Size as a Trait for Understanding the Role of Zooplankton in the Biological Carbon Pump

Living organisms impact carbon transport between the atmosphere and the ocean through the biological carbon pump. Some plankton communities augment carbon export from the ocean’s surface, and are thought to have a major role in global climate. These export communities are often characterized by larger organisms that sink to depths where the carbon they contain is sequestered from the atmosphere. Zooplankton can enhance export by aggregating prey into larger sinking fecal pellets; however fecal pellet flux is a highly variable component of the biological carbon pump. Relating plankton trophic dynamics to changes in particulate carbon flux is an important step in understanding the ocean’s carbon cycle. This research aims to connect plankton ecology with variability in zooplankton fecal pellet carbon flux, using body size as an organizing trait. A copepod fecal pellet carbon flux model is presented and applied to 25 years of copepod data from the Gulf of Maine. This model uses size-based metabolic rates to estimate fecal pellet production, and sinking and decay functions to estimate flux. The results show that copepod community size structure determines fecal pellet carbon flux efficiency, but that flux itself is determined by copepod abundance and size. A second iteration of this model, which includes a temperature-dependent pellet decay function and diel vertical migration behavior, is applied to 55 years of copepod data from the North Atlantic Ocean. Analysis shows that fecal pellet carbon flux is decreasing as a result of declining copepod biomass, coincident with ocean warming. However, these changes vary from region to region, highlighting the importance of local dynamics. A study of local-scale trophic dynamics in the Gulf of Maine tests whether feeding and zooplankton fecal pellet production increases particle size, and therefore flux potential, in plankton communities. These experiments show tight coupling between microplankton and mesozooplankton, and demonstrate the importance of fecal pellet production as a mechanism for aggregating smaller particles into larger, sinking fecal pellets. Collectively this work shows that organism body size can be used as an organizing trait to connect individual-scale biology with variability in the biological carbon pump

Cloud shadows drive vertical migrations of deep-dwelling marine life

Our study provides evidence that, in addition to diel vertical migration, zooplankton residing at \u3e300-m depth during the day perform high-frequency, vertical migrations due to light modulation by clouds. Using a water-following framework and measurements and modeling of the twilight zone light field, we isolated the detailed phototactic response and show that some twilight zone animals are considerably more active than previously thought, with a cumulative distance traveled of more than one-third of that for diel migration. The increased movement increases predation risk and has implications for the metabolic requirements of these animals in the food-limited deep sea

Century-scale changes in phytoplankton phenology in the Gulf of Maine

The phenology of major seasonal events is an important indicator of climate. We analyzed multiple datasets of in situ chlorophyll measurements from the Gulf of Maine dating back to the early 20th century in order to detect climate-scale changes in phenology. The seasonal cycle was consistently characterized by a two-bloom pattern, with spring and autumn blooms. The timing of both spring and autumn blooms has shifted later in the year at rates ranging from ∼1 to 9 days per decade since 1960, depending on the phenology metric, and trends only emerged at time scales of >40 years. Bloom phenology had only weak correlations with major climate indices. There were stronger associations between bloom timing and physical and chemical variables. Autumn bloom initiation correlated strongly with surface temperature and salinity, and spring bloom with nutrients. A later spring bloom also correlated with an increased cohort of Calanus finmarchicus, suggesting broader ecosystem implications of phytoplankton phenology

The Surprising Oceanography of the Gulf of Maine

The oceanography of the Gulf of Maine has changed in ways that have not been seen previously but that are likely to be more common in the future–changes like extreme rapid warming and declines in primary productivity. The changing oceanography has underpinned surprising losses in commercial stocks and endangered species. Because of the rapid rate of change, some have viewed the Gulf of Maine as a window into the ocean’s future, with the idea that lessons learned can be applied in places that have yet to experience similar rapid changes. We can examine the dynamics, origin, and implications of surprising oceanographic conditions–conditions that would have been considered unlikely based on recent prior experience. Based on a formal statistical definition of climate surprises, the frequency of oceanographic surprises in the Gulf of Maine is higher and has increased faster than what would be expected, even given underlying trends. Oceanographic surprises vary in character from one to the next and are often linked to larger scale shifting oceanography across the North Atlantic. The implications for ecological and human communities, industries, and conservation efforts imply a need for policies that consider adaptation to sudden events as well as long-term changes

