Changes in the northern Gulf of St. Lawrence ecosystem estimated by inverse modelling: Evidence of a fishery-induced regime shift?

Mass-balance models have been constructed using inverse methodology for the northern Gulf of St. Lawrence for the mid-1980s, the mid-1990s, and the early 2000s to describe ecosystem structure, trophic group interactions, and the effects of fishing and predation on the ecosystem for each time period. Our analyses indicate that the ecosystem structure shifted dramatically from one previously dominated by demersal (cod, redfish) and small-bodied forage (e.g., capelin, mackerel, herring, shrimp) species to one now dominated by small-bodied forage species. Overfishing removed a functional group in the late 1980s, large piscivorous fish (primarily cod and redfish), which has not recovered 14 years after the cessation of heavy fishing. This has left only marine mammals as top predators during the mid-1990s, and marine mammals and small Greenland halibut during the early 2000s. Predation by marine mammals on fish increased from the mid-1980s to the early 2000s while predation by large fish on fish decreased. Capelin and shrimp, the main prey in each period, showed an increase in biomass over the three periods. A switch in the main predators of capelin from cod to marine mammals occurred, while Greenland halibut progressively replaced cod as shrimp predators. Overfishing influenced community structure directly through preferential removal of larger-bodied fishes and indirectly through predation release because larger-bodied fishes exerted top-down control upon other community species or competed with other species for the same prey. Our modelling estimates showed that a change in predation structure or flows at the top of the trophic system led to changes in predation at all lower trophic levels in the northern Gulf of St. Lawrence. These changes represent a case of fishery-induced regime shift.     

Comparing pristine and depleted ecosystems: The Sørfjord, Norway versus the Gulf of St. Lawrence, Canada. Effects of intense fisheries on marine ecosystems

The Sørfjord, Norway, and the Gulf of St. Lawrence, Canada, are two sub-arctic ecosystems with similar trophic structure. However, in the Gulf of St. Lawrence, severe exploitation of groundfish stocks has lead to important shifts in the trophic structure. In the Sørfjord, the situation is different: fishing pressure is much lighter. Our hypothesis is that overexploitation leads to changes in the trophic structure and severely alters the resilience of ecosystems. Based on the same modelling approach (Ecopath with Ecosim) the food web structure was compared, using different ecosystem indicators. Patterns of food web structure and trophodynamics were contrasted. Cod was the keystone species in both ecosystems, and forage fish were also important. Even after similar environmental changes in both ecosystems, and after a reduction of fishing pressure in the Gulf of St. Lawrence, there is no recovery of cod stocks in this ecosystem. In the Sørfjord, after different perturbations (but not from the fishery), the ecosystem seems to return to its equilibrium.     

Spatio‐temporal variations in demersal fish assemblages and diversity in the northern Gulf of St. Lawrence (Canada)

Species diversity is generally considered one of the key factors of ecosystem resilience in response to anthropogenic pressures, including fishing. In this context, the spatial and temporal changes in demersal fish assemblages and species diversity were investigated in the northern Gulf of St. Lawrence (Canada), over a 20‐year period (1990–2010). Data were obtained from the summer research survey conducted by the Department of Fisheries and Oceans, and include commercial and non‐commercial species. The study covers the period of groundfish fishery collapse, the moratorium period, and the post‐moratorium period, and reflects various modifications in management. Multivariate statistical methods revealed two communities. A coastal community corresponds to strata located above 200 m depth and a deeper community located in the deep channels. Interannual differences in the composition of fish assemblages were observed and are mainly due to the changes in the relative biomass of some dominant species. Three diversity indices (Shannon–Wiener, Simpson's Index of Diversity and Motomura's constant) indicate a slight but significant increase of the diversity over time. This trend is due to the increase of the relative biomass of low‐rank species, which may have been favoured by the prohibition of groundfish trawling after 1997 in that region. The geographical distribution of the Shannon–Wiener index also shows temporal dynamics reflecting the biomass distribution of dominant demersal species.     

