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    Ecological Modelling 114 (1999) 137173

    Spatial and temporal hydrodynamic and water quality modelinganalysis of a large reservoir on the South Carolina (USA)

    coastal plain

    Daniel L. Tufford *, Hank N. McKellar

    Uni6ersity of South Carolina, Department of En6ironmental Health Sciences, 311 Health Sciences Bldg., Columbia, SC 29208, USA

    Accepted 29 June 1998

    Abstract

    Two-dimensional, 31-segment, 61-channel hydrodynamic and water quality models of Lake Marion (surface are

    330.7 km2; volume 1548.3106 m3) were developed using the WASP5 modeling system. Field data from 1985 to 199

    were used to parameterize the models. Phytoplankton kinetic rates and constants were obtained from a related in sit

    study; others from modeling literature. The hydrodynamic model was calibrated to estimates of daily lake volume; th

    water quality model was calibrated for ammonia, nitrate, ortho-phosphate, dissolved oxygen, chlorophyll-a, biochem

    ical oxygen demand, organic nitrogen, and organic phosphorus. Water quality calibration suggested the modcharacterized phytoplankton and nutrient dynamics quite well. The model was validated (Kolmogorov Smirno

    two-sample goodness-of-fit test at PB0.05) by reparameterizing the nutrient loading functions using an independen

    set of field data. The models identified several factors that may contribute to the spatial variability previously reporte

    from other research in the reservoir, despite the superficial absence of complex structure. Sensitivity analysis of th

    phytoplankton kinetic rates suggest that study site-specific estimates were important for obtaining model fit to fie

    data. Sediment sources of ammonia (1060 mg m2 day1) and phosphate (16 mg m2 day1) were importan

    to achieve model calibration, especially during periods of high temperatures and low dissolved oxygen. This sedimen

    flux accounted for 78% (nitrogen) and 50% (phosphorus) of the annual load. Spatial and temporal variability in th

    lake, reflected in the calibrated and validated models, suggest that ecological factors that influence phytoplankto

    productivity and nutrient dynamics are different in various parts of the lake. The WASP5 model as implemented he

    does not fully accommodate the ecological variability in Lake Marion due to model constraints on the specificatio

    of rate constants. This level of spatial detail may not be appropriate for an operational reservoir model, but as research tool the models are both versatile and useful. 1999 Elsevier Science B.V. All rights reserved.

    Keywords: Reservoir; Hydrodynamic model; DYNHYD; Velocity gradients; Water quality; WASP; Eutrophicatio

    model; Phytoplankton productivity; Sensitivity analysis

    * Corresponding author. Tel.: +1-803-7774114; fax: +1-803-7773391; e-mail: [email protected]

    0304-3800/99/$ - see front matter 1999 Elsevier Science B.V. All rights reserved.

    PII: S0304-3800(98)00122-7

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    D.L. Tufford, H.N. McKellar /Ecological Modelling 114 (1999) 137173138

    1. Introduction

    Reservoirs are engineered structures, built to

    benefit human populations. Among these benefits

    are hydropower, recreation, sport and commercial

    fisheries, flood control, and water supply. Any-

    thing that impairs these uses has direct and no-ticeable impact. A common cause of impairment

    to reservoir use is eutrophication, or increase in

    lake productivity. Eutrophication is a natural pro-

    cess that is not inherently negative. It easily be-

    comes a problem in reservoirs because the main

    tributaries import large nutrient loads over time

    scales that are short relative to the ability of the

    aquatic ecosystem to adapt to the loading. The

    nutrient loads facilitate large populations of pri-

    mary producers. These can impact uses such as

    recreation and habitat maintenance or increase

    the cost of uses such as drinking water (Ryding

    and Rast, 1989).

    Study of the causes and consequences of eu-

    trophication and potential mitigating actions is

    complex for any lake or impoundment. These

    studies are often accomplished with the help of

    models of various types. Models are essential

    tools in studies of large reservoirs due to reservoir

    complexity in terms of morphometry, hydrology,

    ecology, and internal and external forcing

    functions.

    Impoundments created by damming major riv-ers may be more complex than natural lakes.

    While streamflow into natural lakes may be of

    limited quantity and impact, rivers import the

    majority of the water, particulates, and dissolved

    substances into most reservoirs. The river may

    define circulation patterns in the reservoir through

    its interaction with lake morphometry and by

    advective transport and density gradients (Ford,

    1990). Many of the external events influencing the

    river (land use practices and weather, for exam-

    ple) are seasonal in nature. This along with themorphometry of reservoirs allows establishment

    of many different functional habitats, in addition

    to the littoral, pelagic, and benthic zones shared

    with natural lakes.

    During the early stages of development of

    mechanistic hydrodynamic models, Orlob (1975)

    stated that circulation is an important determi-

    nant of ecosystem response. This belief was reite

    ated more recently (Falconer et al., 1991

    reflecting its then and current role in motivatin

    development and use of reservoir models wit

    greater hydrodynamic complexity. Ecological sim

    ulation models that include circulation processe

    seek to understand their influence on temperaturzonation and vertical stratification, in-lake an

    outflowing water quality, primary productivit

    turbidity along with the light environment, hab

    tat loss or creation, and hydraulic residence tim

    variation as a result of sedimentation (Kim et al

    1983; Martin, 1988; Riley and Stefan, 1988; Lun

    and Testerman, 1989; Falconer et al., 1991; Ok

    abe et al., 1993; Carrick et al., 1994; Leclerc et al

    1995; Shen et al., 1995; Ziegler and Nisbet, 1995

    Bailey and Hamilton, 1997; Hamilton an

    Schladow, 1997; Schladow and Hamilton, 199

    Soyupak et al., 1997).

    Jorgensen (1994) reviews the role of ecologic

    models in ecosystem understanding and environ

    mental management. He states the case for th

    necessity that models increase in complexity i

    order to understand the system under study an

    its stressors. This understanding is needed t

    provide input to management and political dec

    sions about actions to be taken as a response t

    observed or expected conditions. Straskrab

    (1994) discusses the management role many mod

    els can fulfill. The difference between the use of model for ecological understanding and its use a

    a management or technological tool is often sim

    ply the perspective of the researcher. Ecologic

    results may have management implications an

    vice versa. Thus, understanding the hydrolog

    ecology of a lake or reservoir is an importan

    aspect of research in lacustrine environments.

    Lake Marion, a large reservoir on the Sout

    Carolina Coastal Plain, has been the object o

    much study due to its geographic location an

    economic importance to the state. Though it appears to have little of the morphometric complex

    ity often associated with reservoirs, studies hav

    shown it exhibits spatial zonation (Inabinet, 197

    Pickett and Harvey, 1988). The causes of th

    variability can be only partially understood from

    those empirical studies. The objective of the re

    search reported here is to synthesize availab

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    D.L. Tufford, H.N. McKellar /Ecological Modelling 114 (1999) 137173 1

    morphological, hydrologic, meteorologic, and wa-

    ter quality data to simulate water transport and

    water quality processes in Lake Marion. The

    model was used to identify, characterize, and ex-

    plore in-lake mechanisms that influence lake pri-

    mary productivity. A large database of hydrologic,

    environmental, and water quality data exists forLake Marion. This, coupled with relevant litera-

    ture sources, facilitated simulation of the spatial

    and temporal dynamics of the lake.

    The model selected for this research was

    WASP5, the Water quality Analysis Simulation

    Program (Ambrose et al., 1993a,b,c). WASP5 is a

    group of mechanistic models capable of simulating

    water transport and fate and transport of water

    quality constituents and toxic organics. The mod-

    eling package is distributed and supported by the

    US Environmental Protection Agency (USEPA,

    1995). Various components of WASP5 have been

    used to study a variety of lake, reservoir, and

    estuarine issues including ecological characteriza-

    tion, the effects of anthropogenic activities, and

    the impact of mitigation measures (Ambrose,

    1987; Lung et al., 1993; Nikanorov et al., 1994;

    Bierman et al., 1994; Bierman and James, 1995;

    James and Bierman, 1995; Lung and Larson,

    1995). Reasons given for selecting WASP5 include

    its linkage of hydrodynamics to water quality and

    the broad range of water quality processes in-

    cluded in the modeling framework. The WASP5components used in this study are DYNHYD5 for

    simulation of reservoir hydrodynamics, and

    WASP/EUTRO for water quality simulation.

    2. Study area description

    2.1. Drainage basin hydrology and hydrography

    Lake Marion is in the lower portion of the

    Santee River basin (Fig. 1), one of the largestdrainages on the US east coast. The lake and its

    companion reservoir, Lake Moultrie, are collec-

    tively called the SanteeCooper lake system (Fig.

    2).

    Lake Marion has three principal subdrainages:

    1. the Saluda River, which originates in the

    South Carolina Blue Ridge province;

    2. the Broad River, originating in the North Ca

    olina mountains, and;

    3. the Catawba River, which also originates i

    the North Carolina Mountains.

    The rivers have been extensively dammed fo

    hydropower; at one time more hydropower wa

    generated here than on any other river system ithe world (Savage, 1968). The system now als

    assimilates the wastewater for large and growin

    urban areas, especially Charlotte, NC, th

    Greenville/Spartanburg region of SC, and Colum

    bia, SC.

    The Santee River begins as the confluence o

    the Congaree and Wateree Rivers approximatel

    17.6 km upriver from the lake (Fig. 3). The Con

    garee River begins at the confluence of the Broa

    and Saluda Rivers, 85.5 km upriver in Columbia

    The Wateree River originates 122 km upriver athe dam forming Lake Wateree (its principal trib

    utary is the Catawba River) (Fig. 1).

