Revealing pelagic habitat use: the tagging of Pacific pelagics program

Tagging of Pacific pelagics (TOPP) is a pilot program of the Census of marine life (CoML) that will lead to understanding of pelagic habitat use by marine vertebrates and large squid in the North Pacific. Taking a multispecies approach, the TOPP project will use a range of electronic tag technologies to put the distribution and behavior of pelagic organisms in the context of the oceanography of the North Pacific. Tag-bearing animals will be used as autonomous ocean profilers to enhance sparse oceanographic observations for vast ocean regions. These autonomous ocean samplers will provide unprecedented coverage of the water column structure of the North Pacific. The temporal and spatial data generated by this project will provide an “organism-eye” view of several interactive oceanic regimes in the North Pacific. Twenty target species, including tunas, sharks, pinnipeds, cetaceans, seabirds, and marine turtles, will be monitored with electronic tags. Animal-collected oceanic data will be assimilated into global ocean databases, complement traditional methodologies and be used to help validate nearshore, regional, and basin scale ocean models. As more environmental information is gathered and delivered from the tagged animals, new insights will be obtained about their individual behaviors, as well as how diverse species have separately evolved to forage, reproduce, and survive in the vast pelagic environment. This multi-disciplinary approach will allow a novel merger of biological and physical data to provide a new understanding of the relationship between the movements and behaviors of marine organisms and oceanographic processes in the eastern North Pacific.     

Pacific Basin climate variability and patterns of Northeast Pacific marine fish production

A review of oceanographic and climate data from the North Pacific and Bering Sea has revealed climate events that occur on two principal time scales: a) 2–7 years (i.e. El Niño Southern Oscillation, ENSO), and b) inter-decadal (i.e. Pacific Decadal Oscillation, PDO). The timing of ENSO events and of related oceanic changes at higher latitudes were examined. The frequency of ENSO was high in the 1980s. Evidence of ENSO forcing on ocean conditions in the North Pacific (Niño North conditions) showed ENSO events were more frequently observed along the West Coast than in the western Gulf of Alaska (GOA) and Eastern Bering Sea (EBS). Time series of catches for 30 region/species groups of salmon, and recruitment data for 29 groundfish and 5 non-salmonid pelagic species, were examined for evidence of a statistical relationship with any of the time scales associated with Niño North conditions or the PDO. Some flatfish stocks exhibited high autocorrelation in recruitment coupled with a significant step in recruitment in 1977 suggesting a relationship between PDO forcing and recruitment success. Five of the dominant gadid stocks (EBS and GOA Pacific cod, Pacific hake and EBS and GOA walleye pollock) exhibited low autocorrelation in recruitment. Of these, Pacific hake, GOA walleye pollock and GOA Pacific cod exhibited significantly higher incidence of strong year classes in years associated with Niño North conditions. These findings suggest that the PDO and ENSO may play an important role in governing year-class strength of several Northeast Pacific marine fish stocks.     

Marine ecosystem variability and human community responses: The example of Ghana,West Africa

This study describes variability in the marine ecosystem of Ghana, West Africa, on several temporal and spatial scales and discusses how the human communities using this ecosystem respond to this variability to cope socially and economically. Ghanaian marine waters are part of an upwelling system with strong seasonal and inter-annual variability. Much of this variability is forced at large spatial scales in the tropical Atlantic and by El Niño—Southern Oscillation events in the Pacific Ocean, which influence inter-annual variability of sea surface temperature and pelagic fish landings off Ghana. At decadal scales, Ghanaian marine waters experienced cool sea temperatures and low fishery landings during the 1960s, rapid warming and increases in fishery landings during the late 1970s and 1980s, and variable temperatures and fishery landings during the 1990s. In the late 1990s, pelagic and demersal fish populations appeared to be declining, partly due to over-fishing, although the per capita supply (domestic production plus net imports) of fish was kept high by increased imports. Artisanal fishers and fishing communities in Ghana have devised strategies to deal with variability on seasonal and inter-annual scales. These livelihood strategies include: (i) exploiting marine and terrestrial natural resources more intensively, initially at local scales but expanding to regional scales; (ii) ensuring multiple and diversified income sources; (iii) investing in social relationships and communities for support; and (iv) undertaking seasonal or permanent migrations. In addition, the national government imports fish to deal with shortages. However, these strategies may be less adapted to variability at decadal scales, and may not be sustainable when viewed at the larger scales of environmental change.     

