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Inquiry‐based components of ecology curricula can be valuable, exposing students to what it means to do science, from conceiving of a meaningful question to effectively disseminating results to an audience. Here, we describe two approaches for implementing independent, remote research for undergraduates enacted in the spring semester of 2020 at Reed College in Portland, OR, reporting case studies from an intermediate‐level ecology course and an interdisciplinary environmental science course. We report on both the challenges as well as the novel opportunities for independent research projects in such a setting, the details of how projects were implemented, the tools and resources that may help facilitate such endeavors, as well as perceptions on the effectiveness of this endeavor by students. As institutes of higher education continue to operate in an online learning environment, we hope these materials help spark a discussion about how to engage in meaningful research experiences as part of coursework in the COVID‐19 era and beyond., We report on both the challenges as well as the novel opportunities for independent ecology research projects in a remote learning setting, the details of how projects were implemented, the tools and resources that may help facilitate such endeavors, as well as perceptions on the effectiveness of this endeavor by students.

The majority of organisms on Earth are ectothermic, and their body temperatures largely depend on the surrounding environmental conditions. Body temperature strongly affects fitness. In a spatially variable environment, movement can allow an ectothermic organism to change their body temperature. This project explores body temperature and movement in the context of phytoplankton fitness within lake environments. By modeling movement strategies using lake thermal profile data, we infer the range of temperatures phytoplankton experience and the resulting effect on fitness.

Laboratory measurements of physiological and demographic tolerances are important in understanding the impact of climate change on species diversity; however, it has been recognized that forecasts based solely on these laboratory estimates overestimate risk by omitting the capacity for species to utilize microclimatic variation via behavioral adjustments in activity patterns or habitat choice. The complex, and often context-dependent nature, of microclimate utilization has been an impediment to the advancement of general predictive models. Here, we overcome this impediment and estimate the potential impact of warming on the fitness of ectotherms using a benefit/cost trade-off derived from the simple and broadly documented thermal performance curve and a generalized cost function. Our framework reveals that, for certain environments, the cost of behavioral thermoregulation can be reduced as warming occurs, enabling behavioral buffering (e.g., the capacity for behavior to ameliorate detrimental impacts) and "behavioral rescue" from extinction in extreme cases. By applying our framework to operative temperature and physiological data collected at an extremely fine spatial scale in an African lizard, we show that new behavioral opportunities may emerge. Finally, we explore large-scale geographic differences in the impact of behavior on climate-impact projections using a global dataset of 38 insect species. These multiple lines of inference indicate that understanding the existing relationship between thermal characteristics (e.g., spatial configuration, spatial heterogeneity, and modal temperature) is essential for improving estimates of extinction risk.

Since the inaugural edition of Ecosystems was published in 1998, ecosystem science has undergone substantial changes including the development of new research methods and an increasing emphasis on collaborations across traditional academic boundaries. In response to this transformation, we reflect on the current state of theory in ecosystem science, and make recommendations for training the next generation of Ph.D.-level ecosystem scientists. Specifically, we call for increased integration of theory into ecosystem science and outline the utility of iterating between theory and data generated by observations, experiments, and quantitative models. We recommend exposing graduate students to these three major approaches for generating data and propose strategies that students, advisors, and departments can employ to ensure this exposure. Ultimately, a successful training program will provide students with an understanding of key theories related to ecosystem science and how they interact with data, an appreciation for the interconnectedness of approaches to scientific inference, and a well-developed skill set in at least one approach—thereby empowering them to confidently tackle our pressing environmental problems. Although this is a daunting list of goals, continuing to advance our understanding of how ecosystems function necessitates a rigorous and well-developed training program.

Over the last few decades, biologists have made substantial progress in understanding relationships between changing climates and organism performance. Much of this work has focused on temperature because it is the best kept of climatic records, in many locations it is predicted to keep rising into the future, and it has profound effects on the physiology, performance, and ecology of organisms, especially ectothermic organisms which make up the vast majority of life on Earth. Nevertheless, much of the existing literature on temperature-organism interactions relies on mean temperatures. In reality, most organisms do not directly experience mean temperatures; rather, they experience variation in temperature over many time scales, from seconds to years. We propose to shift the focus more directly on patterns of temperature variation, rather than on means per se, and present a framework both for analyzing temporal patterns of temperature variation and for incorporating those patterns into predictions about organismal biology. In particular, we advocate using the Fourier transform to decompose temperature time series into their component sinusoids, thus allowing transformations between the time and frequency domains. This approach provides (1) standardized ways of visualizing the contributions that different frequencies make to total temporal variation; (2) the ability to assess how patterns of temperature variation have changed over the past half century and may change into the future; and (3) clear approaches to manipulating temporal time series to ask "what if" questions about the potential effects of future climates. We first summarize global patterns of change in temperature variation over the past 40 years; we find meaningful changes in variation at the half day to yearly times scales. We then demonstrate the utility of the Fourier framework by exploring how power added to different frequencies alters the overall incidence of long-term waves of high and low temperatures, and find that power added to the lowest frequencies greatly increases the probability of long-term heat and cold waves. Finally, we review what is known about the time scales over which organismal thermal performance curves change in response to variation in the thermal environment. We conclude that integrating information characterizing both the frequency spectra of temperature time series and the time scales of resulting physiological change offers a powerful new avenue for relating climate, and climate change, to the future performance of ectothermic organisms.

