A Conceptual Approach to Partitioning a Vertical Profile of Phytoplankton Biomass Into Contributions From Two Communities.

We describe an approach to partition a vertical profile of chlorophyll-a concentration into contributions from two communities of phytoplankton: one (community 1) that resides principally in the turbulent mixed-layer of the upper ocean and is observable through satellite visible radiometry; the other (community 2) residing below the mixed-layer, in a stably stratified environment, hidden from the eyes of the satellite. The approach is tuned to a time-series of profiles from a Biogeochemical-Argo float in the northern Red Sea, selected as its location transitions from a deep mixed layer in winter (characteristic of vertically well-mixed systems) to a shallow mixed layer in the summer with a deep chlorophyll-a maximum (characteristic of vertically stratified systems). The approach is extended to reproduce profiles of particle backscattering, by deriving the chlorophyll-specific backscattering coefficients of the two communities and a background coefficient assumed to be dominated by non-algal particles in the region. Analysis of the float data reveals contrasting phenology of the two communities, with community 1 blooming in winter and 2 in summer, community 1 negatively correlated with epipelagic stratification, and 2 positively correlated. We observe a dynamic chlorophyll-specific backscattering coefficient for community 1 (stable for community 2), positively correlated with light in the mixed-layer, suggesting seasonal changes in photoacclimation and/or taxonomic composition within community 1. The approach has the potential for monitoring vertical changes in epipelagic biogeography and for combining satellite and ocean robotic data to yield a three-dimensional view of phytoplankton distribution.

Ocean Lagrangian Trajectories (OLTraj): Lagrangian analysis for non-expert users.

Lagrangian analysis is becoming increasingly important to better understand the ocean's biological and biogeochemical cycles. Yet, biologists and chemists often lack the technical skills required to set up such analyses. Here, we present a new product of pre-computed ocean Lagrangian trajectories (OLTraj) targeting non-expert users, and demonstrate how to use it by means of worked examples. OLTraj is based on satellite-derived geostrophic currents, which allows one to directly compare it with other in-situ or satellite products. We anticipate that OLTraj will foster a new interest in Lagrangian applications in ocean biology and biogeochemistry.

Physical connectivity simulations reveal dynamic linkages between coral reefs in the southern Red Sea and the Indian Ocean.

The southern Red Sea is genetically distinct from the rest of the basin; yet the reasons responsible for this genetic separation remain unclear. Connectivity is a vital process for the exchange of individuals and genes among geographically separated populations, and is necessary for maintaining biodiversity and resilience in coral reef ecosystems. Here, using long-term, high-resolution, 3-D backward particle tracking simulations, we investigate the physical connectivity of coral reefs in the southern Red Sea with neighbouring regions. Overall, the simulation results reveal that the southern Red Sea coral reefs are more physically connected with regions in the Indian Ocean (e.g., the Gulf of Aden) than with the northern part of the basin. The identified connectivity exhibits a distinct monsoon-related seasonality. Though beyond the country boundaries, relatively remote regions of the Indian Ocean may have a substantial impact on the southern Red Sea coral reef regions, and this should be taken into consideration when establishing conservation strategies for these vulnerable biodiversity hot-spots.

Factors Regulating the Relationship Between Total and Size-Fractionated Chlorophyll-a in Coastal Waters of the Red Sea.

Phytoplankton biomass and size structure are recognized as key ecological indicators. With the aim to quantify the relationship between these two ecological indicators in tropical waters and understand controlling factors, we analyzed the total chlorophyll-a concentration, a measure of phytoplankton biomass, and its partitioning into three size classes of phytoplankton, using a series of observations collected at coastal sites in the central Red Sea. Over a period of 4 years, measurements of flow cytometry, size-fractionated chlorophyll-a concentration, and physical-chemical variables were collected near Thuwal in Saudi Arabia. We fitted a three-component model to the size-fractionated chlorophyll-a data to quantify the relationship between total chlorophyll and that in three size classes of phytoplankton [pico- (<2 µm), nano- (2-20 µm) and micro-phytoplankton (>20 µm)]. The model has an advantage over other more empirical methods in that its parameters are interpretable, expressed as the maximum chlorophyll-a concentration of small phytoplankton (pico- and combined pico-nanophytoplankton, C p m and C p , n m , respectively) and the fractional contribution of these two size classes to total chlorophyll-a as it tends to zero (D p and D p,n ). Residuals between the model and the data (model minus data) were compared with a range of other environmental variables available in the dataset. Residuals in pico- and combined pico-nanophytoplankton fractions of total chlorophyll-a were significantly correlated with water temperature (positively) and picoeukaryote cell number (negatively). We conducted a running fit of the model with increasing temperature and found a negative relationship between temperature and parameters C p m and C p , n m and a positive relationship between temperature and parameters D p and D p,n . By harnessing the relative red fluorescence of the flow cytometric data, we show that picoeukaryotes, which are higher in cell number in winter (cold) than summer (warm), contain higher chlorophyll per cell than other picophytoplankton and are slightly larger in size, possibly explaining the temperature shift in model parameters, though further evidence is needed to substantiate this finding. Our results emphasize the importance of knowing the water temperature and taxonomic composition of phytoplankton within each size class when understanding their relative contribution to total chlorophyll. Furthermore, our results have implications for the development of algorithms for inferring size-fractionated chlorophyll from satellite data, and for how the partitioning of total chlorophyll into the three size classes may change in a future ocean.

Remotely sensing harmful algal blooms in the Red Sea.

