Can the subtropical North Atlantic permanent thermocline be observed from space?

The analysis of remotely sensed altimeter data and in situ measurements shows that ERS 2 radar can monitor the ocean permanent thermocline from space. The remotely sensed sea level anomaly data account for similar to 2/3 of the temperature variance or vertical displacement of isotherms at a depth of similar to 550 m in the Subtropical North Atlantic Ocean near 32.5 degree N. This depth corresponds closely to the region of maximum temperature gradient in the permanent thermocline where near semi-annual internal vertical displacements reach 200 to 300 m. The gradient of the altimeter sea level anomaly data correlates well with measured ocean currents to a depth of 750 m. It is shown that observations from space can account for similar to 3/4 of the variance of ocean currents measured in situ in the permanent thermocline over a 2-y period. The magnification of the permanent thermocline displacement with respect to the displacement of the sea surface was determined as - x650 and gives a measure of the ratio of barotropic to baroclinic decay scale of geostrophic current with depth. The overall results are used to interpret an eight year altimeter data tie series in the Subtropical North Atlantic at 32.5 degree N which shows a dominant wave or eddy period near 200 days, rather than semi-annual and increases in energy propagating westward in 1995 (west of 25 degree W). The effects of rapid North Atlantic Oscillation climate change on ocean circulation are discussed. The altimeter data for the Atlantic were Fourier analysed. It is shown how the annual and semi-annual components relate to the seasonal maximum cholorophyll-a SeaWiFS signal in tropical and equatorial regions due to the lifting of the thermocline caused by seasonally varying ocean currents forced by wind stress.

Ocean structure and climate (Eastern North Atlantic): in situ measurement and remote sensing

Structure and climate of the east North Atlantic are appraised within a framework of in situ measurement and altimeter remote sensing from 0 degree - 60 degree N. Long zonal expendable bathythermograph /conductivity-temperature-depth probe sections show repeating internal structure in the North Atlantic Ocean. Drogued buoys and subsurface floats give westward speeds for eddies and wavelike structure. Records from longterm current meter deployments give the periodicity of the repeating structure. Eddy and wave characteristics of period, size or wavelength, westward propagation speed, and mean currents are derived at 20 degree N, 26 degree N, 32.5 degree N, 36 degree N and 48 degree N from in situ measurements in the Atlantic Ocean. It is shown that ocean wave and eddy-like features measured in situ correlate with altimeter structure. Interior ocean wave crests or cold dome-like temperature structures are cyclonic and have negative surface altimeter anomalies; mesoscale internal wave troughs or warm structures are anticyclonic and have positive surface height anomalies. Along the Eastern Boundary, flows and temperature climate are examined in terms of sla and North Atlantic Oscillation (NAO) Index. Longterm changes in ocean climate and circulation are derived from sla data. It is shown that longterm changes from 1992 to 2002 in the North Atlantic Current and the Subtropical Gyre transport determined from sla data correlate with winter NAO Index such that maximum flow conditions occurred in 1995 and 2000. Minimum circulation conditions occurred between 1996-1998. Years of extreme negative winter NAO Index result in enhanced poleward flow along the Eastern Boundary and anomalous winter warming along the West European Continental Slope as was measured in 1990, 1996, 1998 and 2001.

Spatial, seasonal and long-term fluctuations of plankton in relation to hydroclimatic features in the English Channel, Celtic Sea and Bay of Biscay