Changes in trophic flows and ecosystem properties of the Beibu Gulf ecosystem before and after the collapse of fish stocks

Mass-balance models (Ecopath) of the ecosystem before and after collapse (1959-1961 and 1997-1999) of fish stocks were developed with Ecopath software to compare the differences in ecosystem structure, functioning and ecosystem properties of the Beibu Gulf. The model includes 20 functional groups consisting of commercial important fish groups and other ecologically important groups in the ecosystem such as zooplankton, phytoplankton, and detritus. Results indicated that biomass and catches of the system have changed drastically between the 1960s and 1990s, especially for the high trophic levels (TL). The biomass of level V in the early 1960s was 32 times higher than that of the late 1990s, however, the biomass of level I and II in the 1990s was higher than the 1960s. Despite the higher catches in the 1990s, fishing was ecologically less expensive during the 1990s than 1960s due to small fish catches were large. Mean transfer efficiency decreased from for 10.2% in the 1960s to 9.1% in the 1990s periods. According to the summary statistics, the parameters of net system production (NPS) and total primary production to total respiration ratio were increased from 1.013 in the 1960s to 2.184 in the 1990s, however, the connectance index (CI), system omnivore index, Finn’s cycling index and mean path length decreased from the 1960s to the 1990s. The overhead (O) was higher in the 1990s model while the ascendancy (A) decreased nearly 10% in the 1960s. The ‘Keystoneness’ result indicate that zooplankton was identified as keystone species in 1960s, however, the elasmobranches was keystone species in the late 1990s. The average trophic level of the fishery decreased from 3.32 in the 1960s to 2.98 in the 1990s, and exhibits classic symptoms of “fishing down the food web”. All the indices of the system attributes suggests that the Beibu Gulf ecosystem in 1960s was found to be more mature than in the 1990s due to the collapse of demersal ecosystem, and the ecosystem changed from being dominated by long-lived, high trophic level groundfish dominated system toward a system with small-size and low-value species over fifty years.     

An overview of the Oyashio ecosystem

The Oyashio shelf region and the seasonally ice-covered areas north of Hokkaido are highly productive, supporting a wide range of species including marine mammals, seabirds and commercially important species in the western subarctic Pacific. The fishes include gadids, such as walleye pollock and Pacific cod, and subarctic migratory pelagic fishes such as chum salmon and pink salmon. It is also an important summer feeding ground for subtropical migrants such as the Japanese sardine, Japanese anchovy, Pacific saury, mackerels, Japanese common squid, whales and seabirds. In recent decades, some components of the Oyashio ecosystem (i.e., phytoplankton, mesozooplankton, gadid fish, and subtropical migrants) have shown changes in species abundance or distribution that are correlated with environmental changes such as the 1976/1977 and 1988/1989 regime shifts. The First Oyashio Intrusion moved northward from the mid-1960s until the late 1970s, when it moved southward until the 1980s, after which it returned to the north again after the mid-1990s. The sea-surface temperature in spring decreased after the late 1970s, increased after the late 1980s, and remained high during the 1990s. The extent of ice cover in the Sea of Okhostk also decreased during the latest warming in the 1980–1990s but has increased again since the late 1990s. This and other variabilities affect the Oyashio ecosystem and the surrounding region.     

Regime shifts in upwelling ecosystems: observed changes and possible mechanisms in the northern and southern Benguela

A regime shift is considered to be a sudden shift in structure and functioning of a marine ecosystem, affecting several living components and resulting in an alternate state. According to this definition, regime shifts differ from species replacement or alternation of species at similar trophic levels, whereby the ecosystem is not necessarily significantly altered in terms of its structure and function; only its species composition changes. This paper provides an overview of regime shifts, species replacements and alternations that have been observed in the northern and southern Benguela ecosystems over the past few decades. Bottom-up control, initiating and sustaining regime shifts or species replacements via environmental forcing, is documented for both the southern and the northern Benguela ecosystems. Fishing (a case of top-down control) appears to have played an important role in regime shift processes in the Namibian ecosystem. Very low biomass levels of exploited fish stocks associated with less efficient energy transfer in the northern Benguela are indicative of a regime shift. Very high biomass levels have been reached in the southern Benguela in the 2000s. However the alternation between sardine and anchovy that has been observed in the southern Benguela over the last two decades appears not to have had major effects on the overall functioning of the ecosystem. The consequences of regime shifts for exploitation are highlighted, suggesting that fisheries managers should move towards a more effective ecosystem approach to fisheries.     

