© 2005 International Council for the Exploration of the Sea
On distributional responses of North Atlantic fish to climate change
Fisheries Conservation Chair, Fisheries and Marine Institute of Memorial University of Newfoundland St. John's, NL, Canada A1C 5R3
*tel: +1 709 7780482; fax: +1 709 7780669. e-mail: grose{at}mi.mun.ca.
Changes in fish distribution and climate in the North Atlantic have been observed for millennia by seafaring peoples, chronicled in many historical anecdotes, and recently studied systematically. For temperate to Arctic North Atlantic fish, a literature compendium of limits of temperature, salinity, and depth during feeding and spawning was used to investigate factors that influence distribution. Latitude and depth were negatively correlated with species number and density. Peak numbers of species feed at 0–4°C, but spawn at 2–7°C and salinities of 32.5–33.5. Principal components of feeding depths and temperatures suggested four groups of species: (i) small pelagics characterized by shallow habitat and cooler temperatures; (ii) most groundfish in deeper and warmer waters; (iii) warm-water large pelagics; and (iv) deepwater species. Spawning temperatures, salinities, depths, and timing produced groupings consistent with feeding components for pelagics, but differing for distant migrants such as tunas. Principal components (PCA) of spawning characteristics explained 56% of the variance in species resilience (doubling time), while PCA of feeding characteristics explained only 23%. We infer that the small pelagics capelin (Mallotus villosus) and herring (Clupea harengus) react strongly and quickly to climate change because of their physiological limits and potential for fast population growth. Verification comes from Icelandic and Greenland waters, which warmed considerably during 1920–1940, and where capelin, herring, cod (Gadus morhua), and other species shifted north very quickly.
Keywords: capelin, climate change, distribution, migration, North Atlantic
Received 14 June 2004; accepted 13 May 2005.
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Introduction |
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Since time immemorial, seafaring peoples have observed variations in the distributions and migrations of marine animals in association with climate. In terms of their essentially Lagrangian view, there were good and bad days, months, years, and decades. Not all species varied in the same way, and some species varied more than others. Groups of species having similar responses have been identified both from anecdotes and recent systematic study (Murawski, 1993). The earliest reported historical events have ample anecdotes, but lack data or systematic investigation (e.g. Prowse, 1895). In the recent past, however, a major warming event in Icelandic and Greenland waters between 1920 and 1940 was extensively documented (e.g. Sæmundsson, 1932; Ahlmann, 1948; Lysgaard, 1948).
Early works in marine zoogeography addressed some of the historical queries about the comings and goings of fish by explaining the distribution patterns of species and their dynamics in terms of species physiology and ecology. An essential element of this work was to explain how distribution patterns developed over much longer time scales. The seminal works of Ekman (1953) and Briggs (1974) provide rich descriptions of variations in fish distribution and migration in relation to physical factors of the environment, especially sea temperature, salinity, depth, bathymetry, and currents, on both geological and contemporary scales. Similar dynamics are also well known in marine invertebrates (e.g. Petuch, 2004). These works suggest that climate has a dominant influence on the long-term (>>1000 years) distribution and migration patterns of marine fish and invertebrates, and provide comprehensive treatments of how these patterns developed in the world's oceans over geological time scales.
Reports on shorter-term changes in the distribution and migration of many marine phyla, species, and stocks are numerous in the literature. Many works have investigated warming and cooling periods, and describe northward advances of species during warm periods or, alternatively, southward contractions during cold periods. There can be no doubt that the main commercial species of the North Atlantic, including Atlantic cod (Gadus morhua), herring (Clupea harengus), mackerel (Scomber scombrus), haddock (Melanogrammus aeglefinus), and capelin (Mallotus villosus) are subject to north–south movements in response to climate (Ekman, 1953). Cushing (1982) summarized some better known examples, such as North Sea species changes, the effects of cold winters in northwest Europe in the 20th century, and northward movements of many species in the North Atlantic in the warm period of the 1920s to 1940s.
