Plankton communities and summertime declines in algal abundance associated with low dissolved oxygen in the Tualatin River, Oregon
Phytoplankton populations in the Tualatin River in northwestern Oregon are an important component of the dissolved oxygen (DO) budget of the river and are critical for maintaining DO levels in summer. During the low-flow summer period, sufficient nutrients and a long residence time typically combine with ample sunshine and warm water to fuel blooms of cryptophyte algae, diatoms, green and blue-green algae in the low-gradient, slow-moving reservoir reach of the lower river. Algae in the Tualatin River generally drift with the water rather than attach to the river bottom as a result of moderate water depths, slightly elevated turbidity caused by suspended colloidal material, and dominance of silty substrates. Growth of algae occurs as if on a “conveyor belt” of streamflow, a dynamic system that is continually refreshed with inflowing water. Transit through the system can take as long as 2 weeks during the summer low-flow period. Photosynthetic production of DO during algal blooms is important in offsetting oxygen consumption at the sediment-water interface caused by the decomposition of organic matter from primarily terrestrial sources, and the absence of photosynthesis can lead to low DO concentrations that can harm aquatic life. The periods with the lowest DO concentrations in recent years (since 2003) typically occur in August following a decline in algal abundance and activity, when DO concentrations often decrease to less than State standards for extended periods (nearly 80 days). Since 2003, algal populations have tended to be smaller and algal blooms have terminated earlier compared to conditions in the 1990s, leading to more frequent declines in DO to levels that do not meet State standards. This study was developed to document the current abundance and species composition of phytoplankton in the Tualatin River, identify the possible causes of the general decline in algae, and evaluate hypotheses to explain why algal blooms diminish in midsummer. Plankton and water-quality sample data from 2006 to 2008 were combined with parts of a larger discrete-sample and continuous water-quality monitoring dataset and examined to identify patterns in water-quality and algal conditions since 1991, with a particular emphasis on 2003–08. Longitudinal plankton surveys were conducted in 2006–08 at six sites between river miles (RM) 24.5 and 3.4 at 2- to 3-week intervals, or 5–6 per season, and in-situ bioassay experiments were conducted in 2008 to examine the potential effects of wastewater treatment facility (WWTF) effluent and phosphorus additions on phytoplankton biomass and algal photosynthesis. Phytoplankton and zooplankton community composition, streamflow, and water-quality data were analyzed using multivariate statistical techniques to gain insights into plankton dynamics to determine what factors might be most tied to the abundance and characteristics of the phytoplankton assemblages, and identify possible causes of their declines. The connection between low-DO events and algal declines was clearly evident, as bloom crashes were nearly always followed by periods of low DO. Algal blooms occurred each year during 2006–08, producing maximum chlorophyll-a (Chl-a) values in June or July generally in the range of 50–80 micrograms per liter (µg/L). Bloom crashes and absence of sufficient algal photosynthesis in mid- to late-summer contributed to minimum DO concentrations that were less than the State standard of 6.5 milligrams per liter (mg/L) based on the 30-day mean daily concentration, for 62–74 days each year. At times, the absolute minimum State standard (4 mg/L DO) also was not met. To learn more about why low-DO events occurred, specific algal declines during 2003–08 were scrutinized to determine their likely causal factors. From this information, a series of hypotheses were formulated and evaluated in terms of their ability to explain recent declines in algal populations in the river in late summer. Meteorological, streamflow, turbidity, water temperature, and conductance conditions in the Tualatin River during the 2006–08 summer seasons were not atypical. Natural flow comprised the majority (70–80 percent) of flow each year during spring, but then reduced to 38–40 percent during midsummer when WWTF effluent—which contributed as much as 36 percent—and flow augmentation releases comprised a greater fraction of the flow. Summer 2008 was unusual, however, in the prolonged influence from the Wapato Lake agricultural area near Gaston in the upper part of the basin. The previous winter flooding and levee breach at Wapato Lake caused a much greater area of inundation. As a result, drainage from this area continued into July, much later than normal. A subsequent algal bloom in Wapato Lake then seeded the upper Tualatin River, and this drainage had a profound effect on the downstream plankton community. A large blue-green algae bloom developed—the largest in recent memory—prompting a public health advisory for recreational contact for about two weeks. Algal growths and surface blooms are a common feature of the Tualatin River. Most of the dominant algae have growth forms and morphologies that are well suited for planktonic life, employing spines and gas vacuoles to resist settling, forming colonies, and producing mucilage (or toxins) to resist zooplankton grazing. In 2006–08, 143 algal taxa were identified in 117 main-stem samples; diatoms and green algae were more diverse than blue-green, golden, and cryptophyte algae, although these later groups sometimes dominated the overall volumetric abundance (biovolume). The most frequently occurring taxa, occurring in 97–99 percent of samples, were flagellated cryptophytes Cryptomonas erosa and Rhodomonas minuta. Other important algal taxa included centric diatoms Stephanodiscus, Cyclotella, and Melosira species and colonial green algae Scenedesmus and Actinastrum. These taxa comprised the majority of the algal biovolume during much of the growing season. A general seasonal trend in the phytoplankton assemblages was observed, with dominance by filamentous centric diatoms Stephanodiscus and Melosira in spring and early summer, and flagellated cryptophytes and green algae, particularly Chlamydomonas sp., in late-summer; or, in 2008, dominance by blue-green algae Anabaena flos-aquae and Aphanizomenon flos-aquae