Harmful Algal Blooms (HABs) are of global concern, as their presence is often associated with socio-economic and environmental issues including impacts on public health, aquaculture and fisheries. Therefore, monitoring the occurrence and succession of HABs is fundamental for managing coastal regions around the world. Yet, due to the lack of adequate in situ measurements, the detection of HABs in coastal marine ecosystems remains challenging. Sensors on-board satellite platforms have sampled the Earth synoptically for decades, offering an alternative, cost-effective approach to routinely detect and monitor phytoplankton. The Red Sea, a large marine ecosystem characterised by extensive coral reefs, high levels of biodiversity and endemism, and a growing aquaculture industry, is one such region where knowledge of HABs is limited. Here, using high-resolution satellite remote sensing observations (1km, MODIS-Aqua) and a second-order derivative approach, in conjunction with available in situ datasets, we investigate for the first time the capability of a remote sensing model to detect and monitor HABs in the Red Sea. The model is able to successfully detect and generate maps of HABs associated with different phytoplankton functional types, matching concurrent in situ data remarkably well. We also acknowledge the limitations of using a remote-sensing based approach and show that regardless of a HAB's spatial coverage, the model is only capable of detecting the presence of a HAB when the Chl-a concentrations exceed a minimum value of ~ 1 mg m-3. Despite the difficulties in detecting HABs at lower concentrations, and identifying species toxicity levels (only possible through in situ measurements), the proposed method has the potential to map the reported spatial distribution of several HAB species over the last two decades. Such information is essential for the regional economy (i.e., aquaculture, fisheries & tourism), and will support the management and sustainability of the Red Sea's coastal economic zone.

Evaluating tropical phytoplankton phenology metrics using contemporary tools.

The timing of phytoplankton growth (phenology) in tropical oceans is a crucial factor influencing the survival rates of higher trophic levels, food web structure and the functioning of coral reef ecosystems. Phytoplankton phenology is thus categorised as an 'ecosystem indicator', which can be utilised to assess ecosystem health in response to environmental and climatic perturbations. Ocean-colour remote sensing is currently the only technique providing global, long-term, synoptic estimates of phenology. However, due to limited available in situ datasets, studies dedicated to the validation of satellite-derived phenology metrics are sparse. The recent development of autonomous oceanographic observation platforms provides an opportunity to bridge this gap. Here, we use satellite-derived surface chlorophyll-a (Chl-a) observations, in conjunction with a Biogeochemical-Argo dataset, to assess the capability of remote sensing to estimate phytoplankton phenology metrics in the northern Red Sea - a typical tropical marine ecosystem. We find that phenology metrics derived from both contemporary platforms match with a high degree of precision (within the same 5-day period). The remotely-sensed surface signatures reflect the overall water column dynamics and successfully capture Chl-a variability related to convective mixing. Our findings offer important insights into the capability of remote sensing for monitoring food availability in tropical marine ecosystems, and support the use of satellite-derived phenology as an ecosystem indicator for marine management strategies in regions with limited data availability.

Impacts of warming on phytoplankton abundance and phenology in a typical tropical marine ecosystem.

In the tropics, thermal stratification (during warm conditions) may contribute to a shallowing of the mixed layer above the nutricline and a reduction in the transfer of nutrients to the surface lit-layer, ultimately limiting phytoplankton growth. Using remotely sensed observations and modelled datasets, we study such linkages in the northern Red Sea (NRS) - a typical tropical marine ecosystem. We assess the interannual variability (1998-2015) of both phytoplankton biomass and phenological indices (timing of bloom initiation, duration and termination) in relation to regional warming. We demonstrate that warmer conditions in the NRS are associated with substantially weaker winter phytoplankton blooms, which initiate later, terminate earlier and are shorter in their overall duration (~ 4 weeks). These alterations are directly linked with the strength of atmospheric forcing (air-sea heat fluxes) and vertical stratification (mixed layer depth [MLD]). The interannual variability of sea surface temperature (SST) is found to be a good indicator of phytoplankton abundance, but appears to be less important for predicting bloom timing. These findings suggest that future climate warming scenarios may have a two-fold impact on phytoplankton growth in tropical marine ecosystems: 1) a reduction in phytoplankton abundance and 2) alterations in the timing of seasonal phytoplankton blooms.

The Gulf of Aden Intermediate Water Intrusion Regulates the Southern Red Sea Summer Phytoplankton Blooms.

Knowledge on large-scale biological processes in the southern Red Sea is relatively limited, primarily due to the scarce in situ, and satellite-derived chlorophyll-a (Chl-a) datasets. During summer, adverse atmospheric conditions in the southern Red Sea (haze and clouds) have long severely limited the retrieval of satellite ocean colour observations. Recently, a new merged ocean colour product developed by the European Space Agency (ESA)-the Ocean Color Climate Change Initiative (OC-CCI)-has substantially improved the southern Red Sea coverage of Chl-a, allowing the discovery of unexpected intense summer blooms. Here we provide the first detailed description of their spatiotemporal distribution and report the mechanisms regulating them. During summer, the monsoon-driven wind reversal modifies the circulation dynamics at the Bab-el-Mandeb strait, leading to a subsurface influx of colder, fresher, nutrient-rich water from the Indian Ocean. Using satellite observations, model simulation outputs, and in situ datasets, we track the pathway of this intrusion into the extensive shallow areas and coral reef complexes along the basin's shores. We also provide statistical evidence that the subsurface intrusion plays a key role in the development of the southern Red Sea phytoplankton blooms.