Kelp forest

  Occurring worldwide throughout temperate and polar coastal oceans, kelp forests are recognized as one of the most productive and dynamic ecosystems on Earth. ^[1] (In 2007, kelp forests were also discovered in tropical waters near Ecuador.)^[2] Physically formed by brown macroalgae of the order Laminariales, kelp forests provide a unique three-dimensional 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 ^[3] and provide many ecosystem services to humankind.^[4] 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 over-grazing of kelp and other algae.^[5] This can rapidly result in transitions to barren landscapes where relatively few species persist.^[6] The implementation of marine protected areas (MPAs) 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.

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Kelp

Further information: Kelp

  The term 'kelp' refers to marine algae belonging to the taxonomic order Laminariales (Phylum: Heterokontophyta). Though not considered a taxonomically diverse order, kelps are highly diverse structurally and functionally.^[4] The most widely recognized species are the giant kelps (Macrocystis spp.), although there are numerous other genera such as Laminaria, Ecklonia, Lessonia, Alaria and Eisenia.

Frequently considered an ecosystem engineer, kelp provides the physical substrate and habitat about which kelp forest communities are built.^[7] In algae (Kingdom: Protista), the body of an individual organism is known as a thallus rather than as a plant (Kingdom: Plantae). The morphological structure of a kelp thallus is defined by three basic structural units^[6]:

• 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 (usually rock), high nutrients (e.g., nitrogen, phosphorus), and light (minimum annual irradiance dose > 50 E m^{-2 [8]}). Especially productive kelp forests tend to be associated with areas of significant oceanographic upwelling, a process which delivers cool nutrient-rich water from depth to the ocean’s mixed surface layer.^[8] Water flow and turbulence facilitate nutrient assimilation across kelp fronds throughout the water column.^[9] Water clarity affects the depth to which sufficient light can be transmitted. In ideal conditions, giant kelp (Macrocystis spp.) can grow as much as 30-60 centimeters vertically per day. Some species such as Nereocystis are annual while others like Eisenia are perennial, living for more than 20 years.^[10] In perennial kelp forests, maximum growth rates occur during upwelling months (typically spring and summer) and die-backs correspond to reduced nutrient availability, shorter photoperiods and increased storm frequency.^[6]

Kelps are primarily associated with temperate and arctic waters worldwide. Of the more dominant species, 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.^[6] The region with the greatest diversity of kelps (>20 species) 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.^[11][12] This general absence of kelp from the tropics is believed to be mostly due to insufficient nutrient levels associated with warm, oligotrophic waters.^[6] One recent study spatially overlaid the requisite physical parameters for kelp with mean oceanographic conditions has produced a model predicting the existence of subsurface kelps throughout the tropics worldwide to depths of 200m. 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 8 of their sampled sites, all of which had been predicted by the model and thus validated 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.^[2] The importance of this contribution has been rapidly acknowledged within the scientific community and prompts an entirely new trajectory of kelp forest research, particularly emphasizing the potential for a spatial refuge from climate change and explanations to evolutionary patterns of kelps worldwide.^[13]

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^[6]:

• Canopy kelps include the largest species and often constitute floating canopies that extend to the ocean surface (e.g., Macrocystis and Alaria);
• Stipitate kelps generally extend a few meters above the sea floor and can grow in dense aggregations (e.g., Eisenia and Ecklonia);
• Prostrate kelps lie near and along the sea floor (e.g., Laminaria);
• The benthic assemblage is composed of other algal species (e.g., filamentous and foliose functional groups, articulated corallines) 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 there may be a benthic assemblage of foliose red algae. 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.^[6] 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.^[14][15][16] 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.^[14] 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 (the relationships between organisms and their food webs), particularly the understanding of bottom-up 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.^[17][18] This allows kelp to grow and subsequently support herbivores, which in turn support consumers at higher trophic levels.^[19] By contrast, top-down processes are those in which predators limit the biomass of species at lower trophic levels through consumption; in the absence of predation, these lower level species would flourish because the resources that support their energetic requirements are non-limiting. In a well-studied example from Alaskan kelp forests^[20], sea otters (Enhydra lutris) control populations of herbivorous sea urchins through predation. When sea otters are removed from the ecosystem (for example, by human exploitation), 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. The effect of removing one predatory species in this system differs from Alaska because there is redundancy in the trophic levels and other predatory species can continue to regulate urchins.^[15] However, the removal of multiple predators can effectively release urchins from predator pressure and allow the system to follow trajectories towards kelp forest degradation.^[21] Similar examples exist in Nova Scotia^[22], South Africa ^[23], Australia ^[24] and Chile ^[25]. 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.^[26][27][28]

The transition from macroalgal (i.e. kelp forest) to denuded landscapes dominated by sea urchins (or ‘urchin barrens’) is a widespread phenomenon^{[29][30][4][31]}, often resulting from trophic cascades like those described above; the two phases are regarded as alternative stable states of the ecosystem.^[32][33] The recovery of kelp forests from barren states has been documented following dramatic perturbations, such as urchin disease or large shifts in thermal conditions.^[21][34][35] 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 herbivore, others with significant interaction strengths include seastars, isopods, kelp crabs, and herbivorous fishes.^[6][26] 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 to feed on. When there is sufficient drift kelp, herbivorous grazers do not exert pressure on attached plants; when drift subsidies are unavailable, grazers directly impact the physical structure of the ecosystem.^[36][37] Many studies in Southern California have demonstrated that the availability of drift kelp specifically influences the foraging behavior of sea urchins.^[38][39] Drift kelp and kelp-derived particulate matter have also been important in subsidizing adjacent habitats, such as sandy beaches and the rocky intertidal.^[40][41][42]

