Changes in the oceanic northeast Pacific plankton populations; what may happen in a warmer ocean

The Continuous Plankton Recorder has been sampling the northeast Pacific on a routine basis since 2000. Although this is a relatively short time series still, climate variability within that time has caused noticeable related changes in the plankton. The earlier part of the time series followed the 1999 La Nina and conditions were cool, but conditions between 2003 and 2005 were anomalously warm. Oceanic zooplankton have responded to this warming in several ways that are discernible in CPR data. The seasonal cycle of mesozooplankton biomass in the eastern Gulf of Alaska has shifted earlier in the spring by a few weeks (sampling resolution is too coarse to be more accurate). The copepod Neocalanus plumchruslflemingeri is largely responsible as it makes up a high proportion of the spring surface biomass and stage-based determinations have shown an earlier maximum in warmer years across much of the northeast Pacific, spanning nearly 20 degrees of latitude. Summer copepod populations are more diverse than in spring, although lower in biomass. The northwards extension of southern taxa in the summer correlates with surface temperature and in warmer years southern taxa are found further north than in cooler years. These findings support the importance of monitoring the open ocean particularly as it is an important foraging ground for large fish, birds and mammals. Higher trophic levels may time their reproduction or migration to coincide with the abundance of particular prey which may be of a different composition and/or lower abundance at a particular time in warmer conditions.

Shortened duration of the annual Neocalanus plumchrus biomass peak in the Northeast Pacific

The calanoid copepod Neocalan us plumchrus (Marukawa) is a dominant member of the spring mesozooplankton in the subarctic North Pacific and Bering Sea. Previous studies have shown interdecadal and latitudinal variation in seasonal developmental timing, with peak biomass occurring earlier in years and places with warmer upper ocean temperatures. Because N. plumchrus normally has a single dominant annual cohort, its seasonal timing can be indexed from measurements of total population biomass or by following progressive changes in stage composition. Early studies empirically found that peak upper ocean biomass occurred when about half of the pre-dormant population had reached copepodite stage 5 (C5). However, more recent comparisons derived from recent Continuous Plankton Recorder (CPR) data now show peak biomass when a larger fraction (> 80%) of the population is at C5. CPR samples the surface 10 to 15 m, but comparisons to depth-resolved BIONESS data show that this discrepancy is not an artefact of sampling depth. Other causes are either a prolongation of duration of pre-dormant C5 or a narrowing of the age range making up the annual cohort. We assessed changes in cohort width using a modification of Greve's cumulative percentile method, and found that average cohort widths in the Alaska Gyre were significantly narrower in 2000-2007 than in 1957-1965 (1968-1980 were intermediate). Net tow sampling of Strait of Georgia populations showed a similar significant narrowing of cohorts in the 2003-2005 sampling period. This study provides evidence that in addition to the shift to an earlier occurrence of peak biomass reported previously, the duration of the peak has also decreased in the last decade.

Relationships between North Atlantic salmon, plankton and hydroclimatic change in the Northeast Atlantic

The abundance of wild salmon (Salmo salar) in the North Atlantic has declined markedly since the late 1980s as a result of increased marine mortality that coincided with a marked rise in sea temperature in oceanic foraging areas. There is substantial evidence to show that temperature governs the growth, survival, and maturation of salmon during their marine migrations through either direct or indirect effects. In an earlier study (2003), long-term changes in three trophic levels (salmon, zooplankton, and phytoplankton) were shown to be correlated significantly with sea surface temperature (SST) and northern hemisphere temperature (NHT). A sequence of trophic changes ending with a stepwise decline in the total nominal catch of North Atlantic salmon (regime shift in ∼1986/1987) was superimposed on a trend to a warmer dynamic regime. Here, the earlier study is updated with catch and abundance data to 2010, confirming earlier results and detecting a new abrupt shift in ∼1996/1997. Although correlations between changes in salmon, plankton, and temperature are reinforced, the significance of the correlations is reduced because the temporal autocorrelation of time-series substantially increased due to a monotonic trend in the time-series, probably related to global warming. This effect may complicate future detection of effects of climate change on natural systems.

Synchronous response of marine plankton ecosystems to climate in the Northeast Atlantic and the North Sea

Over the last few decades, global warming has accelerated both the rate and magnitude of changes observed in many functional units of the Earth System. In this context, plankton are sentinel organisms because they are sensitive to subtle levels of changes in temperature and might help in identifying the current effects of climate change on pelagic ecosystems. In this paper, we performed a comparative approach in two regions of the North Atlantic (i.e. the Northeast Atlantic and the North Sea) to explore the relationships between changes in marine plankton, the regional physico-chemical environment and large-scale hydro-climatic forcing using four key indices: the North Atlantic Oscillation (NAO), the Atlantic Multidecadal Oscillation (AMO), the East Atlantic (EA) pattern and Northern Hemisphere Temperature (NHT) anomalies. Our analyses suggest that long-term changes in the states of the two ecosystems were synchronous and correlated to the same large-scale hydro-climatic variables: NHT anomalies, the AMO and to a lesser extent the EA pattern. No significant correlation was found between long-term ecosystem modifications and the state of the NAO. Our results suggest that the effect of climate on these ecosystems has mainly occurred in both regions through the modulation of the thermal regime.

