Warmer,deeper, and greener mixed layers in the North Atlantic subpolar gyre over the last 50 years

Shifts in global climate resonate in plankton dynamics, biogeochemical cycles, and marine food webs. We studied these linkages in the North Atlantic subpolar gyre (NASG), which hosts extensive phytoplankton blooms. We show that phytoplankton abundance increased since the 1960s in parallel to a deepening of the mixed layer and a strengthening of winds and heat losses from the ocean, as driven by the low frequency of the North Atlantic Oscillation (NAO). In parallel to these bottom‐up processes, the top‐down control of phytoplankton by copepods decreased over the same time period in the western NASG, following sea surface temperature changes typical of the Atlantic Multi‐decadal Oscillation (AMO). While previous studies have hypothesized that climate‐driven warming would facilitate seasonal stratification of surface waters and long‐term phytoplankton increase in subpolar regions, here we show that deeper mixed layers in the NASG can be warmer and host a higher phytoplankton biomass. These results emphasize that different modes of climate variability regulate bottom‐up (NAO control) and top‐down (AMO control) forcing on phytoplankton at decadal timescales. As a consequence, different relationships between phytoplankton, zooplankton, and their physical environment appear subject to the disparate temporal scale of the observations (seasonal, interannual, or decadal). The prediction of phytoplankton response to climate change should be built upon what is learnt from observations at the longest timescales.     

Trans‐Pacific and trans‐Arctic pathways of the intertidal macroalga Fucus distichus L. reveal multiple glacial refugia and colonizations from the North Pacific to the North Atlantic

Aim We examined the phylogeography of the cold‐temperate macroalgal species Fucus distichus L., a key foundation species in rocky intertidal shores and the only Fucus species to occur naturally in both the North Pacific and the North Atlantic. Location North Pacific and North Atlantic oceans (42° to 77° N). Methods We genotyped individuals from 23 populations for a mitochondrial DNA (mtDNA) intergenic spacer (IGS) (n = 608) and the cytochrome c oxidase subunit I (COI) region (n = 276), as well as for six nuclear microsatellite loci (n = 592). Phylogeographic structure and connectivity were assessed using population genetic and phylogenetic network analyses. Results IGS mtDNA haplotype diversity was highest in the North Pacific, and divergence between Pacific haplotypes was much older than that of the single cluster of Atlantic haplotypes. Two ancestral Pacific IGS/COI clusters led to a widespread Atlantic cluster. High mtDNA and microsatellite diversities were observed in Prince William Sound, Alaska, 11 years after severe disturbance by the 1989 Exxon Valdez oil spill. Main conclusions At least two colonizations occurred from the older North Pacific populations to the North Atlantic between the opening of the Bering Strait and the onset of the Last Glacial Maximum. One colonization event was from the Japanese Archipelago/eastern Aleutians, and a second was from the Alaskan mainland around the Gulf of Alaska. Japanese populations probably arose from a single recolonization event from the eastern Aleutian Islands before the North Pacific–North Atlantic colonization. In the North Atlantic, the Last Glacial Maximum forced the species into at least two known glacial refugia: the Nova Scotia/Newfoundland (Canada) region and Andøya (northern Norway). The presence of two private haplotypes in the central Atlantic suggests the possibility of colonization from other refugia that are now too warm to support F. distichus. With the continuing decline in Arctic ice cover as a result of global climate change, renewed contact between North Pacific and North Atlantic populations of Fucus species is expected.     

Temperature and zooplankton size structure: climate control and basin‐scale comparison in the North Pacific

The global distribution of zooplankton community structure is known to follow latitudinal temperature gradients: larger species in cooler, higher latitudinal regions. However, interspecific relationships between temperature and size in zooplankton communities have not been fully examined in terms of temporal variation. To re‐examine the relationship on a temporal scale and the effects of climate control thereon, we investigated the variation in copepod size structure in the eastern and western subarctic North Pacific in 2000–2011. This report presents the first basin‐scale comparison of zooplankton community changes in the North Pacific based on a fully standardized data set obtained from the Continuous Plankton Recorder (CPR) survey. We found an increase in copepod community size (CCS) after 2006–2007 in the both regions because of the increased dominance of large cold‐water species. Sea surface temperature varied in an east–west dipole manner, showing the typical Pacific Decadal Oscillation pattern: cooling in the east and warming in the west after 2006–2007. The observed positive correlation between CCS and sea surface temperature in the western North Pacific was inconsistent with the conventional interspecific temperature–size relationship. We explained this discrepancy by the geographical shift of the upper boundary of the thermal niche, the 9°C isotherm, of large cold‐water species. In the eastern North Pacific, the boundary stretched northeast, to cover a large part of the sampling area after 2006–2007. In contrast, in the western North Pacific, the isotherm location hardly changed and the sampling area remained within its thermal niche throughout the study period, despite the warming that occurred. Our study suggests that while a climate‐induced basin‐scale cool–warm cycle can alter copepod community size and might subsequently impact the functions of the marine ecosystem in the North Pacific, the interspecific temperature–size relationship is not invariant and that understanding region‐specific processes linking climate and ecosystem is indispensable.     

Reassessing regime shifts in the North Pacific: incremental climate change and commercial fishing are necessary for explaining decadal‐scale biological variability

In areas of the North Pacific that are largely free of overfishing, climate regime shifts – abrupt changes in modes of low‐frequency climate variability – are seen as the dominant drivers of decadal‐scale ecological variability. We assessed the ability of leading modes of climate variability [Pacific Decadal Oscillation (PDO), North Pacific Gyre Oscillation (NPGO), Arctic Oscillation (AO), Pacific‐North American Pattern (PNA), North Pacific Index (NPI), El Niño‐Southern Oscillation (ENSO)] to explain decadal‐scale (1965–2008) patterns of climatic and biological variability across two North Pacific ecosystems (Gulf of Alaska and Bering Sea). Our response variables were the first principle component (PC1) of four regional climate parameters [sea surface temperature (SST), sea level pressure (SLP), freshwater input, ice cover], and PCs 1–2 of 36 biological time series [production or abundance for populations of salmon (Oncorhynchus spp.), groundfish, herring (Clupea pallasii), shrimp, and jellyfish]. We found that the climate modes alone could not explain ecological variability in the study region. Both linear models (for climate PC1) and generalized additive models (for biology PC1–2) invoking only the climate modes produced residuals with significant temporal trends, indicating that the models failed to capture coherent patterns of ecological variability. However, when the residual climate trend and a time series of commercial fishery catches were used as additional candidate variables, resulting models of biology PC1–2 satisfied assumptions of independent residuals and out‐performed models constructed from the climate modes alone in terms of predictive power. As measured by effect size and Akaike weights, the residual climate trend was the most important variable for explaining biology PC1 variability, and commercial catch the most important variable for biology PC2. Patterns of climate sensitivity and exploitation history for taxa strongly associated with biology PC1–2 suggest plausible mechanistic explanations for these modeling results. Our findings suggest that, even in the absence of overfishing and in areas strongly influenced by internal climate variability, climate regime shift effects can only be understood in the context of other ecosystem perturbations.