Par Pond Research Articles

Distribution and biomass of aquatic macrophytes in an abandoned nuclear cooling reservoir

Pond B is an 87-ha abandoned reactor cooling reservoir on the Savannah River Plant, South Carolina. Since abandonment in 1964, the reservoir has reverted to a low conductivity and slightly acidic water chemistry, and has also been colonized by abundant macrophyte vegetation. To determine dominant species and their depth distributions, and to estimate seasonal changes in standing crop, above-sediment biomass of macrophyte vegetation was sampled at four water depths three times during the 1986 growing season. Three floating-leaved species, Nymphaea odorata Ait., Brasenia schreberi Gmel and Nymphoides cordata (Ell.) Fern., comprised 51.8% of biomass harvested, with greatest biomass at 1.5 m and in the October sampling. Free-floating Utricularia spp. (mostly U. floridana Nash) contributed 32% of harvest biomass, were abundant at all sampling depths and had greatest standing crop in May. Submersed rooted species comprised 15.4% of biomass harvest, dominated by Cobomba caroliniana var. pulcherrima Harper in deeper portions of the littoral zone. The species composition found in Pond B was similar to that of aquatic plant communities in other softwater Coastal Plain habitats (such as Carolina Bay ponds and rain-fed swamps), and differed from species composition reported for Par Pond, a neighboring reservoir with higher conductivity. These comparison suggested that differences in water chemistry may have been important in determining establishment of macrophytes species in these two reservoirs.

Dissolved Carbohydrate Distribution and Dynamics in Two Southeastern United States Reservoirs

Dissolved carbohydrate concentrations were measured in the waters of two southeastern United States reservoirs (Pond B and Par Pond) in 1982 and 1983 with attention to the influence of rooted aquatic macrophytes, spatial and temporal variability, and depth. Mean concentrations for all sampling locations were 2.86 ± 1.01 mg∙L−1 in Pond B and 1.72 ± 1.24 mg∙L−1 in Par Pond. There were no consistent spatial or temporal differences within a reservoir, nor were there consistent differences between macrophyte beds and open waters in either impoundment. Dissolved carbohydrate concentrations increased in both reservoirs during the day, reaching maxima in late afternoon, with an accompanying decrease in the dissolved organic carbon (DOC). The dissolved carbohydrates comprised 5.8–20.2% of the DOC in Par Pond and 25.7–42.3% of the DOC in Pond B. In both reservoirs, vertical profiles show the presence of carbohydrates at all depths during aerobic and anaerobic conditions. Midwater concentration maxima were present in Par Pond during periods of holomixis and stratification. The maxima were coincident with an intense pycnocline during periods of stratification; but during holomixis, the maximum was apparently unsupported by physical structure.

The Impact of Preemption on the Zonation of Two Typha Species Along Lakeshores

The objective of this study was to examine the effects of preemption on rates of competitive displacement and on competitive outcomes for two Typha species (T. latifolia, TL, and T. domingensis, TD) growing along gradients of water depth. Evidence from several experiments indicated that the relative competitive abilities of the two species were size dependent. When TL and TD competed as small seedlings, TD was generally favored while TL was the better competitor when the plants were larger before beginning to compete. Because of this effect, initial density was determined to influence strongly the early outcome of competition. When seeds were sown at high densities, TD gained an early advantage; at lower densities TL was the better competitor. At very low densities, such as when the species competed as clones, TL was very much the better competitor. In the absence of preemption, the results showed that differences in germination, seedling growth, and competitive ability act to create an initial segregation of species at the waterline where establishment occurs. TL is upslope and TD downslope initially, but as the species spread downslope, TL progresses faster because of its greater capacity for vegetative spread, and the result is a widespread overlap along the gradient. Studies of the rate of competitive displacement between established plants yielded an estimate of slightly <6 yr required for complete segregation of two species starting from maximum overlap. Preemption had a strong effect on early growth and distribution of the species. After the initial effects of temporal disadvantage, delayed plants were able to recover under certain circumstances. Both species recovered better in shallow water than in deep water, and, in general, TL recovered better than did TD. Experiments on spatial preemption found that the species moved towards the expected segregation of TL upslope and TD downslope regardless of the initial conditions. This process of recovery was somewhat slower for established stands of adults than for seedlings. Comparison of aerial photographs of natural populations of TL and TD in PAR Pond (South Carolina) in 1980 with distributions in 1984 also suggests that recovery from even large—scale spatial preemption can occur over only a few years time. These results suggest that extreme temporal advantage by either species can lead to extinction of the other species and that the magnitude of the effect depends upon the initial density of seedlings. If both species survive the initial phase of competition and the habitat remains undisturbed, subsequent vegetation spread and competitive displacement can over—come the initial effects of spatial and temporal preemption, resulting in a predictable zonation.

