Evaluating top-down, bottom-up, and environmental drivers of pelagic food web dynamics along an estuarine gradient.

Identification of the key biotic and abiotic drivers within food webs is important for understanding species abundance changes in ecosystems, particularly across ecotones where there may be strong variation in interaction strengths. Using structural equation models (SEMs) and four decades of integrated data from the San Francisco Estuary, we investigated the relative effects of top-down, bottom-up, and environmental drivers on multiple trophic levels of the pelagic food web along an estuarine salinity gradient and at both annual and monthly temporal resolutions. We found that interactions varied across the estuarine gradient and that the detectability of different interactions depended on timescale. For example, for zooplankton and estuarine fishes, bottom-up effects appeared to be stronger in the freshwater upstream regions, while top-down effects were stronger in the brackish downstream regions. Some relationships (e.g., bottom-up effects of phytoplankton on zooplankton) were seen primarily at annual timescales, whereas others (e.g., temperature effects) were only observed at monthly timescales. We also found that the net effect of environmental drivers was similar to or greater than bottom-up and top-down effects for all food web components. These findings can help identify which trophic levels or environmental factors could be targeted by management actions to have the greatest impact on estuarine forage fishes and the spatial and temporal scale at which responses might be observed. More broadly, this study highlights how environmental gradients can structure community interactions and how long-term data sets can be leveraged to generate insights across multiple scales.

Governing Ecological Connectivity in Cross-Scale Dependent Systems.

Ecosystem management and governance of cross-scale dependent systems require integrating knowledge about ecological connectivity in its multiple forms and scales. Although scientists, managers, and policymakers are increasingly recognizing the importance of connectivity, governmental organizations may not be currently equipped to manage ecosystems with strong cross-boundary dependencies. Managing the different aspects of connectivity requires building social connectivity to increase the flow of information, as well as the capacity to coordinate planning, funding, and actions among both formal and informal governance bodies. We use estuaries in particular the San Francisco Estuary, in California, in the United States, as examples of cross-scale dependent systems affected by many intertwined aspects of connectivity. We describe the different types of estuarine connectivity observed in both natural and human-affected states and discuss the human dimensions of restoring beneficial physical and ecological processes. Finally, we provide recommendations for policy, practice, and research on how to restore functional connectivity to estuaries.

Five decades (1972-2020) of zooplankton monitoring in the upper San Francisco Estuary.

We present the longest available dataset (by 15 years) of estuarine zooplankton abundance worldwide. Zooplankton have been monitored throughout the upper San Francisco Estuary from 1972 -present due to its status as a central hub of California water delivery and home to commercially important and endangered fishes. We integrated data from five monitoring programs, including over 300 locations, three size-classes of zooplankton targeted with different gears, over 80,000 samples, and over two billion sampled organisms. Over the duration of this dataset, species invasions have driven community turnover, periodic droughts have occurred, and important fishes have declined, likely due in part to reduced food supply from zooplankton. Data from the individual surveys have been used in prior studies on issues related to species invasions, flows, fish diets and population dynamics, zooplankton population dynamics, and community ecology. Our integrated dataset offers unparalleled spatio-temporal scope to address these and other fundamental ecological questions.

Does a complex life cycle affect adaptation to environmental change? Genome-informed insights for characterizing selection across complex life cycle.

Complex life cycles, in which discrete life stages of the same organism differ in form or function and often occupy different ecological niches, are common in nature. Because stages share the same genome, selective effects on one stage may have cascading consequences through the entire life cycle. Theoretical and empirical studies have not yet generated clear predictions about how life cycle complexity will influence patterns of adaptation in response to rapidly changing environments or tested theoretical predictions for fitness trade-offs (or lack thereof) across life stages. We discuss complex life cycle evolution and outline three hypotheses-ontogenetic decoupling, antagonistic ontogenetic pleiotropy and synergistic ontogenetic pleiotropy-for how selection may operate on organisms with complex life cycles. We suggest a within-generation experimental design that promises significant insight into composite selection across life cycle stages. As part of this design, we conducted simulations to determine the power needed to detect selection across a life cycle using a population genetic framework. This analysis demonstrated that recently published studies reporting within-generation selection were underpowered to detect small allele frequency changes (approx. 0.1). The power analysis indicates challenging but attainable sampling requirements for many systems, though plants and marine invertebrates with high fecundity are excellent systems for exploring how organisms with complex life cycles may adapt to climate change.

Costs and compensation in zooplankton pigmentation under countervailing threats of ultraviolet radiation and predation.

