitted to the AEC prior to
publication A f.e .7 theo -, '. ion, which are signiibant to the
understa.:L ~ of the "-'.-r ;..- community and steady state systems in
general may be :'..': I v .
1. Both com~aniuioJ s a&eo very :i..r.'1. .', r~. the coral reef :.i :-C~i r .
(of visible "',' r '.' -~:-i. average commauity .ap thi,
2o Both are "T fr ., *' .. coral reef with production rates
above 75.,000 .. .. .- .. .. :.re per .: r, is among the highest on earth
Silver Springs in Moe produ-ctiv in sui er mtoths but has a lower annual
production
3o Both .,.. are .; atotr. .'' in primary trophic leveo
(John Te:,:1., H.arvard '.vb is .." on ..a.: -.... ry of a small :-i..' .;,
whose primary I'... --ton o 1ci : not light but alloothonous organic mAttero)
Sargent and Austi.n9 '*.. ..:tude. on -' .:." productivity were hi general
confirmedo Gass ." .don 3 "eoskss n both Silver Springs and .;.i.-.1.. :
yield coatings of rather t.an .''" ,; com cities found in inshore
oceaniO waterso
o0 The .
surprisingly similar
ratio of 1.': t7.' produc~ns a:
maintain optimn a s3aiGrcur-o
'and' in .- ;.i..' .-- -
ith a t^Ji'f.'.-.-1" :L ..... '"
Pyramid on one of 5
quadrats
(zone of large hEada)
other i. :.2.:.r' .b..
f.ll -. *.. :. 7. .
:^~ M 1 ..- .. _* .
-i., .- nodv:
Mean
a- end Silver Springs are
in similar, arrestt situaticm a aim xer
.1 ji- ma,,y be t.~*'y*1o
-.' In caloa-rf;os oubstratas vm estim-ted
*Fisjh crabs, cones, annelids
it;- --
;' tjiJ'l" :.Ln "~un :L .jJh;'~7,I? *. i. +-:Y;- in,
U.. u' nivores
of 5 pyrAids
-,ha~Jirbivores
-I ,me *7*~ -- -
_________ L"-;s..-* .1 .
FI'-gwe
on~g 1><.
*. hi v.Pse I. :- caic~
ftpsland~ .~ PFi gwes In dry go
SBoth coam:initi. ; ':; a' close to a stUJr state with production balanced
by respiration in the coral.! within the limits of accuracy of the measuraments
madoo In Silver Springs there is always a contribution of the upstream
organic matter to the do1us 1 c:m coamrnities which the 7. .:c does not have
(except to a v..': small xtont) Curves 'ho.in, production and respiration
are given in Fig ure3, o0 .o : i:'. preYious data of Sargent and Austin
Ai
',
t 6 a,.m.
E.Po Odum
- *~,,
noon
6 pmor
*' -' '. :ial C.r-. of Production I.e5ai'c-.d with tVh
. a".:: ., :,' ::ahsod on Eniwetok Windward P. 7
60 r .- *
state balance for' 7-
systems tend to .':
he7ore tati suggs!
theoretical ''
: ":-' :,o to have been at least r-;mnhl!y in '..r
~: 10.Q.. timne and it is important that coutr. .. ,
vo O;h. ..to taophic structure and characteristic. a in
:\27:,r basic laws of behavior as discusesd in tho
To It is "' that to .r-.'. eafficiencles aire
achieved by a v. .nuint "-7 in K MV:C... 0 .. and by a vary low
nutrient condition n tIh: cornl ."-- 'o '.,n if the -. ,:, -' .: .,. "E', orn..ina
are available the par.' ."... .. Y.. environmental I. .,'.. .t
factors may ba circnM-., td "' a community by -.:o. :'n som part of its
energy in mechanisma '- co oar-' th .: ...e .. : ', -.. so that it is no
?i-;r MI.':..;". -* -. tha it mainta"ns a r1s energetic tax on the
overall i-; t'.:.-,., In t' : case this i$ the conservation of nutrients
in the oalicrou .' -
:o.'L ... ..-,
per :, li.'
1
aa (
-~
e-
: A ,
)
PLAi. COQu :J-Yi UTLBILITY DURING 3 YEAMS
by H. To Odum
In 1"'' X.-,. were made of the distribution of plants in
40 sorinfs. One of tho' e, VanninG Sprli.:r was j~:pcf;jced in the
first process report.
In the foT.':.'.. tr by Dr. NEtelson, entirely inde-
pendent maps are repori:od, ;'.'.: 3 years later in 6 of the sane
springs without having soen but one of the previous ;,i.r.o The
comparison of the earlier maos with the later maps is very
indicative of the de' of stability in these chemostatlcally
and thermostaticaly .regulated natural communities. It is
apparent V.'...t :la gaircl s.me e ecies dominate after three
years alth.'.. h the e..act positions of the various patches -.-.1 ,,
It should o.. ...' *ered !r:nom the work in Lilver that the higher
plants have a r between one and 10 times a year so that
there has been ample time for marked changeSo This is further
evidence along with the pr'sviously reported work on g .1 : and
insects that .. tonss are much more stable than usually found
in nature. This is not to imply that pulses and chlin.. have
not occurred in sone sprsings; n some ..'.- y'le more affected by
surface water, there .i, .io0 a known variation of chemical con-
ditions,
Standing. CGrop and Community Lurvey of
Submerged Vegetation In Seven Springs
By alle atelson
In the first semi.-annual re a.ct of this project in January,
1953, Dr. John HI Davis presented figures on the standing crops
of four springs and their coastal runs and called attention to
changes in density and comp.-,osition of the vegetation, some of
which were correlated w vth changes in turbidity and chlorinity
of the water, i. Is a report on work intended to continue and
extend the above stu.dic.:, .-e present Investioation, which l.y.e.n
in leptL.ic 19'.7, has. for its subject both the quantitative
features of the standing crop of large submer .:ed aquatic plants in
some spr.: and their : uns and the qualitative composition of
their prevalent coryrmm unit.e.
An invent -tion of the submerged vegetation in Wisconsin
lakes revealed no discrete recurr.r, communities, Instead, there
occeMd a pattern of continual c- .. e of community composition
alone a gradient complex .' environmental factors (Natelson, Do,
The phytosociot.., of c:bme -.d aquatic macroohytes in Wisconsin
lakes. Ph.oD Ths"ol.s, Univ;ersity of Wisconsin, 1954). One of the
-principal aims .. the oresent study is to determine if such a
situation exitts in the aquatic vegetation of Florida, and if so,
wh4t is the pattern vegetation here. A knowle .3 of the pattern,
referred to as "veg.eta.tlcona continuum", can be used in constructing
a classification ;-:te, for the communities and facilitates correl-
ations among ve.oetation and environmental factors. i'he Florida
studies here reported are based .i' ~ methods used in the Wisconsin
investigation.