    Fig. 1. Santee River basin with study area indicated. T

    overall Lake Marion watershed is 38000 km2. The study ar

    (4860 km2) consists of the lower subbasins of the Congar

    and Wateree Rivers and the immediate drainage to La

    Marion.

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    D.L. Tufford, H.N. McKellar /Ecological Modelling 114 (1999) 137173140

    Fig. 2. The SanteeCooper lake system. Water enters the system from the Santee River (at the northwest). Most water leaves th

    system via the Tailrace Canal; other outflows are the Santee River at Wilson Dam and the Rediversion Canal.

    The Congaree and Wateree Rivers are large

    alluvial systems while their tributaries within the

    study area all originate in the upper and middle

    coastal plain. Tributary basins exhibit a dendriticdrainage structure. Wharton et al. (1982) describe

    three types of streams originating on the south-

    east coastal plain; blackwater, spring-fed, and

    bog-fed. All three are manifest in the study area

    though blackwater is the more common form.

    Organic matter (humics) dissolves during slow

    flowing floodplain residence and groundwater

    flow through rich floodplain deposits, giving the

    water its characteristic tea color (Wharton et al.,

    1982; Smock and Gilinsky, 1992). Overland flow

    of rainfall runoff produces characteristic irregulardischarge peaks throughout the year. Baseflow is

    maintained by often significant groundwater seep-

    age. These streams typically have floodplains

    though they are not as well developed as those of

    higher order coastal plain rivers.

    Mean annual rainfall in the study area is ap-

    proximately 1200 mm. Although 40% of the an-

    nual precipitation typically occurs in June, Jul

    and August, high evapotranspiration during thos

    months results in annual minima for discharge i

    the streams and rivers of the area (Smock anGilinsky, 1992).

    2.2. Lake Marion

    The SanteeCooper lake system was formed i

    1941 by damming the Santee River (for Lak

    Marion) and with a combination of dikes an

    dams impounding a large floodplain forest and a

    existing canal system (for Lake Moultrie). Th

    principal discharge from Lake Marion is the D

    version Canal to Lake Moultrie. A regulathough relatively small, outflow from Lake Ma

    ion at the Wilson Dam spillway reenters the San

    tee River. Excess flood-flow also discharges from

    the spillway into the Santee River. The princip

    discharge from Lake Moultrie is the Tailrac

    Canal which eventually drains into the Coope

    River. In 1985 the Rediversion Canal was com

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    D.L. Tufford, H.N. McKellar /Ecological Modelling 114 (1999) 137173 1

    Fig. 3. Water quality and discharge sampling station locations in Lake Marion and its immediate drainage.

    pleted which diverts some of Lake Moultrie

    outflow back to the Santee River. Downstream

    control of lake hydrology (lake stage and resi-

    dence time) occurs largely due to power genera-

    tion and flood control requirements at Pinopolis

    Dam. Other power generating stations are at

    Wilson Dam and the Rediversion Canal.The Santee Swamp, at the upriver extreme of

    Lake Marion (Fig. 3), is a remnant of an exten-

    sive floodplain that existed prior to damming the

    river. During extended periods of high flow the

    current swamp is inundated upstream to the Wa-

    teree River. Much of the year, however, only the

    lowest elevations are underwater along with pre-

    existing stream channels (Bates et al., 1992). An

    area just upstream from the Rimini trestle is the

    approximate extent of consistent free flowing

    open lake. For purposes of this study the trestlewas used as the swamp/lake demarcation.

    The Santee Swamp has a complex hydrology

    dependent upon discharge characteristics of the

    Wateree River and lake stage (Fig. 4). At high

    flows some overbank floodflow from the Wateree

    River enters the swamp. Cuts in the levee along

    the Santee River allow additional discharge into

    the swamp; this occurs essentially year round. Th

    effect is that as the swamp and lake converg

    water is flowing both in the main river channel a

    well as along either side of it. The amount o

    water entering the lake from the swamp is highl

    dependent on time of year and antecedent cond

    tions in the swamp. Backwater effects from thlake can inhibit downstream flow from the swam

    (Bates et al., 1992).

    Lake Marion is similar to many reservoi

    formed by major rivers (Ford, 1990). Its upstream

    end is shallow compared to the dam end (mea

    depth 2.2 vs. 4.9 m). Water velocity slows rapid

    as the Santee River enters the lake (Patterson an

    Harvey, 1995). The river is constrained by natura

    levees along its channel until 9 km downstream

    from the trestle. Several cuts in the levees allo

    water to leave the channel prior to their completsubmergence.

    The upper portion of the lake is also the na

    rowest and has several smaller tributary embay

    ments. These factors result in a range of aquat

    environments with varying water velocity, bottom

    morphology, tributary origin, chemistry, and cla

    ity (Inabinet, 1985; Pickett and Harvey, 1988

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    D.L. Tufford, H.N. McKellar /Ecological Modelling 114 (1999) 137173142

    Fig. 4. Hydrologic details of the Santee Swamp and upper Lake Marion. Arrows indicate direction of water flow.

    This portion of the lake is the principal nursery

    for the stripped bass, a major recreational fish in

    the SanteeCooper lake system (Bulak et al.,

    1995).

    Below I-95 the lake widens and deepens, taking

    on more lacustrine features. However, since it isstill relatively shallow and the water is always

    moving, thermal stratification is transient and

    does not occur at all in some years (Inabinet,

    1985). There are several additional smaller tribu-

    tary embayments at this end which are shallower

    than the open lake and, due to the tributary

    inflow, little hydrodynamic mixing occurs with the

    open lake (Patterson and Harvey, 1995).

    A prominent biotic feature in most of the shal-

    low portions of the lake is dense and extensive

    growths of aquatic macrophytes (Inabinet, 1985;Welch et al., 1986; Harvey et al., 1987). The

    dominant macrophyte species have changed over

    time (Inabinet, 1985; Welch et al., 1986), as has

    the distribution (Welch et al., 1986; Harvey et al.,

    1987). The upper portion of the lake receives the

    majority of the river-borne sediment burden

    (Cooney, 1988). As the sediment accumulates and

    reduces water depth, additional macrophyte hab

    tat is created (Harvey et al., 1987). Th

    macrophytes have been the object of intensiv

    biological and chemical control measures.

    Reservoirs typically exhibit marked longitud

    nal gradients in velocity and water quali(Kennedy and Walker, 1990; Thornton, 1990

    Inabinet (1985) and Pickett and Harvey (198

    observed this for Lake Marion in analyses of fu

    annual cycle data from 1984 and 1985/86, respe

    tively. Nutrient concentrations and trophic sta

    indices tended to be higher and primary produc

    tivity lower upstream than downstream. Wate

    clarity as measured by Secchi depth tended t

    increase along the same gradient. These finding

    support the conclusion that like most reservoir

    water clarity in Lake Marion is primarily a function of non-algal turbidity. Trophic state analys

    indicates the lake is eutrophic throughout, thoug

    there is a distinct decrease from the upper- t

    lower-lake (Inabinet, 1985).

    Pickett and Harvey (1988) also found clea

    evidence of latitudinal gradients. At the upper en

    of the lake these were related to complex bas

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    D.L. Tufford, H.N. McKellar /Ecological Modelling 114 (1999) 137173 1

    morphometry associated with the convergence of

    the Santee Swamp, the Santee River, and the

    submerged floodplain. Lateral gradients were also

    associated with tributary embayments at both the

    upper and lower ends of the lake.

    Episodes of large fish kills have been docu-

    mented in the extreme upper lake. Bates andMarcus (1990) believe these were the result of a

    complex series of hydrobiochemical processes that

    ultimately produced oxygen starvation. The kills

    occurred following sudden increases in hydraulic

    flow through the Santee Swamp into the upper

    lake after extended low flow periods. This caused

    a large pulse of anoxic sediment and oxygen

    depleted water into the lake. The resulting pockets

    of water with little oxygen were too large and

    occurred too suddenly for fish to migrate away

    from the area.

    Water quality sampling by the South Carolina

    Public Service Authority (the state owned entity

    that owns and runs the Santee Cooper system)

    generally occurs monthly. Data collected from 29

    stations were used in the research described here

    (Fig. 3). Sixteen stations are from the lake and

    embayments, twelve from tributaries, and one

    from the Santee Swamp.

    3. Model parameterization

    Model parameterization had as an objective

    that the final result would be suitable for studying

    a broad variety of issues. Whenever possible, data

    from a long term interval were used so the model

    would represent an average profile of the lake

    rather than short term conditions. Under these

    criteria the model calibration interval selected was

    July, 1985 through June, 1990. This range was

    chosen because it provided the longest, continu-

    ous record of chlorophyll-a (Chl-a) observations

    for the full complement of lake sample stationsused in the model. July, 1993 through June, 1994

    was used as the water quality model validation

    interval.

    Internally, the hydrodynamic model derives

    daily estimates of variable boundary flows spe-

    cified by the model user. These values are used at

    each time step during the model day. The hydro-

    dynamic equations are solved using a modifie

    Runge-Kutta procedure (Ambrose et al., 1993c

    DYNHYD5 averages results (flows and volume

    to match the time step of the water quality mode

    There are 11 tributary inflows and two outflows i

    the Lake Marion implementation. A code modifi

    cation was required to increase the number ovariable flows from five in the model as it

    distributed by USEPA.