Plankton collected by the Continuous Plankton Recorder (CPR) survey were investigated for the English Channel, Celtic Sea and Bay of Biscay from 1979 to 1995. The main goal was to study the relationship between climate and plankton and to understand the factors influencing it. In order to take into account the spatial and temporal structure of biological data, a three-mode principal component analysis (PCA) was developed. It not only identified 5 zones characterised by their similar biological composition and by the seasonal and inter-annual evolution of the plankton, it also made species associations based on their location and year-to-year change. The studied species have stronger year-to-year fluctuations in abundance over the English Channel and Celtic Sea than the species offshore in the Bay of Biscay. The changes in abundance of plankton in the English Channel are negatively related to inter-annual changes of climatic conditions from December to March (North Atlantic Oscillation [NAO] index and air temperature). Thus, the negative relationship shown by Fromentin and Planque (1996; Mar Ecol Prog Ser 134:111-118) between year-to-year changes of Calanus finmarchicus abundance in the northern North Atlantic and North Sea and NAO was also found for the most abundant copepods in the Channel. However, the hypothesis proposed to explain the plankton/NAO relationship is different for this region and a new hypothesis is proposed. In the Celtic Sea, a relationship between the planktonic assemblage and the air temperature was detected, but it is weaker than for the English Channel. No relationship was found for the Bay of Biscay. Thus, the local physical environment and the biological composition of these zones appear to modify the relationship between winter climatic conditions and the year-to-year fluctuations of the studied planktonic species. This shows, therefore, that the relationship between climate and plankton is difficult to generalise.

Is observed variability in the long-term results of the Continuous Plankton Recorder survey a response to climate change?

In the more than 50 years that the Continuous Plankton Recorder (CPR) survey has operated on a regular monthly basis in the north-east Atlantic and North Sea, large changes have been witnessed in the planktonic ecosystem. These changes have taken the form of long-term trends in abundance for certain species or stepwise changes for others, and in many cases are correlated with a mode of climatic variability in the North Atlantic, either: (1) the North Atlantic Oscillation (NAO), a basin-scale atmospheric alteration of the pressure field between the Azores high pressure cell and the Icelandic Low; or (2) the Gulf Stream Index (GSI), which measures the latitudinal position of the north wall of the Gulf Stream. Recent work has shown that the changes in the GSI are coupled with the NAO and Pacific Southern Oscillation with a 2 year lag. The plankton variability is also possibly linked to changes observed in the distribution and flux of water masses in the surface, intermediate and deep waters of the North Atlantic. For example, in the last two decades, the extent and location of the formation of North Atlantic Deep Water, Labrador Sea Intermediate Water and Norwegian Sea intermediate and upper-layer water has altered considerably. This paper discusses the extent to which observed changes in plankton abundance and distribution may be linked to this basin-scale variability in hydrodynamics. The results are also placed within the context of global climate warming and the possible effects of the observed melting of Arctic permafrost and sea ice on the subpolar North Atlantic.

Variations in the distributions of the Centropages chierchiae and Temora stylifera (Copepoda: Calanoida) in the north-eastern Atlantic Ocean and western european shelf waters.

Centropages chierchiae and Temora stylifera occurred rarely in the Continuous Plankton Recorder (CPR) survey in the Bay of Biscay, Celtic Sea, and English Channel before 1988. By 2000 they were found frequently and in abundance. The seasonal cycles of abundance of these species differ, C. chierchiae occurring mainly in the summer while T. stylifera was found most frequently in late autumn or winter towards the northern limits of its distribution. The increase in abundance of both species is related to temperature. However, in the years when it was found in the samples, the frequency of occurrence of C. chierchiae was correlated positively with the strength of the shelf edge current and negatively with the North Atlantic Oscillation (NAO) while the reverse was true for T. stylifera.

Interregional Biological Responses in the North Atlantic to Hydrometeorological Forcing

Using data from the CPR survey seven case studies are described that document different spatial and temporal responses in the plankton to hydroclimatic events. Long-term trends in the plankton of the eastern Atlantic and the North Sea over the last five decades are examined. Two of the examples revisit correlations that have been described between copepod abundance in the eastern Atlantic and North Sea and indices of atmospheric variability, the North Atlantic Oscillation index and the Gulf Stream North Wall index. Evidence for an increase in levels of Phytoplankton Colour (a visual index of chlorophyll) on the eastern and western sides of the Atlantic is presented. Changes in three trophic levels and in the hydrodynamics and chemistry of the North Sea circa 1988 are outlined as a regime shift. Two of the case studies emphasise the importance of variability in oceanic advection into shelf seas and the role of western and eastern margin currents at the shelf edge. The plankton appear to be integrating hydrometeorological signals and reflecting basin scale changes in circulation of surface, intermediate and deep waters in part associated with the NAO. The extent to which climatic variability may be contributing to the observed changes in the plankton is discussed with a forecast of potential future ecosystem effects in a climate change scenario.