Patch Dynamics

Another major area of kelp forest research has been directed at understanding the spatial-temporal patterns of kelp patches. Not only do such dynamics affect the physical landscape, but they also affect species that associate with kelp for refuge or foraging activities.^[14][19] Large-scale environmental disturbances have offered important insights concerning mechanisms and ecosystem resilience. Examples of environmental disturbances include the following:

• Acute and chronic pollution events have been shown to impact southern California kelp forests, though the intensity of the impact seems to depend on both the nature of the contaminants and duration of exposure.^{[43][44][45][46][47]} Pollution can include sediment deposition and eutrophication from sewage, industrial byproducts and contaminants like PCBs and heavy metals (for example, copper, zinc), runoff of organophosphates from agricultural areas, anti-fouling chemicals used in harbors and marinas (for example, TBT and creosote) and land-based pathogens like fecal coliform bacteria.
• Catastrophic storms can remove surface kelp canopies through wave activity but usually leave understory kelps intact; they can also remove urchins when little spatial refuge is available.^[37][32] Interspersed canopy clearings create a seascape mosaic where sunlight penetrates deeper into the kelp forest and species that are normally light-limited in the understory can flourish. Similarly, substrate cleared of kelp holdfasts can provide space for other sessile species to establish themselves and occupy the seafloor, sometimes directly competing with juvenile kelp plants and even inhibiting their settlement.^[48]
• El Niño-Southern Oscillation (ENSO) events involve the depression of oceanographic thermoclines, severe reductions of nutrient input, and changes in storm patterns.^[32][49] Stress due to warm water and nutrient depletion can increase the susceptibility of kelp to storm damage and herbivorous grazing, sometimes even prompting phase shifts to urchin-dominated landscapes.^[38][50][35] In general, oceanographic conditions (that is, water temperature, currents) influence the recruitment success of kelp and its competitors, which clearly affect subsequent species interactions and kelp forest dynamics.^[51][32]
• Overfishing higher trophic levels that naturally regulate herbivore populations is also recognized as an important stressor in kelp forests.^[5][52][28] As described in the previous section, the drivers and outcomes of trophic cascades are important for understanding spatial-temporal patterns of kelp forests.^[21][20][26]

In addition to ecological monitoring of kelp forests before, during, and after such disturbances, scientists try to tease apart the intricacies of kelp forest dynamics using experimental manipulations. By working on smaller spatial-temporal scales, they can control for the presence or absence of specific biotic and abiotic factors to discover the operative mechanisms. For example, in southern Australia, manipulations of kelp canopy types demonstrated that the relative amount of Ecklonia radiata in a canopy could be used to predict understory species assemblages; consequently, the proportion of E. radiata can be used as an indicator of other species occurring in the environment.^[53]

Human Use

As exemplified by the Aleut hunting of sea otters, kelp forests have been important to human existence for thousands of years.^[54] Indeed, many now theorise that the first colonisation of the Americas was due to fishing communities following the Pacific Kelp Forests during the last Ice Age. Modern economies are based on fisheries of kelp-associated species like lobster and rockfish. Humans also harvest kelp directly to feed aquaculture species like abalone and to extract the compound alginic acid, which is used in products like toothpaste and antacids.^[55][56] Kelp forests are valued for recreational activities such as SCUBA diving and kayaking; the industries that support these sports represent one benefit related to the ecosystem and the enjoyment derived from these activities represents another. All of these are examples of ecosystem services provided specifically by kelp forests.

Threats & Management

  Given the complexity of kelp forests – their variable structure, geography and interactions – they pose a considerable challenge to environmental managers. It is difficult to extrapolate even well-studied trends to the future because interactions within the ecosystem will change under variable conditions, not all relationships in the ecosystem are understood, and there are can be non-linear thresholds to transitions that are not yet recognized.^[57] With respect to kelp forests, major issues of concern include marine pollution and water quality, kelp harvesting and fisheries, invasive species and climate change.^[4] It has been argued that the most pressing threat to kelp forest preservation is the overfishing of coastal ecosystems, which by removing higher trophic levels facilitates their shift to depauperate urchin barrens.^[5] The maintenance of biodiversity is recognized as a way of generally stabilizing ecosystems and their services through mechanisms such as functional compensation and reduced susceptibility to foreign species invasions.^{[58][59][60][61]}

In many places, managers have opted to regulate the harvest of kelp^[18][62] and/or the taking of kelp forest species by fisheries.^[52][4] While these may be effective in one sense, they do not necessarily protect the entirety of the ecosystem. Marine protected areas (MPAs) offer a unique solution that encompasses not only target species for harvesting but also the interactions surrounding them and the local environment as a whole.^[63][64] Direct benefits of MPAs to fisheries (for example, spillover effects) have been well-documented around the world.^{[65][5][66][67]} Indirect benefits have also been shown for several cases among species such as abalone and fishes in Central California.^[68][69] Most importantly, studies have demonstrated that MPAs can be effective at protecting existing kelp forest ecosystems and may also allow for the regeneration of those which have been impacted.^[32][70][71]

References