Size as the master trait in modeled copepod fecal pellet carbon flux

Zooplankton fecal pellet flux is a highly variable component of the biological carbon pump. While fecal pellets can comprise 0 to nearly 100% of particulate organic carbon collected in sediment traps, mechanisms for this variability remain poorly understood. Fecal pellet carbon flux is a complex function of several variables. We present a model that incorporates individual-scale metabolic processes to determine fecal pellet production rate, the relationship between body size and fecal pellet size, the relationship between fecal pellet size and sinking rate, and a function representing the breakdown of particles in the water column. When applied to copepod communities sampled by the continuous plankton recorder in the Gulf of Maine over 25 years, a seasonal pattern of fecal pellet carbon flux emerges. The interannual flux time series produced by the model reflects known oceanographic perturbations and shows how organism-scale processes can be scaled up to explain ecosystem level variability. We conclude that fecal pellet carbon flux in the Gulf of Maine is driven by copepod community size structure and copepod abundance, and that the fraction of fecal pellet carbon that reaches depth is a function of copepod size, rather than abundance. Changes in the physical environment which alter the size composition of the copepod community lead to variability in fecal pellet carbon flux. Our results indicate that incorporating size composition into biogeochemical models can more accurately constrain zooplankton-mediated carbon flux.This is the publisher’s final pdf. The article is copyrighted by the Association for the Sciences of Limnology and Oceanography and published by John Wiley & Sons, Inc. It can be found at: http://www.aslo.org/lo/index.htm

Rapid climate-driven circulation changes threaten conservation of endangered north atlantic right whales

As climate trends accelerate, ecosystems will be pushed rapidly into new states, reducing the potential efficacy of conservation strategies based on historical patterns. In the Gulf of Maine, climate-driven changes have restructured the ecosystem rapidly over the past decade. Changes in the Atlantic meridional overturning circulation have altered deepwater dynamics, driving warming rates twice as high as the fastest surface rates. This has had implications for the copepod Calanus finmarchicus, a critical food supply for the endangered North Atlantic right whale (Eubalaena glacialis). The oceanographic changes have driven a deviation in the seasonal foraging patterns of E. glacialis upon which conservation strategies depend, making the whales more vulnerable to ship strikes and gear entanglements. The effects of rapid climate-driven changes on a species at risk undermine current management approaches.publishedVersio

What are the type, direction, and strength of species, community, and ecosystem responses to warming in aquatic mesocosm studies and their dependency on experimental characteristics? A systematic review protocol

Background Mesocosm experiments have become increasingly popular in climate change research as they bridge the gap between small-scale, less realistic, microcosm experiments, and large-scale, more complex, natural systems. Characteristics of aquatic mesocosm designs (e.g., mesocosm volume, study duration, and replication) vary widely, potentially affecting the magnitude and direction of effect sizes measured in experiments. In this global systematic review we aim to identify the type, direction and strength of climate warming effects on aquatic species, communities and ecosystems in mesocosm experiments. Furthermore, we will investigate the context-dependency of the observed effects on several a priori determined effect moderators (ecological and methodological). Our conclusions will provide recommendations for aquatic scientists designing mesocosm experiments, as well as guidelines for interpretation of experimental results by scientists, policy-makers and the general public. Methods We will conduct a systematic search using multiple online databases to gather evidence from the scientific literature on the effects of warming experimentally tested in aquatic mesocosms. Data from relevant studies will be extracted and used in a random effects meta-analysis to estimate the overall effect sizes of warming experiments on species performance, biodiversity and ecosystem functions. Experimental characteristics (e.g., mesocosm size and shape, replication-level, experimental duration and design, biogeographic region, community type, crossed manipulation) will be further analysed using subgroup analyses

It's about time: A synthesis of changing phenology in the Gulf of Maine ecosystem