    The time step chosen for the hydrodynam

    model was 120 s; for the water quality model

    was 2 h. The models simulated a 365-day interv

    beginning on January 1. Water quality and stream

    discharge data were from the USEPA STORE

    database (see http://www.epa.gov/owowwtr

    STORET/zip/sthp.html for a description

    Tufford (1996) has additional discussion of mod

    parameterization.

    3.1. Model geometry

    Lake bathymetry is needed in multi-segmen

    models to help establish segment geometry. Eac

    segment must have a surface area and dept

    which approximate the actual geometry of th

    corresponding location in the lake. The number o

    segments a model can have is conceptually con

    strained, in part, by the scale at which reasonab

    bathymetric estimates can be made.

    Bathymetry for this model was derived bplanimetry of a bathymetric map of the lak

    (Patterson and Logan, 1988). A principal obje

    tive for this model was that it facilitate evaluatio

    of spatial variability in the lake. Both lateral an

    longitudinal segmentation was developed; se

    ments are rectangular prisms. Segments were als

    designed so that no more than one of the 1

    monthly lake water quality sampling stations oc

    cur in a segment. Other segment design criter

    included that they should complement the natura

    morphology of the lake and they should be alarge as possible while still meeting other criteri

    This combination of constraints resulted in 3

    model segments, each 50106 m3 at maximum

    lake stage (Fig. 5).

    DYNHYD5 uses a channel/ junction geomet

    (Ambrose et al., 1993c). Channels move wate

    between junctions. Most of the hydrodynam

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    D.L. Tufford, H.N. McKellar /Ecological Modelling 114 (1999) 137173144

    Fig. 5. Configuration of water quality model segments with segment number (underlined) and mean depth at full pool. Shading

    included to help visualize depth gradients.

    junctions correspond to segments in the water

    quality model. The model requires additional

    junctions that serve as sources and sinks for

    boundary flows (Fig. 6). These extra junctions do

    not map to water quality model segments. A

    channel must be defined for each lake inflow and

    outflow and to represent flow within the lake.

    Internal flow paths were estimated based on a

    previous tracer study of the lake (Patterson and

    Harvey, 1995). The total number of channels is 61

    (Fig. 6). Defined channels only indicate potential

    flow paths. Actual advective flow is determined at

    each time step based on hydraulic head differen-

    tial between adjacent junctions.

    3.2. Eutrophication constituents and kinetic rates

    and constants

    The water quality model can simulate kinetic

    transport and transformation for up to eight eu-

    trophication constituents (Fig. 7). Constituent

    concentrations from sixteen lake stations wereused to set initial concentrations (5 year mean for

    January) and to produce calibration charts (5 year

    monthly means91 S.E.). Tributary and swamp

    stations were used to obtain constituent inflow

    concentrations during the model execution. Flow

    weighted mean concentrations were derived for

    segments with more than one tributary inflow.

    Water quality parameters simulated were:

    1. chlorophyll-a (Chl-a);

    2. ammonia/um-N (NH3/4);

    3. nitrite/nitrate-N (NO2/3);

    4. ortho-phosphate-P (OPO4);

    5. 5-day biochemical oxygen demand (BOD5);

    6. dissolved oxygen (DO);

    7. organic nitrogen-N (ON), estimated as tot

    Kjeldahl nitrogen (TKN) minus NH3/4;

    8. organic phosphorus-P (OP), estimated as totphosphorus (TOTP) minus OPO4.

    WASP5 models phytoplankton production an

    nutrient kinetics (Fig. 7) as temperature modifie

    functions of rates entered by the model user (Am

    brose et al., 1993a). Phytoplankton production

    also modified by light and nutrient limitatio

    Nutrient limitation is modeled using Michaelis

    Menton kinetics. Nutrient species are taken u

    during phytoplankton growth and released durin

    respiration or at death according to atomic ratio

    entered by the model user. Water quality constituent transport occurs principally by advection

    Dissolved fractions are also subject to diffusio

    and particulate fractions can settle and resuspend

    Constituent loads enter the system in tributar

    inflows, as point sources, or as nonpoint source

    Inorganic nutrients can also enter by benth

    release.

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    D.L. Tufford, H.N. McKellar /Ecological Modelling 114 (1999) 137173 1

    Fig. 6. Channel/junction geometry of the hydrodynamic lake model. (a) Junctions with dashed outline do not appear in the wat

    quality model; they exist as sources and sinks of water. (b) Channels are potential flow paths between junctions.

    The initial values for most of the reaction rates

    and constants were taken from water quality

    modeling literature (Ambrose et al., 1993a; Bowie

    et al., 1985), and when changed during calibra-

    tion, the literature ranges were used as guides

    (Table 1). Exceptions were the phytoplankton

    growth rate, the temperature correction coefficient

    to phytoplankton growth, and the half-saturatio

    constants for nitrogen and phosphorus limitatio

    to phytoplankton growth. Initial values for thes

    were taken from work in Lake Marion by Suda

    shan (1995) (Table 2).

    The ammonia and phosphorus benthic flu

    functions were determined by calibration. The

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    D.L. Tufford, H.N. McKellar /Ecological Modelling 114 (1999) 137173146

    Fig. 7. Energy diagram of the processes in the eutrophication model of Lake Marion. Temperature (left out to simplify the diagram

    modifies all processes except settling, sediment nutrient release, reaeration, extinction, and loading /export. The WASP5 model

    capable of additional, and in some cases different, processes. Those indicated are implemented in the model described her

    Acronyms are defined in the text.

    were needed as additional sources of nutrientsinto the system. There was not a deterministic

    derivation of the values entered, but they gener-

    ally follow the rule that flux increases as tempera-

    ture increases and dissolved oxygen decreases.

    Their ranges are within those found in the

    literature.

    3.3. En6ironmental forcing functions

    Four water temperature specifications (to facili-

    tate spatial differentiation within the model) weredeveloped by cluster analysis (SAS/STAT PROC

    CLUSTER, AVERAGE method; SAS, 1988) us-

    ing the 5-year monthly mean temperatures at each

    sample station. Segments without a sample station

    were assigned a cluster based on proximity to a

    segment with a station. Downstream proximity

    was given the highest consideration.

    Light extinction coefficient specifications wealso derived by cluster analysis. The monthly valu

    was derived with Eq. (1) (Williams, 1980) from th

    field data. These were modified by subtracting a

    estimate of light extinction due to phytoplankto

    (see Eq. 5.9, Ambrose et al., 1993a; Kirk, 1994

    This was done because internally the water qualit

    model adds this estimate to the extinction coeffi

    cient entered by the model user.

    Ke=1.1Secchi depth0.73

    (Secchi depth in meters). (

    Daily total solar radiation (TSR in W m2) wa

    derived from Eq. (2), fitted from a long ter

    record along the South Carolina coast (Summe

    et al., 1980). The daylight fraction of the 24-

    interval was obtained by proportion using sunrise

    sunset for Charleston, SC (United States Nav

    Observatory, 1977).

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    D.L. Tufford, H.N. McKellar /Ecological Modelling 114 (1999) 137173 1

    Table 1

    Reaction rates and constants (and range of values tried during calibration) for the WASP eutrophication model

    RangeValue SourceFunctionName

    Water column dispersive transport Bowie et al. (1985)Dispersion 10 m2 s1

    Bowie et al. (1985)Solids flow specification0.00.50.00.4 m day1Settling

    Ambrose et al.1.03.0SOD1D 3.0 gO2 m2 Sediment oxygen demand

    (1993a)day1

    0.991.04 Temperature correction factor (TCF) for SOD1DSODTA Bowie et al. (1985)1.04

    0.050.15 Bowie et al. (1985)Nitrification rate0.075 day1K12C

    1.08 1.081.20 TCF for K12C Bowie et al. (1985)K12T

    Half-saturation constant for O2 limit. on nitrification Bowie et al. (1985)KNIT 2.0 mg O2 l1

    1.42.6 Maximum phytoplankton (PP) growth rateK1C 2.24 day1 Sudarshan (1995)

    0.981.072 TCF for K1C Sudarshan (1995)1.03K1T

    Ambrose et al.XKC 0.016 mg CHL-a Coefficient for extinction due to chlorophyll

    m3 (1993a)

    Sudarshan (1995)0.0150.0250.015 mg N l1 Half-sat. constant for N limitation to PP growthKMNG1

    0.00320.02 Half-sat. constant for P limitation to PP growthKMPG1 Sudarshan (1995)0.005 mg P l1

    0.050.35 PP respiration rateK1RC 0.2 day1 Bowie et al. (1985)

    Bowie et al. (1985)TCF for K1RC1.0451.11.05K1RT

    0.020.1 Non-predatory PP death rateK1D Bowie et al. (1985)0.04 day1

    P-to-C ratio in PP Bowie et al. (1985)PCRB 0.025 mg P

    mgC1

    N-to-C ratio in PP Bowie et al. (1985)0.25 mg N mgNCRB

    C1

    Half-sat. constant for PP effect on mineralization Ambrose et al.0.0 mg C l1KMPHYT

    (1993b)

    Bowie et al. (1985)BOD deoxygenation rate0.10.30.3 day1KDC

    Bowie et al. (1985)TCF for KDCKDT 1.04

    Ambrose et al.Half-sat. constant for O2 limit. to BOD decomp.KBOD 0.5 mg O2 l1

    (1993a)

    O-to-C ratio in PP Default2.667 mg O2 mgOCRB

    C1

    0.020.2 Mineralization rate of DONK71C 0.06 day1 Bowie et al. (1985)

    TCF for K71C Bowie et al. (1985)K71T 1.02 1.021.3

    0.25,0.5 Fraction of dead PP recycled to DONFON 0.5 Ambrose et al.

