Journal Journal

ortheFisheries iCe
0",'Off
Research 0",l'eCherches
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Board pecheries
sur,es
",Canada o,Canada
1975
Volume32,No.9, September V olume32, no9, septembr e1975

A Simple Method for Predicting the Capacity of a Lake

for DevelopmentBasedon Lake Trophic Status

P. J. Dtt-t-oN

Ontario Ministry of the Environment,Ilater Resources Branch,

For personal use only.

Limnology and Toxicity Section, Rerdale, Ont.

F. H. RIclen

Department of Zoology, University of Toronto, Toronto, Ont'

D r r - L o N , P . J . , e N o F . H . R r c l p r . 1 9 7 5 .A s i m p l e m e t h o d f o r p r e d i c t i n gt h e c a p a c i t y o f a l a k e f o r
developmentbasedonlaketrophicstatus.J.Fish.Res.BoardCan.32:1519-1531.

A general technique is presented for calculating the capacity of a lake for development based
on quantifiable relationships between nutrient inputs and water quality parameters reflecting lake
t r o p h i c s t a t u s . U s e o f t h e t e c h n i q u e f o r s o u t h e r n O n t a r i o l a k e s i s d e s c r i b e d .F r o m t h e l a n d u s e
and geological formations prevalent in a lake's drainage basin, the phosphorus exported to the
l a k e i n r u n o f f w a t e r c a n b e c a l c u l a t e d ,w h i c h , w h e n c o m b i n e d w i t h t h e i n p u t d i r e c t l y t o t h e l a k e ' s
surface in precipitation and dry fallout, gives a measure of the natural total phosphorus load.
From the population around the lake, the maximum artificial phosphorus load to the lake can be
c a l c u l a t e da n d , i f n e c e s s a r y ,m o d i f i e d a c c o r d i n gt o s e w a g ed i s p o s a lf a c i l i t i e su s e d . T h e s u m o f t h e
natural and artificial loads can be combined with a measure ofthe lake's morphometry expressed
as the mean depth, the lake's water budget expressed as the lake's flushing rate, and the
phosphorus retention coefficient of the lake, a parameter dependent on both the lake's mor-
phometry and water budget, to predict springtime total phosphorus concentration in the lake.
Long-term average runoff per unit of land area, precipitation, and lake evaporation data for
Ontario provide a means of calculating the necessary water budget parameters without expensive
and time-consuming field measurements. The predicted spring total phosphorus concentratlon
can be used to predict the average chlorophyll a concentration in the lake in the summer, and this,
in turn, can be used to estimate the Secchi disc transparency. Thus, the effects ofan increase in
development on a lake's water quality can be predicted. Conversely, by setting limits for the
"permissible" summer average chlorophyll d concentration or Secchi disc transparency, the
"permissible" total phosphorus concentration at spring overturn can be calculated. This can be
translated into "permissible" arlificial load, which can then be expressed as total allowable
development. This figure can be compared to the current quantity of development and recom-
mendations made concerning the desirability of further development on the lake.

Printed in Canada (J3681)

Imprim6 au Canada (J3681)
1519
 1520 J. FrsH. RES. BOARD cAN., vol-. 3z(9), tgi5

D I I - r o N , P . J . , e N o F . H . R I c r - E n . 1 9 7 5 .A s i m p l e m e t h o d f o r p r e d i c t i n g t h e c a p a c i t y o f a l a k e f o r
developmentbasedonlaketrophicstatus.J.Fish.Res.BoardCan.32:1519-1531.

Les auteurs d6crivent une m6thode g6n6rale permettant de calculer la capacit6 de

developpement d'un lac. Cette mdthode est fond6e sur les relations quantifiables entre I'apport
d'€l6ments nutritifs et les parambtres de qualit6 de I'eau refl6tant la statut trophique de lac. Ils
J. Fish. Res. Bd. Can. Downloaded from www.nrcresearchpress.com by PENNSYLVANIA STATE UNIVERSITY on 05/11/12

expliquent comment la m6thode peut Otre appliqu6e d des lacs du sud de I'Ontario. A partir de
I'utilisation des terres et des formations 96ologiques qui p16valent dans le bassin hydrographique
d'un lac, on peut calculer le phosphore qui p6nbtre dans le lac avec I'eau de ruissellement, lequel,
combine d l'apport qui tombe directement A la surface du lac par pr6cipitation ou retomb6e sbche,
donne une mesure de la charge naturelle totale de phosphore. En se basant sur la population qui
vit autour du lac, on peut calculer la charge artificielle maximale que peut recevoir le lac et, si
n6cessaire, la modifier en fonction des systbmes d'6go[its existants. La somme des charges
naturelles et artificielles peut Atre combin6e h Ia morphom6trie du lac, exprim6e en profondeur
moyenne, au budget hydrologique du lac, exprim6 en terme de vitesse de vidange du lac, et au
coefficient de r6tention du phosphore du Iac, parambtre d€pendant i la fois de la morphom6trie et
du budget hydrologique du lac, pour pr6dire la concentration printannibre totale de phosphore
dans le lac. Le ruissellement moyen d long terme, par unit6 de superfcie de terre, la pr6cipitation
e t l ' 6 v a p o r a t i o n d a n s I ' O n t a r i o s o n t d e s d o n n 6 e s q u i p e r m e t t e n t d e c a l c u l e r l e s p a r a m d t r e sd e
budget hydrologique sans avoir recours i des mesures longues et dispendieuses sur le terrain. La
concentration printannibre totale pr6dite de phosphore peut servir ir pr6dir Ia concentration
m o y e n n e d e c h l o r o p h y l l e a d u l a c e n 6 t 6 e t c e l l e - c i ,d s o n t o u r , p e u t 6 t r e u t i l i s 6 ep o u r e s t i m e r l a
transparence au disque de Secchi. On peut, de cette fagon, pr6dire les effets d'un d6veloppement
accru sur la qualitd de I'eau du lac. Inversement, en 6tablissant des limites d la concentration
e s t i v a l e m o y e n n e d e l a c h l o r o p h y l l e a o u d e l a t r a n s p a r e n c ea u d i s q u e d e S e c c h i n p e r m i s s i b l e sr ,
on peut calculer la concentration totale de phosphore au moment du brassage
"permissible"
general de printemps. Ceci peut 6tre traduit en terme de charge artificielle .permissibler, pour
enfin 6tre exprim6 sous forme de d6veloppement total permissible. Ce chiffre peut 6tre compar6
au degre de d6veloppement pr6sent et servir de base d des recommandations quant aux avantages
For personal use only.

d e d e v e l o p p e m e n t sp l u s p o u s s 6 ss u r l e l a c .