The reality of climate change in the 21st century has refocused concerns about the potential effects of environmental change on the fish populations of the North Atlantic (e.g. Murawski, 1993; Drinkwater, 2002; Godø, 2003). In addition to the stresses on fish stocks imposed by overfishing (Sinclair and Murawski, 1997), climate change may have particular impacts on the North Atlantic, because of a dependence on north–south oceanic heat transfer (Pflaumann et al., 2003). A key question is, how changes in sea temperatures, salinities, and currents may affect both the fish and the fisheries (McGinn, 2002). There is also the likelihood that environmental effects and overfishing can interact (e.g. Rose and Kulka, 1999), and that recent depletions whose primary cause is recruitment overfishing have been amplified and prolonged by changes in climate (e.g. Cushing, 1982; Rose et al., 2000; Drinkwater, 2002).
All fish have limits in terms of the sea temperatures and salinities that they inhabit, and these limits may change with age and be very different during feeding and spawning (Ekman, 1953). Whether broad or narrow, these limits form the large-scale boundaries of the distribution of species. Within these boundaries, the specifics of stock structure and life cycles have developed as adaptations to the scales and patterns of currents and seasonal variations in temperature, salinity, and the availability of food (Robichaud and Rose, 2004). In the North Atlantic, there are many variations in such life cycles. For example, some fish spawn only in warm subtropical waters but migrate into colder North Atlantic waters to feed (e.g. bluefin tuna, Thunnus thynnus). Other species spawn and carry out their full life cycle within fairly constant physical conditions and locations (e.g. some of the eelpouts, Zoarcidae). Still others may spawn in relatively warm waters, spend part of the juvenile cycle in cool waters, then move to very cold waters as adults to feed (e.g. capelin). There are many variants with many spatial and temporal overlaps. All contribute to and influence what we call "ecosystems".
In this paper, I pose several questions. What factors influence the distribution of species in the temperate to Subarctic regions of the North Atlantic? Can the different responses of species be quantified? Do groups of species respond similarly to and differently from other groups? To address these questions, I examined the feeding and spawning limits of North Atlantic fish for which data were available. As a qualitative test of the outcomes of this broad-scale analysis, I examine historically documented changes in distribution and migration patterns that occurred during the warming period of 1920–1940 in Iceland and Greenland.
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Methods |
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The literature was searched for data on fish species of the temperate to Subarctic North Atlantic (Table 1). In all, 141 species were examined. The list is not comprehensive since data were not available for all known species. Variables recorded for each species both during feeding and spawning periods of adults were minimum and maximum temperature of occurrence in the wild, minimum and maximum salinity, minimum and maximum depth, spawning start and end months, and a measure of species resilience [population doubling time, categorized from literature values of "r", the intrinsic rate of increase, as very short (1 year), short (to 3 years), medium (to 9 years), or long (>9 years)]. All variables were available for only a subset of the total species. In cases where literature values for the measured variables differed, judgements were made to determine which values to use. For some species, especially those that are poorly described, values may not be strictly comparable, because it was evident that some workers described where fish were most often found, while others described the full limits of even minor distribution. In general, feeding limits describe the range where most feeding occurs, but spawning limits are full limits beyond which little or no spawning has been observed. No account has been taken of the success of the spawning. In cases where data were thought not to be reliable, they were not used. Principal component analyses were used to describe associations among the variables measured, and then to identify groups of species with similar variable loadings. These analyses did not include species that were poorly known because insufficient data were available.
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Surface air temperature data were extracted from the NASA database available through the Goddard Institute for Space Studies, New York for Norway, Iceland, Greenland, and Newfoundland (http://www.giss.nasa.gov/data/update/gistemp/station_data/).
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Results |
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The number of species declined exponentially with increasing latitude from approximately 400 at 25°N to 50 at 60°N (Figure 1a). Mean water temperature also declined with increasing latitude (Figure 1b). Hence, there is a general positive association between average temperature and the number of species.
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The number of species declined exponentially with depth, from approximately 60 in shallow shelf waters to very few in waters >2000 m deep (Figure 2a). The same trend is evident in both the present literature-based survey and in original data from the Challenger cruise from 1872 to 1876. A similar trend is evident in species density, which takes into account the total areas of the depth zones (Figure 2b). Increasing depth is also associated with declines in light (Figure 2c). However, species number and density declined with depth at a much slower rate than did light intensity.
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Minimum and maximum temperatures were available for 90 species. These ranged from near –2 to 25°C (Table 1). However, the majority of species had minimum or maximum temperatures between 3 and 4°C, and the ogives were steepest in this region (Figure 3a). Spawning temperatures were somewhat higher (Figure 3b). The temperature range that included the most feeding species was approximately 0–4°C (Figure 3c). For spawning, the temperature range of 2–7°C included the most species. The salinity range that included the most spawning species was approximately 32.5–33.5 (Figure 4).