Piscine macrophage aggregate parameters as health monitors: effect of age, sex, relative weight, season and site quality in largemouth bass (Micropterus salmoides)

The utility of macrophage aggregate (MA) parameters as indicators of fish health and/or environmental stress was tested in largemouth bass ( Micropterus salmoides) collected from Par Pond, a cooling reservoir for the Savannah River nuclear power plant. Initially, the effects of sex, age and season were evaluated using relative weight ( W r) in both healthy ( W r > 80) and stressed ( W r < 80) bass. There was no sex effect. A number of MA parameters, primarily hepatic and splenic no./mm 2 increased linearly with age in the healthy fish. These correlations were not apparent in the stressed bass. A comparison of stressed and healthy fish of the same age showed that stressed bass had significantly more aggregates in liver and spleen as well as higher relative amounts of hepatic iron. The only seasonal difference noted in healthy bass was that fish collected in the spring had significantly less relative iron than those collected in the fall. We then tested the sensitivity of these parameters by comparing fish of similar age and W r from two sites — the Hot Dam area which is thermally impacted and the Cold Dam arca, an unimpacted site. Fish collected from the impacted area had significantly more aggregates in the liver than those collected from the Cold Dam area. Thus, we believe these parameters to be very sensitive indicators of environmental stress.

BIOCHEMICAL GENETICS OF MOSQUITOFISH. IV. CHANGES OF ALLELE FREQUENCIES THROUGH TIME AND SPACE.

Gene frequency data from samples of Gambusia affinis populations at 76 localities across the Savannah River drainage were used to investigate temporal and spatial patterns in population genetic structure. Localities in the Par Pond system on the Savannah River Plant were sampled in 1971, 1977, and 1979. Allelic frequencies in these populations were generally stable through time, although significant temporal changes were observed among samples from Pond C, an impoundment receiving thermal effluent. Significant spatial heterogeneity in allele frequencies was observed on both microgeographic and regional scales. Populations within the Par Pond system were spatially subdivided at four of the five loci surveyed (mean FST = 0.051). Subdivision was even more pronounced when samples from across the Savannah River drainage were compared (mean FST = 0.196). A hierarchial analysis of gene diversity (GST ) demonstrated that most of the genic diversity across the drainage exists as within-subdivision diversity. Even when populations from such contrasting habitats as rivers, creeks, ponds, and reservoirs are compared, an average of only 13% of the total gene diversity was attributed to between-group diversity. Greatest between-group gene diversity was observed when reservoirs were compared with one another. This general pattern of low between-habitat diversity suggests that differential selection pressures are not playing a major role in producing the observed levels of subdivision. In the Par Pond system, neither single locus nor multilocus genetic distances were significantly associated with geographic distance or with its reciprocal. For samples from over the Savannah River drainage, significant correlations between genetic and geographic distance were observed only for the Gpi-2 and Pgm-2 loci. Thus, there was a general lack of concordance between genetic and geographic distances. Spatial autocorrelation demonstrated patterns consistent with Wright's isolation by distance model. Significant positive correlations in allelic frequencies among neighboring populations were observed for five of six alleles; allelic frequencies in more distantly separated populations were typically not correlated.

Winter-Spring Depth Distribution of Chironomidae in a Southeastern Reservoir

Distribution and abundance of midges (Diptera: Chironomidae) along a water depth gradient were studied in thermal and ambient temperature areas of a reservoir in the southeastern United States. Benthic samples were taken at 10 depths (0.16-5 m) during the winter and spring of 1978. Density, genera richness, and genera diversity increased progressively from the shore line to moderately shallow depths (1-2 m) and then decreased at greater depths. This pattern of benthos distribution in Par Pond is compared with those in other lentic systems. Causes for this pattern are multiple but probably relate to habitat complexity (vegetation) and substrate characteristics.