ReceivedDecemberl'1. 1974 Regule 17dlcembre 1974

AcceptedMay 6, 1975 Accent6le 6 mai 1975

THe demand for summer cottases and other
recreational facilities in OntariJ is increasrng
rapidly, with about 10,000 new cottages Llnde;
construction in each year (Department of Tour-
ism and Information 197 1). Governmental plan-
ners, biologists, and engineers must evaluate and
subsequently approve or reject proposed develop-
ments without suidelines that quantitativelv de-
termine the eniironmental imoict of sucir de-
velopment. Suitable guidelines must be based on
predictive ecological and social theories that, to
date, have not been formulated in terms of prac-
tical management tools. It is the aim of this paper
to provide a means of determining the capacity
of a lake for cottage or permanent home develop-
ment, assuming that specific acceptable limits can
be established for some parameters that reflect
the quality of the aquatic environment. Transla-
tion of these limits to permissible cottage numbers
or other developmenf schemes can bi achieved
through application of techniques described in
this report. Conversely, if a decision is made to
allow the development of a given number of
cottages or large-scale condominiums, then the
governmental agency responsible will be com-
pletely cognizant of the environmental ramifica-
tions of its decision.
At this time, only the problem of lake trophic
status is considered, with the equally important
concerns of the effects of development on fisheries,
wildlife, and on human health not taken into
account (there is, of course considerable relation-
ship between trophic state and fisheries potential,
as well as health hazards). It should be stressed
that the approach described herein is limited by
its inability to consider these alternate parameters.
In understandins lake conditions. it is impor-
tant to realize thai the entire watershed and not
just the lake or the lake and its shoreline, is the
basic ecosystem unit. The terrestrial and aquatic
portions of any watershed are inherentlv linked
with the gravitltional movements of minerals in
drainage waters flowing from the land to the
water (i.e. lakes, oceans), as the major terrestrial-
aquatic linkage (Likens and Bormann 1974). It
is important to note that the aquatic portion of
a watershed is "downhill" from the terrestrial
portion; that is, dissolved and particulate mate-
rials from the land are transported by geological
process to the water and eventually to the less
accessible sites of the sediments in the aquatic
portions of the ecosystem. There is, of course,
some "uphill" movement of material by meteoro-
logical and biological transport, e.g. release of N,
 DILLON AND RIGLER: CAPACITY OF LAKES FOR DEVELOPMENT
from a lake through denitrification and sas diffu-
sion, followed by precipitation of the same to
the land for the former process and movement
of material through the food chain for the latter.
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However, the net-movementis always "downhill"
to the aquatic system,with the deficil made up by
geological weatheringof rocks. The implication
of this in terms of a lake's capacity for develop-
ment is that any alteration in a watershedulti-
mately affects the lake. Thus, any management
approachbasedsolely on the lake and its shore-
line is both simplisticand inaccurate.
Although a number of attemptsto managelake
fisheries by the empirical approach have been
tried (Moyle 1946; Rawson 1952, 1955; Ryder
1965), veiy little has been accomplishedin terms
of managing lakes for developmentwith regard
to trophic statusby a similar method. An excep-
tion was Seppanen'sproposal(1972) to determine
a lake's summer cottaging capacity. The author
suggestedthat a suitableformula for the recom-
mendednumber of shorelinecottaseswas
AVD;
For personal use only.
10
where A is the lake area in hectares, Dy, is the
development of shoreline equal to the shore-
length divided by the circumference of a circle of
the same area, and 10 is a figure representing the
minimum area of lake surface (in hectares)
needed per cottage for a round lake with no
islands. While providing a formula that takes
into account both lake area and shoreline length,
many other important factors are neglected limit-
ing the technique. An entirely different, although
somewhat indirect, approach has recently been
used in a lake management sense.In 1968, Vollen-
weider published a plot of total annual phos-
phorus loading L (the amount of material added
per unit surface area of lake per annum) vs. mean
depth (z). He found that bands could arbitrarily
be drawn separating the lakes into the three
standard lake types in terms of degree of eutrophy:
oligotrophic, mesotrophic, and eutrophic. In 1973,
Vollenweider modified his relationship to take
hydraulic flushing rate into account. This was
accomplished by incorporating the mean residence
time of the water in the lake (t,,,). By plotting
L vs. 2/ 7,,., a more realistic representation was
achieved. The parameter 2/ r,,. is equivalent to the
areal water loading (i.e. the height of the water
load over the lake's area that is supplied in I yr).
The L vs. 2/ r,u model has been used for predicting
changes in trophic status of several Muskoka
Lakes (Michalski et al. 1973) that would result
from chanqes brought about by the reduction of
phosphorus in sewage effiuent. As effected by
Michalski et al. (1973), the model can be em-
ployed by environmental managers in the same
manner as the L vs. Z olot: the maximum addi-
tional loading to a lake that will not surpass the
"permissible" loading can be calculated and in-
terpreted in terms of cottage development. Al-
though a significant improvement over the L vs. Z
plot, this method still lacks quantitative pre-
dictibility in terms of water quality parameters.
Basis for a New Approach
It has been demonstrated on numerous occa-
sions (Schindler et al. l97l; Schindler 1974;
Fuhs et al. 1972) that phosphorus is the nutrient
most frequently controlling production and there-
fore troohic status in north temDerate lakes.
Thereforl, any approach to predict water quality
from a troohic status ooint of view must take
into account the imporiance of phosphorus. As
early as 1947, Sawyer (1947) recognized that
phosphorus concentration in the lake water was
the factor controlling eutrophication. Although
Vollenweider's L vs. Z plot relates loading rather
than concentration to trophic state, his improved
second relationship, L vs.2/ r., relates phosphorus
concentration, and not loading, to trophic state.
This may seem at first to be a contradiction but
the rationale is as follows: the lines separatins
Iakes into distinct trophic tvpes have the dimen--
s i o n so f L Z r u . o r g m - ' 2 ' y r t / m y r I , i . e .
g m-3. Thus, the lines are independent of time
and have units of concentration (the lines of the
L vs. Z plot have units of g m-3 yr-1, i.e. of
volumetric rather than areal loading). Since the
parameters with which one subjectively evaluates
a lake (chlorophyll a, oxygen in the hypolimnion)
are also expressed in terms of concentration, the
above inteforetation is most reasonable. Conse-
quently, predictions -relating the impact of develop-
ment on the phosphorus concentration of a lake
and subsequently on parameters describing the
trophic state are central to a predictive manage-
ment scheme.
The overall approach employed here is shown
in Fig. 1. From consideration of the geology and
land use of a lake's drainage basin, it is possible
to estimate the total phosphorus exported or
washed out Der unit of watershed. which. com-
bined with th! drainage area, provides an estimate
of the total phosphorus supplied to the lake from
the land. Addition of the input of phosphorus in
precipitation directly falling on the lake allows
calculation of the natural phosphorus load to
the lake. Development existins on the lake is
then measured (a6rial suruey oifield counts) and
 J . F I S H . R E S . B O A R D C A N . , V O L . 3 2 ( 9 ) ,1 9 7 5

the lake's capacityis outlined for use by planners,

managers,etc. Although the researchwork that
the soeciflcmodels are basedon was largely car-
ried iut in southern Ontario and the plrticular
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SPRING values suggestedfor different parameters apply