A comparison of biological trends from four marine ecosystems: Synchronies, differences, and commonalities

Major features of four marine ecosystems were analyzed based on a broad range of fisheries-associated datasets and a suite of oceanographic surveys. The ecosystems analyzed included the Gulf of Maine/Georges Bank in the Northwest Atlantic Ocean, the Norwegian/Barents Seas in the Northeast Atlantic Ocean, and the eastern Bering Sea and the Gulf of Alaska in the Northeast Pacific Ocean. We examined survey trends in major fish abundances, total system fish biomass, and zooplankton biomasses. We standardized each time series and examined trends and anomalies over time, using both time series and cross-correlational statistical methods. We compared dynamics of functionally analogous species from each of these four ecosystems. Major commonalities among ecosystems included a relatively stable amount of total fish biomass and the importance of large calanoid copepods, small pelagic fishes and gadids. Some of the changes in these components were synchronous across ecosystems. Major differences between ecosystems included gradients in the magnitude of total fish biomass, commercial fish biomass, and the timing of major detected events. This work demonstrates the value of comparative analysis across a wide range of marine ecosystems, suggestive of very few but none-the-less detectable common features across all northern hemisphere ocean systems.     

Wasp-waist populations and marine ecosystem dynamics: Navigating the “predator pit” topographies

Many marine ecosystems exhibit a characteristic “wasp-waist” structure, where a single species, or at most several species, of small planktivorous fishes entirely dominate their trophic level. These species have complex life histories that result in radical variability that may propagate to both higher and lower trophic levels of the ecosystem. In addition, these populations have two key attributes: (1) they represent the lowest trophic level that is mobile, so they are capable of relocating their area of operation according to their own internal dynamics; (2) they may prey upon the early life stages of their predators, forming an unstable feedback loop in the trophic system that may, for example, precipitate abrupt regime shifts. Experience with the typical “boom-bust” dynamics of this type of population, and with populations that interact trophically with them, suggests a “predator pit” type of dynamics. This features a refuge from predation when abundance is very low, very destructive predation between an abundance level sufficient to attract interest from predators and an abundance level sufficient to satiate available predators, and, as abundance increases beyond this satiation point, decreasing specific predation mortality and population breakout. A simple formalism is developed to describe these dynamics. Examples of its application include (a) a hypothetical mechanism for progressive geographical habitat expansion at high biomass, (b) an explanation for the out-of-phase alternations of abundances of anchovies and sardines in many regional systems that appear to occur without substantial adverse interactions between the two species groups, and (c) an account of an interaction of environmental processes and fishery exploitation that caused a regime shift. The last is the example of the Baltic Sea, where the cod resource collapsed in concert with establishment of dominance of that ecosystem by the cod’s ‘wasp-waist” prey, herring and sprat.     

Modeling environmental effects on the size-structured energy flow through marine ecosystems. Part 2: Simulations

Numerical simulations using a physiologically-based model of marine ecosystem size spectrum are conducted to study the influence of primary production and temperature on energy flux through marine ecosystems. In stable environmental conditions, the model converges toward a stationary linear log–log size-spectrum. In very productive ecosystems, the model predicts that small size classes are depleted by predation, leading to a curved size-spectrum.It is shown that the absolute level of primary production does not affect the slope of the stationary size-spectrum but has a nonlinear effect on its intercept and hence on the total biomass of consumer organisms (the carrying capacity). Three domains are distinguished: at low primary production, total biomass is independent from production changes because loss processes dominate dissipative processes (biological work); at high production, ecosystem biomass is proportional to primary production because dissipation dominates losses; an intermediate transition domain characterizes mid-production ecosystems. Our results enlighten the paradox of the very high ecosystem biomass/primary production ratios which are observed in poor oceanic regions. Thus, maximal dissipation (least action and low ecosystem biomass/primary production ratios) is reached at high primary production levels when the ecosystem is efficient in transferring energy from small sizes to large sizes. Conversely, least dissipation (most action and high ecosystem biomass/primary production ratios) characterizes the simulated ecosystem at low primary production levels when it is not efficient in dissipating energy.Increasing temperature causes enhanced predation mortality and decreases the intercept of the stationary size spectrum, i.e., the total ecosystem biomass. Total biomass varies as the inverse of the Arrhenius coefficient in the loss domain. This approximation is no longer true in the dissipation domain where nonlinear dissipation processes dominate over linear loss processes. Our results suggest that in a global warming context, at constant primary production, a 2–4 °C warming would lead to a 20–43% decrease of ecosystem biomass in oligotrophic regions and to a 15–32% decrease of biomass in eutrophic regions.Oscillations of primary production or temperature induce waves which propagate along the size-spectrum and which amplify until a “resonant range” which depends on the period of the environmental oscillations. Small organisms oscillate in phase with producers and are bottom-up controlled by primary production oscillations. In the “resonant range”, prey and predators oscillate out of phase with alternating periods of top-down and bottom-up controls. Large organisms are not influenced by bottom-up effects of high frequency phytoplankton variability or by oscillations of temperature.     