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In all, 43 species had data available on depth and temperature range and time of feeding (approximated as the non-spawning time in months) (Table 1). Principal components of these factors indicated that the species fell into four overlapping groups (Figure 5). The first principal component was loaded most heavily by depth, the second by temperature, and the third by feeding time (Table 2a). The four groups can be characterized as cold-water pelagic species, warm-water pelagics, cool-water shallow-depth demersals, and deeper water demersal species.
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Principal components of spawning characteristics (spawning temperatures, depths, salinities, and timing) were estimated for 17 species (with four capelin stocks) for which data were available (Figure 6). The first component was most heavily loaded by salinity, with species requiring fresher water separating from others (Table 2b). The second component was loaded most heavily by depth, and the third by temperature. A group of low salinity shallow-water spawners composed of capelin and herring was clearly isolated from other groups. A more diffuse group of groundfish and some warmer-water pelagics included the most species. Northeast Atlantic and Icelandic capelin were grouped with the groundfish, although at the fringes of the group, and not with herring (Clupea harengus) and Northwest Atlantic and Greenland capelin. Greenland halibut (Reinhardtius hippoglossoides) was widely separated from the main groupings largely because of their deepwater habitat.
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The principal components of feeding habitat and spawning requirements were related to species resilience (Table 3). Resilience was indexed by the estimated doubling time of the population (Table 1). The first component, which was heavily loaded by depth distribution, indicates that pelagic species have faster doubling times and are likely to react quickly to changes in the environment when feeding. The spawning distribution model was more complex, with the first component of distribution (loaded heavily by salinity), unimportant, but the second and third components loaded by depth and temperature, respectively, highly significant. This confirms that pelagic species are more likely to respond substantially to changes in the environment, evidently because of their shallow-water spawning habits, and can do so more quickly because of faster doubling times.
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The 1920–1940 warming event
From approximately 1920 to 1940, North Atlantic Waters from Greenland to Norway warmed significantly, by as much as 3–4°C (Tåning, 1948). Although the causes of this event are not well understood, there is no doubt of its authenticity or widespread occurrence in temperature records (Figure 7).
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The predictions of the principal components analyses were generally supported by changes in fish distribution documented during the warming event. Species expected to react quickly, such as capelin, and to a lesser extent Atlantic herring and cod (Gadus morhua), expanded their ranges northwards. Some southern migrants became more numerous, and other resident species increased their densities, although evidence of distribution change is lacking. Species expected not to change distribution, such as Greenland halibut, did not, but may have increased in density in some areas. Tåning (1948) summarized earlier accounts and identified several types of changes: (i) species typically found farther south were more frequently observed in Greenland waters (e.g. tuna; Atlantic skipper or saury, Scomberesox saury; mackerel; and others), (ii) local species extended their distributions northwards (e.g. Atlantic cod, capelin, and herring), (iii) local species increased in density (many commercial species including Atlantic cod and herring; salmon, Salmo salar; haddock, Melanogrammus aeglefinus), (iv) spawning grounds changed (e.g. capelin; herring; redfish, Sebastes spp.), (v) local species decreased in density where waters became too warm, but increased in colder areas (e.g. capelin, Greenland halibut). Similar events occurred in Icelandic waters at the same time (Sæmundsson, 1932; Fridriksson, 1948).
From the present analyses, species in the top left of Figure 5 are warm-water species that would be expected to be new arrivals to Greenland and Iceland waters in a warm period. These include Atlantic saury, mackerel, and tuna, as observed (Tåning, 1948). Local species that would be expected to shift north are found in the lower left of Figure 5, and include Atlantic cod, capelin, and several other species not mentioned by Tåning (1948). Changes in spawning grounds are expected from species in the lower left of Figure 6, which includes herring, capelin, and perhaps cod but not redfish (Sebastes spp.), which is consistent with the observations reported by Tåning (1948).
This warming event, which so influenced Greenland and Iceland as well as Norwegian waters and the Barents Sea, did not extend to the Northwest Atlantic (Figure 7a). Temperatures declined during the 1920s at St. John's, in direct contrast to temperatures at Norway, Iceland, and Greenland, although there were increases during the 1930s. No great changes in species in Newfoundland and Labrador waters were noted in an extensive work on capelin published by Templeman (1948), although the same author does recall some changes in the Grand Bank haddock stock during the 1920s in a much later publication (Templeman, 1966).