PHOSPHORUS to southern Ontario lakes, the overall approach
ONCENTRATION
PRECIPITATION
in lake is general enough that with the appropriate data
ARTIFICIAL
available,a similar sequenceof predictionscould
POPU LATION+PHOSPHORUS
DENSITY LOAD be madefor lakesof otherregions.

LAKE Theory for Each Step of the Management

Scheme for Southern Ontario Lakes

r'B LaxE
?.','#:.'j:3,*'?.t3.ElJiSJ9N
Ftc. 1. Scheme of simple empirical models used to 7. Natural phosphorus load lrom land (L.) -
assesseffects of development on trophic status of The geological formations of southern Ontario
lakes. can, as a first approximation, be classified as Pre-
cambrian igneous rock of plutonic origin (Cana-
dian Shield) or sedimentary rock. The former is
the phosphorus loading from artificial sources
typically composed of granites, gneisses, Pegma-
calculated. The total loading, natural plus arti-
tites, syemite, migmatites, diorite, gabbro, horn-
ficiaI, may then be combined with the lake's
blendite, amphibolite, and pyroxenites, the latter
morphometry and water budget to predict a
of limestone, dolomite, shale and basal clastics.
For personal use only.

phosphorus concentration that is subsequently re-

A very large proportion of Ontario's recreational
lated to the average summer chlorophyll c con-
lakes are situated on the Shield, with only the
centration. From this latter calculation, one may
southern portion of the Kawartha-Trent system,
determine the mean Secchi disc visibility. This
Lake Simcoe, and a few smaller lakes on sedi-
method will most often be used in the opposite
mentary material. In addition, the watersheds of
sense. For example, maximum acceptable average
most of the Shield lakes are entirely or almost
summer chlorophvll d concentration (or minimum
entirelv forested. with the remainder being either
Secchi disc reaOing; will be established by the
marshiand or pastureland that, although u-sed for
appropriate governmental agency. From these
agricultural purposes, is not chemically fertilized.
limits, decision-making personnel will be able to
Even south of the Shield on the sedimentary bed-
calculate a maximum permissible phosphorus
rock, there are few areas where intensive agricul-
concentration, which can be interpreted in terms
ture (i.e. chemical fertilization) is practiced in
of a maximum "permissible" toial phosphorus
prime recreational land; some areas around Lake
load, because the lake morphometry and water
Simcoe are a notable exception. Dillon and
budget are essentially fixed. That is, although
Kirchner (1975) have developed a phosphorus
there are vear-to-vear variations in the lake's
export scheme (Table 1) that is based on classifi-
water budget, the long-term average budget is
cation of geology as either "igneous" or "sedi-
used. Finally, the maximum permissible artificial
mentary" and land use as "forest" or "forest plus
phosphorus loading can be estimated and expressed
as the maximum allowable development (i.e.
numbers of cottages, etc.). Although hypolimnetic Tlnrr 1. Ranges and mean values for export of total
phosphorus(E) from 43 watersheds.Resultsin mg m-2
oxygen deficits have been related to phosphorus
yr-1 (from Dillon and Kirchner 1975).
loading for the Great Lakes (Gilbertson et al.
1972), a direct relationship applicable to a wide
Geologicalclassification
variety of lakes and especially to lakes of the
Precambrian Shield is not yet available. Mortimer Land use Igneous Sedimentary
(194142) suggested a limit of 0.25 mg Os
cm-2 day- r for oligotrophic lakes, but the Forest
quantitative link to nutrients is, as yet, unformu- Range 0.7-8.8 6.7-18.3
lated. Mean 4.7 1 1. 7
In the following section the heory and method Forest + Pasture
Range 5 . 9 - 1 6. 0 1 1 . 1 - 3.70
for each step of-the scheme are described and
Mean lo.2 23.3
subsequently a stepwise procedure for calculating
 DILLON AND RIGLER: CAPACITY OF LAKES FOR DEVELOPMENT
pasture"; the latter categoryimplies that 157o or
more of the watershedis clearedbut unfertilized
land. This export schemeis applicableto almost
the entire recreational lake area of Ontario. The
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results of Dillon and Kirchner are based on a
combinationof a study of the phosphorusexport
(E) of 34 watershedsin southern Ontario and
all additional phosphorusexport studiesreported
in the literature where watershedsfall into the
above-mentionedcategories.It is important to
note that the natural export from sedimentarv
materials(off the Shield) is almost exactly double
that from igneousbedrock (on the Shield); there-
fore, lakes such as the Kawarthas have a hisher
natural loading than Precambrianlakes. In aaAl-
tion, a change from a land use of forest to forest
plus pasturedoublesthe export within a geological
classification;for example, from 4.7 to 10.2 mg
6-2 y1-1 for igneouswatersheds and from 11.7
and23.3mg m-2 yr-1in sedimentary watersheds.
If a development significantly alters the amount
of cleared land in a watershed,then an appropri-
ate changein the export value used must be made.
It is also important to note that a range of
values was measuredfor each catesorv.Forested
For personal use only.
igneous watersheds,for example,ixforteO 2.5-
7.7 mg P m-2 yr-l in the southern Ontario
region alone, while Schindleret al. (1974) mea-
sured a range of 3-7 mg rn-2 yp-r in the Ex-
perimental Lakes Area. The total nutrient budget
can obviously be significantly affected by the
choiceof export figure.
To calculate the natural phosphorus loading to
a lake from its drainagearea, one must know the
area of the watershed(Ad) of each tributary to
the lake, and be able to classify each as to
geologyand land use so that the export coemcient
can be determined.Should the location be one of
the few where intensive farming is undertaken,
the reader should consult Vollenweider (1968),
Dillon and Kirchner (1975), and Loehr (1974).
The total amount of phosphorussupplied to the
lake from the land is therefore calculatedas the
sum of the area of each drainagebasin times its
phosphorusexport coefficient:
J ' ( m g ) - > ( A d . E r )w i t h A a i n m ' g . ( 1 )
The areal loading of phosphorussupplied to the
lake from the land Lr, is equal to (I Aa E)/A"
and is equivalentto the supply divided by the lake
a r e a( A o ) .
A complicatingfactor arisesif any tributary or
watershed of the lake in question has an ad-
ditional lake or lakes in its course. These lakes
undoubtedly act as traps for phosphorus and
other nutrients, decreasingthe actual amount of
material transported from the drainage area to
1523
the lake in question.A means of accountingfor
this is describedin a later section.
2. Natural phosphorusload lrom precipitation
(Lpn) - Phosphorusinput via precipitation (wet
and dry fallout) has been virtually ignored until
recent years. Studies by Schindler and Nighs-
wander (1970), Armstrong and Schindler