Each run was 4 sa ipled at several stations scattered al.;!,
the length of the river. Lome subjectivity was used inasmuch as
care was taken to ... characteristic, rather than atypical or
disturbed areas, but selection of stations was otherwise objective,
with two exceptions: 1) in some instances, for comparison pur-
poses, an attempt v.as made to sample at the same station used by
Dr. Davis or Mr. Sc.an, (bloans, Wm., The dstribution of ..--tic
insects in two Florida Springs, M.oB Thesis, Univo of Fla,, 1954);
2) stations at Salt ', s run were located re ulo.rly at two-
mile intervals since the water was too turbid for reliable selec-
tion.
The s..i l tranucts made across the river frequently
traversed two or more o:-.. ously different communities and such
stations were divided into substations. Each of the latter were
sampled individually, so that the data could be used for generall
community analysis as well as for st':din,; crop and species
composition estimates for each river. For the latter purpose,
the several substations at each station were weighted according
to the area of the station which th, ... occupied, and they were
then combined. This teechique was also used for pools where a
mosaic of communities occurred
Substations or stations with homogeneous vegetation will
be referred to as stands. Each stand was sampled by 5 to 25
quadrats, according to the homogeneity of the vegetation and/or
the area of the stand, The quadrat was a heavy wire frame one
square foot in area. With the aid of a face oi mask, the rooted
plants within the quadr.ate were uprooted and brought to the sur-
face. But sometimes, where the depth and substrate were suitable,
a rake was used to den:de one square foot of the bottom (estimated),
instead of uprooting 1 -I.o
To determine the volume of the plants, the displac;er.'.+Lt
of water by the plants removed from the quadrats was measured
using the method described by Dr. Davis (cited above), and an
average volume per squ.iure foot was calculated for each stand,
Since the specific l r ".ity of submerged plants is close to one,
these p figures were u. -:. as an estimate of the wet weight of the
plants. The percent'.'.: cover of the vegetation in each section of
river was multiplied by the weight of the vegetation+ of the stand
sampled in that section, and the results were converd to lbSo/acre
wet weight for each section,, Wet weight of the vegetation in
each section was ;'..:hted by the estimated relative area of the
section in the river and the resulting figures were combined to
produce an estimate of the average wet weight/acre in the river.
'he percentag- of volume contributed by each species was
estimated for each quadrat, and an average was obtained for each
stand. These figures were then weighted by the percentage cover
and percentage area of each stand in the river, in the same manner
as described above .. the combined wet weights. Thus the per-
centage of the wet '-...:" t of plants in the river contributed by
each species was obtairedo The percentage of water content of
each species as previously determined by Dr. Davis unpublished
data) was used to oU:'. the dry weight contributed by each species
to the average dry r .' ;ht/acre for the river
Estimates of total s' .'n'fig crops, both as wet and dry
weight, and the percent., ". .C the total dry weight which was con-
tributed by each species are presented in Table 1.
Figures 1 and 2 illustrate the spring-river systems whose
standing crops were a..i..-.:.:'-Ied in Table 1, and the locations of
stations and the area estimated as representative of each station
are shown.
Across a river, shore to shore,' different communities
often occur within small areas, even at the same depth. Frequently,
an environmental correl..tion is obvious, e.g,, different substrates.
However, in other situ.t.io.ns no reason for the differences is
apparent and it is prol. '... that historical factors such as dis-
turbance, availability of propagules, and conditions conducive
to clone formation were largely responsible for the non-uniformity
of the vegetation.
In contrast, a regular trend in vegetation change downstream
occurs in some rivers, superimposed upon the more random localized
variability. In this investigation, such trends were noted in
the springs with coastal runs. In Weekiwachee River, Chara occurred
near the head, often in great density, but was not found in the
middle or lower parts of the river, Ceratophyllum demersum
likewise occurred in greatest abundance near the head, but extended
much further downstream than @har. As GharA decreased in impor-
tance, Ngjgs ~eadalusenis which was absent from the upper part
of the river, appeared and rapidly became the most abundant species,
Sagittaria was more prevalent in the upper part while Potamogetn
peotinatus and Vallisneria neotropicalis were apparently restricted
to the lower part of the river,
Chassahowitzka River was similar to Weekiwachee River in
some of its vegetational features. Sagittaria was most abundant
hear the head while Vallianeria neotropicalis, aae guadalugensi s,
and Potamogeton gecSinatus reached their maxima in the lower part of
the river.
Such trends were not apparent in Homosassa River, except for
the occurrence of large amounts of filamentous green algae in the
middle section of the river although it was rare in the upper and
lower regions. However, a distinct change in the character of the
vegetation occurred near the Gulf where tidal waters introduce
salinity. There Po~fSo3Sgon pectinatus and algae of marine type
occurred, while the common upstream species were rare.
These three rivers run from their head springs to the Gulf
and thus contain a gradation in chlorinity, as was shown by data
presented by Dr. H. T. Odum in the January, 1953 report of this
project. Tables 2 to 5 show the qualitative changes in vegetation
which occur in some instances from the head of a river to its mouth,
and also show how equally great or even greater variation often
occurs among substations at the s.me general location. Thus it
seems that excluding brackish waters, changes in chlorinity are
probably not as much the cause of community differences as are the
changes in substrate and turbidity.
Changes in community composition in the Salt Springs run,
which does not flow into salt water but into Lake George, showed
no consistent trends, except for the fact that Potamogeton pectin-
atu was the most abundant species in the pool area and the beginning
of the run, and was absent or rare elsewhere.
Hart Springs run had essentially the same plant composition
throughout its short length.
Sources of error in this work fall into two principal
categories. The first results from the mosaic arrangement of
the communities which is revealed by Tables 2 to 5. Because of
this, the error can be considerable when the standing crop or
species composition estimate for a river is derived from a fairly
small number of stations. The second source of error arises from
the necessity for estimating the plant cover of each section of
the river and the extent of the section represented by each stand.
In addition, there are small errors which result from
difficulties in sampling, i.e., the current effects which often
bend the vegetation horizontally and do not permit the quadrat
to be dropped over the top of the plants so that one square foot
of bottom can be denuded. However, these errors may often com-
pensate for e ch other. For example, the excess weight contri-
buted by soil particles, which usually come up with the roots,
is to some extent compensated for by the fact that frequently a
large part of the root system is left in the substrate.
