Decadal Variability in the Northeast Pacific in a Physical-Ecosystem Model: Role of Mixed Layer Depth and Trophic Interactions

A basin-wide interdecadal change in both the physical state and the ecology of the North Pacific occurred near the end of 1976. Here we use a physical-ecosystem model to examine whether changes in the physical environment associated with the 1976-1977 transition influenced the lower trophic levels of the food web and if so by what means. The physical component is an ocean general circulation model, while the biological component contains 10 compartments: two phytoplankton, two zooplankton, two detritus pools, nitrate, ammonium, silicate, and carbon dioxide. The model is forced with observed atmospheric fields during 1960-1999. During spring, there is a similar to 40% reduction in plankton biomass in all four plankton groups during 1977-1988 relative to 1970-1976 in the central Gulf of Alaska (GOA). The epoch difference in plankton appears to be controlled by the mixed layer depth. Enhanced Ekman pumping after 1976 caused the halocline to shoal, and thus the mixed layer depth, which extends to the top of the halocline in late winter, did not penetrate as deep in the central GOA. As a result, more phytoplankton remained in the euphotic zone, and phytoplankton biomass began to increase earlier in the year after the 1976 transition. Zooplankton biomass also increased, but then grazing pressure led to a strong decrease in phytoplankton by April followed by a drop in zooplankton by May: Essentially, the mean seasonal cycle of plankton biomass was shifted earlier in the year. As the seasonal cycle progressed, the difference in plankton concentrations between epochs reversed sign again, leading to slightly greater zooplankton biomass during summer in the later epoch.

Kelp Beds as Habitat for American Lobster Homarus Americanus

The American lobster Homarus americanus and kelp Laminaria longicruris and L. saccharina are prominent and often intimately associated members of the subtidal community in the western North Atlantic Ocean. However, no one has identified the nature of this relationship or specifically investigated whether kelp beds are a superior habitat for lobsters. We conducted field studies in 1990 and 1991 at a coastal site centrally located along the Gulf of Maine, USA, to determine how lobsters use kelp beds as habitat. Identically sized and spaced plots of live and artificial (plastic) kelp were established and monitored for lobster population densities. Adjacent featureless sediment plots of identical size served as controls. Lobster population density and biomass were significantly higher in both real and artificial kelp treatments than in non-kelp control plots (p < 0.0001). The change in lobster density was apparent the day following placement of the experiment, so a secondary trophic effect such as attracting prey into treatments is unlikely to have occurred. Thus, kelp beds can affect local lobster population densities by providing shelter for lobsters, thereby concentrating individuals and increasing the local carrying capacity of potential lobster habitats. The effect of kelp beds on the local carrying capacity of lobster habitats was further explored by testing how lobsters respond to differing patch sizes. A graded size series of circular patches of artificial kelp was established, in which kelp blade density and total area were held constant for each treatment. Treatments were subdivided into four 1 M2, two 2 M2, or one 4 m2 patches. Experiments were surveyed for lobster population density and size structure to determine ff statistical differences existed among treatments. Lobster density was significantly greater in the smallest patches (p < 0.001). Moreover, lobsters typically occupied the edges of kelp beds, and their abundance within kelp patches corresponded to the patch's perimeter-to-area relationship. This suggests that edge effects' influence the local carrying capacity for lobsters by influencing the lobsters' choice of kelp beds as habitat.

Assessment of Skill and Portability in Regional Marine Biogeochemical Models: Role of Multiple Planktonic Groups