Evolutionary responses to opposing directions of natural selection include trade-offs, where the phenotype balances selective forces, and compensation, where other traits reduce the impact of one selective force. Zooplankton pigmentation protects from ultraviolet radiation (UVR) but attracts visual predators. This trade-off is understudied in the ocean where planktonic larvae in surface waters face ubiquitous UVR and visual predation threats. We tested whether crab larvae can behaviorally reduce UVR risk through downward swimming or expansion of photoprotective chromatophores. Then we examined whether more pigmented larvae are more heavily predated by silverside fish under natural sunlight in the tropics in three UVR treatments (visible light, visible + UVA, visible + UVA + UVB). Lastly, we tested the behavioral chromatophore response of larvae to predation threats in two light treatments. Armases ricordi avoided surface waters after exposure to sunlight with UVR. Armases ricordi, Armases americanum, and Eurypanopeus sp. consistently expanded chromatophores in UVR or visible light, while Mithraculus sculptus and Mithraculus coryphe showed no response. Fish preferred pigmented larvae on sunnier days in visible light lacking UVR. Lastly, both M. coryphe and M. sculptus unexpectedly expanded chromatophores in fish cues, but responses were inconsistent over trials and across light treatments. The more consistent larval responses to UVR than to predator cues and the lack of predator preferences in natural light conditions suggest that UVR may have a stronger influence on pigmentation than predation. This study improves our understanding of planktonic adaptation to countervailing selection caused by visual predation and exposure to UVR.

Photoprotective benefits of pigmentation in the transparent plankton community: a comparative species experimental test.

Plankton live under the countervailing selective pressures of predation and ultraviolet radiation (UVR). In lakes, zooplankton are transparent reducing visibility to predatory fishes but are pigmented in the absence of fishes, hypothetically reducing UVR damage. In the sea, planktivorous fishes are widespread, so plankton typically are transparent and ascend to productive surface waters at night to forage and descend during the day to reduce visibility to predators. However, larvae of some species face the unique constraint of traveling in surface currents in the daytime during migrations between adult and larval habitats. We would expect these larvae to be transparent since companion studies demonstrated increased predation risk of pigmented larvae under strong sunlight. Paradoxically, larvae range from being darkly to lightly pigmented. We hypothesize that some larvae are more heavily pigmented to reduce UVR damage, while other species travelling in subsurface currents with low UVR might be more transparent. Linking larval morphology to depth-dependent selective pressures would add a key element to help improve predictions of larval vertical distributions, which are important for simulating larval transport trajectories. We quantitatively tested the hypothesis that selection may have favored photoprotective pigmentation for larvae in the predominantly transparent plankton community while testing the differential effects of UVA and UVB radiation. We measured larval pigmentation of 12 species of crabs and exposed them to visible light only, visible + UVA, or visible + UVA + UVB in the tropics. Controlling for phylogeny, more pigmented species survived UVR better than less pigmented species, especially on sunnier days, though intraspecific comparisons for four species were equivocal. Most species died even from UVA exposure, which has long been regarded as relatively harmless despite penetrating deeper underwater than UVB. Thus, we demonstrate with a phylogenetically controlled analysis that crab larvae are pigmented in the predominantly transparent planktonic community to protect from UVR, improving our understanding of the selective forces acting on animal coloration and the factors determining planktonic distributions, survival, and dispersal. This linkage of morphology with susceptibility will be important for developing mechanistic models of environmental stress responses to better predict larval dispersal in current and future climates.

Growing Safe: Acute Size Escape from Desiccation in Juvenile Crepipatella peruviana (Mollusca: Gastropoda).

Desiccation is an important limiting factor in the intertidal zone. Generally decreasing seaward, desiccation stress can also be alleviated in wet microhabitats. Juvenile snails are generally more susceptible to desiccation than adults, and, for some species, juveniles must therefore hide in microhabitats to survive emersion. The transition from hiding in safe microhabitats to being able to survive fully exposed for the duration of low tide is not well documented. In this study, we investigated the influence of size on desiccation tolerance in juveniles of the calyptraeid gastropod Crepipatella peruviana to determine the size at which they can first survive exposure to air. Juveniles 2-13 mm long were exposed to 75% or 100% relative humidity for 0.5-6.5 hours. Juveniles smaller than 5 mm in shell length did not survive emersion at 75% relative humidity for even 0.5 hours; surprisingly, most also perished after short exposures to air at 100% relative humidity, suggesting that something other than desiccation stress may also be at play. In marked contrast, 82% of juveniles larger than 6 mm in shell length survived exposure to 75% relative humidity for the full 6.5 hours. In a field survey, no juveniles smaller than 9 mm were found on exposed rock but rather were found only in wet microhabitats. We suggest that the clearly defined size escape from desiccation may reflect a change in gill functioning or a newfound ability to retain water more effectively within the mantle cavity at low tide.