Lorenzo Ciannelli

NONADDITIVE EFFECTS OF THE ENVIRONMENT ON THE SURVIVAL OF A LARGE MARINE FISH POPULATION

Climate can affect population dynamics in indirect ways via nonadditive forcing by external variables on internal demographic rates. Current analytical techniques, employed in population ecology, fail to explicitly include nonadditive interactions between internal and external variables, and therefore cannot efficiently address indirect climate effects. Here, we present the results of an analysis, employing specifically developed statistical methodology, on density-dependent survival of walleye pollock (Theragra chalcogramma) prerecruitment stages in relation to background environmental variables in the Gulf of Alaska. We found that spring winds and water temperature mediate the intensity of density-dependent survival from the eggs to the age-0 stage. Fall water temperature and juvenile pollock predator abundance mediate density dependence from the age-0 to the age-1 stage. The inclusion of such nonadditive and nonlinear effects in a population dynamics model improved our ability to simulate pollock recruitment. Our results point to the importance of understanding nonadditive and nonlinear interactions between external (climate) and internal factors in the presence of underlying environmental variation. These topics are discussed in the context of current research priorities in population ecology and conservation biology.

DENSITY DEPENDENCE AND DENSITY INDEPENDENCE DURING THE EARLY LIFE STAGES OF FOUR MARINE FISH STOCKS

Recruitment variability caused by density-dependent and density-independent processes is an important area within the study of fish dynamics. These processes can exhibit nonlinearities and nonadditive properties that may have profound dynamic effects. We investigate the importance of population density (i.e., density dependence) and environmental forcing (i.e., density independence) on the age-0 and age-1 abundance of capelin (Mallotus villosus), northeast Arctic cod (Gadus morhua), northeast Arctic haddock (Melanogrammus aeglefinus), and Norwegian spring spawning herring (Clupea harengus) in the Barents Sea. We use statistical methods that explicitly account for nonlinearities and nonadditive interactions between internal and external variables in the abundance of these two pre-recruitment stages. Our results indicate that, during their first five months of life, cod, haddock, and herring experience higher density-dependent survival than capelin. The abundance of age-0 cod depends on the mean age and biomass of the spawning stock, a result which has implications for the management of the entire cod stock. Temperature is another important factor influencing the abundance at age-0 and age-1 of all four species, except herring at age-1. Between age-0 and age-1, there is an attenuation of density-dependent survival for cod and herring, while haddock and capelin experience density dependence at high and low temperatures, respectively. Predation by subadult cod is important for both capelin and cod at age-1. We found strong indications for interactions among the studied species, pointing to the importance of viewing the problem of species recruitment variability as a community, rather than as a population phenomenon.

SPATIAL ANATOMY OF SPECIES SURVIVAL: EFFECTS OF PREDATION AND CLIMATE‐DRIVEN ENVIRONMENTAL VARIABILITY

The majority of survival analyses focus on temporal scales. Consequently, there is a limited understanding of how species survival varies over space and, ultimately, how spatial variability in the environment affects the temporal dynamics of species abundance. Using data from the Barents Sea, we study the spatiotemporal variability of the juvenile Atlantic cod (Gadus morhua) survival. We develop an index of spatial survival based on changes of juvenile cod distribution through their first winter of life (from age-0 to age-1) and study its variability in relation to biotic and abiotic factors. Over the 25 years analyzed (1980-2004), we found that, once the effect of passive drift due to dominant currents is accounted for, the area where age-0 cod survival was lowest coincided with the area of highest abundance of older cod. Within this critical region, the survival of age-0 cod was negatively affected by its own abundance, by that of older cod, and by bottom depth. Furthermore, during cold years, age-0 cod survival increased in the eastern and coldest portion of the examined area, which was typically avoided by older conspecifics. Based on these results we propose that within the examined area top-down mechanisms and predation-driven density dependence can strongly affect the spatial pattern of age-0 cod survival. Climate-related variables can also influence the spatial survival of age-0 cod by affecting their distribution and that of their predators. Results from these and similar studies, focusing on the spatial variability of survival rates, can be used to characterize species habitat quality of marine renewable resources.