An idea of the relatively small size of error or variability
in results obtained is shown by a comparison between Dr. Davis'
results (cited above) and those of this investigation.
Lbs./acre Lbs,/acre
Dry Wt.* Dry Wt Error
Weekiwachee Springs and River 3941 4686
Chassahowitzka Sprin:s and River 4620 3667 26%
Homosassa Sprines and River 4000 3774
Average error =16
*From Dr. Davis' data.
#From the present work, converted from wet weights by
conversion factors,
rhis comparison of results of work done entirely independently
by two persons leads to the conclusion that standing crop estimates
are relatively good first approximations of the productivity of
spring-river ecosystems. However, it should be emphasized that
it has not been possible to determine from this study what part
of the differences between the two estimates fS the same river is
attributable to chance errors in sampling and which to possible
actual differences in the vegetation when the estimates were made.
the pools of some Of the spring boils were mapped and are
shown as diagrams in Figures 3 to 7. The areas of similar veg-
etation (stands) in each were delineated and the communities analyzed
in a manner similar to the substations of the runs. Each species
is represented by a symbol whose frequency in each stand on the
map indicates its relative importance in the community as determined
by frequency calculations and volume measurements or estimates.
These maps i.,. .i ,,e ,J .d veral different comiauniti..es
often occur withinn a r,. 11 ;rea. Work on communities will con-
time fth. the epri, 1:'55 probably will shed .li.h~ upon
some of the cormmuni.:ty a .::il ite relationships,
Notes:
1) !'. aP !S mentio.od .: r
their tax -. aut .
f. ,- A.i"r* .
n ;V ,';. .. ) A.D Hito
fl-v 1.'1- 4 qI Marie-ViC
report are listed below with.
che
t.
2) -.. loc tic. A-,: riotions of the spr&D s.3 and ther rive
discussed here r. ,. n F.. .. --on, G.E., C. W. Linuch.am,, F.
Love, and R 0.O V ....:, p.ngs of Floril- Geoloical Bu.lleln
No. 31, State of -.. cart ment .' o Cons0-ca:.ctlon, Tallahnorea,:
19470
Table 1. Esti ..- 1 '::'.. c rop and percent .:. con-
tribute d. b: v, : eceis in 5 springs and their rv-.: .
Date
Wet wt. lb!4/acr:>,
Dry wt. lbs.J/areb
35272
4 D
'1')~
''f" :Y flfl
10/23
2375
3437
H ono.
10/5
37926
3774
Salt
10/1
20137
2045
H rt
10/5
Percent- > thr tot:.
species:
Algae (fiilamentous)
Ano ch n. is carnSre u
Cabomba op.
Cerator'Illui r,, .zA,
Chara : ...
Hydrocotylo -, ,
Ludwigia n atia;a
Naji. n e ..-.. is
Pot ,_r cton illi. oi n)o '
P, ... ,"'i" natus t
Rorippa s4-s5.fo ra
Sagittaria Oppa
Vallisne t
Others
il'- t which was contributed by each
3.3
9.0
7' 33
1.,8
303
6.5
2.9
13306
33.8
4.3
3.3
22.9
40.9
0.0
16.5 o0. 1
1.1
66.4 ;2
1.3
7.9
27,6
9 1
2,2
o7
22 4
.6
11,0
6 .1
36,2
4.3
Fanning
10/7
66%
802
Table 2. H-..... .'a .i-i.v: Composition and standing crop of
... action n at t .he :,*. station community level.*
Importance**
1 B i B2 B3
S .'10 .
C B1 D2
E F G
Sc i ttaria s
VvI11s.. -ria rrot-ic.: ... 3Y
Others
lbs /acre wet UI.:tl b W
z 20
S 18
2
53 19
48 A'.
23
3
67 65
42 3 21 33 19 .1
"' .5 10 717 604 6'. C4 95 20
* Communte iit n:..n :
are subte tat:on t
* Imaportanc- e V. .l.V. 1
quency and V
# Al' f moY iwe i
17. the same letter .::.. diff:ery
no. ':;a Stiow
1 ; "- Vx .. d. i -l
1.: o 4 as .* &.u*. d by dis" .
r ':; eripts
117 C.i fre-
cut of water
Tc.l.n 3.
veg t ...a
'f:1'er 0*...;:,-: 'ition and standing; ::.Co of
" ...u. t.ton community levels
i7worance
A ?_ ... o f
,i "S l "; '...
N, g- .'; '" 1 7 ,
Pa. 11100t'u V1
P. pectinatusa
Rorippa aesa i.lo-.ra
Saglttaria sp,
Vo neotrop an -l., :
lbs acree wet y, .
A B C G2 D 1 D2
' 7 10 1
S8 3
.: 15 1
6 15
37 22
El E2
10 "'
; 100 ", 41 76 59 54 9
;' 13 10
. 1 ,.'i 210 340 330 204 120 320 952 150
.-1 G2
52 82
Weekiwache. n .v:r Composition and standinR crop of
v-,:_ .., tion at ti.e substation community level.
species
Algae
Ceratophyllum .2"sum
Uhara sp.
Ludwigia natana
Lajas guadalupenasi
Potamogeton pectinatus
Sagittaria sp.
Vallisneria neotroplcalis
Lbs./acre wet wei'ht
Importance
A B C E E
7
24 49
9
100 59
1000
20 37
675 320 180 440 180
1 180
Table 5.
Salt vor: C opposition .An.-
at the subat.ation community
1 standing crop
level.
of 'c.L station
Importance
Algae
Ceratophyllum demersum
Ghara sp.
Najas ar: .t:, i'ensil
Pot.ear:, [,eton y.v. tinatus
Vallisneria neotropical s
Lbs /acre wet weight
41 A2 A3 B
4 11 4 16
18
24 60 23 49
11 49 2
15 71 24 16
C Dl D2 El E2
8
21 41 2 4
2
79 51 82 19 41
14 76 59
A0 200 300 828 193
Table 4.
8 83 230 40
Figure 1. Locations of stations and section of
river represented by each station.
1 inch 1 mile
c station
i,'.OSASSA SPRINGS
Gulf
CHASSAHC',9 '" KA INGS
Giulf I
WEEKIWACHEE SPI ''i"'
F
Gulf
of river
A1 25
Al 2.5
A2 7o Substations in Weesiwac-e
A3 8-6 River are estimations for
B 3o7 sections of the river which
B1 25-0 were obviously not represented
e 18o8 by the sampled stations.