Top predators in the Benguela ecosystem — implications of their trophic position

Many top predators in the Benguela ecosystem feed on prey species targeted by commercial fisheries. Their roles as indicators of the state of exploited prey resources, as competitors with commercial fisheries for resources, and as susceptible to impact from commercial fishing on those resources are briefly considered. Trends in the occurrence of anchovy Engraulis capensis and pilchard Sardinops ocellatus in the diet of Cape gannets Morus capensis off South Africa's west coast are related significantly to survey estimates of the abundance of these fish species, and they provide useful confirmation of those estimates. In the 1980s, anchovy decreased in the diet of Cape gannets, but pilchard increased. In both the northern and southern portions of the Benguela system, groundfish were thought to eat most (66–73%) of the total quantity of cephalopods and vertebrates consumed by predators and man in the 1980s. South African fur seals Arctocephalus pusillus pusillus, predatory pelagic fish and man removed roughly equal amounts, with squids, seabirds and cetaceans having a smaller impact. In the 1980s, man and seals removed about two million tons live mass more than in 1930. Indices of the rate of natural mortality of anchovy and pilchard attributable to Cape gannets are not related to biomass of the prey species. That for anchovy was high in 1989 when a poor anchovy year-class was formed. Decreased abundance of anchovy led to poor breeding by Cape cormorants Phalacrocorax capensis in 1989 and 1990. A model linking the Cape cormorant population with anchovy is used to explore the impact of possible exploitation strategies for anchovy on Cape cormorants.     

Seals, cod and forage fish: A comparative exploration of variations in the theme of stock collapse and ecosystem change in four Northwest Atlantic ecosystems

The facts: four Northwest Atlantic ecosystems, three cod stock collapses 15 years ago (plus one severely depleted), seals now top predator in all ecosystems, all had cod as a top predator before collapse, groundfish declines in all areas, forage base increased in most systems. No recovery in any system. Have these ecosystems fundamentally changed? Why? The challenge: compare and contrast these four ecosystems. The answer: using mass balance models, empirical data and a suite of ecosystem indicators, we explore how and why these systems have changed over time. At the ecosystem and community level, we see broad similarities between ecosystems. However, structurally and functionally these systems have shifted to an alternate state, with changes in predator structure, trophic structure and flow.     

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.     

The importance of cephalopods as prey for hake and other groundfish in South African waters

Literature on trophic relationships in the Benguela ecosystem has stressed the importance of cephalopods as prey of groundfish. The groundfish community of the shelf and upper slope of southern Africa is dominated by the Cape hakes, and the results presented (1984–1991) confirm that both species of hake are important predators of cephalopods, especially taking into consideration the abundance of hake in the ecosystem. However, geographic, seasonal and species variability are evident in the patterns observed. The main prey species are Sepia spp. (predominantly Sepia australis), Loligo vulgaris reynaudii, Todaropsis eblanae and Lycoteuthis ?diadema. The last-named is an important food organism for fish. Its systematic status needs revision, however. Qualitative results of studies of cephalopod predation are also provided for kingklip and monkfish.     

Long-term impacts of coral bleaching events on the world's warmest reefs

The southern Arabian Gulf houses some of the most thermally tolerant corals on earth, but severe bleaching in the late 1990s caused widespread mortality. More than a decade later, corals still dominated benthos (mean: 40 ± 3% cover on 10 sites spanning > 350 km; range: 11.0-65.6%), but coral communities varied spatially. Sites to the west generally had low species richness and coral cover (mean: 3.2 species per transect, 31% cover), with Porites dominated communities (88% of coral) that are distinct from more diverse and higher cover eastern sites (mean: 10.3 species per transect, 62% cover). These patterns reflect both the more extreme bleaching to the west in the late 1990s as well as the higher faviid dominated recruitment to the east in subsequent years. There has been limited recovery of the formerly dominant Acropora, which now represents <1% of the benthos, likely as a result of recruitment failure. Results indicate that severe bleaching can have substantial long-term impacts on coral communities, even in areas with corals tolerant to environmental extremes.     