Application of biogeochemical models to the study of marine ecosystems is pervasive, yet objective quantification of these models' performance is rare. Here, 12 lower trophic level models of varying complexity are objectively assessed in two distinct regions (equatorial Pacific and Arabian Sea). Each model was run within an identical one-dimensional physical framework. A consistent variational adjoint implementation assimilating chlorophyll-a, nitrate, export, and primary productivity was applied and the same metrics were used to assess model skill. Experiments were performed in which data were assimilated from each site individually and from both sites simultaneously. A cross-validation experiment was also conducted whereby data were assimilated from one site and the resulting optimal parameters were used to generate a simulation for the second site. When a single pelagic regime is considered, the simplest models fit the data as well as those with multiple phytoplankton functional groups. However, those with multiple phytoplankton functional groups produced lower misfits when the models are required to simulate both regimes using identical parameter values. The cross-validation experiments revealed that as long as only a few key biogeochemical parameters were optimized, the models with greater phytoplankton complexity were generally more portable. Furthermore, models with multiple zooplankton compartments did not necessarily outperform models with single zooplankton compartments, even when zooplankton biomass data are assimilated. Finally, even when different models produced similar least squares model-data misfits, they often did so via very different element flow pathways, highlighting the need for more comprehensive data sets that uniquely constrain these pathways.

Phytoplankton Scales of Variability in the California Current System: 1. Interannual and Cross-Shelf Variability

In the California Current System, strong mesoscale variability associated with eddies and meanders of the coastal jet play an important role in the biological productivity of the area. To assess the dominant timescales of variability, a wavelet analysis is applied to almost nine years (October 1997 to July 2006) of 1-km-resolution, 5-day-averaged, Sea-viewing Wide Field-of-view Sensor (SeaWiFS) chlorophyll a (chl a) concentration data. The dominant periods of chlorophyll variance, and how these change in time, are quantified as a function of distance offshore. The maximum variance in chlorophyll occurs with a period of similar to 100-200 days. A seasonal cycle in the timing of peak variance is revealed, with maxima in spring/summer close to shore (20 km) and in autumn/winter 200 km offshore. Interannual variability in the magnitude of chlorophyll variance shows maxima in 1999, 2001, 2002, and 2005. There is a very strong out-of-phase correspondence between the time series of chlorophyll variance and the Pacific Decadal Oscillation (PDO) index. We hypothesize that positive PDO conditions, which reflect weak winds and poor upwelling conditions, result in reduced mesoscale variability in the coastal region, and a subsequent decrease in chlorophyll variance. Although the chlorophyll variance responds to basin-scale forcing, chlorophyll biomass does not necessarily correspond to the phase of the PDO, suggesting that it is influenced more by local-scale processes. The mesoscale variability in the system may be as important as the chl a biomass in determining the potential productivity of higher trophic levels.

Kelp Forest Ecosystems: Biodiversity, Stability, Resilience and Future

Kelp forests are phyletically diverse, structurally complex and highly productive components of cold-water rocky marine coastlines. This paper reviews the conditions in which kelp forests develop globally and where, why and at what rate they become deforested. The ecology and long archaeological history of kelp forests are examined through case studies from southern California, the Aleutian Islands and the western North Atlantic, well-studied locations that represent the widest possible range in kelp forest biodiversity. Global distribution of kelp forests is physiologically constrained by light at high latitudes and by nutrients, warm temperatures and other macrophytes at low latitudes. Within mid-latitude belts (roughly 40-60degrees latitude in both hemispheres) well-developed kelp forests are most threatened by herbivory, usually from sea urchins. Overfishing and extirpation of highly valued vertebrate apex predators often triggered herbivore population increases, leading to widespread kelp deforestation. Such deforestations have the most profound and lasting impacts on species-depauperate systems, such as those in Alaska and the western North Atlantic. Globally urchin-induced deforestation has been increasing over the past 2-3 decades. Continued fishing down of coastal food webs has resulted in shifting harvesting targets from apex predators to their invertebrate prey, including kelp-grazing herbivores. The recent global expansion of sea urchin harvesting has led to the widespread extirpation of this herbivore, and kelp forests have returned in some locations but, for the first time, these forests are devoid of vertebrate apex predators. In the western North Atlantic, large predatory crabs have recently filled this void and they have become the new apex predator in this system. Similar shifts from fish- to crab-dominance may have occurred in coastal zones of the United Kingdom and Japan, where large predatory finfish were extirpated long ago. Three North American case studies of kelp forests were examined to determine their long history with humans and project the status of future kelp forests to the year 2025. Fishing impacts on kelp forest systems have been both profound and much longer in duration than previously thought. Archaeological data suggest that coastal peoples exploited kelp forest organisms for thousands of years, occasionally resulting in localized losses of apex predators, outbreaks of sea urchin populations and probably small-scale deforestation. Over the past two centuries, commercial exploitation for export led to the extirpation of sea urchin predators, such as the sea otter in the North Pacific and predatory fishes like the cod in the North Atlantic. The largescale removal of predators for export markets increased sea urchin abundances and promoted the decline of kelp forests over vast areas. Despite southern California having one of the longest known associations with coastal kelp forests, widespread deforestation is rare. It is possible that functional redundancies among predators and herbivores make this most diverse system most stable. Such biodiverse kelp forests may also resist invasion from non-native species. In the species-depauperate western North Atlantic, introduced algal competitors carpet the benthos and threaten future kelp dominance. There, other non-native herbivores and predators have become established and dominant components of this system. Climate changes have had measurable impacts on kelp forest ecosystems and efforts to control the emission of greenhouse gasses should be a global priority. However, overfishing appears to be the greatest manageable threat to kelp forest ecosystems over the 2025 time horizon. Management should focus on minimizing fishing impacts and restoring populations of functionally important species in these systems.