Arctic boreal plankton species in the Northwest Atlantic

The Continuous Plankton Recorder (CPR) survey has sampled regularly in the Northwest Atlantic since the early 1960s. Over the last decade there has been a dramatic increase in the abundance of a number of arctic boreal plankton species, notably Calanus hyperboreus (Kroyer), Calanus glacialis (Jaschnov), and Ceratium arcticum, and a southerly shift of the copepod C. hyperboreus in the CPR survey. In 1998, C. hyperboreus was recorded at its farthest position south in the survey, 39 degrees N, off the Georges Bank shelf edge. Other studies have reported similar parallel biological responses on three trophic levels. During the late 1990s, production of Labrador Sea Water (LSW) has been at a high, a direct response to the phase of the North Atlantic Oscillation (NAO). The increase in abundance of these species, up to four standard deviations from the long-term mean, is linked to variability in the hydrography of the area and the driving climatic processes of the North Atlantic.

Climate-related increases in jellyfish frequency suggest a more gelatinous future for the North Sea

Data obtained since 1958 from the continuous plankton recorder show an increasing occurrence of jellyfish in the central North Sea that is positively related to the North Atlantic Oscillation (NAO) and Atlantic inflow to the northern North Sea. Since 1970, jellyfish frequency has been also significantly negatively correlated with mean annual pH, independent of NAO trends. Jellyfish frequency increased in the mid-1980s, coincident with the reported regime shift in the North Sea and tracking trends in phytoplankton color. As models produced under all climate-change scenarios indicate a move toward a positive NAO, and pH of the oceans is predicted to decrease with rising CO2, we suggest that jellyfish frequency will increase over the next 100 yr.

North Atlantic climatic signals and the plankton of the European Continental Shelf

Climatic variability on the European Continental Shelf is dominated by events over the North Atlantic Ocean, and in particular by the North Atlantic Oscillation (NAO). The NAO is essentially a winter phenomenon, and its effects will be felt most strongly by populations for which winter conditions are critical. One example is the copepod Calanus finmarchicus, whose northern North Sea populations overwinter at depth in the North Atlantic. Its annual abundance in this region is strongly dependent on water transports at the end of the winter, and hence on the NAO index. Variations in the NAO give rise to changes in the circulation of the North Atlantic Ocean, with additional perturbations arising from El Ni (n) over tildeo - Southern Oscillation (ENSO) events in the Pacific, and these changes can be delayed by several years because of the adjustment time of the ocean circulation. One measure of the circulation is the latitude of the north wall of the Gulf Stream (GSNW index). Interannual variations in the plankton of the Shelf Seas show strong correlations with the fluctuations of the GSNW index, which are the result of Atlantic-wide atmospheric processes. These associations imply that the interannual variations are climatically induced rather than due to natural fluctuations of the marine ecosystem, and that the zooplankton populations have not been significantly affected by anthropogenic processes such as nutrient enrichment or fishing pressure. While the GSNW index represents a response to atmospheric changes over two or more years, the zooplankton populations correlated with it have generation times of a few weeks. The simplest explanation for the associations between the zooplankton and the GSNW index is that the plankton are responding to weather patterns propagating downstream from the Gulf Stream system. It seems that these meteorological processes operate in the spring. Although it has been suggested that there was a regime shift in the North Sea in the late 1980s, examination of the time-series by the cumulative sum (CUSUM) technique shows that any changes in the zooplankton of the central and northern North Sea are consistent with the background climatic variability. The abundance of total copepods increased during this period but this change does not represent a dramatic change in ecosystem processes. It is possible some change may have occurred at the end of the time-series in the years 1997 and 1998.