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Discussion |
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The present analyses indicate that both spawning and feeding distributions of North Atlantic fish are influenced by environmental factors partially regulated by climate variability, and likely, by climate change. Not all species may be impacted equally. Species whose spawning is limited to the cold waters of the Arctic or boreal regions (lower left in principal components plot, Figure 5) are expected to be most affected, and to respond relatively quickly to environmental change. Among these species are the small pelagic boreal-Subarctic species, in particular capelin. Cold-water species such as winter flounder, several species of sculpin, the eelpouts, Arctic cod, daubled shanny, and the alligators may be the most vulnerable. The gadoids and several flatfish and other more boreal species are mid-range in vulnerability and could benefit from warmer conditions, subject to any effects on their prey. Species that utilize the North Atlantic only for feeding (e.g. tunas and mackerel) but spawn further south, even in subtropical waters, may be influenced primarily through migration patterns that are based on temperatures and feeding opportunities. Deepwater species such as Greenland halibut, whose requirements are based on more stable hydrographic conditions in those habitats, may be least impacted (to right of plot, Figure 5), although prey changes (e.g. capelin) may modify their response. An examination of survey data from the northeast coast of the US led to similar conclusions (Murawski, 1993).
The strongest predictors of species richness are latitude and depth. In general, the warmer tropical waters sustain a higher number of species than do the colder waters farther north. If climate change results in a warming of northern waters, it is expected that more species will invade northern waters. There may be concurrent contraction of the range of a fewer number of cold-water species. More worrisome would be the climate scenario that indicates a cooling of northern oceans with a slowing of the North Atlantic heat pump (Häkkinen and Rhines, 2004). This scenario would result in a decline in species number in northern seas and an increase in range of only a few species, such as the cold-adapted eelpouts, sculpins, Arctic cod, and winter flounder. In addition, most climate models predict an increase in sea levels with a warming trend. Such changes might result in some increase in habitat for shallow-water coastal spawners such as herring and capelin. A cooling would have the opposite effect, as during the last ice age, when capelin retreated southwards (Carscadden et al., 1989).
The groupings of species in the present analyses are, by and large, consistent with observed historical responses of species during 1920–1940 from Greenland to Norway for most, but not all, species. A highlight is the expectation and verification by observation that capelin will be among the most likely to respond to climate in terms of changes in distribution and spawning location. Warming will result in capelin shifts to the north as Arctic fronts relax or bend towards the pole (Tåning, 1948). Cooling would have the opposite effect, as during the last ice age, when capelin retreated southwards (Carscadden et al., 1989). During the same years, the temperatures in the Northwest Atlantic did not increase, and there is little evidence of changes in the fisheries. A summary of the known excursions and shifts of capelin with respect to climate changes and variations is given by Rose (2005).
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Conclusions |
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- Latitude and depth provide good predictors of species richness in the temperate to Subarctic North Atlantic. Latitude is a proxy for temperature.
- North Atlantic fish feed and spawn over a wide range of temperatures from –2 to 20°C, but most species are within their limits for feeding at 0–4°C, and for spawning at 2–7°C.
- Groups of species may be expected to react differently to climate change, with species that spawn in shallow, relatively low salinity waters most affected (e.g. capelin), and species that inhabit deeper, hydrographically more stable waters less so. Seasonal migrants that feed in the North Atlantic, but spawn further south (e.g. bluefin tuna), may undergo migration shifts. Note that changes in key forage fish, such as capelin and herring, are likely to have major influences on ecosystem structure and productivity of species that feed on them.
- Climate variability and change may not be uniform over the North Atlantic. History shows that differences are likely. Not all areas should be expected to warm or cool equally, and species responses must be judged accordingly.
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Acknowledgements |
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I am indebted to the fishery researchers who have collected data on the fish of the North Atlantic for over 150 years. Nevertheless, Table 1 is still far from complete and, for many species, poorly known or contested. Additions or improvements to this fundamental knowledge are welcome. Funding was provided by the NSERC Chair in Fisheries Conservation at Memorial University and my NSERC discovery grant. I thank S. B. Fudge for help with the data compilation and two anonymous reviewers for helpful comments.
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References |
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