Interannual Variability in Timing of Bloom Initiation in the California Current System

In the California Current System the spring transition from poleward to equatorward alongshore wind stress heralds the beginning of upwelling-favorable conditions. The phytoplankton response to this transition is investigated using 8 years ( 1998-2005) of daily, 4-km resolution, Sea-viewing Wide Field of view Sensor ( SeaWiFS) chlorophyll a concentration data. Cluster analysis of the chlorophyll a time series at each location is used to separate the inshore upwelling region from offshore and oligotrophic areas. An objective method for estimating the timing of bloom initiation is used to construct a map of the mean bloom start date. Interannual variability in bloom timing and magnitude is investigated in four regions: 45 degrees N - 50 degrees N, 40 degrees N - 45 degrees N, 35 degrees N - 40 degrees N and 20 degrees N - 35 degrees N. Daily satellite derived wind data ( QuikSCAT) allow the timing of the first episode of persistently upwelling favorable winds to be estimated. Bloom initiation generally coincides with the onset of upwelling winds ( +/- 15 days). South of similar to 35 degrees N, where winds are southward year-round, the timing of increased chlorophyll concentration corresponds closely to timing of the seasonal increase in upwelling intensity. A 1-D model and satellite derived photosynthetically available radiation data are used to estimate time series of depth- averaged irradiance. In the far north of the region (> 46 degrees N) light is shown to limit phytoplankton growth in early spring. In 2005 the spring bloom in the northern regions (> 35 degrees N) had a "false start''. A sharp increase in chl a in February quickly receded, and a sustained increase in biomass was delayed until July. We hypothesize that this resulted in a mismatch in timing of food availability to higher trophic levels.

Seasonal Variability in the Bacteriolytic Capacity of the Deposit Feeder Arenicola Marina: Environmental Correlates

Although deposit-feeding macrofauna consume and digest sedimentary bacteria, it is unclear whether feeding rates and digestion efficiencies are high enough to significantly impact the composition and abundance of bacteria in marine sediments. It is likely that both feeding rates and efficiency of digestion vary markedly through space and time. We used a turbidimetric assay to compare the rate of bacteriolysis by digestive fluids collected seasonally from the deposit-feeding polychaete Arenicola marina. Under standardized, experimental conditions, bacteriolytic rates represent concentrations of lytic agents. This concentration was found to vary significantly throughout the year (p = 0.001), showing greater than a 2x range. Lytic agent concentration was positively correlated with bioavailable amino acid concentrations in the surface sediment (r = 0.85, p = 0.03) but showed no apparent relationship to other proxies for food resources (e.g, chl a), sediment temperature, or gut throughput time. In vitro, temperature has been shown to have a strong positive influence on bacteriolytic rate. Temperature has no influence, however, on the in situ concentration of lytic agent in gut fluids, thus it appears that compensation for this temperature dependence is unimportant. These findings, combined with previous kinetics studies with A. marina gut fluids, predict that the quantitative influence of deposit feeding on the microbial ecology of sediments will exhibit clear seasonal variation.