The marine ecosystem off Peru: What are the secrets of its fishery productivity and what might its future hold?

The marine ecosystem located off the coast of central and northern Peru has stood as the “world’s champion” producer, by far, of exploitable fish biomass, generally yielding more than 20 times the tonnage of fishery landings produced by other comparable regional large marine ecosystems of the world’s oceans that operate under similar dynamic contexts and are characterized by comparable, or even greater, basic primary production. Two potentially contributing aspects are discussed from a framework of interregional comparative pattern recognition: (1) the advantageous low-latitude situation that combines strong upwelling-based nutrient enrichment with low wind-induced turbulence generation and relatively extended mean “residence times” within the favorable upwelling-conditioned near-coastal habitat and (2) the cyclic “re-setting” of the system by ENSO perturbations that may tend to interrupt malignant growth of adverse self-amplifying feedback loops within the nonlinear biological dynamics of the ecosystem.There is a developing scientific consensus that one of the more probable consequences of impending global climate changes will be a general slowing of the equatorial Pacific Walker Circulation and a consequent weakening of the Pacific trade wind system. Since the upwelling-favorable winds off Peru tend to flow directly into the Pacific southeast trade winds, a question arises as to the likely effect on the upwelling-producing winds that power the productivity of the regional coastal ecosystems of the Peru–Humboldt Current zone. It is argued that the effects will in fact be decoupled to the extent that upwelling-favorable winds will actually tend to increase off Peru. Data demonstrative of this decoupling are presented. A tendency for less intense El Niño episodes in the future is also suggested. These conclusions provide a framework for posing certain imponderables as to the future character of the Peruvian marine ecosystem and of the fisheries it supports.     

Strengths and weaknesses of the global ocean conveyor: Inter-basin freshwater disparities as the major control

According to the current paradigm of modern climatology and oceanography, the global ocean thermohaline circulation works as the so-called “global ocean salinity conveyor belt” – a system of currents connecting different ocean basins and most notably – the northern North Atlantic and northern North Pacific Oceans – the most distant regions of the world ocean. It is shown here that a slight disparity in freshwater redistribution between the Atlantic and Pacific oceans can be sufficient for building up and maintaining a global conveyor-type ocean thermohaline circulation. On the other hand, relatively small changes in this disparity leading to change in sea surface salinity contrasts between and in the north-south within the northern parts of these two oceans can easily change the conveyor.     

Have there been recent changes in climate? Ask the fish

It is generally accepted that a climate shift occurred about 1977 that affected the dynamics of North Pacific marine ecosystems. Agreement on the possibility of further climate shifts in 1989 and the late 1990s is yet to be achieved. However, there have been changes in the dynamics of key commercial fishes that indicate changes in their environment occurred in the early 1990s, and possibly around 1998. One method of measuring climate change is to observe the dynamics of species that could be affected.Several studies have described decadal-scale changes in North Pacific climate–ocean conditions. Generally, these studies focus on a single index. Using principal components analysis, we use a composite index based on three aspects of climate ocean conditions: the Aleutian Low Pressure Index, the Pacific Atmospheric Circulation Index and the Pacific Interdecadal Oscillation Index. We link this composite index (Atmospheric Forcing Index) to decadal-scale changes in British Columbia salmon and other fish populations. Around 1989 there was a change from intense Aleutian Lows (above average south-westerly and westerly circulation patterns and warming of coastal sea surface temperatures) to average Aleutian Lows (less frequent south-westerly and westerly circulation and slightly cooler coastal sea surface temperatures in winter). These climate–ocean changes were associated with changes in the abundance and ocean survival of salmon (Oncorhynchus spp.), distribution and spawning behaviour of hake (Merluccius productus) and sardines (Sardinops sagax) and in recruitment patterns of several groundfish species.     