during the Wapato Lake bloom event. There were 99 zooplankton taxa identified from the Tualatin River in 2006–08, composed primarily of cladocerans, copepods, and rotifers. A seasonal increase in zooplankton abundance was observed in early summer just as or shortly after the phytoplankton population began to increase, with populations growing to 15,000−120,000 organisms per cubic meter in the lower river. Zooplankton abundance showed a predictable and distinct longitudinal downstream increase, particularly downstream of Highway 99W (RM 11.6). Although grazing rates were not measured, the data suggest that, at times, zooplankton grazing may affect algal abundance and species composition in the Tualatin River, with diatoms becoming relatively less abundant and flagellated cryptophytes and green algae relatively more abundant during periods when zooplankton densities were highest. Multivariate statistical analyses identified soluble reactive phosphorus (SRP), natural flow, flow augmentation, and WWTF effluent as important factors influencing Tualatin River phytoplankton populations, with zooplankton density (particularly rotifers and copepods), specific conductance, chloride, and water temperature also having an important influence. Although SRP was highly correlated with the plankton communities, that correlation was likely the result of high or low algal activity (uptake) as SRP concentrations were often reduced to low levels during blooms. While previous studies have already established that phosphorus, among other factors such as flow, places a theoretical cap on the size of the phytoplankton population in the river, sometimes algal declines occur when SRP concentrations are apparently sufficient. To identify alternative causal factors, additional analyses were performed without SRP to focus on other water-quality parameters, zooplankton density, and flow factors. Considering data for all 3 years and including just those samples from the lower Tualatin River not affected by the 2008 Wapato Lake drainage event, three factors (percentage of reservoir flow augmentation, total natural flow, and rotifer density) best explained variations in the phytoplankton assemblages. Analyses focusing on the possible causes of algal declines included the above multivariate analyses, scrutiny of 10 specific instances of declines in algal populations during 2003–08 including several bloom–crash sequences, and analyses of historic routine watershed monitoring data from Clean Water Services. Six factors were hypothesized to be important in causing bloom crashes or impeding blooms from rebounding in August: (1) light limitation from cloudy weather, (2) a reduction in the plankton inocula or “seed” entering the lower river from upstream sources, (3) increased summer streamflows, (4) changes in the dominant sources of flow as the percentage of flow augmentation and WWTF discharges have increased, (5) zooplankton grazing, and (6) low concentrations of bioavailable phosphorus (<0.015 milligram per liter). All of these hypotheses are supported in some fashion by the available data and statistical analyses. Zooplankton grazing, short-term declines in photosynthesis from cloudy weather, total flow as it affects residence time, and the dominant source of flow are primary factors responsible for the low-DO events caused by declines in algae in the lower Tualatin River during late summer. Cloudy weather and increased turbidity are known to inhibit algal growth in the Tualatin River, and slight increases in turbidity in recent years may be a problem. Upstream sources of algae are critical in determining the characteristics and size of downstream populations, as illustrated by the Wapato Lake bloom in 2008, but more data are needed from upstream to fully define the importance of this connection. The sources of flow, through their differential contribution of plankton inocula (quality and amount), were, at times, important factors affecting phytoplankton populations. While SRP concentrations were often most highly correlated with phytoplankton species community, the bioavailability of phosphorus is still somewhat unknown and there are several sources to consider. Preliminary bioassay tests suggested that while treated wastewater effluent may stimulate algae at 30 percent concentrations, negative effects (or decreased stimulation) on Chl-a and DO production may occur at concentrations of 50 percent. Targeted data collection and future research will be needed to further understand the importance of these factors on Tualatin River phytoplankton. While the data and analysis completed for this report provide insights into future research and monitoring that would be useful to continue, additional monitoring of turbidity, Chl-a, and plankton abundance and species composition in the upper part of the basin would enhance our understanding of plankton dynamics and factors affecting phytoplankton abundance in the lower river. Assessment of the key upstream sources of algal inocula via surveys of the major flow sources as well as tributaries and wetlands would provide useful information for the management of river water quality. Other studies that could prove useful for developing management strategies include targeted experiments to evaluate the bioavailability of phosphorus from a variety of sources. New research on phytoplankton–zooplankton interactions, and studies of planktivorous fish, might also provide insight about food web dynamics and potential “top-down” effects of fish predation on the plankton communities. In addition, further development of neural-network or other water-quality models would help to evaluate management strategies and provide forecasts of water-quality conditions. Finally, periodic future reassessments of the available data with the multivariate statistical tools used in this study would be helpful to assess whether and how plankton communities are changing, and to continue to shed light on the importance of factors shaping the plankton. Although certain types and sizes of algal blooms are undesirable, minimum phytoplankton populations are an important part of aquatic food webs and are needed to maintain healthy levels of DO in the river. By understanding the sources, characteristics, causal factors, and responses of the plankton communities, management strategies can be developed to improve DO conditions in the lower Tualatin River during the important summer low-flow period.