Study Location:
The Coeur d’Alene Basin, USA is centrally located in the northern Idaho panhandle and encompasses several sub-watersheds including the Upper Coeur d’Alene River, South Fork Coeur d’Alene River, Coeur d’Alene Lake, and the Lower St. Joe River (USGS, 2013). Collectively, these sub-watersheds drain an area 5,225 km2 in size (USGS, 2013). The drainage basin extends west from the Idaho-Montana border before reaching the Spokane River, which drains the 129 square km Coeur d’Alene Lake (Woods & Beckwith, 1997).
We collected data at Thompson Lake, Black Lake, and Chatcolet Lake. Thompson and Black lakes are eutrophic based on water column characteristics. Both lakes are in the floodplain of the metals contaminated Coeur d’Alene River (Bookstrom et al., 2013) and are connected to the river by dredged channels and a vast wetland complex. Thompson and Black lakes have maximum depths of ~7 m and ~6 m, respectively, and are often thermally stratified between late May and early October. Seasonal pumping of adjacent converted wetlands into Black Lake has increased nutrient concentrations and decreased water quality through eutrophication (Kann & Falter, 1987). Chatcolet Lake is located in the southern half of Coeur d’Alene Lake and was chosen as a non-metals contaminated reference site. It has a maximum depth of ~13 m, is mesotrophic, and thermally stratified from June to early October (CDA Tribe & Avista Corporation, 2017).
Sampling Regime and Experimental Design:
This study examined the phenology of aquatic macrophyte communities from May 1, 2017 to November 15, 2017. Phenology was assessed using submerged macrophyte biomass, biovolume generated from hydroacoustic sonar surveys, and water quality measurements collected twice per month. Survey sites at Black, Chatcolet, and Thompson lakes were 3.3, 2.8, and 1.6 ha in size, respectively. Macrophyte biomass samples were collected from a 50 m point intercept grid (Madsen & Wersal, 2017) overlain on the sample sites. Grid points were located in the field using a Lowrance Point-1 GPS with 3 m horizontal accuracy. At each point, a biomass sample was collected using the rake twirl method (Johnson & Newman, 2011). Species present, dominant species, and water depth were noted before placing the sample in a plastic bag for laboratory analysis. Depth of water was determined by 0.25 m increments marked on the exterior of the rake handle. A total of 406 biomass samples were collected.
Macrophyte biovolume has been measured with sonar previously (Duarte, 1987; Radomski & Holbrook, 2015; Valley & Drake, 2005) and in this study, we used a consumer- grade sonar system. Sonar data was recorded using a Lowrance HDS-7 Gen 2 with a transom mounted 200 kHz transducer with a 20° beam angle using the BioBase configuration as recommended by Navico (2014). Sonar data was collected along transects perpendicular to shore and traversed the biomass grids to achieve ~25 m spacing between sonar paths. After collection, the sonar data was uploaded to the BioBase website for processing. Biovolume, or the percent of water column occupied by plant biomass, was calculated by dividing plant height by water column depth and was estimated across a five-meter grid spanning the sampling area (Navico, 2014). After processing by BioBase, an ASCII text file was downloaded with latitude, longitude, and biovolume estimates for each sampling event. To minimize false detections, biovolume was not estimated at sonar depths < 0.73 m and when plant detections had lengths < 5% of the water column depth (Valley et al., 2015). Sonar was recorded within a day of when biomass samples were collected at each site.
Because biomass and biovolume measurements could not be taken simultaneously, an assessment of positional sampling error was made. Sonar transects were similar between sampling events, but not identical. Overall mean GPS accuracy when navigating to biomass sampling points was 5.2 ± 3.1 m (mean ± standard deviation), which was approximately 2 m greater than the manufacturer specified accuracy of the GPS receiver. Overall mean matchup distances from pairwise biomass points and biovolume estimates was 1.3 ± 0.5 m. Thus, we feel confident that the biovolume and biomass estimates are representative for each sampling site.
Water Quality and Degree Days:
Vertical profiles of water temperature, pH, dissolved oxygen, specific conductance, and photosynthetic active radiation (PAR) at 0.5 m increments were measured and recorded with a Hach Hydrolab DS5 multi-sensor sonde. A standard 20 cm diam. Secchi disk was used to assess water transparency. Water temperatures in each lake were also recorded at hourly intervals with HOBO Water Temp Pro v2 sensors deployed in approximately 1.0 m of water. These temperature data were used to calculate cumulative growing degree days (GDD), which represent an accumulation of "heat units" (Zalom & Goodell,1983). Heat units are accumulated when the mean of the minimum and maximum daily temperature fall between the lower and upper bounds of an acceptable temperature for growth of a specific organism (Zalom & Goodell, 1983). For this study, GDD values were calculated with the “pollen” package in R (Nowosad, 2018) using minimum (15 °C) and maximum (35 °C) growth thresholds for Myriophyllum spicatum as reported by Smith and Barko (1990).
Analysis of Macrophyte Biomass:
Macrophytes collected for the analysis of biomass were processed following method 10400 D Population Estimates as described by APHA et al. (2012). Macrophytes were first rinsed to remove all sediment and then placed in a salad spinner to remove excess water. Samples were then sorted by species using the keys of Hamel et al. (2001) and Crow and Hellquist (2005a, 2005b). Please note that Elodea nuttallii is occasionally found in Coeur d’Alene Lake, however, we denote all Elodea spp. as Elodea canadensis (Lamb, 2006). Similarly, we have referred to all Myriophyllum spp. as M. spicatum. However, most Myriophyllum spp. encountered in Chatcolet Lake is hybrid watermilfoil comprised of a cross between M. spicatum and M. sibiricum (Thum, 2016). Black and Thompson lakes had mostly M. spicatum, but some hybrid watermilfoil was also present (Thum, 2016).
Once macrophytes were sorted by species, wet mass was determined to the nearest hundredth of a gram with an electronic balance. Sorted plants were then air dried for 4-6 hours. Following this initial drying, samples were transferred to a 105 °C drying oven (Lindberg/Blue M G01350A) and dried until a constant mass was reached. Dried samples were allowed to cool in a desiccator for 15 minutes before the final dry mass was determined.
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