D 6,2
E 16.2
F 2.5
A,B,G C m
sa 7% C. P57 -33%3%
oj river
L o0c atis fsa ?'o sTations
StatoC-S are in reJ.
Hart Sprias
So Feet
t--~-----------I
o-nj, small
scatTered 4j1l a ats
on bo0tte-m
) This side o
i\ has:;
t57. Nasit evedalumtn u
S Ceraltepogilw
s57. Char
5%1 4se- e
Scirpvs conryoidCe f0
Pflar -o-mmynit recordC- i-n TblCe .
A-rea 'asp ed is ou'iinae in red.
5alt Springs d oe trn
I ch a Imile
Fi 4re 2.
t
o'
\c Feo 'To
31V .ifS V
N~ D R"''-s /.- --"
'OO
Nephalr Sa, H ~droof /
I ,' .:
Lem je- tl. ?ede a c I
C~ ~~~~~~i &'< .'* e-*s^^t!'v "*
^ ** i ., ; ^'
^em*-.* s!'1 c.~aa~a ~iyis~a.^;
i -
if, "~:
k-.I.
f/
I -' ...... ---:f .
-. f i- ^ .'- /
if.
-if-u-~c )
re
*i
'if-
,;\ .. s" --,,, ^-,.,, \..
'if,
i''-.,.< N",F \ "
-' N .-
-'y .: ^;i i^ \- v
-|if|% 3
3~
C~Lvd;3aa vnta-hs
A:_O'fp SB~ij~
C
0-~
: I II -
~1
i.\ \
33
e 5 11 f we 'a
cot yle sp.
L aaw dt a r$ s T
Fdia -nTi ll s sp.
Chara
NadJs dv uada fvpen rsi
MA rl '-i-llum
1 '
Bethkir Area
--
e
*" .) o-, ^ A **.
*i c '''^ ^ e: ^
o, :o
so 0e4t
--H^~--
/ 1
0 e
/ .3 s
N.
*o:;0 tha~tuckte Headspri^Q Poo!
);, ~SeTevmber 21 '5
'SO -e t.
Afc-io~f't o^ea o-ac ? S~aty r
fcse y o _iTeatv T f jen h
Figure 5
Gr a Creek oof
at. Chassahowi tz ka Spr s
October 17, ?S1t
so Feet
------------,---
iva Njas udlpns
L~q~iri&r Sp*
2Aksln!2 eer-i -ctr,? i ca
A a3 e
pIvi CC~'toph/Ilaw dCperustis
2ikn;cheli~a paItstris
a -
N- -' d?1
-Ar'- .. .1
-t .--,'g ~
5' -)
House
- Chassa
\-
40.^ I .f<
u^^ Bwt ---Y -- ----- --~
Goat fat
d Revials
F% ure b
fleaJsprno' ok Ctassahowdzka
P4o ata-n eIanoensi5
o~t CeratoehlIntl c~ erxPsuvaj
fiver
/)To c Ier f 16 t
2'o Alqte
FIcan l i sa of alg ae w-th
ius saa'TTaTia Pistia s raeles aind Eichanor-a cvtssipes
NadJas 5uabeIElanisl
p-N 'I ';
r~; ----
e't~3 .0? i'e -0
--. -'i~ "-- -.4r
-- ''r- JI I) beGfl
: .Ci-, 't- i
'SJ ri-" t~~, "
1 9 r (C
-t on r :-
; '4: ~ I;`
P~staa
Ir
S.. I ^n.
-.-.---4-.- -- -j -
P --
0 -'
Fi
j \ ^ ^, -- -'"-l~li^ '/^
V -
5,pT tmer 3a 0 19S1'
*;, Po ost os eaon- ,: ;.'ates
V a l SW b V i0 i e'l 0 S
t<.a' sjas ,~ ad p si
pIus a l a
C50 F-e
I^_r__--P9^L~II-- \
J y
:: ":~, .: IN SILVER SPRINGS
By David K. r ''' .i'j -.ck H, Berry, and Howard T. Odum
A fish '. ': program in Silver Springs was begu n on '.i, ch 12,
1954, and has been continued by various workersto date. A total of
19 trips have been made Jwith the assistance of a number of volunteer
helpers. To date, I?.'. fish have been '.,.:-nO in Silver Springs. Two
types of tags have been used. Plastic Peterson disc tags of varying
colors and sizes have been used on the larger fish, and small metal
clamp tags have been placed on the jaw or opercle of smaller specimens.
The latter are .: I 1 while the unnumbered plastic tags have been
used in various color combinations for each fish, or have been used
cut to different shapes w;.ith the same color combinations. Most of
the fish 1.- have been Largemouth black bass ([J-ot-,i- salmoides)
and Stumpknockers (T puinctatusy, with some other ..;-;;. i.d-.:.lds
and a few other nuibbers of each species -'. .-:d and the
type of tag are summarized as T. :.. 1. As summarized in T ,b)., 2, ten
fish have been reca:ptired. although enough time elapsed between
tagging and recapture for soEe growth to occur, some fish have not
shown any increase, and others show an apparent decrease, Whether
these growth values a-re ty 1 or whether an artifact or injury
is involved is not '.clear. bass show the most growth, and
all of these specimens are jui'.enil.es. It is interesting to note
that all recaptured fish were retaken where they were initially
caught and t. .'
p,-- r .. 7 .. Stulpknockers (the dominant r',-...:p ) have
been taken each month s : 'in "i.,- : and in '=.:-:.ah, Length-
frequency curves and lengit-weight curves have been constructed
with these. The length-eight ratio does not vary materially from
month to ... :t at least for the sizes measured. A typical r.'1.. -
length curve is shown as 'e 1. Enough specimens of other
species have not beae taken :. the construction of such curves,
Monthly length.-: ,y gr 1.. for the Stumpknocker do not show
an distinct age ,; .. ('- .P 2). A -::. long breeding period
is thus indicated. '-. -' is also evidence to .;',r:, this from
observations of :..-. .' .beds and r,.. (or nearly so) adults dur:.
most of the warm months. 1'0 I an entire winter period has not
yet been .: !'i .", evidence indicates that spawning, although rare
during this period in Silver Springs, does occur. Individuals with
developed gonads v:ere taken on October 15 and on ':...:1.r 15. Also,
small specimens were taken ".;& : the winter (Figure 2).
