Inferred support for disturbance-recovery hypothesis of North Atlantic phytoplankton blooms

Analyses of satellite-derived chlorophyll data indicate that the phase of rapid phytoplankton population growth in the North Atlantic (the ‘spring bloom') is actually initiated in the winter rather than the spring, contradicting Sverdrup's Critical Depth Hypothesis. An alternative disturbance-recovery hypothesis (DRH) has been proposed to explain this discrepancy, in which the rapid deepening of the mixed layer reduces zooplankton grazing rates sufficiently to initiate the bloom. We use Bayesian parameter inference on a simple Nutrient-Phytoplankton-Zooplankton (NPZ) to investigate the DRH and also investigate how well the model can capture the multiyear and spatial dynamics of phytoplankton concentrations and population growth rates. Every parameter in our NPZ model was inferred as a probability distribution given empirical constraints, this provides a more objective method to identify a model parameterisation given available empirical evidence, rather than fixing or tuning individual parameter values. Our model explains around 75% of variation in the seasonal dynamics of phytoplankton concentrations, 30% of variation in their population rates of change, and correctly predicts the phases of population growth and decline. Our parameter-inferred model supports DRH, revealing the sustained reduction of grazing due to mixed layer deepening as the driving mechanism behind bloom initiation, with the relaxation of nutrient limitation being another contributory mechanism. Our results also show that the continuation of the bloom is caused in part by the maintenance of phytoplankton concentrations below a level that can support positive zooplankton population growth. Our approach could be employed to formally assess alternative hypotheses for bloom formatio

Status, Change, and Futures of Zooplankton in the Southern Ocean

In the Southern Ocean, several zooplankton taxonomic groups, euphausiids, copepods, salps and pteropods, are notable because of their biomass and abundance and their roles in maintaining food webs and ecosystem structure and function, including the provision of globally important ecosystem services. These groups are consumers of microbes, primary and secondary producers, and are prey for fishes, cephalopods, seabirds, and marine mammals. In providing the link between microbes, primary production, and higher trophic levels these taxa influence energy flows, biological production and biomass, biogeochemical cycles, carbon flux and food web interactions thereby modulating the structure and functioning of ecosystems. Additionally, Antarctic krill (Euphausia superba) and various fish species are harvested by international fisheries. Global and local drivers of change are expected to affect the dynamics of key zooplankton species, which may have potentially profound and wide-ranging implications for Southern Ocean ecosystems and the services they provide. Here we assess the current understanding of the dominant metazoan zooplankton within the Southern Ocean, including Antarctic krill and other key euphausiid, copepod, salp and pteropod species. We provide a systematic overview of observed and potential future responses of these taxa to a changing Southern Ocean and the functional relationships by which drivers may impact them. To support future ecosystem assessments and conservation and management strategies, we also identify priorities for Southern Ocean zooplankton research

Restricted regions of enhanced growth of Antarctic krill in the circumpolar Southern Ocean

Food webs in high-latitude oceans are dominated by relatively few species. Future ocean and sea-ice changes affecting the distribution of such species will impact the structure and functioning of whole ecosystems. Antarctic krill (Euphausia superba) is a key species in Southern Ocean food webs, but there is little understanding of the factors influencing its success throughout much of the ocean. The capacity of a habitat to maintain growth will be crucial and here we use an empirical relationship of growth rate to assess seasonal spatial variability. Over much of the ocean, potential for growth is limited, with three restricted oceanic regions where seasonal conditions permit high growth rates, and only a few areas around the Scotia Sea and Antarctic Peninsula suitable for growth of the largest krill (>60 mm). Our study demonstrates that projections of impacts of future change need to account for spatial and seasonal variability of key ecological processes within ocean ecosystems

Circumpolar patterns in Antarctic krill larval recruitment: an environmentally-driven model

Larval recruitment in Antarctic krill is known to be episodic and regional. We consider the importance of a range of recruitment factors using an environmentally driven model of larval development from spawning to post-larvae. Our model examines the timing of spawning, interaction with bathymetry, susceptibility to cold temperatures, temperature-driven development and the seasonal cycle of sea ice to identify those factors with the greatest impact. The model predicts that the seasonal location of sea ice is the main limiting factor for successful larval recruitment. Spawning in January leads to the greatest area of viable larval recruitment habitat. Dense sea ice cover, which we assume that adult krill do not spawn under, prevents spawning in large areas early in the breeding season (December). Nevertheless, later spawning in February, when sea ice is at a minimum, means there is often insufficient time for the larvae to reach a viable developmental stage before the sea ice advances. Meanwhile, although spawning is possible in more northerly areas throughout the breeding season, these are generally remote from winter sea ice, which is assumed to be necessary for larvae to overwinter. Interaction with bathymetry before hatching further limits suitable habitat. Over a 12 yr period, the model predicted larval re - cruitment from January spawning in all years in the Cooperation, Ross and Weddell Seas, with episodic larval recruitment in the Bransfield Strait in 9 of the 12 years. Additional understanding of the overwintering requirements of larvae, together with regional studies at higher spatial resolution, particularly in shelf regions, will better constrain the uncertainties in the model

