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The Ecology of Signal Crayfish in Two Large Ultra-Oligotrophic Ecosystems: Crater Lake and Lake Tahoe
AuthorUmek, John William
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Invasive species have become an increasing problem in the Western United States particularly when there are multiple stressors (e.g., invasive species and eutrophication) occurring to ecosystems. Invasive omnivores can present unique problems for aquatic ecosystems by having both direct and indirect impacts on native benthic invertebrates and vertebrates. Omnivorous crayfish, for example, strongly influence littoral habitats and biota with their foraging habits, creating both direct and indirect effects on trophic interactions in aquatic systems. Once they invade, these crayfish can ultimately dominate freshwater ecosystems. This dissertation investigates the distribution, density changes, and the direct and indirect impacts of the invasive signal crayfish (Pacifastacus leniusculus) in two oligotrophic lentic ecosystems in the western United States; Lake Tahoe (CA-NV) and Crater Lake (OR). In chapter 1, I investigate the distribution, movement, and feeding behavior of invasive signal crayfish in Crater Lake. This lake population presents a unique opportunity to understand the movement of crayfish in a recently expanding population. I used minnow traps and snorkeling to determine crayfish distribution and stable isotope ratios of δ13C and δ15N to determine the flow of organic matter through the food web, trophic position, and percent benthic reliance. Depth gradient minnow traps demonstrate that crayfish densities can live as deep as 250 m. Trap and snorkel surveys from 2008 to 2013 indicate an expansion of crayfish from 44% to 78% of the littoral zone. Summer water temperature in Crater Lake has been warming, which may increase the recruitment of individuals and expand habitat availability for growth. Between 1965 and 2014 the nearshore surface temperature increased by 3.5°C. Principal component analysis revealed a positive relationship between crayfish occupation and cobble and boulder habitats of the lake. Crayfish in the littoral zone rely heavily (97.4%) on littoral-benthic carbon sources indicating their potential for impacting native invertebrate communities and the overall dynamics of Crater Lake’s ecosystem. Our findings indicate, however, that deeper water crayfish also may rely on littoral benthic energy resources. Crayfish movement to deeper waters may be subsidizing generally nutrient poor, deep-water habitats with littoral energy through excretion and egestion, where physical conditions are stable and natural perturbation is low.In Chapter 2, I quantify the influence of this early, expanding invasion in Crater Lake to littoral zone ecology by evaluating their influence on zoobenthic consumer biomass and basal algal biomass. Benthic invertebrate biomass was 77% lower in hard substrate and 78% lower in soft substrate areas with crayfish present than in crayfish-absent locations. Using Bayesian, stable isotope mixing models, dietary preferences of crayfish at three locations with varying crayfish densities were quantified. Only slight variations in crayfish diet were detected between the three locations where crayfish have been established, the outer boundary of crayfish expansion, and the middle of the crayfish population indicating that crayfish. Despite differing densities, crayfish are feeding on similar food sources, particularly benthic invertebrates. At low crayfish densities (0 to 10), benthic invertebrate numbers were 222.3±36.6 individuals m-2, while chlorophyll a was 16.8±5.8 mg m-2. At high densities of crayfish (>50), benthic invertebrates had low mean density 3.0±4.2 individuals m-2, while chlorophyll a biomass was high 226.7±48.1 mg m-2. Crayfish are impacting native invertebrate communities and periphyton biomass in Crater Lake by changing trophic interactions in the lake’s littoral zone and altering the lake’s food web.In Chapter 3, I focus on the benthic environment and biodiversity of Lake Tahoe and regional lakes (Donner Lake, Marlette Lake, and Fallen Leaf Lake. Signal crayfish were introduced into the Central Sierra Nevada region of the United States in the late 19th to early 20th century. I used a long-term data set to document highly variable crayfish densities in the littoral zone of Lake Tahoe, showing an increase during the summer months linked to an increase in water temperature (R2 = 0.69, P<0.001). Crayfish responded to site-specific characteristics of the nearshore rather than to lake-wide characteristics; local stream discharge was the only factor that explained a positive increase in lake densities (P< 0.04). Trophic niche models developed from stable isotope measurements of crayfish and nongame fish indicate that crayfish influence the dietary breadth (e.g. niche area) of nongame fish consumers. Crayfish feeding behavior may be forcing nongame fish to feed on a broader set of food resources when crayfish are present. Stable isotope analysis also indicates an overlap of crayfish niche area with other nongame fish and amphibians, indicating interspecific competition between organisms. Our study highlights that local factors influence cold-water crayfish movement and densities in large lakes, as well as potential direct and indirect influences on nongame fish consumers in the littoral region, potentially affecting native biota and ecosystem function. This research has significant implications for understanding the direct and indirect impacts of signal crayfish in oligotrophic food webs, particularly on benthic invertebrate densities. It expands on the current understanding of expansion of signal crayfish and the factors that influence crayfish density. Future research will need to focus on better understanding the life history and mechanisms controlling this species if they are to be controlled in lakes of the Western United States.