(1971), Barica and Armstrong (1971), Dillon
and Rigler (1974a), Lee and Kluesener (1971)
and Shannonand Brezonik (1972) have demon-
strated that, for many lakes,precipitation can be
a maior source of nutrients. From consideration
of the above-mentionedstudies, which found
loadingsrangingfrom 24 to 102 mg m-2 of lake
surface Jr-l, a value of 75 mg -*z yr-1 is
recommendedas applicable to southern Ontario
lakes. Some of the lower values did not include
a measure of phosphorus in dry fallout, while
others were measuredin more remote areas far
from urban and agricultural centres (Schindler et
al. 1974) and the highest value, 102 mg m-2
vr-1. was measuredin an area where much of
ihe land was used for agriculture. Furthermore,
the figure of 77 mg r.l-2 yr-t found by Dillon
(1974a) was measuredin the Haliburton High-
lands area of the province, an area of prime con-
cern for recreational development, although one
close to much of the province's agricultural and
industrial development.Thus, a figure of 75 mg
m-2 vr-1. for southern Ontario. although rea-
sonabie may be in error by as much as"1.OOVo.
The year to year variation alone was greaterthan
100% in certain areas (Schindler et al. 1974).
Additional research on this topic is obviously
needed.A review has been provided by Chapin
a n d U t t o r m a r k( 1 9 7 3 ) .
With LpR - 75 mg 6-2 y1-r, the natural
phosphorussupplyto a lake (Jn) is given as:
Jn:Ju+Jpp
: I ( A o ' E ) + 7 5 .A , (mgyr-') (2)
or theloading(Lx)
(Ao'rJ
I -
,1x. -_.: - T * ,, (mgm-z yr-,)
3. Artificial phosphorusload (L) - The cal-
culation of the phosphorussuppliedto a lake by
the population in its drainage basin is a difficult
task and must necessarilybe based on a supply
per capita-year figure with several assumptions
inherent in this method. Although numerous
figures are quoted in the literature (e.g. see Vol-
lenweider1968or Vollenweiderand Dillon 1974,
summary) great care must be taken in selecting
 t524 J, FISH. RES. BOARD CAN.. VOL. 32(9'1,1975
the appropriate value. The following points must
be remembered:
a) Values measured in studies such as that of
Johnson and Owen 1n l97 l (1.5 kg capita-r
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y-1) are no longer applicable because of legisla-
tion to reduce the phosphorus content of laundry
detergents. High phosphorus content dishwashing
detergents, however, remain legal and a survey of
the Muskoka Lakes found that 30% of the
cottages in the area employed automatic dish-
washers.
b) The waste disposal technique for most de-
velopments that are already established or will
be established in the near future is the conven-
tional septic tank-tile field system. The efficiency
of this treatment as far as phosphorus removal is
concerned is dependent on the type and depth of
soil surroundins the tile bed and between the tile
bed and the laie. In Precambrian areas typically
having very shallow, coarse-textured sandy or
muck soils there is no satisfactory evidence which
indicates that phosphorus is retained in the soils.
Therefore, it must be assumed that all phosphorus
discharged to soils of a tile bed area eventually
gains access to the lake. In sedimentary areas,
For personal use only.
septic tank-tile field systems located in sand,
gravel, or muck areas are likely to be as ineffec-
iive as far as phosphorus retention is concerned
as those systems located on the Shield. Lakes
surrounded by silt, clay, or clay-loam soil, how-
ever, will be provided with some measure of pro-
tection. Representativevalues for the fraction of
phosphorus retained in some soils are given in
Table 2.
c) Additional sources of phosphorus, (e.9.
fertilizer applied to lawns in cottage areas), are
Tasrr 2. The retentioncoefficientsof total phosphorus
for septic tile filter beds of different characteristics.
Results are based on 4 yr of data (after Brandes er a/.
1974J.
Filter bed R"
l. 22 in. sand(Dro : 0.24mm) 0.76
8 in. mixture 4/o red mud,96ft sand
2. 30 in. sand(D1e : 0.30 mm) 0.34
3. 30 in. sand(Dts : 0.60 mm) o.22
4. 30 in. sand(D1e : 0.24 mm) 0 .4 8
5. 30 in. sand(D1e : 1.0mm) 0.01
6. 30 in. sand (D.o : 2.5 mm) 0.04
7. 15 in. sand(D1q : 0.24 mm) 0 .8 8
l5 in. mixture 7O/"redmud,90f sand
8. 15 in. sand(D1s : 0.24 mm) 0.73
15 in. mixture 50f limestone,50% sand
9. 30 in. silty sand 0 .6 3
10. 15 in. sand(D16 : 0.24 mm) o.74
l5 in. mixture 50f clay-silt, 50ft sand
impossible to evaluate quantitatively without a
lake bv lake survev.
With the above factors in mind, a reasonable
phosphorussupply per capita-yearcan be calcu-
lated. According to the data of Bucksteeg (de-
scribed in Vollenweider 1968), the annual per
capita amount of phosphorus uPtake in food in
Germany in 1960 was 0.55 kg.; a similar amount
is assumedto be excreted.In addition, the yearly
per capitaphosphorussuppliedas domesticsewage
?excrerneniplui householdwastes)in 13 studiesin
North Ameiica and Europe averaged 0'80 kg'
year-r, the studies having been carried out
before use of high phosphate detergents was
common. Thus, the two resultsare in basic agree-
ment, and with food consumption in North
America being greater than that in Europe and
there being continued usage of high phosphate
dishwashing detergentsin North America, the
figure of O.SOtg phosphoruscapita-l year-1 is
a good estimate of the phosphorus supplied from
domesticsources.
For all lakes situated on the Precambrian
Shield the artificial supply of phosphorus can
thereforebe calculatedas:
Ja : 0.8(kg capita-1yr-1) x N (cottages)
x T
(capita-yrcottage-l yr-l) (3)
where N : the number of cottages or dwelling unit
equivalents
T : the averagenumber of capita years spent at
each cottage in each year.
The same formula can be used for areas south of
the Shield that consist of sand, gravel, or muck
soils. In cases where holding tanks are employed
and all wastes are removed to a treatment plant
outside the lake's watershed, the supply of phos-
phorus from such dwellings should not be in-
cluded in the calculations. Recent investigations
(Brandes et al. 1974) involving the use of a red
mud (aluminum mill waste) fill around the tile
bed have demonstrated a potential phosphorus
removal mechanism. In cases where such modi-
fications to conventional tile bed systems are used
and known to be in working order, a per capita
phosphorus supply figure of 0.10 kg is suggested
(88% removal). A factor (1 - R*), can be in-
cluded in the model to allow for alterations in the
phosphorus removal capacity of the system, with
some values for R, given in Table 2. The number
of cottages or permanent dwellings (N) situated
on a lake can be obtained by aerial photography
or by field count. The time (in capita-years)
spent at each dwelling per year (T) can be calcu-
lated using data gathered by the Department of
Tourism and Information (1971). For cottages
 DILLON AND RIGLER: CAPACITY OF LAKES FOR DEVELOPMENT 1525

