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.

From mice to elephants: overturning the 'one size fits all' paradigm in marine plankton food chains.

It is widely believed that consumer control is a weak regulator of marine phytoplankton communities. It remains unclear, however, why this should be the case when marine consumers routinely regulate their prey at higher trophic levels. One possibility is that the weak consumer control of phytoplankton communities results from the inability of field researchers to effectively account for consumer-prey trophic relationships operating at the scale of the plankton. We explored this issue by reviewing studies of trophic control in marine plankton. Experimental studies indicate that size is a critical determinant of feeding relationships among plankton. In sharp contrast, of the 51 field studies reviewed, 78% did not distinguish among the sizes or species of phytoplankton and their consumers, but instead assumed a general bulk phytoplankton-zooplankton trophic connection. Such an approach neglects the possibility that several trophic connections may separate the smallest phytoplankton (0.2 µm) from the larger zooplankton (~ 1000 µm), a remarkable size differential exceeding that between a mouse (~10 cm) and an elephant (~2500 cm). The size-based approach we propose integrates theory, experiments and field observations and has the potential to greatly enhance our understanding of the causes and consequences of recently documented restructuring of plankton communities.

Spatial patterns and predictors of trophic control in marine ecosystems.

A key question in ecology is under which conditions ecosystem structure tends to be controlled by resource availability vs. consumer pressure. Several hypotheses derived from theory, experiments and observational field studies have been advanced, yet a unified explanation remains elusive. Here, we identify common predictors of trophic control in a synthetic analysis of 52 observational field studies conducted within marine ecosystems across the Northern Hemisphere and published between 1951 and 2014. Spatial regression analysis of 45 candidate variables revealed temperature to be the dominant predictor, with unimodal effects on trophic control operating both directly (r(2) = 0.32; P < 0.0001) and indirectly through influences on turnover rate and quality of primary production, biodiversity and omnivory. These findings indicate that temperature is an overarching determinant of the trophic dynamics of marine ecosystems, and that variation in ocean temperature will affect the trophic structure of marine ecosystems through both direct and indirect mechanisms.

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.

Breaking Bergmann's rule: truncation of Northwest Atlantic marine fish body sizes.

A strictly species-centric view of human impacts on ecological communities may conceal important trait changes key to ecosystem functioning and stability. Analyses of body size and community composition data for 326 Northwest Atlantic fish species sampled across > 900000 km2 over three decades revealed a rapid and widespread reduction of body sizes driven by declines within species and changes in relative abundances. The changes were unrelated to species richness but of sufficient magnitude to eliminate biogeographic scale gradients of increasing body size with latitude commonly characterized as Bergmann's rule. These changes have persisted despite reduced potential for intraspecific competition and favorable bottom water temperatures, both of which should lead to increased growth rates. The aggregate body sizes in these Northwest Atlantic fish communities may now represent a mismatch between the environmental variability characteristic of the Northwest Atlantic and the historical body size, life history traits, and productivity of species across this region. We discuss how these changes may jeopardize the potential for recovery of these important temperate/subarctic ecosystems.

Maternal influences on population dynamics: evidence from an exploited freshwater fish.

We used a field experiment, population modeling, and an analysis of 30 years of data from walleye (Sander vitreus; a freshwater fish) in Lake Erie to show that maternal influences on offspring survival can affect population dynamics. We first demonstrate experimentally that the survival of juvenile walleye increases with egg size (and, to a lesser degree, female energy reserves). Because egg size in this species tends to increase with maternal age, we then model these maternal influences on offspring survival as a function of maternal age to show that adult age structure can affect the maximum rate at which a population can produce new adults. Consistent with this hypothesis, we present empirical evidence that the maximum reproductive rate of an exploited population of walleye was approximately twice as high when older females were abundant as compared to when they were relatively scarce. Taken together, these results indicate that age- or size-based maternal influences on offspring survival can be an important mechanism driving population dynamics and that exploited populations could benefit from management strategies that protect, rather than target, reproductively valuable individuals.

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.

Effects of maternally transferred organochlorine contaminants on early life survival in a freshwater fish.

Laboratory research has shown that female fish can pass toxic organochlorines (OCs) from their bodies to their eggs, killing their offspring if sufficient quantities are transferred. We conducted a controlled incubation study using gametes from a wild, OC-contaminated walleye (Sander vitreus) population (Bay of Quinte, Lake Ontario, Canada) in order to assess among-female variation in offspring early life survival in relation to ova concentrations of planar OCs (polychlorinated dibenzo-p-dioxins and furans and planar polychlorinated biphenyls) and a suite of other maternal and ova characteristics. Equal volumes of ova from each female were fertilized, pooled, and incubated together as an experimental cohort. Relative survival of each female's offspring was estimated as the proportion of surviving larvae (at approximately 5 d posthatch) that she contributed to the cohort as determined by microsatellite DNA parentage assignment. Total planar OC concentration (expressed as toxic equivalency of 2,3,7,8-tetrachlorodibenzo-p-dioxin) of ova was positively related to maternal age and size and to ova lipid content. However, early life survival did not decline with increasing ova planar OC concentrations. Similarly, we observed no significant relationships between early life survival and ova thiamine content, ova fatty acid composition, or maternal age or size. Early life survival was more strongly correlated with date of spawn collection, thyroid hormone status of the ova, and ovum size. Maternally transferred planar OCs do not appear to negatively influence female reproductive success in this walleye population.

Temporal changes in mercury bioaccumulation by predatory fishes of boreal lakes following the invasion of an exotic forage fish.

We evaluated the prediction that mercury concentrations of predatory fishes in boreal lakes would rise following the invasion of an exotic forage fish species (rainbow smelt, Osmerus mordax) that was believed to feed at a higher trophic position than native forage fishes. We compared temporal trends (postinvasion minus preinvasion values) in fish mercury bioaccumulation between lakes experiencing recent smelt invasions and reference lakes of central Canada. Piscivore mercury concentrations in this region have remained stable or declined during approximately the last 20 years. These trends were not strongly influenced by the smelt invasion, despite the fact that smelt were a major prey item for all piscivore species examined. The effect of smelt invasion on mercury bioaccumulation in the predator species reflected the importance of smelt in their respective diets (lake trout, Salvelinus namaycush > walleye, Stizostedion vitreum > northern pike, Esox lucius). However, these effects were not statistically significant for any piscivore species. The impact of rainbow smelt invasion on mercury bioaccumulation in native piscivores of this region has been much less than previous food-web studies have predicted.

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.