Regulation of Phytoplankton Carbon to Chlorophyll Ratio By Light, Nutrients and Temperature in the Equatorial Pacific Ocean: A Basin-Scale Model

The complex effects of light, nutrients and temperature lead to a variable carbon to chlorophyll (C:Chl) ratio in phytoplankton cells. Using field data collected in the Equatorial Pacific, we derived a new dynamic model with a non-steady C:Chl ratio as a function of irradiance, nitrate, iron, and temperature. The dynamic model is implemented into a basin-scale ocean circulation-biogeochemistry model and tested in the Equatorial Pacific Ocean. The model reproduces well the general features of phytoplankton dynamics in this region. For instance, the simulated deep chlorophyll maximum (DCM) is much deeper in the western warm pool (similar to 100 m) than in the Eastern Equatorial Pacific (similar to 50 m). The model also shows the ability to reproduce chlorophyll, including not only the zonal, meridional and vertical variations, but also the interannual variability. This modeling study demonstrates that combination of nitrate and iron regulates the spatial and temporal variations in the phytoplankton C:Chl ratio in the Equatorial Pacific. Sensitivity simulations suggest that nitrate is mainly responsible for the high C:Chl ratio in the western warm pool while iron is responsible for the frontal features in the C:Chl ratio between the warm pool and the upwelling region. In addition, iron plays a dominant role in regulating the spatial and temporal variations of the C:Chl ratio in the Central and Eastern Equatorial Pacific. While temperature has a relatively small effect on the C:Chl ratio, light is primarily responsible for the vertical decrease of phytoplankton C:Chl ratio in the euphotic zone.

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.

Physical and Biological Controls on the Latitudinal Asymmetry of Surface Nutrients and pCO(2) in the Central and Eastern Equatorial Pacific

Surface nutrients and dissolved inorganic carbon (DIC) in the central (CEP) and eastern equatorial Pacific (EEP) show much higher concentrations to the south than to the north of the equator. In this study, the physical and biological controls on this asymmetry are investigated using a coupled physical-biogeochemical model. Two numerical experiments are conducted to examine the effects of asymmetrical photosynthetic efficiency (a) due to asymmetrical iron supply about the equator. The experiment with asymmetrical photosynthesis produces improved results as compared with historical observations. A nitrate budget analysis suggests that in the EEP the divergence of upwelling waters controls the surface nitrate asymmetry with additional contribution from the South Equatorial Current (SEC) carrying nutrient-rich Peru upwelling water. The changes of a affect the surface nitrate distribution but not the overall asymmetry. The SEC further carries excess nitrate to the west and thus extends the asymmetry in the east to the CEP. In the CEP, however, stronger northward than southward transport tends to reduce the nitrate asymmetry, while the asymmetrical photosynthesis would help to maintain it. Similar processes also control the distributions of surface silicate and DIC in the equatorial Pacific, which is also affected by the air-sea CO(2) exchange. The asymmetrical photosynthesis influences the distribution of surface DIC, pCO(2), and the air-sea CO(2) flux, by redistributing about 20% CO(2) flux from the north to the south of the equator. Owing to the adjustment of air-sea CO(2) flux, however, the net surface DIC change is smaller than the direct change associated with primary production.