Exploitation drives an ontogenetic-like deepening in marine fish.

Virtually all studies reporting deepening with increasing size or age by fishes involve commercially harvested species. Studies of North Sea plaice in the early 1900s first documented this phenomenon (named Heincke's law); it occurred at a time of intensive harvesting and rapid technological changes in fishing methods. The possibility that this deepening might be the result of harvesting has never been evaluated. Instead, age- or size-related deepening have been credited to interactions between density-dependent food resources and density-independent environmental factors. Recently, time-dependent depth variations have been ascribed to ocean warming. We use a model, initialized from observations of Atlantic cod (Gadus morhua) on the eastern Scotian Shelf, where an age-dependent deepening of ∼60 m was observed, to assess the effect of size- and depth-selective exploitation on fish distribution. Exploitation restricted to the upper 80 m can account for ∼72% of the observed deepening; by extending exploitation to 120 m, all of the deepening can be accounted for. In the absence of fishing, the model indicated no age-related deepening. Observations of depth distributions of older cod during a moratorium on fishing supported this prediction; however, younger cod exhibited low-amplitude deepening (10-15 m) suggestive of an ontogenetic response. The implications of these findings are manifold, particularly as they relate to hypotheses advanced to explain the ecological and evolutionary basis for ontogenetic deepening and to recent calls for the adoption of evidence of species deepening as a biotic indicator or "footprint" of warming seas.

Large scale, synchronous variability of marine fish populations driven by commercial exploitation.

Synchronous variations in the abundance of geographically distinct marine fish populations are known to occur across spatial scales on the order of 1,000 km and greater. The prevailing assumption is that this large-scale coherent variability is a response to coupled atmosphere-ocean dynamics, commonly represented by climate indexes, such as the Atlantic Multidecadal Oscillation and North Atlantic Oscillation. On the other hand, it has been suggested that exploitation might contribute to this coherent variability. This possibility has been generally ignored or dismissed on the grounds that exploitation is unlikely to operate synchronously at such large spatial scales. Our analysis of adult fishing mortality and spawning stock biomass of 22 North Atlantic cod (Gadus morhua) stocks revealed that both the temporal and spatial scales in fishing mortality and spawning stock biomass were equivalent to those of the climate drivers. From these results, we conclude that greater consideration must be given to the potential of exploitation as a driving force behind broad, coherent variability of heavily exploited fish species.

Transient dynamics of an altered large marine ecosystem.

Overfishing of large-bodied benthic fishes and their subsequent population collapses on the Scotian Shelf of Canada's east coast and elsewhere resulted in restructuring of entire food webs now dominated by planktivorous, forage fish species and macroinvertebrates. Despite the imposition of strict management measures in force since the early 1990s, the Scotian Shelf ecosystem has not reverted back to its former structure. Here we provide evidence of the transient nature of this ecosystem and its current return path towards benthic fish species domination. The prolonged duration of the altered food web, and its current recovery, was and is being governed by the oscillatory, runaway consumption dynamics of the forage fish complex. These erupting forage species, which reached biomass levels 900% greater than those prevalent during the pre-collapse years of large benthic predators, are now in decline, having outstripped their zooplankton food supply. This dampening, and the associated reduction in the intensity of predation, was accompanied by lagged increases in species abundances at both lower and higher trophic levels, first witnessed in zooplankton and then in large-bodied predators, all consistent with a return towards the earlier ecosystem structure. We conclude that the reversibility of perturbed ecosystems can occur and that this bodes well for other collapsed fisheries.

Fishing, predation, and temperature drive herring decline in a large marine ecosystem.

Since 1960, landings of Atlantic herring have been the greatest of any marine species in Canada, surpassing Atlantic cod and accounting for 24% of the total seafood harvested in Atlantic Canada. The Scotian Shelf-Bay of Fundy herring fisheries (NAFO Division 4VWX) is among Canada's oldest and drives this productivity, accounting for up to 75% of the total herring catch in some years. The stocks' productivity and overall health have declined since 1965. Despite management measures to promote recovery implemented since 2003, biomass remains low and is declining. The factors that drive the productivity of 4VWX herring are primarily unresolved, likely impeding the effectiveness of management actions on this stock. We evaluated potential drivers of herring variability by analyzing 52 time-series that describe the temporal and spatial evolution of the 4VWX herring population and the physical, ecological, and anthropogenic factors that could affect them using structural equation models. Variation in herring biomass was best accounted for by the exploitation rate's negative effect and the geographic distribution of fishing and recruitment. Thermal phenology and temperature adversely and egg predation positively impacted the early life stage mortality rate and, ultimately, adult biomass. These findings are broadly relevant to fisheries management, but particularly for 4VWX herring, where the current management approach does not consider their early life stage dynamics or assess them within the ecosystem or climate change contexts.

Decline in top predator body size and changing climate alter trophic structure in an oceanic ecosystem.

