Multiple sampling trips during calendar years 2013 through 2015 were coordinated to provide measurements of interdependent benthic processes that potentially affect contaminant transport in Upper Klamath Lake (UKL), Oregon. The measurements were motivated by recognition that such internal processes (for example, solute benthic flux, bioturbation and solute efflux by benthic invertebrates, and physical groundwater-surface water interactions) were not integrated into existing management models for UKL. Up until 2013, all of the benthic-flux studies generally had been limited spatially to a number of sites in the northern part of UKL and limited temporally to 2–3 samplings per year. All of the benthic invertebrate studies also had been limited to the northern part of the lake; however, intensive temporal (weekly) studies had previously been completed independent of benthic-flux studies. Therefore, knowledge of both the spatial and temporal variability in benthic flux and benthic invertebrate distributions for the entire lake was lacking. To address these limitations, we completed a lakewide spatial study during 2013 and a coordinated temporal study with weekly sampling of benthic flux and benthic invertebrates during 2014. Field design of the spatially focused study in 2013 involved 21 sites sampled three times as the summer cyanobacterial bloom developed (that is, May 23, June 13, and July 3, 2013). Results of the 27-week, temporally focused study of one site in 2014 were summarized and partitioned into three periods (referred to herein as pre-bloom, bloom and post-bloom periods), each period involving 9 weeks of profiler deployments, water column and benthic sampling. Partitioning of the pre-bloom, bloom, and post-bloom periods were based on water-column chlorophyll concentrations and involved the following date intervals, respectively: April 15 through June 10, June 17 through August 13, and August 20 through October 16, 2014. 

To examine dissolved-solute (0.2-micrometer [μm] filtered) benthic flux, sets of nonmetallic pore-water profilers (U.S. Patent 8,051,727 B1) were deployed. In 2013, the deployment of profilers at 21 UKL sites occurred at the beginning of the annual cyanobacterial bloom of Aphanizomenon flos–aquae (AFA), in the middle of the bloom period, and at the peak of the bloom. Coordinated benthic invertebrate collections also were made. Based on results from 2013, weekly deployments of profilers and collection of benthic invertebrate samples from late spring to early autumn were used to estimate temporal trends in solute flux and benthic invertebrate densities. Estimates of nutrient efflux by benthic invertebrates were determined in the spring and autumn from 2011 through 2013 and three times (spring, summer, and autumn) in 2015. This work extends UKL studies that began in 2006 to quantify the importance of benthic solute sources in the lake. In 2015, piezometers and thermistor sets were deployed to quantify potential groundwater exchange with the lake water column. 

Analysis of the 2013 soluble reactive phosphorus (SRP) benthic flux indicated no effect of location (lake region), habitat, or sampling period, and the average lakewide flux values were consistent with earlier studies that had been confined to the northern region of UKL and adjacent wetlands. The 2014 study therefore focused on estimating temporal trends at a site within Ball Bay. During both 2013 and 2014 field studies, fluxes of macronutrients (soluble reactive phosphorus (SRP) and ammonia) and micronutrients (iron [Fe] and manganese [Mn]) were consistently positive and increased prior to the initial AFA bloom, varied or lagged with water-column chlorophyll during the summer bloom period, then decreased after the cyanobacterial blooms, only to rebound toward pre-bloom conditions in the final weeks of sampling. These four solutes exhibited benthic loads greater than maximum riverine loads estimated during the spring and early summers of 2013 and 2014. However, consistently detectable concentrations for all four solutes provide no evidence that they consistently serve as the limiting nutrient for primary production in the lake. In contrast to the four solutes (SRP, ammonia, Fe, and Mn), benthic fluxes of dissolved arsenic (As) were both negative and positive (that is, the lakebed currently serves as both a source and a sink for dissolved As, depending on season). In a further contrast with SRP, ammonia, dissolved Fe, and Mn, dissolved-As riverine loads to UKL were of similar magnitude to benthic loads. A negative relationship between dissolved-As flux and water-column As over the 2014 temporal study provides a potential advantage for the management of water-quality in contrast to solutes, like SRP or ammonia, with consistently positive flux. 