Climate, fishery and society interactions: Observations from the North Atlantic

Interdisciplinary studies comparing fisheries-dependent regions across the North Atlantic find a number of broad patterns. Large ecological shifts, disastrous to historical fisheries, have resulted when unfavorable climatic events occur atop overfishing. The “teleconnections” linking fisheries crises across long distances include human technology and markets, as well as climate or migratory fish species. Overfishing and climate-driven changes have led to a shift downwards in trophic levels of fisheries takes in some ecosystems, from dominance by bony fish to crustaceans. Fishing societies adapt to new ecological conditions through social reorganization that have benefited some people and places, while leaving others behind. Characteristic patterns of demographic change are among the symptoms of such reorganization. These general observations emerge from a review of recent case studies of individual fishing communities, such as those conducted for the North Atlantic Arc research project.     

Contrast in life histories of exploited fishes and ecosystem structures in coastal waters off west Canada and east Korea

By reviewing the history of fishery exploitation in the coastal waters of west Canada and east Korea, related with contrasting life history strategies of the dominant species, the fishery management challenges that each country would face in the upcoming decades were outlined. In the ecosystem of the Canadian western coastal waters, the dominant oceanographic feature is the coastal upwelling domain off the west coast of Vancouver Island, the northernmost extent of the California Current System in the eastern North Pacific. In the marine ecosystem of the eastern coasts of Korea (the Japan/East Sea), a major oceanographic feature is the Tsushima Warm Current, a branch of the Kuroshio Current in the western North Pacific. Fishes in the Canadian ecosystem are dominated by demersal, long-lived species such as flatfish, rockfish, sablefish, and halibut. During summer, migratory pelagic species such as Pacific hake, Pacific salmon, and recently Pacific sardine, move into this area to feed. In the late 1970s, Canada declared jurisdiction for 200 miles from their coastline, and major fisheries species in Canadian waters have been managed with a quota system. The overall fishing intensity off the west coast of Vancouver Island has been relatively moderate compared to Korean waters. Fishes in the ecosystem of the eastern Korean waters are dominated by short-lived pelagic and demersal fish. Historically, Korea has shared marine resources in this area with neighbouring countries, but stock assessments and quotas have only recently (since the late-1990s) been implemented for some major species. In the Korean ecosystem, fisheries can be described as intensive, and many stocks have been rated as overfished. The two ecosystems responded differently to climate impacts such as regime shifts under different exploitation histories. In the future, both countries will face the challenge of global warming and subsequent impacts on ecosystems, necessitating developing adaptive fisheries management plans. The challenges will be contrasting for the two countries: Canada will need to conserve fish populations, while Korea will need to focus on rebuilding depleted fish populations.     

Long-term changes in the fish community structure from the Tsushima warm current region of the Japan/East Sea with an emphasis on the impacts of fishing and climate regime shift over the last four decades