Samples of scales have been taken throughout the study and an
effort is now being made to determine if these can be used in
determining .,. and rate of ...:-. This study is primarily being
done on the St-;-.-- -, K. some attempt will be made to study
the scales of the other Centrarchids, particularly the bass.
Preliminary studies show the presence of r;w-.i., but fuj ;h::'.! study is
36
necessary to determine if these rings represent true annuli. Scales
from this constant t'...:-, ,i:..; spring will be compared with scales
from the same species from other (non constant temperature) Florida
waters, and if pon.. i.?~l with scales from northern waters.
A straight line ratio has been shon to exist between standard
length and total 1..;! :..1 the Stumpknocker. This ratio (S.L./TL.*.79)
exists throughout the entire size range for this species as encountered
in Silver E ::..i.;s, and will be helpful in comparing the work done in
Silver Springs with that of other workers on the same species in other
areas,
Coincident with the fishery work on larger fishes, a general
collection of small fr' :, invertebrates, and algae has been made
for further study of seasonal periodicity of reproduction in this
constant temperature environment.
NUMBER WITH NUMBER WITH
SPECIES METAL TAG PLASTIC TAG TOTAL
Lepomis j. punctatus 83 147 230
Lepomis macrochirus 12 12 24
Legomis auritus 3 0 3
Lepomis marginatus 1 0 1
Lepomis megalois 25 9 34
Lepomis microlophus 1 1 2
Micropterus salmoides 40 19 59
Chaenobryttus coronarius 11 17 28
Esox americanus 1 1 2
Pomoxis niromaculatus 0 1 1
Leiosteus platyrhincus 0 3 3
Anguilla rostrata 0 1 1
Erigzon sucetta 0 1 1
Ameiurus natalis O 3 3
Grand totals 177 215 392
Table 1. Summary of fish tagged at Silver Springs, Florida, between
March 12, and December 29, 1954.
Species
Micropterus salmoides
Micropterus salmoides
Miropterus salmoides
.icropterus it, c.l.:
_.r---. to salmoides
Chaenobz ttus coronarius
w:w-: _:- .. ..
Lenomis n _nr___*ct_
T... I .- "L i .; '
Tag
Type
Metal
Metal
Metal
real
Plastic
Metal
Plastic
Plastic
Date
Tagged
VI-11-54
VII-25-54
IX-24-54
VII-25-54
IX-24-54
:7--2-54
VI-18-54
VIII-31-54
XI-24-54
Date
Recaptured
IX-24-54
X-15-54
X-15-54
XI-24-54
XII-15-54
XII-29-54
X I -54
X-15-54
XI-24-54
XII-15-54
Days
Out
105
82
21
122
82
96
119
119
85
21
St. Len. (mm)
When tagged
124
112
116
110
120
105
116
112
142
214
Table 2. Recaptures of tagged fish at Silver Springs, Florida.
St. Len.
at Eecap.
134
131
115
128
124
107
116
117
139
214
Growth
(mm)
10
19
-1
18
4
2
0
-3
0
80-
75-
70-
65--
60-
55"-
50-
45-
40-
35-
30-
25-
20-"
15-
10-
5-
I I I I I I
10 20 20 40 50 60 70 80 90 100 110
Standard Length (mm.)
Figure 1. Length-weight, Lepomis gnctatus punctatus, Silver
Springs, Florida, October 15, 1954.
120 130
--
--
L
'T4
t
..O
Figure 2. Length-frequency graphs for bE mis aunctatus, Silver
4U
Springs, Florida.
t
I \
I\
, / / I I
I' 1
*ft i
MARCH
1954 -
I
j? D
.-.. 4
ft 8, '
NO APRIL OR MAY SAMPLE
'4--
,- ,' JUNE 1954
/ a --. '" ..
JULY
0 '4
19g4
N /\
0 %"
10- --
SAUGUST 1954
0 - ---- ---*---- -.1- <-- --
15-
10- / SEPTEMBER 1954 -
5- \ ... -*
.- o--^ e------.a -
10- OCTOBER 1954 -
0-
--" ^ x-^---^-- ----- :-`-`---
10.- /- NOVEMBER 1954
5-A $ DEE\ R/ 19541
__--a-B ~t>~~I-----D--ECE-MBER- ~ 1954
"-' DECEMBER 1954
'i 0-__
I I I I I I I I I I I I I I I I I I I it I I I Ir- r-I I I
", A r-1 r A 0- A -- -4 H- H -H r-H H r-1 r-4
STANDARD LENGTH (mm.)
STANDARD LENGTH (mm.) H ? 4 4 H HH4HHHHHr
40-
35-
30-
' 25-
20-
15-
10-
5-
0-
!I~ii~t
10-
G-
2(-
10-
r5-
/ N 0 /
0
q .
0I
To
^.(
U____^______IY___II___
- -~-- ------ -~----CIII-~-*IUII-~ --L---ZIWIII-I.I.IC~LI .-LlltlWIIIII-YI- I__
HTT.rviTV T55o-
HoT,:, 0''."
A., p .'iTi D tlu t.. :,.. --.. .'. ", ;.r Principle Applied to itasyNiTh.;*,!L
SIn --prcr.....s :;.,*., ::i;,. tha principle has been stated that open steady state
systems tend to be adjusted at an optimum but low .[f. cl :.acy that corresponds
to the maximum power -....;.. (" ., a.nd Pinkertonp American Scienti f.., in press)
S That this pr:inY.. is ,.,L .' -*:,: ~A~.vi.. systems seems indicated in
Figure 1$ below:
Snatural or high light intensities
(kW -Ao -.'I photosynthesis
per fir quantum controversy
'.. ;;.!.1 ..;.t ./ -.
low but,.;.. i-, -""e. associated with maximum output
This graph shows that '.,-, ..,,.i. .'. such as have been achieved in
all the work associated P .0, the quantum controvercey have all been r.-'-v.!
light intensities so that the .:.: output of glucose has been .'.n.