Growth and mineral nutrition of cattails inhabiting a thermally‐graded South Carolina reservoir: I. Growth and the macronutrients1

Abstract Growth and macronutrient uptake in two species of cattail, Typha latifolia and T. domingensis, were determined during an annual growth cycle in Par Pond, a thermally‐graded nuclear reactor cooling reservoir in South Carolina. Shoots of the same age class were sampled periodically from the initiation of growth in early spring to senescence in late autumn. Although differences between thermal regimes were mostly statistically non‐significant, growth and macronutrient uptake, especially of N, P, and K, tended to be enhanced by elevated water temperatures. The pattern for growth and N, P, and K uptake in decreasing order were: “Warm”; > “Intermediate”; > “Ambient”; regimes. The non‐significant statistics were attributed to large between‐stand and between‐temperature regime variations. Both Typha species exhibited sharp declines in N, P, and K concentrations in the shoots, and some decline in the rhizomes, early in the growing season, followed by either a more gradual decline or leveling off of concen...

A thermal effluent as a sporadic cornucopia: effects on fish and zooplankton

We report effects of a thermal effluent on fish and zooplankton in an impoundment in South Carolina. The effects are not directly due to heated effluent but are indirect effects on the trophic dynamics of the system and depend on the physical structure of the system, season, and geography. As effluent from a nuclear reactor cooling pond (Pond C) enters the larger Par Pond it sporadically carries with it zooplankton and/or dead fish. Pond C produces high zooplankton densities in two situations: (1) high and hot effluent flow during cold winter weather; and (2) low (cool) effluent flow during warmer seasons. High zooplankton densities attract Blueback herring, Alosa aestivalis, which attract largemouth bass, Micropterus salmoides. Dying and dead bluegills, Lepomis macrochirus from Pond C (heat killed) are eaten by Par Pond bass which swim into effluent temperatures as high as 46°C to take these easy prey. Blueback herring and largemouth bass distribution are affected by the presence of food rather than by temperature. Par Pond blueback herring appear to be available to bass mainly when herring are near the effluent from Pond C and when they spawn in spring in the littoral zone. Because Blueback herring in Par Pond live only one year and because their presence in bass habitat is seasonal there is a strong seasonal component to bass food abundance. A previously reported annual oscillation in bass condition (K), with a peak in winter, occurs throughout Par Pond but is extreme in the vicinity of the effluent. The strong seasonality of food abundance for bass at the effluent correlates the winter peak in bass condition.

Regulation of Freshwater Community Structure at Multiple Intensities of Dragonfly Predation

We examined the role played by predaceous dragonfly nymphs, Celithemis fasciata (Odonata: Libellulidae), in the regulation of the community structure of a benthic macroinvertebrate assemblage in Par Pond, an 1100—ha reservoir in South Carolina. Effects of predation intensity on species richness, evenness, and density wer evaluated by adding zero, two, four, and eight large dragonfly nymphs (antepenultimate and penultimate instars) to previously sieved (0.85—mm mesh) bottom sediment containing benthic macroinvertebrates. Predator and prey assemblages were then placed in individual field microcosms that consisted of polyethylene trays surrounded by underwater screens (mesh &lt;2 mm diagonally) and suspended 15 cm below floating platforms. Twelve replicates of each treatment level were run during each of three 6—wk experimental periods: April—May 1980, August—October 1980, and January—February 1981. In addition, colonization of microcosms by invertebrates was quantified, and samples from natural, unenclosed benthic fauna were collected seasonally along a transect for comparison with experimental assemblages. We tested whether predator enhanced, depressed, both increased and decreased, or had no effect on the complexity of the community structure. The dual effect of predation on community structure is predicted by Connell's "intermediate disturbance hypothesis." Results showed that dragonfly nymphs can significantly influence the structure of the benthic community. However, the results did not show that invertebrate predation is the sole or even the primary regulator of community structure. Species richness was significantly greater at intermediate treatment levels (thus supporting Connell's general hypothesis), but the increase was not great (a range of °10%). The mechanisms by which species richness is maximized at intermediate intensities of predation are not entirely evident, but are probably a combination of prey refuges and nonselective predation with patch switching. In contrast, species evenness, as measured by equitability and by Simpson's index, was greatest at the highest predation level (which does not support the intermediate disturbance hypothesis). Dragonflies appeared to exert a greater influence on prey density than on community diversity.