Climate change causing phase transitions of walleye pollock (Theragra chalcogramma) recruitment dynamics

In 1976 the North Pacific climate shifted, resulting in an average increase of the water temperature. In the Gulf of Alaska the climate shift was followed (i.e. early 1980s) by a gradual but dramatic increase in the abundance of groundfish species that typically prey on pre-recruitment stages of walleye pollock. In the present study we used a previously parameterized model to investigate the effect of these climate and biological changes on the recruitment dynamics of walleye pollock in the Gulf of Alaska. Simulations covered the 1970–2000 time frame and emphasized the medium-to-long temporal scale (i.e. about 5–10 years) of environmental variability. Results showed that during periods characterized by high sea surface temperature and high predation on juvenile pollock stages, recruitment variability and magnitude were below average, and recruitment control was delayed to stages older than the 0-group. Opposite dynamics (i.e. high abundance and variability, and early recruitment control) occurred during periods characterized by low temperature and predation. These results are in general agreement with empirical observations, and allowed us to formulate causal explanations for their occurrence. We interpreted the delay of recruitment control and the reduction of variability as an effect of increased constraint on the abundance of post age-0 stages, in turn imposed by high density dependence and predation mortality. On the other hand, low density-dependence and predation favoured post age-0 survival, and allowed for an unconstrained link between larval and recruitment abundance. Our findings demonstrate that the dominant mechanisms of pollock survival change over contrasting climate regimes. Such changes may in turn cause a phase transition of recruitment dynamics with profound implications for the management of the entire stock.

Small-scale genetic structure in a marine population in relation to water circulation and egg characteristics

Until the last decade it was assumed that most marine species have pronounced gene flow over vast areas, largely because of their potential for dispersal during early life stages. However, recent genetic, modeling, and field studies have shown that marine populations may be structured at scales that are inconsistent with extensive dispersal of eggs and larvae. Such findings have stimulated the birth of new studies explaining the mechanisms that promote population structure and isolation in the oceans, in the face of high potential for dispersal. Here we study the vertical and horizontal distribution of cod (Gadus morhua) eggs in relation to small-scale circulation and water column hydrography in a coastal location of southern Norway. Previous studies conducted in this region have shown that cod populations inhabiting fjord locations, which are on average 30 km apart, are genetically differentiated, a remarkable outcome considering that Atlantic cod have pelagic egg stages and long pelagic larval duration. We document that cod eggs are found in greater abundance in shallow water layers, which on average are flowing up the fjord (away from the open ocean), and in the inner portion of the fjord, which is subject to lower current speeds compared to the outer or mouth of the fjord. Eggs were found to be neutrally buoyant at shallow depths, a trait that also favors local retention, given the local circulation. The same patterns held during two environmentally contrasting years. These results strongly suggest that population structure of Atlantic cod is favored and maintained by a balance between water circulation and egg characteristics.

Ontogenetic and sex‐specific differences in density‐dependent habitat selection of a marine fish population

The spatial dynamics of species are the result of complex interactions between density-independent and density-dependent sources of variability. Disentangling these two sources of variability has challenged ecologists working in both terrestrial and aquatic ecosystems. Using a novel spatially explicit statistical model, we tested for the presence of density-independent and density-dependent habitat selection in yellowfin sole (Limanda aspera) in the eastern Bering Sea. We found specificities in the density-dependent processes operating across ontogeny and particularly with gender. Density-dependent habitat expansion occurred primarily in females, and to a lesser degree in males. These patterns were especially evident in adult stages, while juvenile stages of both sexes exhibited a mix of different dynamics. Association of yellowfin sole with substrate type also varied by sex and to a lesser degree with size, with large females distributed over a wider range of substrates than males. Moreover, yellowfin sole expanded northward as cold subsurface waters retracted in summer, suggesting high sensitivity to arctic warming. Our findings illustrate how marginal habitats can play an important role in buffering density-dependent habitat expansion, with direct implications for resource management. Our spatially explicit modeling approach is effective in evaluating density-dependent spatial dynamics, and can easily be used to test similar hypotheses from a variety of aquatic and terrestrial ecosystems.

Seasonal plankton dynamics along a cross-shelf gradient.

Much interest has recently been devoted to reconstructing the dynamic structure of ecological systems on the basis of time-series data. Using 10 years of monthly data on phyto- and zooplankton abundance from the Bay of Biscay (coastal to shelf-break sites), we demonstrate that the interaction between these two plankton components is approximately linear, whereas the effects of environmental factors (nutrients, temperature, upwelling and photoperiod) on these two plankton population growth rates are nonlinear. With the inclusion of the environmental factors, the main observed seasonal and inter-annual dynamic patterns within the studied plankton assemblage also indicate the prevalence of bottom-up regulatory control.