    (1993a)

    Mineralization rate of DOP Bowie et al. (1985)0.22 day1 0.10.4K83C

    TCF for K83C Bowie et al. (1985)K83T 1.08 1.081.2

    Fraction of dead PP recycled to DOPFOP Set equal to FON0.5

    TSR=((39451695cos((2pi/365)(day+11)))

    27.9). (2)

    3.4. Other parameterization issues

    Constituent mass transfer into and out of seg-

    ments in the water quality model occurs principally

    by advection as specified by the hydrodynamic

    model. Transfer also occurs as a result of dispersive

    exchange and by settling; both were determine

    during calibration of the hydrodynamic model.

    dispersion coefficient of 10 m2 s1 was specifiebetween segment pairs 1/2, 5/6, 22/26, 25/27, an

    27/29.

    Reaeration coefficients were derived for th

    model using estimates of wind speed (Banks, 1975

    Estimates of wind speed (at 10-day intervals) wer

    the 36-year mean of observations kept by NOAA

    for Columbia and Charleston (USEPA, n.d.).

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    D.L. Tufford, H.N. McKellar /Ecological Modelling 114 (1999) 137173148

    Two source code modifications were made to

    the water quality model. The first modification

    resulted from concern that in the model as dis-

    tributed by USEPA the daily derivation of the

    carbon-to-chlorophyll ratio underestimated the

    light in the water column. Conversations with

    the model developer resulted in adjustments tothe derivation to increase the light estimate.

    The second code modification implemented an

    estimate of saturating light intensity as a linear

    function of incident light developed by Sudar-

    shan (1995) during her work in Lake Marion.

    This replaced the derivation already in the water

    quality model; a function of water temperature

    and phytoplankton physiological state. Sudar-

    shan developed her equation using incident solar

    radiation as photosynthetically active radiation

    (PAR) in mol m2 day1 Eq. (3).

    Is=0.12+(0.47Id) r2=0.63 (3)

    where Is and Id are saturating and total daily

    PAR, respectively, in mol m2 day1.

    The model uses total solar radiation as Ly

    day1. Documentation from LI-COR (1979) in-

    dicated that 1 mol m2 day1 (400700 nm

    wavelength)=5.733 Ly day1 (photosynthetic

    radiant exposure). The proportion of total inci-

    dent radiation that is PAR is about 0.48 (Kirk,

    1994). With these conversion factors the equation

    developed by Sudarshan (1995) was modified for

    the model.

    4. Surface water balance model

    The Diversion Canal is the principal surfac

    outflow from Lake Marion. It is ungauged sinc

    its discharge is under direct control of Lak

    Moultrie. To estimate its discharge a surface wa

    ter budget model was developed. Flow measurments or estimates with daily frequency we

    entered into the model as 10-day arithmet

    means. This was because of a limitation in th

    DYNHYD5 model. All other flows had monthl

    measured or estimated values.

    4.1. Stream discharge sampling

    Discharge estimates for the Congaree (nea

    Columbia) and Wateree (near Camden) Riv

    were daily averages from US Geological Surve(USGS) continuous stage/discharge records. Th

    accounts for drainage from 87% of the overa

    basin area. The net effect of additional sma

    tributary inflow, loss to extensive floodplain

    and groundwater exchange downstream from

    these recorders was probably small; it was no

    included in these discharge estimates. Discharg

    estimates from USGS stations near the Congare

    and Wateree confluence and on the Santee Rive

    upstream from the lake have been discontinue

    due to uncertainty over backwater effects.Small tributary discharge estimates were ob

    tained from monthly sampling by the South Ca

    olina Department of Health and Environment

    Control (SCDHEC) and the South Carolin

    Public Service Authority (also known as Santee

    Cooper, the state owned entity that owns an

    operates the SanteeCooper lakes). Santee

    Cooper also estimates daily discharge from th

    spillway at Wilson Dam.

    4.2. Santee swamp discharge estimate

    It is estimated that Wateree River discharg

    above 283.2 m3 s1 spills over into the Sante

    Swamp (Pickett, 1992). It is also estimated th

    10% of the Santee River discharge flows into th

    Santee Swamp through cuts in the natural levee

    which form the main channel upriver from th

    Table 2

    Estimated phytoplankton growth parameters and 95% confi-

    dence intervals

    Estimate 95% CIParameter

    2.24 1.722.76Maximum growth rate

    1.031.071.05Temperature correction

    0.00430.0250.015Ks for Na

    0.0010.0075Ks for Pa 0.0032

    16.7623.8220.2Is (mol m2 day1)b

    See text for discussion. From Sudarshan (1995).a MichaelisMenton half-saturation constant.b Saturating light intensity.

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    D.L. Tufford, H.N. McKellar /Ecological Modelling 114 (1999) 137173 1

    lake (Patterson and Harvey, 1995) (Fig. 4). Pick-

    ett (1992) estimated that the annual average

    travel time from the USGS gauging stations at

    both Camden (Wateree River) and Columbia

    (Congaree River) to the confluence is about 1

    day. An additional lag of 1 day was introduced

    to account for travel from the confluence to thelake. The estimated discharge into the lake from

    the Santee River main channel is given by Eq.

    (4).

    Discharge=0.9(2 days prior to Congaree River

    at Colombia+2 days prior to Wateree River

    discharge at Camdenportion of Wateree

    discharge\283.2 m3 s1)). (4)

    Bates et al. (1992) estimated the annual mean

    hydraulic travel time through the Santee Swamp

    is 40 days. A travel time of 5 days for SanteeRiver levee cut water was assigned to introduce a

    lag. The estimated flow of water from the swamp

    into the lake is 10% of the 5 days prior Santee

    River discharge, as described above, plus the

    portion of the 40 days prior Wateree River dis-

    charge \283.2 m3 s1, if any. The Santee River

    main channel and the two Santee Swamp sources

    were entered into the model as three separate

    flows to facilitate modification to them to evalu-

    ate research or management questions.

    4.3. Small tributary discharge estimates

    The natural log transform of the monthly

    mean discharge (m3 s1) from each of the mea-

    sured small streams was regressed against the log

    transformed watershed area (m2) using SAS

    PROC REG (SAS, 1988). This provided a

    method to estimate the monthly discharge of two

    unmeasured creeks Eq. (5). Of the six small lake

    tributaries which were measured, the measure-

    ment was taken upstream from the lake. Thedischarge regression model provided a method to

    estimate their full basin discharge into the lake

    (Table 3).

    ln(q)=a+bln(A) where

    Q=flow(m3 s1) A=subbasin area (m2).

    (5)

    4.4. Precipitation and e6aporation

    Direct precipitation onto, and evaporatio

    from, the lake surface was estimated based o

    National Climate Data Center (NCDC) record

    (NCDC, 1985a,b, 1986a,b, 1987a,b, 1988a,b

    1989a,b, 1990a,b). Precipitation and evaporatioare the monthly mean of measurements fro

    stations near Lake Marion. Estimated lake evap

    oration was taken as 70% of the measured pa

    evaporation. This is a standard open-water co

    rection factor from pan measurements (Veih

    meyer, 1964). Data are from 1985 to 1990. Th

    net effect was evaporative loss every mont

    (Table 4).

    4.5. Lake 6olume estimates

    The bathymetry map of the lake used for seg

    ment geometry (Patterson and Logan, 1988) in

    cluded a lake stage-volume curve which was use

    to fit a 5th order polynomial to estimate volum

    from stage Eq. (6). Volume has units of acr

    feet; stage has units of feet. The stage interv

    that can be used to predict volume with th

    expression is 5077.5 feet (15.2423.62 m).

    Volume=(16500.629481525.16547stage

    +55.67088

    stage

    2

    1.01081

    stage

    3

    +0.00915stage4

    0.0003stage5)1000, r2=0.999.

    (6

    4.6. Di6ersion canal discharge estimates

    The estimate of daily diversion canal outflo

    was then the net difference of the daily volum

    change9all other measured or estimated flow

    Eq. (7).Discharge=(previous day lake volume

    current day lake volume)

    +Santee River main channel discharge

    +Santee River discharge through swamp

    +Wateree River discharge through swamp

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    D.L. Tufford, H.N. McKellar /Ecological Modelling 114 (1999) 137173150

    Table3

    Drainagebasinareas(km2)andpredictedmonthlymeandischarges(m3

    s1)forthe

    smalldirecttributariesofLakeMarion(upperpan

    el)andstatisticsforthemonthlyregressionmodels(allP-valuesB0.0

    001)usedto

    predictthedischarges(lowerpanel)