Input of metals to the North Atlantic from two large Canadian estuaries

The freshwater discharge from the Gulf of St. Lawrence and Hudson Bay accounts for more than 80% of the total freshwater discharge from eastern Canada and approximately 70% of the total discharge from the eastern seaboard of North America, from Baffin Island to Florida. If the Gulf of Mexico is included, the Gulf of St. Lawrence/Hudson Bay discharge is still more than half of the total.Both the Gulf of St. Lawrence and Hudson Bay have some distinctly estuarine characteristics and discharges of metals can be estimated using estuarine models. For Al, Mn, Fe, Co, Ni, Cu and Zn, the distributions can be described by simple, highly significant relationships with salinity. These regressions can be used in conjunction with knowledge of water discharges to estimate net metal inputs to the North Atlantic from these two systems.Most of the discharge from the Gulf of St. Lawrence flows along the Scotian Shelf to the Gulf of Maine. Metal-salinity relationships can also be used to estimate metal transports through this region for some metals. The applicability of this estuarine model, however, tends to break down as the oceanography becomes more complex and metal distributions are determined more by oceanic rather than estuarine processes.     

Comparing trophic flows in the southern Benguela to those in other upwelling ecosystems

A balanced trophic flow model of the southern Benguela ecosystem is presented, averaging the period 1980–1989 and emphasizing upper trophic levels. The model is based largely on studies conducted within the framework of the Benguela Ecology Programme and updates the results of an expert workshop held in Cape Town in September 1989. Small pelagic fish other than anchovy Engraulis capensis and sardine Sardinops sagax, mainly round herring Etrumeus whiteheadi and mesopelagic fish, were important components of the food web in the southern Benguela. Severe balancing difficulties were encountered with respect to the semi-pelagic resources (hake Merluccius spp.) and demersal top predators (sharks), indicating the need for further research on the interaction of these groups with their ecosystem. The model is compared to other existing trophic flow models of ecosystems in major upwelling areas, i.e. the northern Humboldt Current (4–14°S), the California Current (28–42°N) and the southern Canary Current (l2–25°N), and to two independently constructed models of the northern Benguela ecosystem. These models are compared using network analysis routines of the ECOPATH software, focusing on the interactions between the five dominant fish species (anchovy, sardine, horse mackerel Trachurus trachurus capensis, chub mackerel Scomber japonicus and hake) that support important fisheries in all systems. The upwelling systems rank by size rather than species dominance. The ratio of catches and primary production differs between systems, partly because of differences in fishing regimes. Predation on the five dominant fish groups by other fish in the system was the most important cause of fish mortality in all models. Fishery catches are generally a larger cause of mortality for these groups than predation by mammals. The ecological cost of fishing appears to be comparatively low in the southern Benguela, because catches are low compared with the primary production, but also because the fishery is relatively low in the foodweb. However, in view of the very tight foodweb demonstrated in the model. it is likely that an increase in fishing pressure would cause severe trade-offs with respect to other components of the southern Benguela ecosystem.     

Interannual fluctuations in recruitment of walleye pollock in the Oyashio region related to environmental changes

The Japanese Pacific walleye pollock (Theragra chalcogramma) stock is the largest stock of this species in Japanese waters. It is a key component of the Oyashio ecosystem. In southern Hokkaido waters, these fish spawn mainly during January and February near the mouth of Funka Bay (FB), and most eggs and larvae are transported into FB. During midsummer juvenile pollock migrate along the southern coast of Hokkaido to a nursery ground on the continental shelf off eastern Hokkaido (Doto area). However, some eggs and larvae are transported southward to the Tohoku region (TR). Transport depends largely on the Oyashio, which generally flows southward along the eastern coasts of Hokkaido and Tohoku. Thus, this stock has two different recruitment routes: FB–Doto and FB–TR. In the 1980s, when the southward flow of the Oyashio was strong, the number of age-2 pollock estimated from a virtual population analysis (VPA) indicated that recruitment to the entire stock remained at a medium level. In the 1990s, when the Oyashio weakened, strong year-classes occurred in 1991, 1994, and 1995, but not in the latter half of the 1990s. Juvenile catches in the TR by commercial fisheries, which can be taken as indices of recruitment level via FB–TR, were high during the 1980s and decreased in the 1990s. Although there was no significant difference in the average number of recruits between the 1980s and the 1990s as estimated from a VPA, the recruitment patterns differed between the two decades. Here, we propose that recruitment routes of this stock shifted in response to environmental changes.     