Japanese fisheries production in the Japan/East Sea between 1958 and 2003 increased to their peak (1.76 million tons) in the late 1980s and decreased abruptly with the collapse of Japanese sardine. Catch results for 58 fisheries and various environmental time-series data sets and community indices, including mean trophic level (MTL) and Simpson’s diversity index (DI), were used to investigate the impacts of fishing and climate changes on the structure of the fish community in the Tsushima warm current (TWC) region of the Japan/East Sea. The long-term trend in fisheries production was largely dependent on the Japanese sardine that, as a single species, contributed up to 60% of the total production in the Japanese waters of the Japan/East Sea during the late 1980s. Excluding Japanese sardine, production of the small pelagic species was higher during 1960s and 1990s but lower during 1970s and 1980s. This variation pattern generally corresponds with the trend in water temperature, warmer before early 1960s and after 1990s but colder during 1970s and 1980s. The warm-water, large predatory fishes and cold water demersal species show opposite responses to the water temperature in the TWC region, indicating the significant impact of oceanic conditions on fisheries production of the Japan/East Sea. Declines in demersal fishes and invertebrates during 1970s and 1980s suggested some impact of fishing. MTL and DI show a similar variation pattern: higher during 1960s and 1990s but lower during 1970s and 1980s. In particular, the sharp decline during the 1980s resulted from the abundant sardine catches, suggesting that dominant species have a large effect on the structure of the fish community in the Japan/East Sea. Principal component analysis for 58 time-series data sets of fisheries catches suggested that the fish community varied on inter-annual to inter-decadal scales; the abrupt changes that occurred in the mid-1970s and late 1980s seemed to correspond closely with the climatic regime shifts in the North Pacific. These results strongly suggest that the structure of the fish community in the Japan/East Sea was largely affected by climatic and oceanic regime shifts rather than by fishing. There is no evidence showing “fishing down food webs” in the Japan/East Sea. However, in addition to the impacts of abrupt shifts that occurred in the late 1980s, the large predatory and demersal fishes seem to be facing stronger fishing pressure with the collapse of the Japanese sardine.     

Interdecadal climate regime dynamics in the North Pacific Ocean: theories, observations and ecosystem impacts

Basin-scale variations in oceanic physical variables are thought to organize patterns of biological response across the Pacific Ocean over decadal time scales. Different physical mechanisms can be responsible for the diverse basin-scale patterns of sea-surface temperature (SST), mixed-layer depth, thermocline depth, and horizontal currents, although they are linked in various ways. In light of various theories and observations, we interpret observed basinwide patterns of decadal-scale variations in upper-ocean temperatures. Evidence so far indicates that large-scale perturbations of the Aleutian Low generate temperature anomalies in the central and eastern North Pacific through the combined action of net surface heat flux, turbulent mixing and Ekman advection. The surface-forced temperature anomalies in the central North Pacific subduct and propagate southwestwards in the ocean thermocline to the subtropics but apparently do not reach the equator. The large-scale Ekman pumping resulting from changes of the Aleutian Low forces western-intensified thermocline depth anomalies that are approximately consistent with Sverdrup theory. These thermocline changes are associated with SST anomalies in the Kuroshio/Oyashio Extension that are of the same sign as those in the central North Pacific, but lagged by several years. The physics of the possible feedback from the SST anomalies to the Aleutian Low, which might close a coupled ocean–atmosphere mode of decadal variability, is poorly understood and is an area of active research. The possible responses of North Pacific Ocean ecosystems to these complicated physical patterns is summarized.     

Implications of variability on many time scales for scientific advice on sustainable management of living marine resources

The conceptual basis for understanding and management of living marine resources is built on three basic ecological principles developed in the first half of the past century: the law of the minimum, competitive exclusion, and succession. This paper highlights aspects of these principles that make them insufficient as a sound foundation for understanding and managing marine ecosystems, points out dangers of continuing to use approaches built on them, and presents alternatives which might be more appropriate and of lower risk.To do this, the paper considers variability of marine ecosystems on annual, medium and long-term time scales, highlighting that these scales correspond to less than, approximately equal to, and much greater than, the generation times of dominant predators in the systems. It also considers how each interval of variability may affect directly ecosystems which are controlled from the bottom up, top down, and middle outward, and how position and duration of forcing affect five types of responses: growth, maturation, recruitment, predation, and competition. Generally these five processes have manifestations at the scale of individuals, populations, and ecosystems, attention is drawn to which manifestations are the most significant for each duration and position of forcing.Effects of some combinations of duration of forcing and position of forcing can be explained reasonably well by conventional ecological theory. For other combinations, particularly forcing at time scales of predator generations on top-down or middle-out ecosystems, theory based on contest competition and equilibria are likely to be misleading. In these systems the major dynamics are transients, when many ecosystems are far from their carrying capacities, so scramble competition dominates, and the carrying capacity is not helpful in explaining the system dynamics. This review clarifies the sorts of questions that we should be asking, in order to begin to understand the transient behaviour of these non-equilibrium ecosystems. The answers to the new classes of questions may lead to great improvements in how ecosystems are managed, as well as how their variation is explained.     