Non-additive and non-stationary properties in the spatial distribution of a large marine fish population

Density-independent and density-dependent variables both affect the spatial distributions of species. However, their effects are often separately addressed using different analytical techniques. We apply a spatially explicit regression framework that incorporates localized, interactive and threshold effects of both density-independent (water temperature) and density-dependent (population abundance) variables, to study the spatial distribution of a well-monitored flatfish population in the eastern Bering Sea. Results indicate that when population biomass was beyond a threshold a further increase in biomass-promoted habitat expansion in a non-additive fashion with water temperature. In contrast, during years of low population size, habitat occupancy was affected positively only by water temperature. These results reveal the spatial signature of intraspecific abundance distribution relationships as well as the non-additive and non-stationary responses of species spatial dynamics. Furthermore, these results underscore the importance of implementing analytical techniques that can simultaneously account for density-dependent and density-independent sources of variability when studying geographical distribution patterns.

Interannual and regional variability in distribution and ecology of juvenile pollock and their prey in frontal structures of the Bering Sea

Abstract The distribution, size, length-specific weight, growth, and feeding of age-0 walleye pollock ( Theragra chalcogramma ) were examined along with their prey distribution patterns in two contrasting transects over a 4-year period (1994–1997) in relation to biophysical properties of frontal regions around the Pribilof Islands, Bering Sea. There were significant interannual differences in catch of age-0 pollock, but transect and habitat differences (inshore vs. front vs. offshore) were not significant for either catch or size of pollock. There were significant variations in length-specific weight and growth of pollock, but the trends were inconsistent. Copepods dominated the zooplankton biomass in all habitats and years; there were no consistent differences in the densities of the dominant zooplankton taxa among the habitats. There were, however, strong habitat and transect differences in juvenile pollock diet, particularly for the larger and presumably rarer prey taxa (euphausiids, chaetognaths, fish). We did not find any evidence that occupying a particular habitat was beneficial to young pollock, although other factors (e.g. bioenergetic advantage and predation refuge) that we did not examine here could have been more variable and critical to pollock survival. In a physically dynamic system such as the Pribilof Islands, age-0 pollock may need to continuously search for optimal conditions of high prey availability and low predation pressure.

BOUNDARIES OF OPEN MARINE ECOSYSTEMS: AN APPLICATION TO THE PRIBILOF ARCHIPELAGO, SOUTHEAST BERING SEA

We applied ecosystem energetics and foraging theory to characterize the spatial extent of the Pribilof Archipelago ecosystem, located in the southeast Bering Sea. From an energetic perspective, an ecosystem is an area within which the predatory demand is in balance with the prey production. From a foraging perspective, an ecosystem boundary should at least include the foraging range of the species that live within it for a portion of their life cycle. The Pribilof Islands are densely populated by species that adopt a central place foraging strategy. Foraging theory predicts that the area traveled by central place foragers (CPF) should extend far enough so that their predatory demands are in balance with prey production. Thus, the spatial extent of an ecosystem, as defined by energetics and the foraging range of constituent species, will require a similar energy balance, and independent assessments should yield similar results. In this study, we compared the area of maximum energy balance, estimated with a food web model during the decade 1990- 2000, with estimates of the foraging range of northern fur seals (the farthest traveling CPF in the Pribilof Islands community) obtained from the literature. From the food web sim- ulations, we estimated that a circle of 100 nautical miles (NM), or 185.2 km, radius encloses the area of highest energy balance and lowest biomass import and that it represents a switch from a piscivorous-dominated (smaller areas) to a zooplanktivorous-dominated (larger ar- eas) community. The distance from the breeding site to locations recorded at sea for lactating female fur seals, during the years 1995-1996, ranged from 5.0 to 172.2 NM (9.3-318.9 km), with a median of 97 NM (179.6 km). Thus, ;50% of the locations recorded for lactating fur seals occurred beyond the area of energy balance estimated by the model, indicating that additional factors can motivate their foraging extent. We propose that en- ergetic constraints set the minimum extent of the Pribilof ecosystem, while the foraging distance of fur seals may indicate the maximum extent. In discussing these results, we highlight the limitations of current definitions of the spatial extent in ecosystems, when related to open oceanic environments, and discuss viable alternatives to characterize bound- aries of aquatic systems that are not physically separated from adjacent areas. We believe that these arguments, though controversial, are very timely given the increased emphasis currently placed on the management and protection of entire marine ecosystems.