Kelp forest


Kelp forests are underwater areas with a high density of kelp, which covers a large part of the world's coastlines. Smaller areas of anchored kelp are called kelp beds. They are recognized as one of the most productive and dynamic ecosystems on Earth. Although algal kelp forest combined with coral reefs only cover 0.1% of Earth's total surface, they account for 0.9% of global primary productivity. Kelp forests occur worldwide throughout temperate and polar coastal oceans. In 2007, kelp forests were also discovered in tropical waters near Ecuador.
File:Kelp forest Cojo Anchorage.PNG|thumb|350px|A kelp forest at Cojo Anchorage near Point Conception, California.
Physically formed by brown macroalgae, kelp forests provide a unique habitat for marine organisms and are a source for understanding many ecological processes. Over the last century, they have been the focus of extensive research, particularly in trophic ecology, and continue to provoke important ideas that are relevant beyond this unique ecosystem. For example, kelp forests can influence coastal oceanographic patterns and provide many ecosystem services.
However, the influence of humans has often contributed to kelp forest degradation. Of particular concern are the effects of overfishing nearshore ecosystems, which can release herbivores from their normal population regulation and result in the overgrazing of kelp and other algae. This can rapidly result in transitions to barren landscapes where relatively few species persist. Already due to the combined effects of overfishing and climate change, kelp forests have all but disappeared in many especially vulnerable places, such as Tasmania's east coast and the coast of Northern California. The implementation of marine protected areas is one management strategy useful for addressing such issues, since it may limit the impacts of fishing and buffer the ecosystem from additive effects of other environmental stressors.

Kelp

The term kelp refers to marine algae belonging to the order Laminariales. Though not considered a taxonomically diverse order, kelps are highly diverse structurally and functionally. The most widely recognized species are the giant kelps, although numerous other genera such as Laminaria, Ecklonia, Lessonia, Nereocystis, Alaria, and Eisenia are described.
A wide range of sea life uses kelp forests for protection or food, including fish. In the North Pacific kelp forests, particularly rockfish, and many invertebrates, such as amphipods, shrimp, marine snails, bristle worms, and brittle stars. Many marine mammals and birds are also found, including seals, sea lions, whales, sea otters, gulls, terns, snowy egrets, great blue herons, and cormorants, as well as some shore birds.
Frequently considered an ecosystem engineer, kelp provides a physical substrate and habitat for kelp forest communities. In algae, the body of an individual organism is known as a thallus rather than as a plant. The morphological structure of a kelp thallus is defined by three basic structural units:
  • The holdfast is a root-like mass that anchors the thallus to the sea floor, though unlike true roots it is not responsible for absorbing and delivering nutrients to the rest of the thallus.
  • The stipe is analogous to a plant stalk, extending vertically from the holdfast and providing a support framework for other morphological features.
  • The fronds are leaf- or blade-like attachments extending from the stipe, sometimes along its full length, and are the sites of nutrient uptake and photosynthetic activity.
In addition, many kelp species have pneumatocysts, or gas-filled bladders, usually located at the base of fronds near the stipe. These structures provide the necessary buoyancy for kelp to maintain an upright position in the water column.
The environmental factors necessary for kelp to survive include hard substrate, high nutrients, and light. Especially productive kelp forests tend to be associated with areas of significant oceanographic upwelling, a process that delivers cool, nutrient-rich water from depth to the ocean's mixed surface layer. Water flow and turbulence facilitate nutrient assimilation across kelp fronds throughout the water column. Water clarity affects the depth to which sufficient light can be transmitted. In ideal conditions, giant kelp can grow as much as 30–60 cm vertically per day. Some species, such as Nereocystis, are annuals, while others such as Eisenia are perennials, living for more than 20 years. In perennial kelp forests, maximum growth rates occur during upwelling months and die-backs correspond to reduced nutrient availability, shorter photoperiods, and increased storm frequency.
Kelps are primarily associated with temperate and arctic waters worldwide. Of the more dominant genera, Laminaria is mainly associated with both sides of the Atlantic Ocean and the coasts of China and Japan; Ecklonia is found in Australia, New Zealand, and South Africa; and Macrocystis occurs throughout the northeastern and southeastern Pacific Ocean, Southern Ocean archipelagos, and in patches around Australia, New Zealand, and South Africa. The region with the greatest diversity of kelps is the northeastern Pacific, from north of San Francisco, California, to the Aleutian Islands, Alaska.
Although kelp forests are unknown in tropical surface waters, a few species of Laminaria have been known to occur exclusively in tropical deep waters. This general absence of kelp from the tropics is believed to be mostly due to insufficient nutrient levels associated with warm, oligotrophic waters. One recent study spatially overlaid the requisite physical parameters for kelp with mean oceanographic conditions and produced a model predicting the existence of subsurface kelps throughout the tropics worldwide to depths of. For a hotspot in the Galapagos Islands, the local model was improved with fine-scale data and tested; the research team found thriving kelp forests in all eight of their sampled sites, all of which had been predicted by the model, thus validating their approach. This suggests that their global model might actually be fairly accurate, and if so, kelp forests would be prolific in tropical subsurface waters worldwide. The importance of this contribution has been rapidly acknowledged within the scientific community and has prompted an entirely new trajectory of kelp forest research, highlighting the potential for kelp forests to provide marine organisms spatial refuge under climate change and providing possible explanations for evolutionary patterns of kelps worldwide.