    Area

    Jan

    Feb

    Mar

    Ap

    r

    May

    Jun

    Jul

    Aug

    Sep

    Oct

    Nov

    Dec

    WybooSwampCreek

    45

    0.4

    9

    0.6

    7

    0.6

    1

    0.4

    4

    0.3

    0

    0.2

    4

    0.25

    0.2

    6

    0.4

    2

    0.3

    3

    0.3

    5

    0.5

    5

    103

    1.5

    1

    1.8

    2

    1.6

    3

    1.2

    9

    0.9

    7

    PotatoCreek

    0.9

    3

    0.85

    1.0

    0

    1.2

    3

    1.1

    4

    1.4

    6

    1.6

    6

    TawcawCreek

    102

    1.1

    6

    1.4

    4

    1.3

    0

    1.0

    1

    0.7

    4

    0.6

    8

    0.64

    0.7

    4

    0.9

    6

    0.8

    5

    1.0

    5

    1.2

    8

    97

    0.1

    2

    0.1

    9

    0.1

    8

    0.1

    2

    0.0

    7

    JacksCreek

    0.0

    5

    0.06

    0.0

    5

    0.1

    1

    0.0

    7

    0.0

    6

    0.1

    4

    61

    1.1

    1

    1.3

    8

    1.2

    4

    0.9

    6

    0.7

    0

    SpringGroveCreek

    0.6

    5

    0.61

    0.7

    0

    0.9

    2

    0.8

    1

    0.9

    9

    1.2

    3

    12

    0.4

    2

    0.5

    8

    0.5

    3

    0.3

    8

    0.2

    6

    ChapelBranchCreek

    0.2

    0

    0.22

    0.2

    2

    0.3

    6

    0.2

    8

    0.2

    9

    0.4

    8

    39

    0.6

    8

    BigPoplarCreek

    0.8

    9

    0.8

    1

    0.6

    0

    0.4

    2

    0.3

    6

    0.36

    0.3

    9

    0.5

    7

    0.4

    7

    0.5

    3

    0.7

    6

    225

    2.6

    8

    3.0

    3

    2.6

    8

    2.2

    3

    1.7

    6

    HalfwaySwampCreek

    1.8

    6

    1.58

    1.9

    9

    2.1

    2

    2.1

    3

    3.0

    3

    2.9

    0

    34

    0.3

    7

    0.5

    1

    0.4

    7

    0.3

    3

    0.2

    2

    0.1

    7

    H.S.unnamedtriba

    0.19

    0.1

    9

    0.3

    2

    0.2

    4

    0.2

    4

    0.4

    2

    8.5

    4

    10.5

    2

    9.4

    6

    7.3

    6

    5.4

    5

    5.1

    4

    Totaldischarge

    4.76

    5.5

    3

    7.0

    1

    6.3

    2

    8.0

    2

    9.4

    1

    0.7

    59

    0.5

    35

    0.6

    89

    0.6

    83

    0.6

    22

    r2

    0.6

    64

    0.507

    0.6

    42

    0.5

    55

    0.7

    43

    0.8

    46

    0.6

    67

    19.2

    3

    16.9

    2

    16.6

    1

    18.4

    4

    20.4

    9

    23.5

    8

    21.27

    Intercept

    23.3

    4

    18.4

    5

    21.3

    7

    24.4

    4

    18.6

    1

    1.0

    51

    0.9

    37

    0.9

    15

    1.0

    01

    1.0

    95

    1.2

    59

    1.130

    Coefficient

    1.2

    50

    0.9

    98

    1.1

    51

    1.3

    28

    1.0

    23

    a

    HalfwaySwampCreekunnamed

    tributary.

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    D.L. Tufford, H.N. McKellar /Ecological Modelling 114 (1999) 137173 1

    Table 4

    Precipitation (P), evaporation (E), and net (PE) used in the hydrodynamic model. Also shown are comparisons of evaporatio

    among the current model and three literature estimates

    Net (PE) E (current) EbEaP E Ea

    (mm year1)(mm year1)(mm day1)(mm day1) (mm month1)(mm day1) (mm month1)

    41Jan 342.2 2.8 0.2

    61Feb 2.3 2.9 0.2 3181Mar 2.5 4.6 0.7 36

    114Apr 361.8 6.5 1.5

    38 137May 2.3 7.1 1.6

    160Jun 3.2 7.5 1.4 37

    168Jul 392.7 7.8 1.7

    Aug 5.2 6.0 0.3 37 152

    140Sep 3.9 4.9 0.3 35

    112Oct 353.9 4.3 0.1

    33 76Nov 2.3 3.2 0.3

    Dec 2.0 2.2 0.1 33 48

    425 1290 1325Total 1092

    a

    Van der Leeden et al. (1990).b Morton (1983).

    +(precipitationevaporation)

    +small tributary discharges

    Wilson Dam spillway discharge. (7)

    5. Methods

    5.1. Calibration and6

    alidation

    Hydrodynamic model calibration was per-

    formed by adjusting the estimated, 5-year mean

    boundary flows. The calibration objective was

    that the daily modeled volume would be within

    5% of the estimated 5-year daily mean volume.

    Hydrologic estimates are discussed in a subse-

    quent section. The calibrated model specifically

    simulated hydrologic conditions during the cali-

    bration interval. Under this implementation the

    only meaningful validation would be of the datacollection and derivation methods; none was

    performed.

    The water quality model was calibrated by

    comparing measured monthly mean concentra-

    tions (91 S.E.) to the modeled monthly mean.

    This was done for each of the eight constituents

    for each model segment with a corresponding

    water quality sampling station. Calibration wa

    accomplished by adjusting rates and constan

    within the limits of literature values for biochem

    cal processes in the lake. Rates controlling phyto

    plankton kinetics were kept within ranges derive

    from field work in Lake Marion by Sudarsha

    (1995) (Table 2).

    For validation the calibrated model boundar

    loads were set to field values for January, 199through August, 1994. The validation interval wa

    July, 1993 to June, 1994. This corresponds to th

    field work interval from which Sudarshan (1995

    estimated the phytoplankton growth kinetics. Th

    model was parameterized to run from Januar

    1993 because the hydrodynamic flows and vo

    umes simulate a January 1 begin date.

    Each of the 16 segments with correspondin

    sample stations were individually assessed for va

    idation using the KolmogorovSmirnov two-sam

    ple goodness-of-fit test (Conover, 1980; Reckhoet al., 1990). Parameters evaluated were Chl-a an

    the inorganic nutrient species. The model pr

    dicted values corresponding to station samplin

    days were used as one sample. The station sam

    ples themselves were the second sample. Statio

    sampling did not occur every month at all station

    during the validation interval (range was 5 12

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    Table 5

    Phytoplankton production kinetic constants, their values in the calibrated model, and values in the sensitivity analysis

    Constant Calibrated Values in sensitivity analysis

    2.24Max growth 1.51.0 2.0 2.5

    0.080.010.015N half-sat 0.040.02

    0.08P half-sat 0.005 0.0005 0.001 0.01 0.04

    The KolmogorovSmirnov test is a nonparamet-

    ric test that converts the two samples to their

    empirical distribution function (EDF). The princi-

    ple is that if the two samples are from the same

    underlying population, their EDFs should be

    close to each other. The test statistic is derived as

    the maximum vertical distance (y-axis) between

    the two EDFs. A P-value is determined based on

    the sample sizes. The null hypothesis is that the

    two samples are taken from the same underlying

    distribution. For this research the hypothesis was

    rejected at P50.05. The test was performed using

    PROC NPAR1WAY (SAS, 1988) with the EDF

    option.

    5.2. Tracer simulations

    The water quality model can simulate a conser-

    vative chemical that is subject only to transport,

    boundary, and loading processes (Ambrose et al.,

    1993a). Two types of tracer simulations were exe-cuted during this study. In the first, a steady unit

    concentration of 1 mg l1 was included as a

    boundary concentration in all inflows. Significant

    deviations from that concentration anywhere in

    the reservoir would indicate a source or loss of

    chemical not associated with surface advective

    flows.

    The second type of tracer simulation was exe-

    cuted as a way of estimating hydraulic travel time

    through the reservoir during high and low flow

    scenarios. This was accomplished by introducinga pulse (simulated as a point source load) of 50 t

    of tracer into junction 12 (water quality model

    segment 1) on model day 62 (775 m3 s1) and 192

    (222 m3 s1). The day of the occurrence of the

    peak tracer concentration in junction 42 (water

    quality model segment 31) minus the entry day of

    the tracer was considered hydraulic travel time.

    The quantity of tracer in the pulse was est

    mated by trial-and-error. Significant dilution oc

    curs as the pulse move downstream. Determinin

    the location of the peak tracer cloud, versus jun

    tions with steadily diminishing residual concentr

    tions, was not possible during simulations wit

    lesser quantity. The tracer did not move down

    stream as a single pulse because of the comple

    flow structure in the model (Fig. 6).

    5.3. Sensiti6ity analysis

    Model response to calibrated values of con

    stituent boundary loading and rate constants wa

    evaluated by two series of sensitivity analyses. Th

    first series proportionately altered the nitroge

    and phosphorus loads (both organic and ino

    ganic) in the calibrated model by factors of 0.5

    1.5, and 2.0 times. The nitrogen and phosphoru

    simulations were each run separately, then thewere combined. Results were plotted as the pro

    portion change (from the calibrated model) i

    constituent concentration (abscissa; 1.0=cal

    brated model) and the proportion change (from

    the calibrated model) in mean growing seaso

    Chl-a concentration in the segments 1 and 6 (ord

    nate; 1.0=calibrated model). Growing season

    defined as May through September.

    The second series of sensitivity analyses in

    volved altering the values of the phytoplankto

    maximum growth rate and the nitrogen and phophorus half-saturation constants (Table 5). Th

    range of values used corresponds to the rang

    found in Bowie et al. (1985). The effect of bot

    increases and decreases was simulated. Howeve

    the calibrated values for all three constants wer

    near an extreme from the literature source s

    there was some limitation in available choice

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    D.L. Tufford, H.N. McKellar /Ecological Modelling 114 (1999) 137173 1

    Results of the second series were plotted as the

    rate constant value (abscissa) and a sensitivity

    index Eq. (8).

    Sensitivity index (SI)

    =(D Chla)}(calibrated mean Chla)

    (D rate constant)}(value of constant in calibrated model) (8)

    where D Chl-a is the change in growing season

    mean CHl-a, and D rate constant is the change in

    value of the rate constant.

