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Stockwell, J. D.

Publications and source records attributed to Stockwell, J. D..

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Comparing 3-D visual and 2-D tactile encounter rates in benthic and pelagic habitats

Encounter rate models are important tools for evaluating and estimating trophic interactions between species. While encounter rate parameters have been measured for many freshwater pelagic fishes, most benthic fishes remain mostly unstudied. Those few efforts to generate encounter rate models for benthic fishes often hold mathematical assumptions based on visual foraging, despite the many cases in which benthic fishes employ the lateral line to forage. Furthermore, encounter rate models are rarely compared, despite the many cases in which prey animals face predation risk from multiple types of predators. For example, the macroinvertebrate Mysis is exposed to both benthic and pelagic predation risk during diel vertical migration (DVM). Comparing the risks between habitats could help evaluate predation risk as an ultimate cause of their DVM behavior. We created a novel encounter rate model based on lateral line ("tactile") foraging by sculpins (Cottidae) given the saltatory (stop-and-go) nature of their movement. The tactile model demonstrated variation in behavior and peak encounter rate with detection distance, movement velocity, and rest durations. We then directly compared predation risk for Mysis by parameterizing both our tactile benthic (2D) encounter rate model for sculpin and a visual pelagic (3D) for rainbow smelt (Osmerus mordax). Tactile encounter rates were generally lower than visual rates for individual predators. However, population level encounter rates at night were greater in the benthic habitat than the pelagic habitat. Overall, our model estimates of encounter rates were consistent with the long-standing hypothesis that predation is an ultimate driver of DVM behavior.

ecology↗

The role of edible habitat complexity in food webs

Habitat complexity (HC) in part determines the diversity, stability, and behavior of food webs and can influence predation according to a wide variety of functional relationships. Many aquatic species provide habitat complexity and are also consumed by other species (e.g., macrophytes, corals, mussels). However, food web theory does not readily account for these species that act as edible habitat complexity (EHC). Here, we combine existing theory on predator-prey interactions, HC, and prey switching to describe the role of EHC in benthic food web models. We dissect feedback loops in each model to demonstrate how self-regulation of the prey species, mediated by species densities and HC, drives that food webs behavior. HC can stabilize predator-prey interactions by coupling prey self-regulation with HC self-regulation. EHC can further stabilize predator-prey interactions across a wide variety of "HC functions" that relate HC to predation rates. Significance StatementHabitat complexity (HC) plays a critical role in trophic interactions, population dynamics, and food web stability. However, little theory exists to describe edible habitat complexity (EHC), where a species is both consumed and confers habitat complexity for other species. We provide a series of models demonstrating how HC and EHC alter the population dynamics and stability of simple aquatic food webs. HC is strongly stabilizing in food webs by providing safety in rarity for prey. EHC provides safety in rarity for both prey and the EHC species because their predators are omnivorous. Given the prevalence of EHC species in aquatic systems (e.g., macrophytes, corals, mussels), our models demonstrate the importance of maintaining EHC species in aquatic systems for stable food webs.

ecology↗

Effects of warming winter embryo incubation temperatures on larval cisco (Coregonus artedi) survival, growth, and critical thermal maximum

Freshwater whitefishes, Salmonidae Coregoninae, are cold stenothermic fishes of ecological and socio-economic importance in northern hemisphere lakes that are warming in response to climate change. To address the effect of warming waters on coregonine reproduction we experimentally evaluated different embryo incubation temperatures on post-hatching survival, growth, and critical thermal maximum of larval cisco (Coregonus artedi) sampled from lakes Superior and Ontario. Embryos were incubated at water temperatures of 2.0, 4.4, 6.9, and 8.9{degrees}C to simulate present and increased winter temperatures, and hatched larvae were reared in a common environment. For both populations, larval survival and critical thermal maximum were negatively related to incubation temperature, and larval growth was positively related to incubation temperature. The magnitude of change across incubation temperatures was greater in the population sampled from Lake Superior than Lake Ontario for all traits examined. The more rapid decrease in survival and critical thermal maximum across incubation temperatures for larval cisco in Lake Superior, compared to those from Lake Ontario, suggests that Lake Superior larvae may possess a more limited ability to acclimate to and cope with increasing winter water temperatures. However, the rapid increase in growth rates across incubation temperatures in Lake Superior larvae suggests they could recover better from hatching at a small length induced by warm winters, as compared to Lake Ontario larvae. Our results suggest propagation and restoration programs may want to consider integrating natural habitat preferences and maximizing phenotypic variability to ensure offspring are set up for success upon stocking.

ecology↗

Shining a light on Laurentian Great Lakes cisco (Coregonus artedi): how ice coverage may impact embryonic development

Changes in winter conditions, such as decreased ice coverage and duration, have been observed in the Laurentian Great Lakes for more than 20 years. Such changes have been hypothesized to be linked to low Coregonus spp. survival to age-1 as most cisco (Coregonus artedi) populations are autumn spawners whose embryos incubate under ice throughout the winter. The quantity of light during winter is regulated by ice coverage, and light affects embryo survival and development in some teleosts. We experimentally evaluated how cisco embryos from lakes Superior and Ontario respond to three light treatments that represented day-light intensity under 0-10, 40-60, and 90-100% ice coverage. Embryonic response measures included two developmental factors (embryo survival and incubation period) and two morphological traits (length-at-hatch and yolk-sac volume). Embryo survival was highest at the medium light treatment and decreased at high and low treatments for both populations, suggesting cisco may be adapted to withstand some light exposure from inter-annual variability in ice coverage. Light intensity had no overall effect on length of incubation. Increasing light intensity decreased length-at-hatch in Lake Superior but had no effect in Lake Ontario. Yolk-sac volume was positively correlated with increasing light in Lake Superior and negatively correlated in Lake Ontario. Contrasting responses in embryo development between lakes suggests differences in populations response to light is flexible. Our results provide a step towards better understanding the high variability observed in coregonine recruitment and may help predict what the future of this species may look like under current climate trends.