Coccolithophore surface distributions in the North Atlantic and their modulation of the air-sea flux of CO2 from 10 years of satellite Earth observation data

Coccolithophores are the primary oceanic phytoplankton responsible for the production of calcium carbonate (CaCO3). These climatically important plankton play a key role in the oceanic carbon cycle as a major contributor of carbon to the open ocean carbonate pump (similar to 50 %) and their calcification can affect the atmosphere-to-ocean (air-sea) uptake of carbon dioxide (CO2) through increasing the seawater partial pressure of CO2 (pCO(2)). Here we document variations in the areal extent of surface blooms of the globally important coccolithophore, Emiliania huxleyi, in the North Atlantic over a 10-year period (1998-2007), using Earth observation data from the Sea-viewing Wide Field-of-view Sensor (SeaWiFS). We calculate the annual mean sea surface areal coverage of E. huxleyi in the North Atlantic to be 474 000 +/- 104 000 km(2), which results in a net CaCO3 carbon (CaCO3-C) production of 0.14-1.71 Tg CaCO3-C per year. However, this surface coverage (and, thus, net production) can fluctuate inter-annually by -54/+81% about the mean value and is strongly correlated with the El Nino/Southern Oscillation (ENSO) climate oscillation index (r = 0.75, p < 0.02). Our analysis evaluates the spatial extent over which the E. huxleyi blooms in the North Atlantic can increase the pCO(2) and, thus, decrease the localised air-sea flux of atmospheric CO2. In regions where the blooms are prevalent, the average reduction in the monthly air-sea CO2 flux can reach 55%. The maximum reduction of the monthly air-sea CO2 flux in the time series is 155 %. This work suggests that the high variability, frequency and distribution of these calcifying plankton and their impact on pCO(2) should be considered if we are to fully understand the variability of the North Atlantic air-to-sea flux of CO2. We estimate that these blooms can reduce the annual N. Atlantic net sink atmospheric CO2 by between 3-28 %.

Variation in ocean colour may help predict cod and haddock recruitment

Characteristics of the spring and fall phytoplankton blooms in spawning areas on the Scotian Shelf, Canada, were estimated from remote sensing data. These blooms, along with anomalies in the North Atlantic Oscillation, were used to explain variation in the recruitment of 4 populations of cod and haddock. We tested the effects of the timing of the bloom using the chlorophyll a (chl a) signal, the maximum amount of chl a, the timing of the diatom bloom, and the maximum relative dominance of diatoms on the recruitment (to Age 1) of cod and haddock on the Scotian Shelf. Models were run separately for the effects of the spring and fall blooms. Only 3 of 10 models tested (0-lag) explained significant (80 to 92%) variation in recruitment. However, the performance of these models was not consistent across populations or species, suggesting that generalities about how spring and fall phytoplankton blooms affect recruitment cannot yet be made. The differences among models suggest that fish larvae are probably adapted locally to food production and thus indirectly to the characteristics of the phytoplankton bloom, which in turn are influenced by regional (meso-scale) oceanographic conditions.

Changes in significant and maximum wave heights in the Norwegian Sea

This paper analyses 10 years of in-situ measurements of significant wave height (Hs) and maximum wave height (Hmax) from the ocean weather ship Polarfront in the Norwegian Sea. The 30-minute Ship-Borne Wave Recorder measurements of Hmax and Hs are shown to be consistent with theoretical wave distributions. The linear regression between Hmax and Hs has a slope of 1.53. Neither Hs nor Hmax show a significant trend in the period 2000–2009. These data are combined with earlier observations. The long-term trend over the period 1980–2009 in annual Hs is 2.72 ± 0.88 cm/year. Mean Hs and Hmax are both correlated with the North Atlantic Oscillation (NAO) index during winter. The correlation with the NAO index is highest for the more frequently encountered (75th percentile) wave heights. The wave field variability associated with the NAO index is reconstructed using a 500-year NAO index record. Hs and H max are found to vary by up to 1.42 m and 3.10 m respectively over the 500-year period. Trends in all 30-year segments of the reconstructed wave field are lower than the trend in the observations during 1980–2009. The NAO index does not change significantly in 21st century projections from CMIP5 climate models under scenario RCP85, and thus no NAO-related changes are expected in the mean and extreme wave fields of the Norwegian Sea.