in southernOntario, 253 capitadays per year (or 'T-

0.69 capita-yearsper year) are spent at each f P l : z\o '
+ p)
.1
cottage (based on a survey figure of 4.3 people
per cottage). For permanent dwellings, the ap- _ (, _ 2(o *_p)[P],\ .-,o*0,, (6)
J. Fish. Res. Bd. Can. Downloaded from www.nrcresearchpress.com by PENNSYLVANIA STATE UNIVERSITY on 05/11/12

propriate value for T is 1570 capita-daysper year \' L l-

(or 4.3 capita-yearsper year). An actual field
survey for a particular lake would undoubtedly where [P]" is the initial concentration.The steady-
improve these flgures; nevertheless, they will statesolutionto this differentialequationis:
serve as adequate,generallyapplicableestimates.
The total supply to the lake can be calculated pri : --!- (7)
as the sum of the natural and artificial supplies; 7(o t p)
Jr: Jn * Ja
Equation (7) can be used to predict the phos-
: E(Aa.E,) + LpR.Ao + 0.8 N.T (1 -R.) (4)
phorus concentrationin a lake if the parameters
where R" is the retention capacity of the disposal required for this, the loading (L), mean depth
svstem and will be eoual to 0 for most Shield (Z), sedimentationrate (o), and flushing rate
areas. The loading isi of course, equal to the (p), can be predictedor easilymeasured.While
supplyper unit of lake area, the loading is calculatedas describedin the pre-
ceding section and the mean depth is measured
Lr: lr/4". or obtainedfrom previous lake surveys,the flush-
ing rate (p) is approximatedfrom the total out-
flow volume per year (Q) and the lake volume
Prediction of the PhosphorusConcentrationin (V). Sincethe method outlinedhere is intended
a Lake for use without field work if possible,a method
of calculation of Q is required. The long-term
For personal use only.

A review of the theory and derivation of nu- averageannual areal runoff (r, in m yr-l) has
trient budgetmodelspublishedto the end of 1972 been mapped for
Ontario south of the Shield by
has been preparedby Dillon (1974b). Additional Coulson (1967) and for the entire
Great Lakes
work by R. A. Vollenweider (unpublisheddata), Basin by Pentland (1968).
Thus, virtually the
Imboden (1973, 1974), Lerman (1974), Fleming entire area of recreational lakes in
Ontario is
(1974), W. J. Snodgrass and C. R. O'Melia (un- mapped in terms of r, where r representsthe
publisheddata), and Dillon (1975) is also highly differencesbetweenprecipitation and evapotrans-
pertinent to the topic. The model used here de- piration. The total long-term averageinflow via
rived from Vollenweider (1969) is employed surface runoff to a lake can therefore be calcu-
becausea) it is simply derived and can be used lated as Ar'r. The total water balancecan now
with a minimum of field measurementsand b) it be describedby the equation:
alone has been tested for lakes in southern On-
tario and has been found to have good predictive Q = Ao.r* A, (Pr- Ev) (8)
capabilities.
Vollenweider assumedthat the change in the where Ar.Pr representsthe direct -A,.Ev input of water
concentrationof phosphorusin a lake with time to the lake by precipitation and repre-
is equal to the supply added per unit volume sents loss from the lakes bv evaooration. The
minus the loss through sedimentationand the loss average value for Pr for a given area can be
by outflow: obtained from isohyet maps (e.g. Canada Land
Inventory 1966) of Canada or for particular
d[P] J areas,e.g. Ontario (Brown et al. 1968). Maps of
- p[P] (s) Ev (evaporation from lake surface) are given
v- "[P]
for Canada by Bruce and Weisman (1966). If
where [P] representsthe phosphorusconcentra- Au is large comparedto Ao, then Q can be simply
tion, J is the amount of phosphorussuppliedper approximatedas A,,'r. In the few studiesunder-
annum, V is the lake volume, o is the sedimenta- taken in Ontario (e.g. Schultz 1950), ground-
tion rate (yr-t), and p is the flushingrate (yr-l) water has made a negligible contribution to the
which is equal to the total volume of water out- overall budget. The flushing rate, p, is then cal-
flowing per year (Q) divided by the lake volume c u l a t e da s Q ( m 3y r - 1 ) / V ( m a ) .
(v). It must be cautioned that water budgets pre-
The solution to this differential equation de- dicted in this manner may be very inaccurate,
scribing the concentration at a time t is: especially for small lakes and small drainage
 1526 J. FISH. RES. BOARD CAN.. VOL. 32(9), 1975
areas.Thus, Schindler et al. (1974) found mea-
sured water budgets for Rawson Lake to provide
3-5 times more water to the lake than predicted
from precipitation, evaporation, and evapotrans-
J. Fish. Res. Bd. Can. Downloaded from www.nrcresearchpress.com by PENNSYLVANIA STATE UNIVERSITY on 05/11/12
piration figures.Dillon (1974a), however,found
that measuredvalues were generally wirhin 25%
of those predicted using long-term runoff maps,
althoush these watershedswere much larger and
might'be expectedto integrate out localiied hy-
drologic variations. Measurement of the water
budget rather than estimationwould undoubtedly
provide much more accurateresults, and should
be undertakenwherepossible.
Measurementof o, the sedimentationrate co-
efficient, for phosphorus in a lake is, at best,
extremely difficult. Fortunately, an alternate pa-
rameter, the retention coefficient, R, can be
shown to have a functional relationship to o. R
is much more easily measuredthan o; (R is egual
to the fraction of the loading that is not lost via
the outflow) as well. the retention coefficienthas
been shown to be predictable (Kirchner and
-1975).
Dillon Equation (7) used to predict the
steady-statephosphorus concentration was re-
written by Dillon and Rigler (1974a) as:
For personal use only.
pnl : L(1= R)
zP
Kirchner and Dillon (1975) determinedby mul-
tiple regresion analysis that R was highly cor-
relatedwith Q/A, the areal water loading,usually
written as g". Their equation for prediction of R
ls:
R : 0.426exp (-0.271 q,) * 0.574 exp
x ( - 0 . 0 0 9 4 9q " )
This basic formulation is almost identical to that
predicted on theoretical grounds (Snodgrass
r974).
In some cases,the lake in question will have
one or more lakes upstream that are sufficiently
large to retain a significant amount of the total
phosphorusexportedfrom their respectiveportion
of the watershed.This is taken into account by
calculating the supply to the upstream lake, the
lake's retention coefficient and multiplying the
supply by ( t - Ro) to give the fraction transfer
to Lake B.
The Concept of ResponseTime of a Lake
As pointed out earlier, change in the phos-
phorus loading to a lake resultsin a changein the
phosphorusconcentrationand a subsequent change
in water quality.The changeis, of course,not in-
stantaneousbut rather is described in eouation
(6) and shown in Fig. 2. The time required for
E
E 87 5%
I (vr)
Frc. 2. Phosphorus concentration as a function of
tirne for a lake with an increase in loading. The initial
concentration was 8 mg m-t, the final steady-state
concentrationis 18 mg m-", the half-life is 2.1 yr.
a lake having an initial loading of L1 to respond
to a change in loading to a new level L2, (L2 can
be greater or less than Lr) can be described by
the half-life of the change in concentration, /4
(i.e. by the time required for the lake's phos-
phorus concentration to move half-way from the
(e) original steady-state concentration to the final
steady-state concentration. Because equation (6)
is exponential, two halflives (2t+) are required
for the lake to reach 757o of its final concentra-
tion, 3t2 for 871/27o,etc. The halfJife, t2, de-
pends only on the rate coefficients c and p, reP-
resentinq the losses of phosphorus by sedimenta-
tion and outflow; it is independent of the loading
(10) level or the initial phosphorus concentration in
the lake. It can be eaiily sfiown that
t+=ln2/(aI p)
and using the approximation that o = l0/z
(R. A. Vollenweider unpublished data), the half-
life can be estimated as:
t + = 0 . 6 9 / ( p+ l 0 / z ) . (11)
Therefore, one must consider the response time
of a lake when predictins the effects of an in-
creased loading (6.9. resuliing from development)
or a decreased loading (e.9. from improved
sewage treatment facilities). It is suggested that
3-5 times the r+ (i.e. 87.5-96.9% of the way to
the final steady-state concentration) can be used
as an indicator of the lake's response time. Char-
acteristically, lakes with a rapid flushing rate will
have short half-lives and therefore response times,
while lakes that are very slowly flushed may take
a lonq time to respond to a change in loading. For
example, a lake with flushing rate of p : 1.0 yr-1
 DILLON AND RIGLER: CAPACITY OF LAKES FOR DEVELOPMENT r527