On the other hand the .";."': populations of algae d.3ui.:d to h~gi .natural
light intensities run at .., ....t..a.s and h... .light intensities but so
that a much greater :i ,.". of .'-.-.S rrn.s ',-.. If pl nt. are evolutionarily
adapted to maximum output A%"- must sacrifice efficiency for power by
this hypothesis This is a .- of ett ..,': that attempts to increase world
food by raising chlorella at ..,;. .,A"Jci...., must necessarily flopo A
second part of this ..,i;. .."..j may be stated that the optimum efficiency
for maximum power output :.'; -- .. decreases as the light intensity increases
Plants adapted in nature to deep water achieve the optimum adjustment
at a higher effic:-:.m-vr that at the surface "-iv.,v a plant adjusted for
one light intensity can not be moved ir,~,O. *'it.ly to another light intensity
and achieve the ct-n., i', '.... n without internal modification. A car
climbing a hill in .*;:..:'' ,.. *at optimum ..':..irj .nr.cy cannot achieve the
optimum efficiency for a -1,; .t away without -'.n..r Q a-. In the
plant gears may be the concentrate tion gradientso The efficiencies in
Figure 15 are of the same .d a, at optimum adjustment as those
found in Silver Springs and the Coral Raeo?,
Bo Organional Size versus .l .~".: TA.L: in Phototrophe in Optimum Adjustment
It is now well know that amtabolism of heterotrophic organisms is inverse
to body size roughly in a 2/3 o'..- '.... function that is presumed to be related
to the surface/volume 7514:. to V02ll' n processes. Ma the above section
it was concluded that 1 .... .' y, .'; in open steady state tend to
all become adjusted to a similar state of running at maximum power output
because of the survival 4 '''. in : l. ":7*. .":. n'....'..n both in an environmental
and evolutionary sense Xith .r,.- intensitiesA similar steady
state plant ~stfli ..' ...& be ;". .-..-i to similar efficiencies and thus
similar total -.ho.-. .,.:. .. power output of pl-I ....co an onAREA basis
Now if the plants run at the same rate of output per area and if size
effects hold for plants as v:.:.'l as animals, then small plants like 6hlorella
if growing in steady state ~:.I-.'il. achieve the same output per area but with
less biomasso The large climax rain-forest with big plants on the other hand
if adjusted to the sams light and optiImum efficiency will require a much larger
standing crop biomass because of the slower metabolism per pound of tissues
In Figure 16 -~lhcr is sih~ n a graph of photosynthetic rate of plants of
various minimum diameterP under natural light or maximum photosynthetic
adjustments, The V-;-;.. a j. '1.'Ci V :..c'l 3i-,
data from
Verduin
mum o001l- 0- 3
SAnnual
Photosynthesis a Surnover
Gm glucose
per gm dry o.01 -3
weight per
Hro ,
r 01 *'3 / ""300
1o .,.
3000
,o01.01 .1
Size in Cm
Figure 16o Effect of organismal size on Photosynthesis per weight
It is apparent that there is a size effect over a wide range just as in the
heterotrophso Thus, 0.-i1' ,o; the light intensity, the size of the producer,
and the rough offL'.-icJ-'.?.,! f*-.:.'.. in given environments, one can compute
the steady state b.:'l.,C car.. rying .-p.ci. o
Co Pyramid '. '... nd r':-:; .:. ..'z. Size
If as o.tl ,.d above siall sized producers put out the same production
per area with a . r.1.J, biomass as do large producers with a large biomass
both working at a.L.:.e.~,: offici'rl:,. one can visualize two extreme types
of pyramid as c gap .:i.'.; in C..r 17 (For data showing similar
efficiencies and o.'..;.:..i ?.:, mass Shlorella culture and grass plots
see (Burlews Mass Culture of 1~.:...b --.[.t-e ---Wi.e.,r-b..,k et. aloCarnegie@ 1953)
In one the size of the organism decreases as one goes up the food chain as in
grass-grasshopper--o.idA lr.'o In the other the size of the organism increases
as one goes up the food chan as in .L.0i' ulla, paramecium and fish, If the
same energy passes up tbr',^ .':cr:t food chains with the same 10% efficiency
for the higher trophic .?;:'- l, two ~I tir'l different shaped pyramids of
.at~Judy state biomass result because of the different rates of turnover
Some metabolic rate fV. i;::-.. are used to compute Figure I? from H.il';nn s text
Photosynthatic values are taken .:.i..:.i Figure 16o
The pyramids in Figure 17 I'.>ip to visualize the possible relationship of
the tropical ocean to the tropical rain forest. The small size of tropical
plankters as well as the h.;I -..: .'..r: tend to cas o the reversed
pyramid calculated for Chloralla--paramecium-fisho
Spidr 22 Fish
Grasshopper 62 Paramecium
Cas* '810
810 _.6 Chlorella
Figure 17. 'i *... !,.:;-:. : with the Same Energy Flux sho' rinytr the
,p.,.;..c, of A.., ,. : .,d h '. on ,rE:.niaimsl size.
Do Energy Contributions of T'dr C::. '.:' and Curr~nl-, Plankton Size
The efficiencies of primary production so .?--ir estimated for Silver Springs
(8%) and for the ,;.-i.,';,'.. .-E ', are 'o:;.!? .;"i2 nbi.;- higher than many other
natural communities or 1.:.- V-'..:v experiments ran at high light intensities
for maximum production 2:icj there is a ila.s difference in nutrients
between the roof and Silver Springs it seems that some other property
is in part responsible for this high production. It is reasonable to
postulate that hi, e-:.. efficient ies are produced by the strong currents that
serve as a community cir .. :' y.:d, rnintai,'i)g better nutrient
concentrations a ..1: .t to alls and r: p.ovini waste products
The effect of the circulation might be stated in two ways:
Ao The community receives .-;*'.*; ," :'i both the sun and from the current
Bo The energy directly from the sun goes further because of the current condition.