Foraging ecology and movement patterns of jumbo squid (Dosidicus gigas) in the California Current System

From 2002 to 2010, the jumbo squid (Dosidicus gigas) has been regularly encountered in large numbers throughout the California Current System (CCS). This species, usually found in subtropical waters, could affect coastal pelagic ecosystems and fisheries as both predator and prey. Neither the abundance of jumbo squid nor the optimal ocean conditions in which they flourish are well known. To understand better the potential impacts of this species on both commercial fisheries and on food-web structure we collected nearly 900 specimens from waters of the CCS, covering over 20° of latitude, over a range of depths and seasons. We used demographic information (size, sex, and maturity state) and analyzed stomach contents using morphological and molecular methods to best understand the foraging ecology of this species in different habitats of the CCS. Squid were found to consume a broad array of prey. Prey in offshore waters generally reflected the forage base reported in previous studies (mainly mesopelagic fishes and squids), whereas in more coastal waters (shelf, shelf break and slope habitats) squid foraged on a much broader mix that included substantial numbers of coastal pelagic fishes (Pacific herring and northern anchovy, as well as osmerids and salmonids in northern waters) and groundfish (Pacific hake, several species of rockfish and flatfish). We propose a seasonal movement pattern, based on size and maturity distributions along with qualitative patterns of presence or absence, and discuss the relevance of both the movement and distribution of jumbo squid over space and time. We find that jumbo squid are a generalist predator, which feeds primarily on small, pelagic or mesopelagic micronekton but also on larger fishes when they are available. We also conclude that interactions with and potential impacts on ecosystems likely vary over space and time, in response to both seasonal movement patterns and highly variable year-to-year abundance of the squid themselves.     

Honey, I cooled the cods: Modelling the effect of temperature on the structure of Boreal/Arctic fish ecosystems

Historically colder regions of the North Atlantic had fisheries dominated by only a few fish species; principally cod and capelin. Possible population dynamic mechanisms that lead to such dominance are investigated by considering how a charmingly simple published multispecies model of the North Sea would react if the system operated at a lower temperature. The existing model equations were modified to describe temperature effects on growth, fecundity and recruitment and the model was rerun based on typical temperatures for the North Sea and a colder system. The results suggest that total fish biomass in the colder system increases but the community is more vulnerable to a given rate of fishing mortality. In the colder system, within species density dependence is reduced but relative predation rates are higher. Consequently, intermediate-sized species are vulnerable to relatively high levels of predation throughout their life history and tend to be excluded, leading to a system dominated by small and large species. The model helps to explain how temperature may govern coexistence and competitive exclusion in fish communities and accounts for the observed dominance of small and large species in Boreal/Arctic ecosystems.     

Mesopelagic fishes of the Bering Sea and adjacent northern North Pacific Ocean

Fourteen midwater trawl collections to depths of 450 m to 1,400 m were taken at eleven stations in the Bering Sea and adjoining regions of the northern North Pacific by the R/V Hakuho Maru during the summer of 1975. A total of 29 kinds of fishes were identified. Mesopelagic fishes of the families Myctophidae, Gonostomatidae and Bathylagidae predominated in the catches, contributing 14 species (94%) of the fishes caught.Seventeen species of fishes were caught in the Bering Sea, and all of these are known from nearby areas. The mesopelagic fish fauna of the Bering Sea is similar to that in adjoining regions of the northern North Pacific Ocean: endemic species are rare or absent. Stenobrachius nannochir was usually the most common mesopelagic fish in our catches.Stenobrachius leucopsarus is a diel vertical migrant that is usually the dominant mesopelagic fish in modified Subarctic waters of the northeastern Pacific. The change in dominance fromS. nannochir in the western Bering Sea toS. leucopsarus in the eastern Bering Sea is related to differences in oceanographic conditions.     