Volcanic ash fuels anomalous plankton bloom in subarctic northeast Pacific

Using multiple lines of evidence, we demonstrate that volcanic ash deposition in August 2008 initiated one of the largest phytoplankton blooms observed in the subarctic North Pacific. Unusually widespread transport from a volcanic eruption in the Aleutian Islands, Alaska deposited ash over much of the subarctic NE Pacific, followed by large increases in satellite chlorophyll. Surface ocean pCO2, pH, and fluorescence reveal that the bloom started a few days after ashfall. Ship-based measurements showed increased dominance by diatoms. This evidence points toward fertilization of this normally iron-limited region by ash, a relatively new mechanism proposed for iron supply to the ocean. The observations do not support other possible mechanisms. Extrapolation of the pCO2 data to the area of the bloom suggests a modest ∼0.01 Pg carbon export from this event, implying that even large-scale iron fertilization at an optimum time of year is not very efficient at sequestering atmospheric CO2.

Distribution and multi-annual abundance trends of the copepod Temora longicornis in the US Northeast shelf ecosystem

The average spatial distribution and annual abundance cycle are described for the copepod Temora longicornis from samples collected on broadscale surveys (1977-2006) and along continuous plankton recorder transects (1961-2006) of the US Northeast continental shelf ecosystem. After its annual low in winter, T. longicornis abundance begins to increase in coastal waters with the northern progression of spring conditions. Annual maximum shelf concentrations were found in the more southern inshore waters of the region during the summer months. Abundance throughout most of the ecosystem increased sharply in the early 1990s and remained high through 2001. During this period, the copepod became more numerous and widespread in offshore shelf waters. Abundance declined to approximately average levels in 2002 for the remainder of the time series, but its extended offshore range remained intact. Correlation analysis found that the copepods interannual abundance variability had a significant negative relationship with surface salinity anomalies throughout the ecosystem, with higher correlations found in the northernmost subareas. Temora longicornis abundance in the ecosystem's southernmost subarea (Middle Atlantic Bight) did not increase in the 1990s and was found to be negatively correlated to surface temperature, indicating that continued global warming could adversely impact the copepods annual abundance cycle in this region.

A blooming jellyfish in the northeast Atlantic and Mediterranean

A long-term time series of plankton records collected by the continuous plankton recorder (CPR) Survey in the northeast Atlantic indicates an increased occurrence of Cnidaria since 2002. In the years 2007 and 2008, outbreaks of the warm-temperate scyphomedusa, Pelagia noctiluca, appeared in CPR samples between 45° N to 58° N and 1° W to 26° W. Knowing the biology of this species and its occurrence in the adjacent Mediterranean Sea, we suggest that P. noctiluca may be exploiting recent hydroclimatic changes in the northeast Atlantic to increase its extent and intensity of outbreaks. In pelagic ecosystems, Cnidaria can affect fish recruitment negatively. Since P. noctiluca is a highly venomous species, outbreaks can also be detrimental to aquaculture and make bathing waters unusable, thus having profound ecological and socio-economic consequences.

Effects on zooplankton of a warmer ocean: Recent evidence from the Northeast Pacific

The consequences for pelagic communities of warming trends in mid and high latitude ocean regions could be substantial, but their magnitude and trajectory are not yet known. Environmental changes predicted by climate models (and beginning to be confirmed by observations) include warming and freshening of the upper ocean and reduction in the extent and duration of ice cover. One way to evaluate response scenarios is by comparing how "similar" zooplankton communities have differed among years and/or locations with differing temperature. The subarctic Pacific is a strong candidate for such comparisons, because the same mix of zooplankton species dominates over a wide range of temperature climatologies, and observations have spanned substantial temperature variability at interannual-to-decadal time scales. In this paper, we review and extend copepod abundance and phenology time series from net tow and Continuous Plankton Recorder surveys in the subarctic Northeast Pacific. The two strongest responses we have observed are latitudinal shifts in centers of abundance of many species (poleward under warm conditions), and changes in the life cycle timing of Neocalanus plumchrus in both oceanic and coastal regions (earlier by several weeks in warm years and at warmer locations). These zooplankton data, plus indices of higher trophic level responses such as reproduction, growth and survival of pelagic fish and seabirds, are all moderately-to-strongly intercorrelated (vertical bar r vertical bar = 0.25-0.8) with indices of local and basin-scale temperature anomalies. A principal components analysis of the normalized anomaly time series from 1979 to 2004 shows that a single "warm-and-low-productivity" vs. "cool-and-high-productivity" component axis accounts for over half of the variance/covariance. Prior to 1990, the scores for this component were negative ("cool" and "productive") or near zero except positive in the El Nino years 1983 and 1987. The scores were strongly and increasingly positive ("warm" and "low productivity") from 1992 to 1998; negative from 1999 to 2002; and again increasingly positive from 2003-present. We suggest that, in strongly seasonal environments, anomalously high temperature may provide misleading environmental cues that contribute to timing mismatch between life history events and the more-nearly-fixed seasonality of insolation, stratification, and food supply. Crown Copyright (c) 2007 Published by Elsevier Ltd. All rights reserved.