    6. Results and discussion

    6.1. Hydrodynamic model calibration

    The hydrodynamic model was calibrated with amaximum volume difference of 3.3% (Fig. 8). The

    lake starts the year at its annual low volume. Late

    in January it begins a rapid increase to its annual

    maximum in late March. This corresponds to the

    period of highest discharge of the Congaree and

    Wateree Rivers (Fig. 9) and graphically depicts

    the reservoir function as a flood control structure.

    The volume steadily declines after reaching its

    peak. This is due to low tributary inflows, sea-

    sonal high power requirements, and to increase

    storage capacity during the tropical storm season(Inabinet, 1985). Discharges begin to increase

    again in late summer and early autumn, resulting

    in higher volume. During December, even though

    river inflow is relatively high, volume rapidly de-

    Fig. 9. Mean daily discharge of the Congaree and Water

    Rivers during the study period.

    creases in preparation for the flood control r

    quirement after the beginning of the year.In general, the model tracks the significant an

    nual volume trends very closely. Principal di

    crepancies are that it does not increase in volumas quickly as the lake early in the year and once a

    maximum volume the model declines more gradu

    ally. But it is sensitive to even short term fluctuations such as the volume increase at about da

    230 and the brief dip beginning at day 270. Th

    lag is likely an artifact of the 10-day and monthaveraging of the tributary flows.

    The measured volume was estimated from ac

    tual lake stage as recorded by SanteeCooper. Aempirical stage-to-volume relationship in such a

    enormous, irregularly shaped basin has some unknown error in its accuracy. This suggests thcloser correspondence between measured an

    modeled volumes may not necessarily bring th

    model into closer agreement with actual lakvolume.

    Fig. 8. Volume comparison of the calibrated hydrodynamic

    model to the estimated 5-year daily mean volume. Daily

    proportional difference is also shown.

    Fig. 10. Comparison of tracer concentration when the hydr

    dynamic model has evaporation included versus not include

    Data represents junction 23, as an example.

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    D.L. Tufford, H.N. McKellar /Ecological Modelling 114 (1999) 137173154

    6.2. Effect of e6aporation

    During execution of the conservative tracer

    model a deterministic deviation from uniform

    concentration occurred. This resulted from evapo-

    rative loss from the lake surface. Tracer does not

    leave the lake with evaporation so it is concen-trated in the lake by the evaporative loss. A model

    run with evaporation removed from the hydro-

    dynamic model eliminated the evaporative con-

    centration (Fig. 10).

    Table 4 includes some literature values based

    on mechanistic models of lake evaporation for a

    geographic range that includes the study area.

    The evaporation used in this model is an underes-

    timate by at least 2.5 times compared to them.

    James and Bierman (1995) estimated open water

    rainfall as 80% of the land station gauges for theirmodel of Lake Okeechobee; the current model

    used 100%. In combination these literature values

    suggest the net evaporation used in this model

    may be conservative. If so, the evaporative con-

    centration effect may be more pronounced, and

    significant, than the current results indicate.

    Evaporation was left in the hydrodynamic

    model for use with the water quality model. The

    numbers used were empirical estimates of the net

    monthly open water evaporation and rainfall

    (Table 4). The effect is less important at theupriver extreme (maximum of 1% in junction 12

    and 2% in junction 13 versus 6% in junction 23

    and 10% in junction 41). This may reflect the

    influence of residence time on the opportunity fo

    evaporation to occur. The maximum evaporativ

    effect may have little impact on lake water qualit

    constituents. However, since the estimate use

    here was at the low end of literature estimates, th

    effect may be considerably greater than that ob

    served in this model. Under any estimate, it likely the effect would only be significant in th

    downstream portion of the lake.

    6.3. Water 6elocity and settling rates

    The model suggests the existence of longitud

    nal velocity gradients in the lake (Fig. 11). This

    often seen in reservoirs (Ford, 1990) and wa

    observed in Lake Marion during a tracer stud

    (Patterson and Harvey, 1995). Velocity gradien

    can influence sedimentation of tributary particulate burden (Thornton, 1990); heavier particles a

    lost first followed by lighter material as velocit

    further reduces downstream. Settling also can be

    significant loss to water column phytoplankto

    and nutrient stocks (Kennedy and Walker, 1990

    The water quality model component of th

    WASP5 system includes settling flow specific

    tions to account for loss and resuspension o

    sediment, phytoplankton, and particulate nutrien

    fractions. These values were determined by cal

    bration using literature ranges as a guide (Fi12). In the upper- and mid-lake junctions (13 an

    22) there is no settling during the period of max

    mum velocity early in the year (Figs. 11 and 12

    Fig. 11. Comparison of water velocities estimated by the

    hydrodynamic model for three junctions (13, 22, and 41)

    indicating the upstream-to-downstream gradient.

    Fig. 12. Particle settling velocities for three junctions (13, 2

    and 41). Rates were estimated by model calibration.

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    D.L. Tufford, H.N. McKellar /Ecological Modelling 114 (1999) 137173 1

    Fig. 13. Annual modeled velocity profile in junctions just upstream (17) and downstream (18, 20, and 22) from I-95.

    Settling increases as velocity declines in midyear

    before reducing again late in the year. Settling in

    the lower lake is uniform throughout the year.

    Water velocity is also much less than in upstream

    portions of the lake. This result suggests a

    threshold advection velocity, below which most

    particulate material in the lake is capable of set-

    tling out. However, it cannot be inferred from this

    that all suspended sediment is removed from the

    lake water by settling prior to outflow. Residence

    time is a factor. In a study from 1983 to 1985, the

    USGS reported a suspended sediment trap effi-

    ciency of about 73% for Lake Marion (Cooney,

    1988).Turbulence may be a factor in settling as well.

    The model does not numerically account for pro-

    cesses that reduce net sedimentation such as tur-

    bulence and horizontal advection so rate estimates

    must incorporate these effects. The upper portion

    of the lake is relatively shallow with variable

    bottom topography. This along with greater ve-

    locity variability may increase turbulence. These

    factors do not exist in the lower lake.

    The I-95 bridge across the lake restricts open

    flow to a single, main channel that is approxi-mately 47% of lake width at the southern end of

    the bridge. This constriction appears to create a

    velocity gradient (Fig. 13) with a significant in-

    crease in the junction (18) immediately down-

    stream from the bridge. The gradient is greatest in

    magnitude during periods of high flows (compare

    Figs. 9 and 13); the first 70 days of the year are

    especially pronounced. It is uncertain what effec

    this has although it may lead to enhanced particu

    late settling upstream from the bridge and r

    Fig. 14. High flow tracer simulation results. Peak tracer con

    centration indicated by darkest shading.

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    Table 6

    Hydraulic residence time estimates (days) for Lake Marion from the WASP5 model (shown are four hydrologic extremes and annu

    mean)

    Min. flow Min. volMax. flow Max vol. Annual mean

    1519.3 1399.5Vol.106 m3 1495.4 1506.31555.8

    365 n/a22691Day of year 66

    177 522Outflow (m3 s1) 682 384501

    176 410Inflow (m3 s1) 797 511 406

    466177 395506Mean-flow (m3 s1) 739

    Retention

    99 35Whole lake 23 36 44

    843 56168Jacks Creek 105

    18 5Tawcaw Creek 4 5

    9812041609Potato Creek 162

    45Wyboo Swamp 32 46 141

    duced residence times just below the bridge. The

    early year period is also the time of greatest

    sediment transport into the lake from the water-

    shed (Patterson et al., 1996), so enhanced settling

    may exacerbate basin filling in the upstream por-

    tion of the lake. This inference seems to contra-

    dict the discussion of calibrated settling flows,

    immediately above. Both may apply to the results

    observed if differing particle size fractions are

    involved. The heaviest sediment particles may not

    play a significant role in water column nutrient

    dynamics but they are most likely to settle in the

    upstream portion of the lake.

    6.4. Hydraulic tra6el time

    USGS conducted a tracer study in Lake Mar-

    ion in 1984 (Patterson and Harvey, 1995). They

    estimated tracer travel time through the lake (be-

    ginning in the Wateree River upstream from the

    Santee Swamp) during periods of high and low

    inflow from the Congaree and Wateree Rivers.

    For logistical reasons they divided the lake into

    three sections; their lower two sections togethercorrespond to the lake as described for the current

    research. As a comparison, a similar tracer slug

    simulation was run with this model. Travel time

    from the Rimini trestle to the Diversion Canal at

    high flow (775 m3 s1) estimated by the model is

    17 days (Fig. 14). USGS (810 m3 s1) estimated

    13 days. The low flow (222 m3 s1) travel time for

    the model is 53 days; for USGS (240 m3 s1) it

    40 days. The differences are fairly small conside

    ing the significant differences in method betwee

    the two estimates. This result is a confirmation o

    the volume calibration discussed above; the mod

    appears to lag the lake in its hydrologic respons

    The tracer result also provides evidence of lim

    ited hydrologic exchange between the open lak

    and the small tributary embayments (Fig. 14

    also found by Patterson and Harvey (1995). Th

    embayment toward the upstream end of the lak

    receives less tracer and retains it well after th

    peak tracer concentration has moved downstream. Similarly, the embayment just past th

    middle of the lake retains tracer after the pea

    concentration has moved past it. The embaymen

    at the downstream end of the lake has not r

    ceived any appreciable amount of tracer as th

    peak concentration first reaches the diversio

    canal.