Heterogeneity is increasingly recognized as a foundational characteristic of ecological systems. Under global change, understanding temporal community heterogeneity is necessary for predicting the stability of ecosystem functions and services. Indeed, spatial heterogeneity is commonly used in alternative stable state theory as a predictor of temporal heterogeneity and therefore an early indicator of regime shifts. To evaluate whether spatial heterogeneity in species composition is predictive of temporal heterogeneity in ecological communities, we analyzed 68 community data sets spanning freshwater and terrestrial systems where measures of species abundance were replicated over space and time. Of the 68 data sets, 55 (81%) had a weak to strongly positive relationship between spatial and temporal heterogeneity, while in the remaining communities the relationship was weak to strongly negative (19%). Based on a mixed model analysis, we found a significant but weak overall positive relationship between spatial and temporal heterogeneity across all data sets combined, and within aquatic and terrestrial data sets separately. In addition, lifespan and successional stage were negatively and positively related to temporal heterogeneity, respectively. We conclude that spatial heterogeneity may be a predictor of temporal heterogeneity in ecological communities, and that this relationship may be a general property of many terrestrial and aquatic communities.

Understanding the processes that influence range expansions during climate warming is paramount for predicting population extirpations and preparing for the arrival of non-native species. While climate warming occurs over a background of variation due to cyclical processes and irregular events, the temporal structure of the thermal environment is largely ignored when forecasting the dynamics of non-native species. Ecological theory predicts that high levels of temporal autocorrelation in the environment– relatedness between conditions occurring in close temporal proximity– will favor populations that would otherwise have an average negative growth rate by increasing the duration of favorable environmental periods. Here, we invoke such theory to explain the success of biological invasions and evaluate the hypothesis that sustained periods of high environmental temperature can act synergistically with increases in mean temperature to favor the establishment of non-native species. We conduct a 60-day field mesocosm experiment to measure the population dynamics of the non-native cladoceran zooplankter Daphnia lumholtzi and a native congener Daphnia pulex in ambient temperature environments (control), warmed with recurrent periods of high environmental temperatures (uncorrelated-warmed), or warmed with sustained periods of high environmental temperatures (autocorrelated-warmed), such that both warmed treatments exhibited the same mean temperature but exhibited different temporal structures of their thermal environments. Maximum D. lumholtzi densities in the warmed-autocorrelated treatment were three- and eight-fold higher relative to warmed-uncorrelated and control treatments, respectively. Yet, D. lumholtzi performed poorly across all experimental treatment relative to D. pulex and were undetectable the end of the experiment. Using mathematical models we show that this increase in performance can occur alongside increasing temporal autocorrelation, and should occur over a broad range of warming scenarios. These results provide both empirical and theoretical evidence that the temporal structure of the environment can influence the performance of species undergoing range expansions due to climate warming. This article is protected by copyright. All rights reserved.

Environmental variability is ubiquitous, but its effects on populations are not fully understood or predictable. Recent attention has focused on how rapid evolution can impact ecological dynamics via adaptive trait change. However, the impact of trait change arising from plastic responses has received less attention, and is often assumed to optimize performance and unfold on a separate, faster timescale than ecological dynamics. Challenging these assumptions, we propose that gradual plasticity is important for ecological dynamics, and present a study of the plastic responses of the freshwater green algae Chlamydomonas reinhardtii as it acclimates to temperature changes. First, we show that C. reinhardtii 's gradual acclimation responses can both enhance and suppress its performance after a perturbation, depending on its prior thermal history. Second, we demonstrate that where conventional approaches fail to predict the population dynamics of C. reinhardtii exposed to temperature fluctuations, a new model of gradual acclimation succeeds. Finally, using high-resolution data, we show that phytoplankton in lake ecosystems can experience thermal variation sufficient to make acclimation relevant. These results challenge prevailing assumptions about plasticity's interactions with ecological dynamics. Amidst the current emphasis on rapid evolution, it is critical that we also develop predictive methods accounting for plasticity.

Thermal variation through space and time are prominent features of ecosystems that influence processes at multiple levels of biological organization. Yet, it remains unclear how populations embedded within biological communities will respond to climate warming in thermally variable environments, particularly as climate change alters existing patterns of thermal spatial and temporal variability. As environmental temperatures increase above historical ranges, organisms may increasingly rely on extreme habitats to effectively thermoregulate. Such locations desirable in their thermal attributes (e.g., thermal refugia) are often suboptimal for resource acquisition (e.g., underground tunnels). Thus, via the expected increase in both mean temperatures and diel thermal variation, climate warming may heighten the trade-off for consumers between behaviors maximizing thermal performance and those maximizing resource acquisition. Here, we integrate behavioral, physiological, and trophic ecology to provide a general framework for understanding how temporal thermal variation, mediated by access to a thermal refugium, alters the response of consumer-resource systems to warming. We use this framework to predict how temporal variation and access to thermal refugia affect the persistence of consumers and resources during climate warming, how the quality of thermal refugia impact consumer-resource systems, and how consumer-resource systems with fast vs. slow ecological dynamics respond to warming. Our results show that the spatial thermal variability provided by refugia can elevate consumer biomass at warmer temperatures despite reducing the fraction of time consumers spend foraging, that temporal variability detrimentally impacts consumers at high environmental temperatures, and that consumer-resource systems with fast ecological dynamics are most vulnerable to climate warming. Thus, incorporating both estimates of thermal variability and species interactions may be necessary to accurately predict how populations respond to warming.