Variability in northwards extension of warm water copepods in the NE Pacific

The Continuous Plankton Recorder has been deployed in the NE Pacific on two intersecting transects since 2000. Many deployments included a temperature sensor providing in situ temperature data to supplement the species abundance data for 1300 samples. Twenty-nine copepod taxa were sufficiently abundant to examine their temperature-related distributions. Groups of warm- and cold-water species were identified, with overlapping distributions between 48 and 588N. Recent fluctuations in ocean climate, from the warmest year on record in 2005 to one of the coldest in decades in 2008, provided ideal conditions to observe temperature-related interannual variability. The abundance and northwards extension of warm water species were significantly positively correlated with mean annual temperature and the Pacific Decadal Oscillation. The cold water species showed no correlations with temperature/Pacific Decadal Oscillation (PDO) within the study region; however, if the 4 years of sampling that extended south to 398N were examined separately, there was a strong relationship between temperature/PDO and the southern extent of subarctic copepods. Under warm ocean conditions, the range overlap of the two groups will increase as warm water species extend northwards, causing an increase in copepod diversity. Since warm water species are generally smaller and nutritionally poorer, this has implications for higher trophic levels

Acquisition of continuous plankton recorder data

The Continuous Plankton Recorder has been deployed on a seasonal basis in the north Pacific since 2000, accumulating a database of abundance measurements for over 290 planktonic taxa in over 3,500 processed samples. There is an additional archive of over 10,000 samples available for further analyses. Exxon Valdez Oil Spill Trustee Council financial support has contributed to about half of this tally, through four projects funded since 2002. Time series of zooplankton variables for sub-regions of the survey area are presented together with abstracts of eight papers published using data from these projects. The time series covers a period when the dominant climate signal in the north Pacific, the Pacific Decadal Oscillation (PDO), switched with unusual frequency between warm/positive states (pre-1999 and 2003-2006) and cool/negative states (1999-2002 and 2007). The CPR data suggest that cool negative years show higher biomass on the shelf and lower biomass in the open ocean, while the reverse is true in warm (PDO positive) years with lower shelf biomass (except 2005) and higher oceanic biomass. In addition, there was a delay in plankton increase on the Alaskan shelf in the colder spring of 2007, compared to the warmer springs of the preceding years. In warm years, smaller species of copepods which lack lipid reserves are also more common. Availability of the zooplankton prey to higher trophic levels (including those that society values highly) is therefore dependent on the timing of increase and peak abundance, ease of capture and nutritional value. Previously published studies using these data highlight the wide-ranging applicability of CPR data and include collaborative studies on; phenology in the key copepod species Neocalanus plumchrus, descriptions of distributions of decapod larvae and euphausiid species, the effects of hydrographic features such as mesoscale eddies and the North Pacific Current on plankton populations and a molecularbased investigation of macro-scale population structure in N. cristatus. The future funding situation is uncertain but the value of the data and studies so far accumulated is considerable and sets a strong foundation for further studies on plankton dynamics and interactions with higher trophic levels in the northern Gulf of Alaska.

Unanticipated biological changes and global warming

Evidence of global warming is now unequivocal, and studies suggest that it has started to influence natural systems of the planet, including the oceans. However, in the marine environment, it is well-known that species and ecosystems can also be influenced by natural sources of large-scale hydro-climatological variability. The North Atlantic Oscillation (NAO) was negatively correlated with the mean abundance of one of the subarctic key species Calanus finmarchicus in the North Sea. This correlation was thought to have broken down in 1996, however, the timing has never been tested statistically. The present study revisits this unanticipated change and reveals that the correlation did not break down in 1996 as originally proposed but earlier, at the time of an abrupt ecosystem shift in the North Sea in the 1980s. Furthermore, the analyses demonstrate that the correlation between the NAO and C. finmarchicus abundance is modulated by the thermal regime of the North Sea, which in turn covaries positively with global temperature anomalies. This study thereby provides evidence that global climate change is likely to alter some empirical relationships found in the past between species abundance or the ecosystem state and large-scale natural sources of hydro-climatological variability. A theory is proposed to explain how this might happen. These unanticipated changes, also called ‘surprises’ in climatic research, are a direct consequence of the complexity of both climatic and biological systems. In this period of rapid climate change, it is therefore hazardous to integrate meteo-oceanic indices such as the NAO in models used in the management of living resources, as it has been sometimes attempted in the past.