    6.5. Hydraulic residence time

    Hydraulic residence time can influence reservomixing processes, nutrient availability, and phyto

    plankton dynamics (Thornton, 1990). A short re

    idence time (rapid flushing rate) can decrease th

    opportunity for phytoplankton production. It ca

    also result in additional nutrient import from th

    watershed resulting in increased production afte

    a temporal lag. If longitudinal zonation develop

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    Fig. 15. Segment 02 calibration charts. Monthly monitoring data (5-year mean91 S.E.) and monthly mean model results.

    due, in part, to long residence times, it can be

    reduced during periods of increasing flushing

    rates.

    Previous estimates of Lake Marion residence

    time are variable. Inabinet (1985) estimated an

    annual mean residence time of 31.1 days using a

    mean outflow of 577 m3 s1 and volume of

    1548.31106 m3. Patterson and Harvey (1995

    as part of their tracer experiments, made tw

    estimates of residence time, one each during hig

    and low flow periods. Both used a lake volume o

    1664.55106 m3. The estimate based on mea

    inflow during the tests was 73 days at low flo

    (266 m3 s1) and 23 days during high flow (82

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    Fig. 16. Segment 10 calibration charts. Monthly monitoring data (5-year mean91 S.E.) and monthly mean model results.

    m3 s1). Retention based on their actual tracer

    results was 60 days at low flow and 20 days at

    high flow.

    With WASP5 a modeled estimate of lake vol-

    ume and discharges are known at any interval.

    Internally, WASP5 estimates retention time for

    each segment using its current volume and mean

    of segment inflow and outflow. Retention times a

    the whole-lake scale were derived using tha

    method (Table 6). Because of the extreme time

    for the embayments their volume was not in

    cluded in the whole lake estimates. Estimate

    given are at principal hydrologic extremes and a

    annual mean. Times range from 23 days at hig

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    Fig. 17. Segment 30 calibration charts. Monthly monitoring data (5-year mean91 S.E.) and monthly mean model results.

    flow during March to 99 days at low flow dur-

    ing August. The annual mean is 44 days. Peri-

    ods of high flow and low volume tend to

    coincide resulting in rapid flushing in the cool

    months. Discharge is high (and residence time is

    low) even at maximum volume, however, indi-

    cating the longest residence times occur only

    during periods of very low flow. This result sug

    gests that tributary discharge is a dominan

    forcing function in Lake Marion. From the ea

    lier discussion this effect is subject to modific

    tion spatially and temporally, creating

    heterogeneous hydrologic environment in th

    reservoir.

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    Fig. 18. Incident light curve used in the water quality model and comparison of three estimates of saturating light intensity. See te

    for discussion.

    6.6. Water quality model calibration

    The water quality model calibration demon-

    strated the capability of the model to simulate the

    spatial heterogeneity that occurs in the lake. (See

    Table 1 for summary of reaction rates and coeffi-

    cients.) Figs. 1517 are example calibration charfor segments 2 (upstream), 10 (mid-lake), and 3

    (downstream), respectively. The model tracks Ch

    a dynamics well in each segment even though th

    annual pattern was different for all three. Th

    model also simulates the annual cycle of the re

    maining constituents well. Discrepancies are on

    of two types: a divergent month or two occurs i

    an otherwise close correspondence (NH3/4 in seg

    ment 10, for example) or the model diverges i

    magnitude but the annual pattern was general

    correct (NO2/3 in segment 2, for example).Some discrepancies may reflect the existence o

    a process in the lake that was not conceptualize

    in the model. For example, DO simulation tend

    to be very accurate. This probably indicates tha

    DO dynamics are controlled more by water tem

    perature and reaeration than by water colum

    biochemistry. In segment 2, however, the mod

    misses a significant dip in September. Fish kill

    believed to be caused by DO exhaustion, we

    recorded in that area of the lake at that time o

    year during the study period (Bates and Marcu1990). The kills occurred after an extended dr

    period followed by a heavy rain and flood surg

    through the Santee Swamp. A large pulse o

    reduced organic material entering the upper lak

    from the swamp may have caused the DO dip.

    The DO depression can be simulated with th

    model by introducing a BOD5 load of 350

    Fig. 19. Comparison of Chl-a results when using three differ-

    ent estimates of saturating light intensity. The calibrated

    model was used as the base. Shown are data for segment 2

    (upper), segment 10 (middle), and segment 30 (lower).

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    D.L. Tufford, H.N. McKellar /Ecological Modelling 114 (1999) 137173 1

    day1 from the swamp into segment 2 during the

    September interval. At the flow rate into segment

    2 specified in the hydrodynamic model (mean

    discharge of 34.41 m3 s1) this BOD5 load

    corresponded to a concentration of 117.7 mg l1.

    The mean concentration of BOD5 into the seg-

    ment based on measured boundary conditionswas 1.4 mg l1. A simulation run in which the

    inflowing water had no DO did not have an

    observable effect on model results.

    This BOD5 test result raised a question as to

    whether or not a load from outside the lake could

    have been the sole cause of the DO reduction

    observed in the field data. Even assuming a flood

    surge of 10-times the modeled swamp discharge

    (an unrealistically high value) the field data for

    BOD5 should approach 11.8 mg l1. Decaying

    vegetation, carried as bed-load and undetected as

    BOD5, may be a partial explanation for these

    results. In-lake effects also seem likely. Senescence

    of extensive macrophyte beds in the segment is

    one possibility. That a large BOD5 flux had oc-

    curred in the lake was suggested by the observa-

    tion of BOD5 spikes during September in

    segments 10 and 30 (Figs. 16 and 17).

    Another possible source of discrepancies in the

    calibrated model was that most of the biochemical

    rates and constants have a single value for the

    entire lake. This does not allow for possible eco-

    logical differences among regions. Differencesmay result from physical characteristics such as

    depth or water velocity, or biochemical character-

    istics such as phytoplankton assemblages or ex-

    tent of macrophyte coverage.

    Phytoplankton, for example, adapt their physi-

    ology to differing physical environments (Kirk,

    1994) so even assuming community homogeneity

    throughout the lake there still may be some varia-

    tion based on adaptation. Inabinet (1978) found

    evidence of longitudinal variability in species oc-

    currence in the lake. The significance of his resultsare difficult to interpret because he only noted

    presence or absence so relative densities cannot be

    assessed. But this does suggest differences among

    areas in the lake that may be reflected in the rates

    and constants in a mechanistic model.

    The hydrodynamic model suggests a cause for

    the discrepancy in the phytoplankton results in

    downstream segments 10 and 30 during Februar

    Approximately 50% of the I95 crossing of th

    lake is a built on a land bridge, resulting in

    constriction. The constriction may cause poolin

    upstream from the bridge and a significant in

    crease in velocity just downstream from th

    bridge. This increase may have a washout effect ithat area of the lake; the effect is greatest durin

    the highest flows early in the year. If this effec

    exists, as suggested by the hydrodynamic model,

    would seem the water quality model should r

    spond to it. A possible explanation is that th

    actual velocity increase may be much greater i

    the lake than in the model. This could be e

    plained by the 10-day averaging of flows used t

    parameterize the hydrodynamic model.

    6.7. Phytoplankton kinetics

    Phytoplankton production is dependent o

    availability of nutrients and favorable environ

    mental conditions, light and temperature (Fig. 7

    The water quality model takes the supplied phyto

    plankton maximum growth rate and modifies

    with a temperature correction factor and light an

    nutrient limitation factors Eq. (9).

    Rg=GmaxFtF1Fn (9

    where: Rg is the effective phytoplankton growt

    rate, Gmax is the maximum growth rate, Ft is thtemperature correction factor, Fl is the light lim

    tation factor and Fn is the nutrient limitatio

    factor.

    The nitrogen and phosphorus nutrient limit

    tion factors are derived using the MichaelisMen

    ton construct with half-saturation constan

    supplied in the model input file. The mod

    derives the light limitation factor as a function o

    incident light, light extinction in the wat

    column, water column depth, saturating light in

    tensity, and the daylight fraction of the currenday. Temperature correction may range above o

    below unity; the other three factors are all b

    tween 0 and 1. The model selects the lowe

    nutrient limitation factor for adjustment of th

    growth rate. This reflects the empirical and labo

    ratory observation that the growth rate is dete

    mined by the availability of the nutrient i

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    D.L. Tufford, H.N. McKellar /Ecological Modelling 114 (1999) 137173162

    Fig. 20. Annual curve showing sediment OPO4 flux determined by calibration. DO curve included to show inverse relationshi

    Fig. 21. Annual curve showing sediment NH3/4 flux determined by calibration. DO curve included to show inverse relationshiRates for segment 16 are 1.5 times those indicated here.

    shortest supply (Hecky and Kilham, 1988). The

    phytoplankton growth rate for a given model day

    was adjusted below maximum as a result of nutri-

    ent or environmental limitations.

    Kinetic rates and constants in models of differ-

    ent aquatic systems vary due to physical and

    biochemical differences among environments

    (Bowie et al., 1985). If rates are not known for a

    specific environment they are estimated duringmodel calibration. Recognition of the variability

    in, and uncertainty of, key rates was the prime

    motivating factor in a related project that evalu-

    ated phytoplankton growth kinetics in situ (Su-

    darshan, 1995). She estimated the phytoplankton

    maximum growth rate, the temperature correction

    factor, and the nitrogen and phosphorus half-sat-

    uration constants using a nonlinear curve fittin

    technique. The results of that work provided guid

    ance within which calibration of other parameter

    in the model proceeded. The final model was at o

    within the 95% confidence interval of all ra

    estimates developed by Sudarshan (1995).