Seasonal responses of lentic midge assemblages to environmental gradients

We examined biotic and abiotic factors associated with density and diversity of larval midges (Diptera: Chironomidae) in the littoral zone of Par Pond, an 1100 ha cooling reservoir located near Aiken, South Carolina, USA. A total of 144 core samples were removed from 12 areas during each of 3 sample periods. Principal component analysis. Mantel analysis, and various univariate statistical tests were used to detect seasonal differences in (1) associations among areas and genera of Chironomidae, and (2) responses of midges to a multi‐faceted environmental gradient. Water temperature, which in this case was a function of both season and thermal effluents, was the factor most strongly associated with relative changes in density and diversity within the chironomid assemblages. Plant biomass and/or plant diversity were significant factors correlated with midge density and diversity. Both particle size and organic content of the substratum, which varied little across stations, were relatively unimportant factors. Despite the presence of a multifaceted environmental gradient, the composition and relationships within the midge assemblages were remarkably similar across areas although total abundance and diversity varied.On the basis of their similarity in structure and function, we classify the midge assemblages within the reservoir as members of the same community. However, in response to dissimilar or unequal selective forces (principally the area‐specific effects of thermal effluents), these assemblages appear to behave environmentally as multiple units rather than as a single component of the benthic community in Par Pond.

Phytoplankton Distribution in Three Thermally Distinct Reactor Cooling Reservoirs

Phytoplankton community structure was studied in relation to physicochemical characteristics of three South Carolina reservoirs in close proximity and of similar age and bottom type. Thermal alteration, resulting from the input of cooling water from a nuclear reactor, was substantially different in each reservoir. This provided an opportunity to compare water temperature effects separated from season. Water temperature (when examined independently in statistical models) appeared to be less important than other environmental variables in determining phytoplankton community structure. Pond C, a reservoir receiving intensely heated effluent (> 20/sup 0/C ..delta..T), displayed low species diversity (Shannon-Weaver H < 2.0), except in winter. Unexpectedly, more than 50% of the total phytoplankton density in this reservoir was consistently comprised of green algae (principally Chlamydomonas sp.) despite water temperatures that averaged 47.3/sup 0/C in summer. Par Pond, having a maximum ..delta..T of 5/sup 0/C, displayed no temperature-induced alteration of phytoplankton community structure.

Evaluating the constraints of temperature, activity and consumption on growth of largemouth bass

We present a bioenergetics model for largemouth bass (Micropterus salmoides) which simulates growth as a function of body size, temperature, activity and consumption level. We apply the model to investigate seasonal changes in condition factor exhibited by bass in Par Pond, South Carolina, a reservoir receiving heated effluent. Previous authors have suggested that these changes occur due to bioenergetic constraints, primarily the effects of heated effluent on metabolic rate. Model simulations were used to evaluate the hypotheses that seasonal changes in condition factor were caused by the heated effluent, seasonally variable activity, seasonally variable consumption, or reproductive costs.

Field experiments on interactions between vertebrate predators and larval midges (Diptera: Chironomidae) in the littoral zone of a reservoir.

A field experiment was designed to test a frequent assumption in the literature that vertebrate predators (in this case, fish and turtles) are capable of regulating the seasonal abundance and diversity of benthic communities in the littoral zone of lentic environments. Effects of thermal effluents from a nuclear reactor on predator-prey relationships were also examined. Benthic samples were removed after each of three, 3-month test periods from 36 predator exclusion cages (4m2) and 36 control plots located along a thermal gradient in Par Pond, an 1,100 ha freshwater reservoir in the southeastern United States.Results of our field experiments provide little evidence to suggest that either a single "keystone" species or vertebrate predators as a group were capable of regulating the abundance, diversity or productivity of chironomids in Par Pond. The relationship between predator treatment and community response (changes in density and species richness) was generally unaffected by either plot location or temperature fluctuations. When data from caged and control plots were pooled, however, both location and water temperature individually had significant impacts on the chironomid community. Alternative hypotheses are proposed to explain the lack of regulatory control of the benthic community by individual species or guilds of predators.