(i.e. the lake's volume is replacedonce Per year

by flushing) and a mean depth of 10 m has a f4
of 0.69/(l + 10/ 10) - 0.35 yr. Therefore,be-
tween 1.1 and 1.8 yr are required for the lake to
J. Fish. Res. Bd. Can. Downloaded from www.nrcresearchpress.com by PENNSYLVANIA STATE UNIVERSITY on 05/11/12

approach a new steady-statefollowing a change

in loading.

Relationship of the Spring Phosphorus Concen-

tration to the Summer Chlorophyll a Concen-
tration and Water Transparency (Secchi Disc) G
;6
Basedon the work of Sakamoto(1966), Dillon 0)
and Rigler (1974b) developeda predictive rela- E
tionship suitablefor estimatingthe averagesum-
mer chlorophyll a concentration in lakes with a
o5
springN:P ratios )12, a conservative estimateof
the lower limit for proportionate uptake. The
"+
data comprising this relationship were based on =
a combination of information presented by I J J
Sakamoto for 30 Japaneselakes, results for a uJ
wide variety of North American and European
lakes reported by various authors, and informa- 2
tion for 19 lakes in southernOntario studied by
Dillon and Rigler. The equationfor the prediction
of the summer average chlorophyll d concentra-
For personal use only.

tion from the spring phosphorusconcentrationis:

Iog.[chla] = 1.45log.[Pl - 1.14 152025
o510
with [chl a] and [P] in mg m-3. a q)s/l)
CHLOROPHYLL

1000 Frc. 4. The relationshipbetweenSecchidisc depth

and chlorophyll c concentrationfor a number of
lakesin southernOntario.Valuesfor eachlake are
basedon meansof valuescollectedover the period
(June-September).
of stratification

Additional phosphorus-chlorophyll a data

ol have been supplied by Scheiderand Rigler (un-
J
published studies) for six lakes in Algonquin
t Park, by Oglesby and Shaffner for some of the
I
(L1
o Finser Lakes in New York State. and by Bach-
mari and Jones ( 1974) for some Iowi lakes.
J
I These data are also shown in Fig. 3 and it is
O apparent that in all casesthe fit to the model is
tr very good.
L I T T L EO I T E R L A K E
>'t.o . ALGONQUINPARK L It is expectedthat chlorophyll a concentration
. IOWA LAKES will be inverselyrelated to ih6 Secchi disc read-
l . FINGERLAKES
U)
"
RILEY LAKE ing, a measureof the water's transparency.Col-
. GRAVENHURSTBAY
lection of data for a large number of lakes is
shown in Fig. 4. From the predicted chlorophyll
0.1 a concentration,a prediction of the likely average
1000
TOTAL PHOSPHORUSmg m-3 Secchi disc reading is possible. Care must be
FIc. 3. Total phosphorus concentration at spring
taken in interpreting this relationshipin the case
overturn vs. summer averagechlorophyll d concentra- of dystrophic (brown water) lakes, whose Secchi
tion for lakes not included by Dillon and Rigler disc readingsare lower than would be expected
( 1974b). on the basis of chlorophyll a concentration alone.
 1528 J , F I S H . R E S . B O A R D C A N . . V O L . 3 2 ( 9 ) t975