Globally, overfishing large-bodied groundfish populations has resulted in substantial increases in their prey populations. Where it has been examined, the effects of overfishing have cascaded down the food chain. In an intensively fished area on the western Scotian Shelf, Northwest Atlantic, the biomass of prey species increased exponentially (doubling time of 11 years) even though the aggregate biomass of their predators remained stable over 38 years. Concomitant reductions in herbivorous zooplankton and increases in phytoplankton were also evident. This anomalous trophic pattern led us to examine how declines in predator body size (approx. 60% in body mass since the early 1970s) and climatic regime influenced lower trophic levels. The increase in prey biomass was associated primarily with declines in predator body size and secondarily to an increase in stratification. Sea surface temperature and predator biomass had no influence. A regression model explained 65 per cent of prey biomass variability. Trait-mediated effects, namely a reduction in predator size, resulted in a weakening of top predation pressure. Increased stratification may have enhanced growing conditions for prey fish. Size-selective harvesting under changing climatic conditions initiated a trophic restructuring of the food chain, the effects of which may have influenced three trophic levels.

Temporal dynamics within a contemporary latitudinal diversity gradient.

Poleward declines in species diversity [latitudinal diversity gradients (LDG)] remain among the oldest and most widespread of macroecological patterns. However, their contemporary dynamics remain largely unexplored even though changing ecological conditions, including global change, may modify LDG and their respective ecosystems. Here, we examine temporal variation within a temperate Northwest Atlantic LDG using 31 years of annual fisheries-independent surveys and explore its dynamics in relation to a dominant climate signal [the wintertime North Atlantic Oscillation (NAO)] that varies interannually and alters the latitudinal gradient of Northwest Atlantic continental shelf bottom water temperatures. We found that the slopes of the annual LDG vary dramatically due to changes in geographic distributions of 100+ species, variations that are concealed within the cumulative, static LDG. These changes are strongly associated with changes in NAO sign and strength. This is the first illustration of temporal dynamics in a contemporary LDG and the first demonstration of the speed at which local environmental variations can alter an LDG. Our findings underscore the need to investigate factors that modify LDG separately from those that contribute to their origins.

Environmental structuring of marine plankton phenology.

Seasonal cycles of primary production (phenology) critically influence biogeochemical cycles, ecosystem structure and climate. In the oceans, primary production is dominated by microbial phytoplankton that drift with currents, and show rapid turnover and chaotic dynamics, factors that have hindered understanding of their phenology. We used all available observations of upper-ocean phytoplankton concentration (1995-2015) to describe global patterns of phytoplankton phenology, the environmental factors that structure them, and their relationships to terrestrial patterns. Phytoplankton phenologies varied strongly by latitude and productivity regime: those in high-production regimes were governed by insolation, whereas those in low-production regimes were constrained by vertical mixing. In eight of ten ocean regions, our findings contradict the hypothesis that phytoplankton phenologies are coherent at basin scales. Lastly, the spatial organization of phenological patterns in the oceans was broadly similar to those on land, suggesting an overarching effect of insolation on the phenology of primary producers globally.

Reconciling differences in trophic control in mid-latitude marine ecosystems.

The dependence of long-term fishery yields on primary productivity, largely based on cross-system comparisons and without reference to the potential dynamic character of this relationship, has long been considered strong evidence for bottom-up control in marine systems. We examined time series of intensive empirical observations from nine heavily exploited regions in the western North Atlantic and find evidence of spatial variance of trophic control. Top-down control dominated in northern areas, the dynamics evolved from bottom-up to top-down in an intermediate region, and bottom-up control governed the southern areas. A simplified, trophic control diagram was developed accounting for top-down and bottom-up forcing within a larger region whose base state dynamics are bottom-up and can accommodate time-varying dynamics. Species diversity and ocean temperature co-varied, being relatively high in southern areas and lower in the north, mirroring the shifting pattern of trophic control. A combination of compensatory population dynamics and accelerated demographic rates in southern areas seems to account for the greater stability of the predator species complex in this region.

The ups and downs of trophic control in continental shelf ecosystems.

Traditionally, marine ecosystem structure was thought to be determined by phytoplankton dynamics. However, an integrated view on the relative roles of top-down (consumer-driven) and bottom-up (resource-driven) forcing in large-scale, exploited marine ecosystems is emerging. Long time series of scientific survey data, underpinning the management of commercially exploited species such as cod, are being used to diagnose mechanisms that could affect the composition and relative abundance of species in marine food webs. By assembling published data from studies in exploited North Atlantic ecosystems, we found pronounced geographical variation in top-down and bottom-up trophic forcing. The data suggest that ecosystem susceptibility to top-down control and their resiliency to exploitation are related to species richness and oceanic temperature conditions. Such knowledge could be used to produce ecosystem guidelines to regulate and manage fisheries in a sustainable fashion.

Trophic cascades in a formerly cod-dominated ecosystem.

Removal of top predators from ecosystems can result in cascading effects through the trophic levels below, completely restructuring the food web. Cascades have been observed in small-scale or simple food webs, but not in large, complex, open-ocean ecosystems. Using data spanning many decades from a once cod-dominated northwest Atlantic ecosystem, we demonstrate a trophic cascade in a large marine ecosystem. Several cod stocks in other geographic areas have also collapsed without recovery, suggesting the existence of trophic cascades in these systems.