Agglutinating Antibody to Aeromonas hydrophila in Wild Largemouth Bass

Abstract Among largemouth bass Micropterus salmoides in Par Pond, South Carolina, a significantly larger percentage of those with red-sore disease were positive for anti-Aeromonas hydrophila agglutinin than of uninfected fish. Highest titers occurred during summer and fall, when the prevalence of the disease was declining. Most agglutinin activity was associated with a single serum fraction; the agglutinin has an apparent molecular weight of >340,000 daltons, suggesting it may be a macroglobulin-like antibody. Homologous agglutinin reacted better with A. hydrophila than heterologous agglutinin. Differences in severity and duration of red-sore epizootics in the southeastern United States may be due to differing virulence among strains of A. hydrophila.

Field Experiments on Responses of a Freshwater, Benthic Macroinvertebrate Community to Vertebrate Predators

We examined the seasonal importance of vertebrate predators in potentially regulating the abundance and diversity of the benthic macroinvertebrates in the littoral zone of a soft—bottom reservoir that receives thermal effluent from a nuclear production reactor. Thirty—six predator (fish and turtle) exclusion cages (4 m2) were placed in shallow water at six locations along a thermal gradient in Par Pond, a 1100—ha cooling reservoir on the Savannah River Plant near Aiken, South Carolina, USA. An additional 36 control plots (4 m2) were also set up. Cages were in place during three, 3—mo test periods beginning in September 1977. Estimates of benthic density, taxon richness, and distribution within functional groups (defined by feeding mechanism) were calculated for each test period. Effects of temperature on predator—prey relationships were also determined. Experimental results of this study suggest that vertebrate predation was not the fundamental parameter organizing the benthic macrionvertebrate community in the littoral zone of this reservoir. Neither taxon richness nor density of total macroinvertebrates was conclusively related to predator treatment. Relationships between predator treatment and community response (changes in density and taxon richness) were generally unaffected by either plot locality, temperature fluctuations from thermal effluent, or seasonal changes. When data from caged and control plots were pooled, however, both location and water temperature individually had direct impacts on the benthic community. From our results and other field studies we hypothesize that individual species of "keystone" benthic predators (Paine 1969b) do not occur in the littoral zone of freshwater lentic environments with soft bottoms. This hypothesis is based on the following four arguments. First, environmental heterogeneity should reduce predator efficiency relative to that found in rocky intertidal systems where keystone predation may be present. Second, relative food web complexity should reduce the community importance of an individual predator species. Third, the apparent relatively greater resource partitioning of space and food in the freshwater littoral zone hinders monopolization of rate—limiting environmental resources by a single dominant prey species. Fourth, the effects of predation may not be extensive enough in time or area to provide sufficient resource space (and thus food) for exploitation by fugitive species. Possible regulation by a guild of predators and cases where the general hypothesis might be falsified are discussed.

Stress and Body Condition in a Population of Largemouth Bass: Implications for Red-Sore Disease

Abstract The body conditions, K = 105(weight, g) ÷ (standard length)3, and various hematological characters were examined for largemouth bass (Micropterus salmoides) taken from Par Pond, a reservoir heated by effluent from a nuclear production reactor at the Savannah River Plant near Aiken, South Carolina. Largemouth bass with K less than 2.0 had significantly lower (P < 0.05) hematocrits, hemoglobin concentrations, total red blood cell counts, total white blood cell counts, and lymphocyte fractions, and significantly higher granulocyte fractions and cortisol concentrations, than those with K greater than 2.0; monocyte, thrombocyte, and reticulocyte fractions were not different between the two K-factor groupings. When data were pooled, all blood variables except the reticulocyte fraction were significantly correlated with K. Hematocrit, the lymphocyte fraction, and cortisol concentration account for 20.5% of the variation in K. These data support a previous hypothesis that elevated water temperature promo...

Sun Compass Orientation of Immature Bluegill

Abstract Immature bluegills, Lepomis macrochirus (N = 389), were collected from Par Pond, a 1,120-hectare impoundment on the United States Department of Energy's Savannah River Plant near Aiken, South Carolina, and tested individually by observing their directional movement from the center of a circular testing facility. Bluegills captured 10–20 m from shore in areas lacking extensive submerged vegetation swam in a direction which would have taken them to shore at the area of their capture, whereas those taken from areas having submerged vegetation offshore swam in the direction opposite from shore. The directional responses of fish that were phase-shifted by 6 hours demonstrated that they were using time-compensated sun orientation. Fish exposed to a predator oriented in the direction of shore in their training facility, but if no predators had been present during training, moved in the direction of deep water. Present evidence indicates that immature bluegills use their sun compass mechanism in oriented...