StepwiseProcedurefor Calculatingthe Step 3: Determine the lake surface area (Ao

DevelopmentCapacityof a SouthernOntario in m2), mean depth (z in m), and volume (V in
Lake mB) from available information if possible. If
such data are not available, the lake must be
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Step1. Basedon long-range plansfor the lake, sounded and a contour maD drawn. The area
decide what the maximum permissibleaverage (A,) is obtained by planimitry from an aerial
summer chlorophyll a concentration will be: photograph of known scale.
Level l: 2 mg m-3' for lakes to be used Step 4: Outline the lake's drainage area on a
primarily for body contact water 1:50,000 scale topographic map or on aerial
recreation, and where it is desirable photographs and calculate the area (Aa in m2)
to maintain hvpolimnetic concentra- by planimetry.
tions of oxyg6n in excessof 5 mg Step 5: From Plate 13 in Pentland (1968)
liter-l to ^Dreservecold water fish- determine the total annual unit runoff (r) in cfs
eries. The lake will be extremely mi-2 and convert f6 6.r y1-1 m-2 or m yr-1 by
clear with a mean Secchi disc visi- multiplying by 0.345.
bility of 5 m and will be very un- Step 6: If Ad > 10 A, calculate Q, the total
productive. (Note - the Secchidisc outflow volume as:
visibility may be lower in brown
water [dystrophic]lakes). Aa'r(m"yr-')
Level 2: 5 mg m-3' for lakesto be used for
and calculate the flushing rate (p) as Q,/V or
water recreation but where the
preservation of cold water fisheries (Aa ' r)V (yr')
is not imperative. The lake will be
moderatefi productive and corre- If Ao is <10 Ao, determine the mean annual
spondingly less clear, with a mean precipitation (Pr) from figure 32 in Brown et al.
For personal use only.

Secchidisc visibility of 2-5 m.
Level 3: 10 mg m-3; for lakes where body-
contact recreationis of little impor-
tance, but emphasis is placed on
fisheries (bass, walleye, pickerel,
pike, maskinonge, bluegill, yellow
perch). Hypolimnetic oxygen deple-
tion will be common. Secchi disc
depths will be low (1-2 m), and
there is a danser of winterkill of fish
in shallowlakEs.
Level 4: 25 mg m-3; suitable only for
warmwater fisheries. Secchi disc
depth (1.5 m, hypolimnetic oxygen
depletion early in .sum-
.beginning
mer, considerabledanger of winter-
kill of fish exceptin deep lakes.The
planning agencymay pick any inter-
mediatelevel shouldit so desire.
Step 2: From the chosen summer average
chlorophyll a concentration, calculate the per-
missible spring phosphorus concentration, [P]
from:
Iogle [chl a] : 1.45logle tPl - Ll4
i.e. logto [P] :
loglo [chl 4] + 1.14
1.45
lchl al [Pl
2 mg m-3 9.9 mgm-3
5 r8.5
l0 29.9
25 56.3
(1968), and the mean annual lake evaporation
(Ev) from figure 9 in Bruce and Weisman
(1966), convert to m yr-1 by multiplying by
0.0254 and calculate Q:
(Pr-Ev)
Q:Aa'r*A
p : (A;r + A, (Pr-Ev))/V (vr-')
Step 7: Calculate the areal water load as (9,) as
Q/A. (m yr-t)
Step 8: Calculate retention coefficient (R) as
R = 0 . 4 2 6e x p ( - 0 . 2 7 1 q , ) - f 0 . 5 7 4e x p ( - 0 . 0 0 9 4 9g " )
Step 9: Calculate the response time of the lake
to a change in phosphorus loading:
Responsetime : (3 + 5) /1
: (3 + s) 0.69/(p + 10lz) (yr)
This will provide an indication of the time re-
quired foi a lake to "respond" to development
and will give an idea of when follow-up studies
(if any) should be carried out. Conversely, the
response time can assist in the interpretation of
present water quality. For example, a lake with
250 new cottages may appear to be in good con-
dition, but if one can calculate that it has a re-
sponse time of 6-lO yr, then caution is necessary
before additional develooment is allowed.
Step 10: Calculate the permissible phosphorus
load (Lo""-) to the lake:
 DILLON AND RIGLER: CAPACITY OF LAKES FOR DEVELOPMENT 1529

Lpe,m:f+i
and the permissiblesupply (Jp"..)
J. Fish. Res. Bd. Can. Downloaded from www.nrcresearchpress.com by PENNSYLVANIA STATE UNIVERSITY on 05/11/12
Ncr:0.69XNc+4.3XND
(mgm-2vr-l)
i.e. one permanentunit : 6.2 cottageunits.
Step 15: calculate the phosphorussupplied to
the lake from the cottageunits (artificial supply)

Jperm : (kgvr-')
Step 11: Divide the watershedof the lake (on
the topographic map) into subunits for all inflows
and determinethe area for each one (Aon). From
the most recent aerial photographs available de-
termine if )L5Vo of the area of each watershed
is either clearedland or marsh.Determinewhether
the watershed(not just the lake) is on Precam-
brian igneousrock or on sedimentaryrock, and
estimatethe export (E mg m-z yr-t) for each
subwatershedfrom Table 1. Calculate the total
supply of phosphorusfrom the land to the lake:
Jt - (2, Aar ' E)
and the load
lp = (2r Aar . E)/4"
1mgm-'yr-t)
Take into consideration an upstream lake by missiblenumberof cottaseunits:
reducing the supply to the downstreamlake from
For personal use only.
as:
Jr = 0.8 X Ncy (l - R") (kgyr-)
where R* = 0 for conventional septic tank-tile
field systemson the PrecambrianShield. If there
is firm evidencethat holding tanks are used for
all householdwastesand the systemsare pumped
and removed to a treatment plant outside of the
watershed,neglectsuch cottage(s) in the calcula-
tions. If the septic tile filter beds are situatedoff
of the Shield on soils that correspondto those of
-lable
2, use the appropriatevalies for R,.
Step 16: Calculate the present total of phos-
phorusto the lake:
Jr=Ix*Je (kg yr-)
If I\ > Jp,,"n,,
allow no further development.
Step 17: If Jr < Jp""-, calculatethe total per-