The latitudinal and bathymetric ranges of marine fishes: a global analysis to test the application of Rapoport′s Rule

We evaluated the applicability of Rapoport’s Rule (RR) to the marine bony fishes of the world. The biogeographical pattern predicted by RR has been the subject of a large number of studies, some supporting it and some not. In this exercise, we attempted to generate results free of biases from taxonomy or geographic scale. The study area encompassed all world oceans. Our analysis was based on secondary data. We tested the relationship of the geographic range and the bathymetric range to the latitude gradient, using the method of Stevens (American Naturalist 1989, 133 , 240–256) . We compared all known species of marine bony fishes together, and performed a second analysis limited to the non‐pelagic species. Our results were generated from a databank including 13,957 species. The results indicated that RR (latitudinal range) is valid for 11 regions. However, there were exceptions for the northern part of the Eastern Atlantic and Pacific, the northern part of the Western Indian, and the Arctic Oceans. The analyses of RR (bathymetric range) were supported for 13 regions. We found only one exception, for the northern part of the Western Indian Ocean. The applicability of RR and its relationship to energy supply, represented here by the temperature variability of the marine water masses, was clear. The exceptions appeared when some feature associated with the distribution of water masses was superposed on the latitudinal gradient, or when some life trait characteristic of a taxon, for instance body size, was confounded with the effect of climate variability.     

The evolution of oceanic and atmospheric anomalies in the northeast Pacific during the El Niño and La Niña events of 1995–2001

From late 1995 through early 2001, three major interannual climate events occurred in the tropical Pacific; the 1995–97 La Niña (LN), 1997–98 El Niño (EN), and 1998–2001 LN. We analyze atmospheric and upper oceanic anomalies in the northeast Pacific (NEP) during these events, and compare them to anomalies both elsewhere in the north and tropical Pacific, and to typical EN and LN anomaly patterns. The atmospheric and oceanic anomalies varied strongly on intraseasonal and interannual scales. During the 1995–97 LN and 1997–98 EN, the Northeast Pacific was dominated by negative SLP and cyclonic wind anomalies, and by upper ocean temperature and sea surface height (SSH) anomalies. The latter were positive along the North American west coast and in the NEP thermal anomaly pool (between Hawaii, Vancouver Island, and Baja California), and negative in the central north Pacific. This atmospheric/oceanic anomaly pattern is typical of EN. An eastward shift in the atmospheric teleconnection from east Asia created EN-like anomalies in the NEP during the 1995–97 LN, well before the 1997–98 EN had begun. The persistence of negative sea-level pressure (SLP) and cyclonic wind anomalies in the NEP during the 1997–98 EN intensified pre-existing upper oceanic anomalies. Atmospheric anomalies were shifted eastward during late 1996–early 1998, leading to a similar onshore shift of oceanic anomalies. This produced exceptionally strong positive upper ocean temperature and SSH anomalies along the west coast during the 1997–98 EN, and explains the unusual coastal occurrences of several species of large pelagic warm-water fishes. The growth and eastward shift of these pre-existing anomalies does not appear to have been linked to tropical Pacific EN anomalies until late 1997, when a clear atmospheric teleconnection between the two regions developed. Prior to this, remote atmospheric impacts on the NEP were primarily from east Asia. As the 1998–2001 LN developed, NEP anomalies began reversing toward the typical LN pattern. This led to predominantly negative SLP and cyclonic wind anomalies in the NEP, and upper ocean temperature and SSH anomalies that were mainly negative along the west coast and positive in the central north Pacific. The persistence of these anomalies into mid-2001, and a number of concurrent biological changes in the NEP, suggest that a decadal climate shift may have occurred in late 1998.During 1995–2001, NEP oceanic anomalies tracked the overlying atmospheric anomalies, as indicated by the maintenance of a characteristic spatial relationship between these anomalies. In particular, wind stress curl and SSH anomalies in the NEP maintained an inverse relationship that strengthened and shifted eastward toward the west coast during late 1996–early 1998. This consistent relationship indicates that anomalous Ekman transport driven by regional atmospheric forcing was an important contributor to temperature and SSH anomalies in the NEP and CCS during the 1997–98 EN. Other studies have shown that coastal propagations originating from the tropical Pacific also may have contributed to coastal NEP anomalies during this EN. Our results indicate that at least some of this coastal anomaly signal may have been generated by regional atmospheric forcing within the NEP.