If the usual efficiency at maximma adjustment is about 2%, about 5% might
be postulated as due to the current system This might imply that these
flow systems dlerive wre of their r ':,vrgy f..'<::- primary production from the
current energy than *:..,n tho 1~-.ht received directly
If current is as important as ipi,-i' above~ a suggestion can be made
as to why larger and heavier : r...rkton usually prevail over ~:-..-.~I'r
species with f?: (cr"%:.'s so that theyy may float with the same density as
water Heavier 7: 'e.,"'t.rs are ,;.i;.i 'i::d by the turbulent eddies so that
they are continually iall.l-:,. through the currents that support them This
mechanism provides the ce ll vth a local current which the organism at
water density would not have.o Th relatively low Chlorella efficiencies
obtained in high light ,nt.. 'i.-.': ;.7 mass cultures in spite of high nutrients
may be accounted &.'."r small size of 'ic:a.'l" which decreases the effectiveness
of stirring mechanismso
BE Decrease of Daytime Plant !epi:ation Accounted for by the iypc '~.;i.is of
Plant Respiratory Sy.'~-.rv' as herbivores of the Autotrophic systemai- Arctic
Significance
The data of several ainT.: ,o (ioeo Kok, see Rabinowitch) suggests that
plant respiration in the 2 .~'me is much less than when the light is rii n ; the
compensation point The :;o;,: ;, of pyramids of biomass which may be
supported during - :'..-'- state suggests a reason, Whenever food i'rg,
is passed through an energy '.. ;.::li.l:. :-.s' step a lirg-c percent, perhaps
90, must go into heat as ri.- ..-:'r A, by the second law of th..y--. .:y;ir.ii:': for
these types of reactions under optimu.n officiencyrmaximum power adicij:.;..,:,t.E.;-,
If a plant in the day can drive some of its work systems dir-c.,-,
rather than making glucose first and then b.'niiC the glucose, the
plant can save a step and thus avoid the heat lose from the extra step,
Thus a higher efficiency in the b.,:.'o'"phic system is maintained with
lss respiration, At l1'-.ht, by this view, it becomes necessary for the
plant to fall back on .'. ,,;-. stored thus lengthening the chain of
transformations and :. ,'..'-.".? increased respiration
If true,this becomes : ,-i '. all.y significant when length of day and night
is considered, for a .-.;,: .:.'.? 'ence in day Ien, means increased efficiency
as well as additional .i.,:1'., i!l; .- is less glucose that must be stored for
use at the inefficient ,'*-i-,. rate, By this view Arctic plants during
continuous cla5.,'<. ,, of suaenr should be much more efficient than similar
temperate pla'r: On an aual basis however there would be no gain for
the community would have to store ::..:-i.;y months worth of organic matter
at the 'inefficient ::..(. rate
Fo Anaerobic .i,:,- o .. nd tbh -:. .;r-., ?::. t,-.I
The saprobs system of classifying pollution communities assumes
the association of characteristic indicator species for different degrees
of sewage type pollution A c:.;:-'i4o.t.!t,-n of Florida anaerobic springs
shows that the e'_.' while ": r'1 when properly used has a fallacy that
leads to misl:,'..u.,; cc,.'-. .:..J when :.ei D- on waters in general
Beecher, Orange, Warm 2 11.., and Volusia Co. Blue ?pr)m: are all examples
of large springs with low or: .: content water that is also low in oxygen
The communities that result are both anaerobic and autotrophic in nature
In contrast sewafg communities are anaerobic and heterotrophico In the
springs one gets sulfur bacteria and blue green algae but no ciliateso
In short one gets some of tho saproba system biota in water that is
the extreme opp.s'!it;1 of *.. .:.- polluted water, Thoughtless use of the
saprobe system -:.i to a : .:,*' :.-' misclassification of the
type of primary production in these cases.
Go Some Definitions, S, ~;-! methodology
In agriculture, mans r:.i h..-di labor and supervision guise a complex
community in a direction he desires o However in Ecological Engineering
the outcome of production of a complex community is achieved by proper
selection of components at the start with subsequent hands off thus
permitting the n:.:_nity to reach a unique steady state adjustment.
In the usual scientific experiment, man controls one variable so as to study
the behavior of another dir:.d-irjt variable while holding other conditions constant.
Thus the process is one of analyzing component processes In Microcosm
experimentation on the other hand components are put together and the complex
allowed to make its own trends under observation
The study of ecological ,. :. I: .-I.ng by microcosm *;r,.-:rr.,.ian ,.To,: is
a practical met.ho:_:..1o,-,r for h.-..., the ";[:.l 0.,'i ,, .--.. ,i .l .--,.:.t.',.1.
This ;.'.. n i'.. ; .*..*. ". is ma : .., % ..: of the icrocoom :,' .:.-.,
A hypothesis rei;.eirding dependence of community
structure and density on productivity
by J. L. Yount
The hypothesis is offered that under similar general conditions
the species variety is an inverse function of the community pro-
ductivityo
The salps of a series of plankton samples made by Pacific
Oceanic Fishery Investigations of the United States Fish and
Wildlife Service in epipelagic waters of the central Pacific
Ocean were studied dur:i., 1952-19549 Observations on them led
to the formulation of the hypothesis presented below. The most
pertinent observations were as follows. In most of the samples
studied, many species of salps were taken with little predom-
inance of any one species. In one sample, however, there were
both a far greater total salp quantity and a great predominance
of one species of salp, only a few others being taken and these
in insignificant quantities. All salp species apparently simul-
taneously' occupy L~Ial.l!.r niches (the concept of the niche used
here is that of Elton, 1927, Animal Ecology: 63-4), and appar-
ently also are subject to the same environmental conditions, thus
apparently are ecological equivalents (in impoverished waters;
see below).
Observations ii':r: by other investigators are also pertinent
here. Students of marine plankton of hiLn latitudes have des-
cribed it as "monotonous", consisting predominantly of one species
of organism in each niche .-ppr~arently, although the term niche has
not been applied in these descriptions. Mo-t descriptions of the
plankton of low latitudes, however, emphasize the great variety
of species with little or no predominance by any one species (per
niche) (see Steuer, 1910, Planktonkunde: 601-4; Russell and Yonge,
1936, The c71 ; 123-6; Dakin and Colefax, 1940, The Plankton of
the Australian Coastal Waters off i'ew South Wales, UnvJ _Sdney,
Dept2 ZoolJg Publ. I: 27-34). Another pertinent observation
discussed by Steuer and Dakin is that productivity in the tropics
in waters influenced by land drainage and in regions of upwelling
may equal or even exceed that of high latitudes.
If these observations are considered together, it appears
that in epipelagic waters with relatively great quantities of
nutrient chemicals (the enriched areas), production of the plank-
ton is great in quantity but trends toward few species of organisms--
probably only one dominant species oer niche--and in epipelagic
waters with relatively small quantities of nutrient chemicals (the
impoverished areas), the plankton is small in quantity and trends
toward many species :.' organisms--apparently many species per
niche.
A hypothesis r;,.arding dependence of community
structure and density on productivity
by J. L Yount
The hypothesis is (-'..:.red that under similar general conditions
the species variety is an inverse function of the community pro-
ductivity.
The salps of a series of plankton samples made by Pacific
Oceanic Fishery Investigations of the United States Fisi and
Wildlife Su-.-'ice in epipelagli waters of the central Pacific
Ocean were studied during 1952-1.951., Observations on them led
to the formulation c"., the hypothesis presented below, The most
pertinent observations were as follows,. In most of the samples
studied, many species of salps were taken with little predom-
inance of any one species. In one sample, however, there were
both a far greater total salp quantity and a great predominance
of one species of salp, only a few others being taken and these
in insignificant quantities. All salp species apparently simul-
taneously occupy lmiinlr.r niches (the concept of the niche used
here is that of Elton, 1927, Animal Ecoo: 63-4), and appar-
ently also are subject to the same environmental conditions, thus
apparently are ecological equivalents (in impoverished waters;
see below).
