Patterns and drivers of phytoplankton phenology off SW Iberia: A phenoregion based perspective

Phytoplankton patterns, tightly linked to the dynamics of the ocean surface layer and its atmospheric forcing, have major impacts on ecosystem functioning and are valuable indicators of its response to environmental variability and change. Phytoplankton phenology and its underlying drivers are spatially variable, and the study of its patterns, particularly over heterogeneous regions, benefits from a delineation of regions with specific phenological properties, or phenoregions. The area Southwest off the Iberian Peninsula (SWIP, NE Atlantic) integrates a highly complex set of coastal and ocean domains that collectively challenge the understanding of regional phytoplankton phenology and related forcing mechanisms. This study aims to evaluate phytoplankton phenology patterns over the SWIP area, during an 18-year period (September 1997 – August 2015), using an objective, unsupervised partition strategy (Hierarchical Agglomerative Clustering – HAC) based on phenological indices derived from satellite ocean colour data. The partition is then used to describe region-specific phytoplankton phenological patterns related to bloom magnitude, frequency, duration and timing. Region-specific variability patterns in phenological indices and their linkages with environmental determinants, including local ocean physical-chemical variables, hydrodynamic variables and large scale climate indices, were explored using Generalized Additive Models (GAM). HAC analyses identified five coherent phenoregions over SWIP, with distinctive phytoplankton phenological properties: two open ocean and three coastal regions. Over the open ocean, a single, low magnitude and long bloom event per year, was regularly observed. Coastal phenoregions exhibited up to six short bloom events per year, and higher intra-annual and variability. GAM models explained 50–90% of the variance of all phenological indices except bloom initiation timing, and revealed that interannual patterns in phytoplankton phenology and their environmental drivers varied markedly among the five phenoregions. Over the oceanic phenoregions, large-scale climate indices (Eastern Atlantic Pattern, Atlantic Meridional Oscillation), mixed layer depth (MLD) and nitrate concentration preceding primary bloom events were influential predictors, reflecting the relevance of nutrient limitation. For the Coastal-Slope, a relatively more light-limited phenoregion, North Atlantic Oscillation and wind speed were more relevant, and bloom magnitude was also positively influenced by riverine discharge. This variable was a significant predictor of bloom frequency, magnitude and duration over the Riverine-influenced region. Over the Upwelling-influenced region, upwelling intensity and mean annual MLD showed stronger partial effects on phytoplankton phenology. Overall, our phenology-based unsupervised approach produced a biologically-relevant SWIP partition, providing an evaluation of the complexity of interactions between phytoplankton and multiple environmental forcing, particularly over coastal areas