Variation among northeast Atlantic regions in the responses of zooplankton to climate change: Not all areas follow the same path

Broad scale climate forcing can interact with local environmental processes to affect the observed ecological phenomena. This causes potential problems of over-extrapolation for results from a limited number of sites or the averaging out of region-specific responses if data from too wide an area are combined. In this study, an area similar in extent to the Celtic Biscay Large Marine Ecosystem, but including off-shelf areas, was partitioned using clustering of satellite chlorophyll (chl-a) measurements. The resulting clusters were used to define areas over which to combine copepod data from the Continuous Plankton Recorder. Following filtering due to data limitations, nine regions were defined with sufficient records for analysis. These regions were consistent with known oceanographic structure in the study area. Off-shelf regions showed a progressively later timing in the seasonal peak of chl-a measurements moving northwards. Generalised additive models were used to estimate seasonal and multiannual signals in the adult and juvenile stages of Calanus finmarchicus, C. helgolandicus and the Paracalanus–Pseudocalanus group. Associations between variables (sea surface temperature (SST), phenology and annual abundance) differed among taxonomic groups, but even within taxonomic groups, relationships were not consistent across regions. For example, in the deep waters off Spain and Portugal the annual abundance of Calanus finmarchicus has a weak positive association with SST, in contrast to the pattern in most other regions. The regions defined in this study provide an objective basis for investigations into the long term dynamics of plankton populations and suggest suitable sub regions for deriving pelagic system indicators.

Trophodynamics and stability of regional scale ecosystems in the Northeast Atlantic

This study addresses the long-term stability of three trophic groupings in the Northeast Atlantic at regional scales. The most abundant taxa representing phytoplankton, herbivorous copepods, and carnivorous zooplankton were examined from the Continuous Plankton Recorder database. Multivariate control charts using a Bray–Curtis similarity metric were used to assess whether fluctuations within trophic groupings were within or beyond the expected variability. Two evaluation periods were examined: annual changes between 1960 and 1999 (2000–2009 baseline) and recent changes between 2000 and 2009 (1960–1999 baseline). The trends over time in abundance/biomass of trophic levels were region-specific, especially in carnivorous copepods, where abundance did not mirror trends in the overall study area. The stability of phytoplankton was within the expected limits, although not in 2008 and 2009. Higher trophic levels were less stable, perhaps reflecting the added complexity of interactions governing their abundance. In addition, some regions were consistently less stable than others. Correlations in stability between adjacent trophic levels were positive at large marine ecosystem scale but generally non-significant at regional scales. The study suggests that certain regions may be particularly vulnerable to periods of instability in community structure. The benefits of using the control chart method rather than other multivariate measures of plankton dynamics are discussed.

From silk to satellite: half a century of ocean colour anomalies in the Northeast Atlantic

Changes in phytoplankton dynamics influence marine biogeochemical cycles, climate processes, and food webs, with substantial social and economic consequences. Large-scale estimation of phytoplankton biomass was possible via ocean colour measurements from two remote sensing satellites – the Coastal Zone Color Scanner (CZCS, 1979-1986) and the Sea-viewing Wide Field-of-view Sensor (SeaWiFS, 1998-2010). Due to the large gap between the two satellite eras and differences in sensor characteristics, comparison of the absolute values retrieved from the two instruments remains challenging. Using a unique in situ ocean colour dataset that spans more than half a century, the two satellite-derived chlorophyll-a (Chl-a) eras are linked to assess concurrent changes in phytoplankton variability and bloom timing over the Northeast Atlantic Ocean and North Sea. Results from this unique re-analysis reflect a clear increasing pattern of Chl-a, a merging of the two seasonal phytoplankton blooms producing a longer growing season and higher seasonal biomass, since the mid-1980s. The broader climate plays a key role in Chl-a variability as the ocean colour anomalies parallel the oscillations of the Northern Hemisphere Temperature (NHT) since 1948.

Reply to comment: is climate change the most likely driver of range expansion of a critically endangered top predator in northeast Atlantic waters?

The comment by Votier et al. (2008) on our recently published article (Wynn et al. 2007) contains two main criticisms: (i) that our analytical approach is inappropriate and (ii) that we have failed to acknowledge other factors that may have contributed to the change in Balearic Shearwater numbers recorded throughout northwest European waters. We strongly disagree with both these criticisms.