-JN
the watershed containing the upstream lake by Npc.^ :
multiplying by (t - R1) where R1 is the phos- 0.69 x 0.8 (1 - R")
phorus retention coefficientof the upstreamlake.
Step 18: The additional number of cottase
R1 is calculated as in Step 8, using q, for the
unitspermittedis:
upstreamlake. R1 is calculatedas in Step 8, using
q" for the upstreamlake. Naaa: Np"," - Ncr
Step 12: The phosphorusload from precipita-
tion, Lpn, is 75,mg-2 yr-1.
Calculate the supply from precipitation as: Discussion
The methodologyoutlinedin the previoussec-
, : 75'Ao (kgvr- t)
.rpn tion providesa simplemeansof calculatingthe
10e
capacity for development of southern Ontario
Step 13: The natural supply and natural load- lakes basedon lake trophic status.A number of
ing are: previouslytestedempirical modelsare linked in
a logical sequencerelating watershedcharacteris-
Jn:Jt+JPx tics and utilization to water quality. Although the
L.:Lt+Lt* specificfiguresused are applicableonly to south-
ern Ontario, the approach is general enough to
If J\ > Jn""n,,i.e. if the natural supply is greater be applied to any lake, provided suitabledata are
than the permissiblesupply, allow no (further) available for parameterssuch as the phosphorus
development. export coefficient,E, the loading from precipita-
Step 14: Determine the present number of tion, Lr,r,, the phosphorussoil retentioncoefficient,
cottages (N") and permanent dwellings (Nn) R,, etc. The advantagesof this method are that
within 300 m (1000 ft) of the lake or any of the quantitativepredictionsof the effectsof develop-
inflowing streams or rivers from recent aerial ment on important water quality parameters,
photographsor field surveys.For cottages,assume chlorophyll a concentrationand Secchidisc depth,
253 capita-days per year (0.69 capita-years yr-1). are possible,and that these predictions can be
Assume 4.3 people per dwelling, and calculate made with no field work, provided the lake's
N,.t, the number of capita-yearsyr-1 sPent at morphometry is available.The disadvantages are
the lake: that other important considerations,e.g. location
 l 530 J. FISH. RES. BOARD CAN.. VOL. 32(q. 1975
of fish spawning grounds and wildlife habitat,
microbiological contamination etc., are not taken
into account, and that, although
the individual
-been
models or relationships have tested and
J. Fish. Res. Bd. Can. Downloaded from www.nrcresearchpress.com by PENNSYLVANIA STATE UNIVERSITY on 05/11/12
found to make reasonably accurate ^oredictrons
predictions,
for lakes in southern Ontirio. overall
provided by the modeling sequence must be con-
siderably less accurate. The uncertainty in the
phosphorus export figures and in the loading from
precipitation alone could result in a 1007o error
in the calculation of the natural phosphorus
budget, while factors such as the soil retention
factor are still only approximations. Nevertheless,
these predictions should be valuable to those in-
volved in lake management because of both their
-quantitative
simplicity and their nature, provided
proper caution is taken in interpretation of the
results.
Acknowledgments
We thank all those involved in the various field
studies from which were drawn the individual com-
ponents of this study; W. Scheider, W. Coombs,
P. Cross, D. Andrews, C. Fehr, H. Banks, and
For personal use only.
J. McNight. We also thank Mr M. F. P. Michalski,
Mr M. Fielding, and Mr W. Scheider for helpful
comments on the manuscript.
Anr,rsrnoNc,F. A. J.. nNo D. W. ScurNor_en. 1971.Pre-
liminarychemicaichar:rcterization of watersin the
ExperimentalLakes Area, nofihwesternOntario.J.
F i s h .R e s .B o a r dC a n . 2 8 :l 7 l - 1 8 7 .
Becnr,reN,R. W., rNo J. R. JoNes. 1974.Phosphorus
inputsandalgalbloomsin lakes.IowaStateJ. Res.49:
155-t60.
B a n r c a ,J . , e N o F . A . J . A n v s r n o N c .1 9 7 1C. o n t r i b u t r o n
by snow to the nutrient budgetof some small nofth-
westOntariolakes.Limnol.Oceanogr.l6: 891-899.
B n e N o e sM , . , N . A . C n o w o n y , , c N o WW . . CsrNc.1974.
Experimentalstudy on removalof pollutantsfrom
domesticsewage by underdrained soilfilters.National LrnueN, A. 19'14.Eutrophication and water quality of
HomeSewageDisposalSymposium, Am. Soc.Agric.
Eng.,Chicago,lll., 1914.
B n o w N ,D . M . , G . A . M c K e v . e N o L . J . C H l p r r a e N
The climateof southernOntario.Can. Deo. Trans.
C l i m a t o lS. t u d .N o . 5 .
BnucE,J. P., eNo B. WrlsueN. 1966.Provisional
oration maps of Canada. Manuscript, D.O.T.,
Meterological Branch.
ClNnne LeNo INvp.NroRy. 1966.Theclimatesof Canada M r c n e l s x r , M . F . P . , M . G . J o u l l s o N , , q N o D . M . V e e . l .
for Agriculture,Report No. 3. The Queen'sPrinter,
Ottawa.24 p.
CHeptN,J. D., aNo P. D. Urronuenr. 1973.Atmospheric Monrtprrn, C. H. 194142. The exchange of dissolved
contributions of nitrogenandphosphorus. WaterRe-
sourcesCenter,Univ. Wisconsin,Madison,Wis.
Cour-soN,A. 1967.Estimatingrunoff in SouthernOntario.
Tech. Bull. No. 7. Inland Waters Branch, Dep.
Energy,MinesandResources. 19p.
Dpplnrvr,Nr op Tounrsu aNo INponuerroN. 1971.
Analysis of Ontario cottage survey. Travel Research
R e p o r l . N o . 5 5 : 1 1 4p .
Drrr-oN, P. J. 19'74a.The prediction of phosphorus and
chlorophyll concentrations in lakes. Ph.D. Thesis.
Univ. Toronto, Toronto, Ont. 330p.
1974b. A critical review of Vollenweider's nu-
trient budget model and other related models. Water
Res. Bull. l0:969-991.
1975. The phosphorus budget of Cameron Lake,
Ontario: the imporlance of flushing rate to the degree
of eutrophy of lakes. Limnol. Oceanogr. 20 : 28-39.
D r l r - o N . P . J . . l N o W . B . K I n c s N e n . 1 9 7 5 .T h e e f f e c t s o f
geology and land use on the exporl ofphosphorus from
w a t e r s h e d s .W a t e r R e s . 9 : 1 3 5 - 1 4 8 .
D u - r - o N ,P . J . , l N o F . H . R r c l E n . 1 9 7 4 a .A t e s t o f a s i m p l e
nutrient budget model predicting the phosphorus con-
centration in lake water. J. Fish. Res. Board Can. 31:
1771-1178.
1974b. The phosphorus-chlorophyll relationship
in lakes. Limnol. Oceanogr. 19:767-773.
F r - E u r N c , W . M . 1 9 7 4 .A m o d e l o f t h e p h o s p h o r u s c y c l e
and phytoplankton growth in Skaha Lake, British
Columbia. Ph.D. Thesis. Univ. British Columbia,
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