Ecosystem architecture

The architecture of a kelp forest ecosystem is based on its physical structure, which influences the associated species that define its community structure. Structurally, the ecosystem includes three guilds of kelp and two guilds occupied by other algae:
  • Canopy kelps include the largest species and often constitute floating canopies that extend to the ocean surface.
  • Stipitate kelps generally extend a few meters above the sea floor and can grow in dense aggregations.
  • Prostrate kelps lie near and along the sea floor.
  • The benthic assemblage is composed of other algal species and sessile organisms along the ocean bottom.
  • Encrusting coralline algae directly and often extensively cover geologic substrate.
Multiple kelp species often co-exist within a forest; the term understory canopy refers to the stipitate and prostrate kelps. For example, a Macrocystis canopy may extend many meters above the seafloor towards the ocean surface, while an understory of the kelps Eisenia and Pterygophora reaches upward only a few meters. Beneath these kelps, a benthic assemblage of foliose red algae may occur. The dense vertical infrastructure with overlying canopy forms a system of microenvironments similar to those observed in a terrestrial forest, with a sunny canopy region, a partially shaded middle, and darkened seafloor. Each guild has associated organisms, which vary in their levels of dependence on the habitat, and the assemblage of these organisms can vary with kelp morphologies. For example, in California, Macrocystis pyrifera forests, the nudibranch Melibe leonina, and skeleton shrimp Caprella californica are closely associated with surface canopies; the kelp perch Brachyistius frenatus, rockfish Sebastes spp., and many other fishes are found within the stipitate understory; brittle stars and turban snails Tegula spp. are closely associated with the kelp holdfast, while various herbivores, such as sea urchins and abalone, live under the prostrate canopy; many seastars, hydroids, and benthic fishes live among the benthic assemblages; solitary corals, various gastropods, and echinoderms live over the encrusting coralline algae. In addition, pelagic fishes and marine mammals are loosely associated with kelp forests, usually interacting near the edges as they visit to feed on resident organisms.

Trophic ecology

Classic studies in kelp forest ecology have largely focused on trophic interactions, particularly the understanding and top-down trophic processes. Bottom-up processes are generally driven by the abiotic conditions required for primary producers to grow, such as availability of light and nutrients, and the subsequent transfer of energy to consumers at higher trophic levels. For example, the occurrence of kelp is frequently correlated with oceanographic upwelling zones, which provide unusually high concentrations of nutrients to the local environment. This allows kelp to grow and subsequently support herbivores, which in turn support consumers at higher trophic levels. By contrast, in top-down processes, predators limit the biomass of species at lower trophic levels through consumption. In the absence of predation, these lower-level species flourish because resources that support their energetic requirements are not limiting. In a well-studied example from Alaskan kelp forests, sea otters control populations of herbivorous sea urchins through predation. When sea otters are removed from the ecosystem, urchin populations are released from predatory control and grow dramatically. This leads to increased herbivore pressure on local kelp stands. Deterioration of the kelp itself results in the loss of physical ecosystem structure and subsequently, the loss of other species associated with this habitat. In Alaskan kelp forest ecosystems, sea otters are the keystone species that mediates this trophic cascade. In Southern California, kelp forests persist without sea otters and the control of herbivorous urchins is instead mediated by a suite of predators including lobsters and large fishes, such as the California sheephead. The effect of removing one predatory species in this system differs from Alaska because redundancy exists in the trophic levels and other predatory species can continue to regulate urchins. However, the removal of multiple predators can effectively release urchins from predator pressure and allow the system to follow trajectories towards kelp forest degradation. Similar examples exist in Nova Scotia, South Africa, Australia, and Chile. The relative importance of top-down versus bottom-up control in kelp forest ecosystems and the strengths of trophic interactions continue to be the subject of considerable scientific investigation.
The transition from macroalgal to denuded landscapes dominated by sea urchins is a widespread phenomenon, often resulting from trophic cascades like those described above; the two phases are regarded as alternative stable states of the ecosystem. The recovery of kelp forests from barren states has been documented following dramatic perturbations, such as urchin disease or large shifts in thermal conditions. Recovery from intermediate states of deterioration is less predictable and depends on a combination of abiotic factors and biotic interactions in each case.
Though urchins are usually the dominant herbivores, others with significant interaction strengths include seastars, isopods, kelp crabs, and herbivorous fishes. In many cases, these organisms feed on kelp that has been dislodged from substrate and drifts near the ocean floor rather than expend energy searching for intact thalli on which to feed. When sufficient drift kelp is available, herbivorous grazers do not exert pressure on attached thalli; when drift subsidies are unavailable, grazers directly impact the physical structure of the ecosystem. Many studies in Southern California have demonstrated that the availability of drift kelp specifically influences the foraging behavior of sea urchins. Drift kelp and kelp-derived particulate matter have also been important in subsidizing adjacent habitats, such as sandy beaches and the rocky intertidal.