The mean total benthic invertebrate density during 2013 was 12,610 individuals per square meter (n=63). Although benthic invertebrate density did not change over the study period, it was higher in littoral habitats than open-lake or trench habitats and higher in the northern region compared to the central or southern regions of UKL. Mean total benthic invertebrate density during 2014 was 19,726 individuals m−2 (n=27). Density during the pre-bloom and bloom periods of April 15 to August 13, 2014 (the first two thirds of the 2014 sampling period), were similar to 2013. However, benthic invertebrate density more than doubled during the latter one-third of the study, that is, the post-bloom period between August 20 to October 16, 2014. Oligochaeta, Chironomidae and Hirudinea represented well over 90 percent of the benthic fauna; Oligochaeta were twice as abundant as Chironomidae or Hirudinea, the latter two of which were similar in density. 

Benthic invertebrates may enhance dissolved-nutrient (or toxicant) transport across the sediment-water interface by (1) modifying diffusion-layer thicknesses and permeability through bioturbation, (2) enhancing advective flow across the interface through bioirrigation, and (3) excreting or expelling dissolved or particulate solutes directly into the overlying water column (Boudreau and Jorgensen, 2001). We evaluated SRP efflux via excretion for approximately 15 different major taxa in UKL. Once these measures were scaled, it was evident that benthic invertebrates potentially contribute approximately 1.5 times the amount of SRP to the water column of Upper Klamath Lake as diffusive SRP flux alone, measured in profiler deployments. 

Sets of piezometers and temperature loggers were deployed in UKL to obtain estimates of vertical advective solute flux. The pressure transducer installations, within the piezometers, did not perform as designed, rendering the head gradient data unreliable. However, in terms of future research, this field work did demonstrate the feasibility of collecting vertical gradient data with piezometer deployments. Advective flux estimates herein are based solely on heat-flow modeling based on temperature data from four lake sites, without use of transducer data. Given the magnitudes (both positive or negative) of the heat-transfer fluxes for SRP, relative to diffusive-flux and macroinvertebrate efflux measurements (all positive but spanning the same orders of magnitude), further examination of solute advective flux is recommended as a potential transport process to integrate into existing water-quality (for example, Total Maximum Daily Load [TMDL]) models. 

As a complement to the biogeochemical focus of this study, initial analyses of suspended-particle (floc) characteristics and settling velocities from the water column were derived near the surface and lakebed at two UKL sites. To better understand changing particle characteristics during the AFA-bloom period, suspended particles were examined in 2015 using a LabSFLOC (LF), which is a Laboratory Spectral Flocculation Characteristics version of an In-Situ Settling Velocity instrument (INSSEV-LF). Particle characteristics and settling velocities were analyzed from the water column near the surface (sample dp_10) and lakebed (sample dp_90) at two lake sites (open-lake site ML and littoral site LS01). The term “floc” refers herein to suspended particles that may aggregate or disaggregate to change in size, composition, and settling velocity. 

During pre-bloom (May) conditions, where maximum suspended particulate matter concentration (SPMC) was 140 milligrams per liter (mg L⁻¹) was now observed at site LS01 in close proximity to the bed, where D_mean peaked at 305 μm, and the corresponding Ws_mean was 3.9 millimeters per second (mm s⁻¹). The high near-bed SPMC (828 mg L⁻¹) experienced during post-bloom October 2015 at LS01 formed a benthic nepheloid layer (BNL) above the lake’s bed. Numerous low density, fast settling macrofloc-sized organic aggregates (D >160 μm) were observed (some up to 1 mm in size) near bed at LS01 both during the bloom and post-bloom conditions; many of these flocs displayed fibrous organic structures. In terms of mass settling fluxes, the post-bloom BNL produced a total MSF of 4,139 milligrams per square meter per second (mg m⁻² s⁻¹) (92.1 percent of MSF credited to the macrofloc-sized organic aggregates/cyanobacterial colonies); that was nearly three times the corresponding near-bed settling flux observed during the July 2015 bloom and 360 times greater than the pre-bloom conditions from May 2015 (98.8 percent and 14 percent of MSF credited to the macrofloc-sized fractions for those respective months). Such changes in the near-bed settling flux demonstrate the highly significant seasonal effects that the AFA bloom has on the floc depositional fluxes in UKL and highlights the importance of seasonal monitoring of these conditions in order to correctly parameterize the wide range in depositional characteristics and floc properties measured throughout UKL. 

Collectively, floc populations observed within UKL demonstrated a wide range in settling velocity (Ws) for a given particle size, D. Similarly, a given settling velocity was not associated with a specific particle size. This variability in particle characteristics and properties indicates the influence of varying floc effective density and its effect on mass and mass settling fluxes (MSF). The use of instruments, such as the INSSEV-LF, enables measuring the variability of settling velocity and its relation to particle density and size.