白令海大型底栖动物群落结构及其空间分布格局

Due to its unique geological location, the Bering Sea is an ideal place to investigate the water exchange and ecosystem connectivity of the Pacific Ocean–Arctic Ocean and subarctic–Arctic region. Based on a number of summer surveys(July to September, 2010, 2012 and 2014), macrobenthic communities and their spatial-temporal patterns are exhibited for the majority of the Bering Sea(53°59′–64°36′N). The results show that the macrobenthic communities were dominated by northern cold-water species and immigrant eurythermic species, and the communities assumed a dispersed and patchy distribution pattern. Polychaetes(Scoloplos armiger), crustaceans(Ceradocus capensis) and sea urchins(Echinarachnius parma) were the main dominant groups in the shallow shelves; the sea star(Ctenodiscus crispatus) and the brittle star(Ophiura sarsii) were the main dominant groups in the continental slope; whereas small polychaetes(Prionospio malmgreni) dominated the basin area. Sediment type, water depth, and currents were the major factors affecting the structure and spatial distribution of the macrobenthic communities. Compared with other seas, the shallow areas of the Bering Sea showed an extremely high-standing biomass. In particular, the northern shelf area(north of St. Lawrence Islands and west of 170°W),which is primarily controlled by Anadyr Water, is an undersea oasis. In contrast, a deficiency in the downward transport of particulate organic carbon has resulted in a desert-like seabed in the basin area. By comparing our results to previous studies, we found that macrobenthic communities of the Bering Sea have undergone significant structural changes in recent decades, resulting in a decrease in abundance and an increase in biomass.In addition, populations of amphipods and bivalves in the northern shelves have decreased significantly and have been gradually replaced by other species. These changes might be associated with advanced seasonal ice melting,changes in organic carbon input, and global warming, indicating that large-scale ecosystem changes have been occurring in the Bering Sea.     

胶州湾营养盐结构变化的研究

于１９８５年８月－１９８６年１２月和１９９１年５月－１９９３年１１地胶州湾表层海水营养盐的调查，根据Ｊｕｓｔｉｃ等建立的浮游植物生长的化学计量及可能营养盐限制因素标准，并结合海洋硅藻正常所需的Ｓｉ，Ｎ，Ｐ之间的原子比，用数学统计的方法研究了胶放湾表层海不营养盐结构，结果表明，含有丰富溶我机氮和磷的胶州湾，Ｓｉ：Ｐ的值有显著的下降，同时Ｓｉ：ＤＩＮ和ＤＩＮ：Ｐ的值也表现出下降的趋势，使胶州湾水域营养     

Source regions for recruitment of Calanus finmarchicus to Georges Bank: evidence from molecular population genetic analysis of mtDNA

Molecular population genetic analysis has provided evidence that the copepod, Calanus finmarchicus, of the Labrador Current, Gulf of St Lawrence, Scotian Shelf, Gulf of Maine, and Georges Bank constitute a single, interbreeding population. The DNA sequence of a 350 base pair portion of the mitochondrial large subunit (16S) ribosomal RNA (rRNA) gene was determined for a total of 72 individuals collected in 1992, and 110 individuals collected in 1993 from these regions. There was significant heterogeneity in haplotype frequencies among the samples collected in 1992, but this heterogeneity did not resolve into regional patterns. The only regional differences seen were between pooled samples of the western N. Atlantic and those of the Norwegian Sea. There were no significant differences in haplotype frequencies among the samples collected in 1993, and fewer haplotypes were observed in these samples. Intraspecific molecular variation was typical of other marine species: there were 29 haplotypes among the 182 individuals sequenced. The frequency distribution of the haplotypes was highly skewed: 128 individuals shared one haplotype and 19 individuals were unique. There were 24 variable sites among the 350 bases sequenced; estimated nucleotide diversity was 0.0042. The genetic character of C. finmarchicus populations in the western N. Atlantic was stable over time in that three of the haplotypes (including the most abundant) occurred in both 1992 and 1993. However, haplotype frequencies differed significantly between the two years. The lack of regional structure in the 1992 samples and the genetic homogeneity of samples collected in 1993 across the domain from the Labrador Current to the Gulf of St Lawrence to Georges Bank and the Gulf of Maine indicated that there is significant gene flow across this region. The persistent genetic pattern suggests that the Gulf of St Lawrence may be an important source region for recruitment of C. finmarchicus to Georges Bank. Determination of zooplankton dispersal patterns within high gene flow species will provide information that may not be determined by conventional oceanographic analyses.