© The Author(s), 2019. This article is distributed under the terms of the Creative Commons Attribution License. The definitive version was published in Staudinger, M. D., Mills, K. E., Stamieszkin, K., Record, N. R., Hudak, C. A., Allyn, A., Diamond, A., Friedland, K. D., Golet, W., Henderson, M. E., Hernandez, C. M., Huntington, T. G., Ji, R., Johnson, C. L., Johnson, D. S., Jordaan, A., Kocik, J., Li, Y., Liebman, M., Nichols, O. C., Pendleton, D., Richards, R. A., Robben, T., Thomas, A. C., Walsh, H. J., & Yakola, K. It's about time: A synthesis of changing phenology in the Gulf of Maine ecosystem. Fisheries Oceanography, 28(5), (2019): 532-566, doi: 10.1111/fog.12429.The timing of recurring biological and seasonal environmental events is changing on a global scale relative to temperature and other climate drivers. This study considers the Gulf of Maine ecosystem, a region of high social and ecological importance in the Northwest Atlantic Ocean and synthesizes current knowledge of (a) key seasonal processes, patterns, and events; (b) direct evidence for shifts in timing; (c) implications of phenological responses for linked ecological‐human systems; and (d) potential phenology‐focused adaptation strategies and actions. Twenty studies demonstrated shifts in timing of regional marine organisms and seasonal environmental events. The most common response was earlier timing, observed in spring onset, spring and winter hydrology, zooplankton abundance, occurrence of several larval fishes, and diadromous fish migrations. Later timing was documented for fall onset, reproduction and fledging in Atlantic puffins, spring and fall phytoplankton blooms, and occurrence of additional larval fishes. Changes in event duration generally increased and were detected in zooplankton peak abundance, early life history periods of macro‐invertebrates, and lobster fishery landings. Reduced duration was observed in winter–spring ice‐affected stream flows. Two studies projected phenological changes, both finding diapause duration would decrease in zooplankton under future climate scenarios. Phenological responses were species‐specific and varied depending on the environmental driver, spatial, and temporal scales evaluated. Overall, a wide range of baseline phenology and relevant modeling studies exist, yet surprisingly few document long‐term shifts. Results reveal a need for increased emphasis on phenological shifts in the Gulf of Maine and identify opportunities for future research and consideration of phenological changes in adaptation efforts.This work was supported by the Department of the Interior Northeast Climate Adaptation Science Center (G14AC00441) for MDS, AJ, and KY; the National Science Foundation's Coastal SEES Program (OCE‐1325484) for KEM, ACT, MEH, and AA; the National Aeronautics and Space Administration (NNX16 AG59G) for ACT, KEM, NRR, and KSS; the USGS Climate Research and Development Program for TGH; National Science & Engineering Research Council of Canada, University of New Brunswick, Environment Canada, Sir James Dunn Wildlife Research Centre, and New Brunswick Wildlife Trust Fund for AD. We also thank the Regional Association for Research on the Gulf of Maine for support, and the Gulf of Maine Research Institute for hosting and providing in kind resources for a two day in‐person workshop in August 2016. We greatly appreciate contributions from K. Alexander, G. Calandrino, C. Feurt, I. Mlsna, N. Rebuck, J. Seavey, and J. Sun for helping shape the initial scope of the manuscript. We thank J. Weltzin and two anonymous reviewers for their constructive comments. The contents of this paper are solely the responsibility of the authors and do not necessarily represent the views of the Northeast Climate Adaptation Science Center, U.S. Geological Survey, National Oceanographic and Atmospheric Administration, Fisheries and Oceans Canada or the US Environmental Protection Agency. This manuscript is submitted for publication with the understanding that the United States Government is authorized to reproduce and distribute reprints for Governmental purposes. None of the authors have conflicts of interest to declare in association with the contents of this manuscript

Eggs of the copepod Acartia tonsa Dana require hypoxic conditions to tolerate prolonged embryonic development arrest

Additional file 1. Raw data and calculations for all experiments as well as an overview of experiments in this study