Observations made by other investigators are also pertinent
here. Students of marine plankton of high latitudes have des-
cribed it as "monoonon.i:-", consisting predominantly of one species
of organism in each niche .ppar-r.ly, although the term niche has
not been applied in these descriptions. Most descriptions of the
plankton of low latitudes, however, emphasize the great variety
of species with little or no predominance by any one species (per
niche) (see Steuer, 1910, Planktonkunde: 601-4; Russell and Yonge,
1936, The Seas: 123-6; Dakin and Colefax, 1940, The Plankton of
the Australian Coastal Waters off New South Wales, Unlv Sydney,
Dept. Zoo2001 Pub.l: 27-34). Another pertinent observation
discussed by Steuer and Dakin is that productivity in the tropics
in waters influenced by land drainage and in regions of upwelling
may equal or even exceed that of high latitudes,
If these observations are considered together, it appears
that in epipelagic waters with relatively great quantities of
nutrient chemicals (the enriched areas), production of the plank-
ton is great in quantity but trends toward few species of organisms--
probably only one dominant species oer niche--and in epipelagic
waters with relatively !':,jl.l quantities of nutrient chemicals (the
impoverished areas), the plankton is small in quantity and trends
toward many species of organisms--apparently many species per
niche
If the plankton now be compared to sessile organisms of the
Aufwuchs and benthos, it will be seen that the two groups have
similar natures, in that sessile organisms and planktero are
moderately passive in ability to capture foo., and to move in
their medium (most sessile organisms, of course, move during
some phase of their life cycle, and plankters have some ability
to move, but directed movement is limited for both). NFor this
reason, at least the plankton and sessile organisms probably
should be nonsideri'. together in the hypothesis.
It is ',vi.,.,.l:' .ted from these observations that if productivity
is low and other factors are constant, species of ecologically
passive ori...,al:n at le -"1, may occur together as ecological
equivalents with little of no predominance of one species In each
niche; and conversely if productivity is high and other factors
are constant, species of these organism should not occur together
as ecological equivalents, but rather one species should dominate
in each niche~ X'hia :,!.s': to a further postulate: if productiv-
ity is low and other factors are constant, competition and other
coactions should be reduced for these organisms; whereas if pro-
ductivity is high and other factors are constant, competition and
other coactions should be Increased for them.
the followli.,. evidences tend to support this hypothesis. In
the tropic epipelr., .c holoplankton, productivity is low and quan-
tity is small, except in certain regions mentioned above, and
species numbers are great; in the epiul..'ti.c holoplankton of high
latitudes, product. ity is high .:.nr. quantity is great, but species
numbers are small. Occasionally in tropic waters, swarms of
plankters appear, con.~Mt'_:u, of few species of orgo.icon and a
relatively great quantity--productivity is therefore high and
species numbers few, even in the midst of impoverished waters,
under enriched cond lt:ons In plankton tows from impoverished
waters, many species of u. .in1..l:.ik occur together that apparently
are ecological equivalent, and this is evidently not true of
plankton tows i:' , enrich... waters (evidences for this statement
are based bchefly on studies 0i salpB, but it is postulated as
being true of other pl.ankters : well). In Florida Springs, it
has been noted (H. T. O(.::;, L, A. Whitford, W. C. sloan) that
productivity is high h*-.. the number of species of the various
groups o o organisms is Low,
his byothesls is being tested at present with Aufwuchs
growth under the controlled conditions of the Florida Springs
and, if results warrant, is expected to be tested under other
fresh water and marine conditions. Counts of Aufwuchs species
numbers on slides, relative to current controlled total pro-
ductivity, have been started Counts made on the preliminary
first series are consistent with the hypothesis
U5-7- 1 92 9
46 PF 3/,
If the plankton now be compared to sessile organisms of the /150*f
Aufwuchs and benthos, it will be seen that the two groups have
similar natures, in that sessile organisms and plankters are
moderately passive in ability to capture food and to move in
their medium (most sessile organisms, of course, move during
some phase of their life cycle, and plankters have some ability
to move, but directed mcnc~rtnl is limited for both). f'or this
reason, at leW.:,n the plankton and sessile organisms ,.. .'l-bly
should be considered together in the hypothesis.
It is postulated j't L.I theeose Observations that if productivity
is low and other factors are constant, so:pci.es of ecologically
passive orclr:n..r at least, may occur together as ecological
equivalents with little of no predominance of one species in each
niche; and conversely if productivity is high and other factors
are constant, species of these organisms should not occur together
as ecological equivalents, but rather one species should dominate
in each niche. 'his leads to a further postulate: if productiv-
ity is low and other factors are constant, competition and other
coactions should be reduced for these organisms; whereas if pro-
ductivity is high and other factors are constant, competition and
other coactions should be increased for them.
t'he followvrin evidences tend to support this hypothesis. In
the tropic epipe:.!ic holoplj.nhl; ton, productivity is low and quan-
tity is small, except in certain regions mentioned above, and
species numbers are great; in the epipelagic holoplankton of high
latituderJ, productivity 1j. hi1,h and quantity is great, but species
numbers are small. Occasionally in tropic waters, swarms of
plankters appear, consistin5 of few species of organisms and a
relatively great quantity--productivity is therefore hlgh and
species numbers few, even in the midst of impoverished waters,
under enriched cou.'llUons. In plankton tows from impoverished
waters, many species of .., 0;:if.l",- occur together that apparently
are ecological equivalent;, and this is evidently not true of
plankton tows ,'ru. enriched waters (evidences for this statement
are based chiefly on studies of salps, but it is postulated as
being true of other plankters as well). In Florida Springs, it
has been noted (H. T. Odum, L. A. Whitford, Wo C. sloan) that
productivity is hI.'. and the number of species of the various
groups of organisms is lowo
'his hypothesis is being tested at present with Aufwuchs
growth under the c ,,. tro) c.' condi tions of the Florida Springs
and, if results warrant, is expected to be tested under other
fresh water and marine conditions. Counts of Aufwuchs species
numbers on slides, relative to current controlled total pro-
ductivity, have been started Counts made on the preliminary
first series are consistent with, the hypothesis.
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