    The Sudarshan (1995) estimate of a linear rela

    tionship for predicting saturating light intensit

    from incident solar radiation was coded into thphytoplankton growth kinetics module. She foun

    that the saturating intensity increases as inciden

    radiation increases. Saturating intensity is a con

    struct used to account for the field and laborator

    observation that phytoplankton growth increase

    with increasing light intensity up to a certain leve

    Above that level light toxicity causes growth rat

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    Table 7

    Mineralized nutrient loads in the calibrated model

    Santee River Sediment ProportionAnnual load

    (kg)

    851 883 3 931 503 4.62N as NH3/40.785 018 103N as NH3/4 3 931 503

    and NO2/3765 092 383 561P as OPO4 0.50

    ences to Smith and DiToro during this discussio

    refer to the WASP5 implementation.

    The Sudarshan and Smith derivations of satu

    rating intensity are similar during the growin

    season but diverge during periods of low inciden

    intensity (Fig. 18). The similarity between Smit

    and Sudarshan was reflected in the resultingrowth (Fig. 19). Differences tend to be in th

    months associated with low incident radiation (se

    especially segment 30). This result suggests th

    relationship developed by Sudarshan (1995) wa

    more sensitive to the actual light environment i

    Lake Marion, though this sensitivity did not ap

    pear to have a significant impact on model result

    The default fixed value for saturating intensit

    was always greater than the derived intensity (Fi

    18). The growth results for both Smith and Suda

    shan are different from DiToro (Fig. 19). DiToris not fully comparable since the phytoplankto

    kinetic is different in other respects in addition t

    the saturating light intensity. However, the rela

    tive absence of effect in segment 2 suggests th

    upstream area is not as sensitive to the ligh

    conditions as the downstream area.

    6.8. Internal nutrient loading

    Nutrient calibration required inclusion of ben

    thic sources of both phosphorus (as OPO4) annitrogen (as NH3/4). Rates are comparable t

    literature ranges (Ryding and Forsberg, 197

    Thomann and Mueller, 1987) and increase durin

    midyear (Figs. 20 and 21). The inverse relation

    ship between benthic release rates and DO con

    centration suggests higher sediment flux durin

    periods of low DO and high temperature.

    The relationship between temperature, low DO

    and sediment nutrient release is much studied an

    discussed in the literature. Aerobic sediment flu

    has been reported by several researchers (Rydinand Forsberg, 1977; Premazzi and Provini, 198

    Thomann and Mueller, 1987) although the flu

    during anaerobic conditions is greater. Of thos

    who report values for an annual cycle, relea

    rates are higher during warm seasons (Baccin

    1985; Recknagel et al., 1995). Lee et al. (1977

    explicitly concluded that release appears affecte

    reduction (Kirk, 1994). This effect is referred to as

    photoinhibition. The critical light level is known

    as the saturating light intensity.

    The Sudarshan (1995) formulation replaced the

    derivation already in the module. The Smith

    (1980) derivation is a function of incident light,

    temperature, light available in the water column,

    and the phytoplankton carbon-to-chlorophyll ra-tio. It derives a time-integrated light limitation to

    growth based on the varying amount of light

    available in the water column during the day. An

    alternate method in WASP5 is to use a fixed value

    for saturating light intensity (default was 300 ly

    day1) based on work by DiToro et al. (1971).

    This method develops a light limitation factor as a

    single instantaneous estimate for each day. (See

    Ambrose et al., 1993a for a discussion of the

    WASP5 implementation of these kinetics.). Refer-

    Table 8

    P-values for the KolmogorovSmirnov validation test

    OPO4Segment NH3/4CHL-a NO2/3

    0.541 0.0021 0.5410.006

    0.541 0.1142 0.203 0.056

    0.019 0.0100.5183 0.034

    0.699 0.2064 0.0060.000

    0.758 0.0015 0.010 0.001

    0.518 0.2706 0.002 0.001

    0.270 0.2708 0.964 0.627

    0.699 0.0009 0.076 0.2060.441 0.03110 0.893 0.441

    0.2700.08814 0.9640.270

    0.001 0.13716 0.270 0.270

    0.1640.015 0.001 0.00326

    0.1640.015 0.164 0.05527

    1.0000.0140.40128 0.329

    0.015 0.7590.16429 0.401

    1.0000.0820.32930 0.329

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    D.L. Tufford, H.N. McKellar /Ecological Modelling 114 (1999) 137173 1

    Fig. 23. Segment 02 validation charts. Field data (+) and model results.

    are well described in the model, likely due to the

    field work that estimated important rate values.

    An alternative explanation is that phytoplankton

    production is not sensitive to the difference in

    nutrient loading values used in this validation.

    Validation was especially weak in the upstream

    segments (1,3,4,5). The explanation may be due,

    in part, to the nutrient load into the lake (Fig.26). During the validation run the phosphorus

    load was less and the nitrogen load was greater

    than during calibration (Fig. 26 represents load

    from the Santee River only). Differing nutrient

    levels may have altered system primary produc-

    tion and nutrient cycling. Downstream areas of

    the lake are apparently under less direct riverine

    influence.

    Another view of this issue considers actual

    lake hydrology in the calibration and validation

    intervals. The hydrodynamic model used in bothcalibration and validation of the water quality

    model was developed using hydrologic data from

    1985 to 1990. The annual discharge of the Con-

    garee and Wateree Rivers during the calibration

    interval was 22% greater than during the valida-

    tion interval (Fig. 27). So the nutrient loads dur-

    ing the validation run (Fig. 26) are likely

    overestimates of what actually occurred becau

    the modeled discharge was greater than the mea

    sured discharge during the interval. The use of

    longer term discharge dataset in the hydrody

    namic model would probably not solve th

    problem; it may exacerbate it (Fig. 27). This ha

    implications for future modeling studies of LakMarion. To obtain reasonable results it may b

    necessary, depending on the modeling objective

    to recalibrate the hydrodynamic model represen

    ing the specific hydrologic conditions during th

    model interval.

    The validation results suggest hydrodynamic

    and riverine nutrient loads are dominant forcin

    functions in controlling nutrient status in Lak

    Marion. This is especially significant in light o

    the earlier discussion of the apparent importanc

    of benthic sources of nutrients to lake productivity. Phytoplankton productivity was influence

    by other factors as well; light and temperatur

    based on results discussed earlier. A high degre

    of spatial zonation was also suggested, confirm

    ing results from the model calibration and othe

    research in Lake Marion (Inabinet, 1985; Picke

    and Harvey, 1988).

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    Fig. 24. Segment 10 validation charts. Field data (+) and model results.

    6.10. Sensiti6ity analysis

    Altering nitrogen loading in the model suggests

    that most of the lake has ample nitrogen for

    phytoplankton growth (Fig. 28). Upper-lake (seg-

    ment 2) growing season mean Chl-a concentra-tions are substantially attenuated by nitrogen

    reduction, but at the other simulated levels the

    entire lake was unaffected. The result for segment

    2 may be due to the riverine nature of that

    segment; nutrients are either used by phytoplank-

    ton for growth or they are transported down-

    stream. A reduction in any nutrient load reduces

    the opportunity for photosynthetic assimilation

    and growth.

    The model exhibits greater sensitivity to alter-

    ations in phosphorus loading than to nitrogenloading (Fig. 28). Again this is reflected more in

    segment 2 than in either the mid- or lower-lake

    segments (10 and 30, respectively). This result for

    segment 2 tends to support the above observation

    from the nitrogen loading test, and further sug-

    gests phosphorus limitation in the upstream

    segment.

    The effect of decreasing both nutrients load

    was not different from the phosphorus result, bu

    increasing both had a cumulative effect. In up

    stream segment 2, doubling phosphorus alone r

    sulted in a 60% increase in growing season mea

    Chl-a concentration; when both nutrients wedoubled the effect was an 80% increase. Th

    incomplete utilization of the additional nutrien

    load may result from relatively short residenc

    times. Also that of the total boundary nutrien

    loads, 60% of the nitrogen and 26% of the pho

    phorus are in the immediately bioavailable mine

    alized forms. The result in segments 10 and 3

    (mid- and lower-lake) suggests colimitation t

    phytoplankton growth by both nutrients.

    The validation test discussed earlier raised th

    question of whether or not the changed nutrienload could have produced a detectable differenc

    in Chl-a concentration from the calibrated mode

    Nitrogen loading during validation was propo

    tionately 1.2 times greater than during calibration

    phosphorus loading was 0.55 times the amount i

    the calibrated model. According to the sensitivit

    results (Fig. 28) a significant reduction in Chl

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    D.L. Tufford, H.N. McKellar /Ecological Modelling 114 (1999) 137173 1

    Fig. 25. Segment 30 validation charts. Field data (+) and model results.

    concentration would be expected, especially in

    upstream segment 2. This was the situation (Fig.

    29), suggesting that the model can be used to

    project the effect of changing nutrient loads that

    actually occur, or may occur, in the study area.

    The second sensitivity analysis evaluated modelresults with respect to values of three kinetic rate

    constants that are key to phytoplankton produc-

    tion in the WASP5 model. A negative value of the

    sensitivity index indicates the corresponding rate

    constant value reduces growing season mean Chl-

    a concentration relative to the calibrated model; a

    positive value indicates an increase (Fig. 30).

    (Note that conceptually the SI for the calibrated

    model is zero, but arithmetically it is undefined;

    see Eq. (8).) Results for the analysis of the nitro-

    gen half-saturation constant are not presentedbecause variation from the calibrated model was

    extremely small.

    The three areas of the lake respond to the rate

    changes in varying magnitudes. Segment 2 ex-

    hibits the greatest range, further suggesting the

    highly dynamic environment of this riverine sec-

    tion of the lake. For all three segments and for

    both rate constants, the value of the constant