Myctophid fish (Family Myctophidae) are central consumers in the food web of the Scotia Sea (Southern Ocean)

Myctophid fish are the most abundant and diverse mesopelagic fishes in the Southern Ocean. They are a conduit of energy between primary consumers and higher marine predators, and between the upper surface layers and the mesopelagic depths. However, there remain major uncertainties about their ecology, particularly regarding their role in Southern Ocean food webs, which are often regarded as dominated by Antarctic krill in waters south of the Antarctic Polar Front. Limited data on the feeding ecology of myctophids has made it difficult to assess the importance of myctophids as consumers of krill and how they fit in the traditional view of a krill-dominated system (diatom-krill-higher predator). We provide a new assessment of the role of myctophids in Southern Ocean food webs using information from recent trophodynamic studies of myctophids conducted in the Scotia Sea, one of the most productive regions of the Southern Ocean and a region that sustains both major populations of higher predators (sea birds, seals, whales) and important commercial fisheries (krill, toothfish and mackerel icefish). Collectively, these data show that myctophids have a central role in Southern Ocean food webs as both predators and prey. Large myctophid species are prevalent consumers of krill throughout their distributional range and in different seasons in the Scotia Sea. Moreover, best estimates of both myctophid and higher predator consumption of krill to date indicate that large myctophids are the greatest predators of krill in this region, consuming almost as much krill as all other vertebrate predators of krill. Nevertheless, there are several smaller myctophid species that do not eat krill, instead consuming copepods and other small euphausiids. Myctophids therefore link primary producers to higher predators through both krill-dependent and krill-independent trophic pathways, emphasizing their importance in regional food webs. Consequently, myctophid-based trophic pathways are unlikely to be exempt from the direct consequences of a redistribution and/or reduction in krill population biomass. The extent to which myctophids can maintain food web stability and sustain higher predator populations during periods of prolonged reductions in krill abundance is considered further

Understanding the structure and functioning of polar pelagic ecosystems to predict the impacts of change

The determinants of the structure, functioning and resilience of pelagic ecosystems across most of the polar regions are not well known. Improved understanding is essential for assessing the value of biodiversity and predicting the effects of change (including in biodiversity) on these ecosystems and the services they maintain. Here we focus on the trophic interactions that underpin ecosystem structure, developing comparative analyses of how polar pelagic food webs vary in relation to the environment. We highlight that there is not a singular, generic Arctic or Antarctic pelagic food web, and, although there are characteristic pathways of energy flow dominated by a small number of species, alternative routes are important for maintaining energy transfer and resilience. These more complex routes cannot, however, provide the same rate of energy flow to highest trophic-level species. Food-web structure may be similar in different regions, but the individual species that dominate mid-trophic levels vary across polar regions. The characteristics (traits) of these species are also different and these differences influence a range of food-web processes. Low functional redundancy at key trophic levels makes these ecosystems particularly sensitive to change. To develop models for projecting responses of polar ecosystems to future environmental change, we propose a conceptual framework that links the life histories of pelagic species and the structure of polar food webs

An intercomparison of models predicting growth of Antarctic krill (Euphausia superba): The importance of recognizing model specificity

Antarctic krill (Euphausia superba) is a key species of the Southern Ocean, impacted by climate change and human exploitation. Understanding how these changes affect the distribution and abundance of krill is crucial for generating projections of change for Southern Ocean ecosystems. Krill growth is an important indicator of habitat suitability and a series of models have been developed and used to examine krill growth potential at different spatial and temporal scales. The available models have been developed using a range of empirical and mechanistic approaches, providing alternative perspectives and comparative analyses of the key processes influencing krill growth. Here we undertake an intercomparison of a suite of the available models to understand their sensitivities to major driving variables. This illustrates that the results are strongly determined by the model structure and technical characteristics, and the data on which they were developed and validated. Our results emphasize the importance of assessing the constraints and requirements of individual krill growth models to ensure their appropriate application. The study also demonstrates the value of the development of alternative modelling approaches to identify key processes affecting the dynamics of krill. Of critical importance for modelling the growth of krill is appropriately assessing and accounting for differences in estimates of food availability resulting from alternative methods of observation. We suggest that an intercomparison approach is particularly valuable in the development and application of models for the assessment of krill growth potential at circumpolar scales and for future projections. As another result of the intercomparison, the implementations of the models used in this study are now publicly available for future use and analyses