ecology↗

Influence of warming temperatures on coregonine embryogenesis within and among species

The greatest known global response of lakes to climate change has been an increase in water temperatures. The responses of many lake fishes to warming water temperatures are projected to be inadequate to counter the speed and magnitude of climate change. We experimentally evaluated the responses of embryos from a group of cold, stenothermic fishes (Salmonidae Coregoninae) to increased incubation temperatures. Study groups included cisco (Coregonus artedi) from lakes Superior and Ontario (USA), and vendace (C. albula) and European whitefish (C. lavaretus) from Lake Southern Konnevesi (Finland). Embryos from artificial crossings were incubated at water temperatures of 2.0, 4.5, 7.0, and 9.0{degrees}C, and their responses were quantified for developmental and morphological traits. Embryo survival, incubation period, and length-at- hatch were inversely related to incubation temperature whereas yolk-sac volume increased with incubation temperature within study groups. However, varying magnitudes of responses among study groups suggested differential levels of developmental plasticity to climate change. Differential levels of parental effects indicate genetic diversity may enable all study groups to adapt to cope with some degree of changing environmental conditions. Our results suggest that the coregonines sampled within and among systems may have a wide range of embryo responses to warming incubation conditions.

ecology↗

Rainbow smelt population responses to species invasions and change in environmental condition

Invasive species can have major disruptions on native food webs, yet the impact of species introductions and whether they will become invasive appears to be context-dependent. Rainbow smelt and alewife co-exist as invasive species in the Laurentian Great Lakes and as native species on the Atlantic coast of North America, but in Lake Champlain rainbow smelt is the dominant native forage fish and alewife are invasive. Alewife became abundant by 2007, providing an opportunity to explore the dynamics of these two species in a system where only one is invasive. We used data from a 31-year forage fish survey to compare demographics of rainbow smelt populations in three basins of Lake Champlain with different pelagic volumes, nutrient levels, and predator abundances. Rainbow smelt catch-per-unit-effort (CPUE) remained constant in the large, deep Main Lake before and after alewife invaded, but decreased in the two smaller basins. Declines were primarily a result of increased age-0 and age-1 mortality. Predation by top piscivores, system productivity, and resource competition alone could not explain the patterns in CPUE across the basins. The mechanisms that allow alewife and rainbow smelt to co-exist could be related to system volume and oxythermal habitat availability, and may explain why the two species do not negatively affect each other in other systems. Summer hypoxia in the smaller basins could force individuals into smaller habitat volumes with higher densities of competitors and cannibalistic adult smelt. Our findings suggest that habitat size mediates the impact of invasive alewife on native rainbow smelt.

ecology↗

Under-ice mesocosm experiments reveal the primacy of light but the importance of zooplankton in winter phytoplankton dynamics

Factors that regulate planktonic communities under lake ice may be vastly different than the open-water season. However, under-ice food webs in temperate lakes are poorly understood, despite expected changes in light availability, ice cover, and snowfall associated with climate change. We hypothesized that light limitation (bottom-up control) outweighs zooplankton grazing (top-down control) on phytoplankton biovolume and community structure under ice in a north temperate lake. Using in situ under-ice mesocosms, we found that light had stronger effects on phytoplankton abundance than zooplankton, as expected. Specifically, low light limited growth of diatoms, cryptophytes, chrysophytes, and chlorophytes. Zooplankton, however, also significantly affected phytoplankton by decreasing diatoms and cryptophytes, in contrast to the common assumption that zooplankton grazing has negligible effects under ice. Ammonia and soluble reactive phosphorus decreased in high light treatments presumably through uptake by phytoplankton, whereas ammonia and soluble reactive phosphorus increased in high zooplankton treatments, likely through excretion. In situ experimental studies are commonly applied to understand food web dynamics in open-water conditions, but are extremely rare under ice. Our results suggest that changes in the light environment under ice have significant, rapid effects on phytoplankton growth and community structure and that zooplankton may play a more active role in winter food webs than previously thought. Changes in snow and ice dynamics associated with climate change may alter the light environment in ice-covered systems and significantly influence community structure.

ecology↗

Differential lipid dynamics in stocked and wild juvenile lake trout

After more than 40 years of stocking, lake trout (Salvelinus namaycush) in Lake Champlain have started to exhibit strong, natural recruitment. The abrupt surge in recruitment suggests a change in limiting factors such as prey availability or overwinter survival. The distribution of juvenile wild lake trout varies in relative abundance among regions of Lake Champlain. The differences suggest the prey base, or foraging success, may vary geographically within the lake. Stocked and wild lake trout may differ in their ability to use resources and in overwinter survival. One metric that can indicate differences in resources across regions is lake trout lipid content, which reflects the quality of available food and serves as an important energy reserve for overwinter survival. We quantified total lipid content of stocked and wild juvenile lake trout across spatial (lake regions) and temporal (seasonal) scales. No spatial differences in lipid content were apparent. Wild fish had greater lipid content than stocked fish. Seasonally, stocked fish showed a continuous drop in lipid content from pre-winter levels at stocking to the following autumn. Wild fish showed a cyclical summer increase in lipids following winter depletion, which plateaued by autumn. The high lipid content of hatchery lake trout may be necessary as they acclimate to foraging in the wild. Hatcheries would benefit from evaluating whether post-stocking survival could be improved by altering feeding or rearing regimes.

ecology↗