Ecology ofAeromonas hydrophila in a South Carolina cooling reservoir

Densities ofAeromonas hydrophila were determined monthly from December 1975 to December 1977 in a South Carolina cooling reservoir which receives heated effluent from a single nuclear production reactor. Selected water quality parameters and prevalence of red-sore disease among largemouth bass were monitored simultaneously.Higher densities ofA. hydrophila were observed in areas of the reservoir receiving effluent from the reactor. Densities ofA. hydrophila generally were heterogeneous in the water column. The sediments had lower densities ofA. hydrophila than water immediately above.A. hydrophila could not be isolated from sediments greater than 1 cm from the water interface. Temperature, redox potential, pH, and conductivity were all significantly correlated with densities ofA. hydrophila in the water column. The temporal and spatial distribution and abundance ofA. hydrophila in water were not related to total organic carbon, dissolved organic carbon, particulate organic carbon, inorganic carbon, or dissolved oxygen. High densities ofA. hydrophila were observed in mats of decomposingMyriophyllum spicatum and, enterically, in largemouth bass, several other species of fish, turtles, alligators, and snails. The greatest densities ofA. hydrophila in water occurred during March and June with a second peak in October. The mean monthly densities ofA. hydrophila were positively correlated with the incidence of infection in largemouth bass. Largemouth bass from thermally altered parts of the reservoir had a significantly higher incidence of infection. It is concluded that thermal effluent significantly affects the ecology ofA. hydrophila and the epizootiology of red-sore disease within Par Pond.

BIOCHEMICAL AND MORPHOLOGICAL EFFECTS OF TEMPERATURE ON TYPHA LATIFOLIA L. (TYPHACEAE) ORIGINATING FROM DIFFERENT ENDS OF A THERMAL GRADIENT. I. CONTROLLED ENVIRONMENTAL STUDIES

T. latifolia grows along most of the shoreline of Par Pond, a cooling reservoir for the reactors at the Savannah River Plant in Aiken, South Carolina. A thermal gradient exists because of the movement of hot water from one end of the pond to the other. This study was undertaken to determine the biochemical and morphological effects of temperature on Typha latifolia clones originating from different ends of this thermal gradient. The data indicated that both “hot” and “cold” end plants react similarly to temperature. Forty‐eight rhizome plants from both ends of Par Pond were transplanted into tanks at 20 C and 30 C. Total levels of malic dehydrogenase activity, proportions of mitochondrially bound MDH and growth rates were significantly different in plants grown in the two tanks, but no significant differences were observed within each tank between individuals from the “hot” and “cold” end. The relation of these results to hypotheses concerning quantitative strategies of enzyme adaptation are discussed in the paper.

The freshwater littoral meiofauna in a South Carolina reservoir receiving thermal effluents*

SUMMARY. The freshwater littoral meiofauna along a temperature gradient in Par Pond (a cooling reservoir receiving thermal effluents from a nuclear reactor) was sampled from September 1975 to October 1976. Monthly samples were taken at three stations; ‘hot‐water’ (15–40°C), ‘warm‐water’ (13–37°C) and ambient or ‘cold‐water’ (8–37°C) sites. Total numbers of individuals at the ambient site ranged from 971 to 3674 per 10 cm2 (mean = 2263), approximating the density reported from productive estuarine environments. Nematodes, rotifers, ostracods, cladocerans and mites comprised 80% of overall density. When compared to the ambient site, thermally affected sites demonstrated reduced faunal density.Contrary to other environmental perturbation studies, the Shannon‐Weaver diversity index (H') did not reflect alteration of structural complexity within the rotifer taxocene when hot‐water, warm‐water and cold‐water sites were compared. Though there was a significant reduction in number of species at the thermally altered sites, high ‘equitability’ among the reduced species resulted in H’ values comparable to those in natural communities. Distinct winter‐spring and summer‐autumn assemblages were evident in the ambient community while no seasonal clustering was apparent at the thermally affected sites. It is hypothesized that the maintenance of temperatures above ambient normals masks environmental cues that normally would elicit seasonal changes in the rotifer fauna at those two sites.