Post-glacial redistribution and shifts in productivity of giant kelp forests

Quaternary glacial–interglacial cycles create lasting biogeographic, demographic and genetic effects on ecosystems, yet the ecological effects of ice ages on benthic marine communities are unknown. We analysed long-term datasets to develop a niche-based model of southern Californian giant kelp (Macrocystis pyrifera) forest distribution as a function of oceanography and geomorphology, and synthesized palaeo-oceanographic records to show that late Quaternary climate change probably drove high millennial variability in the distribution and productivity of this foundation species. Our predictions suggest that kelp forest biomass increased up to threefold from the glacial maximum to the mid-Holocene, then rapidly declined by 40–70 per cent to present levels. The peak in kelp forest productivity would have coincided with the earliest coastal archaeological sites in the New World. Similar late Quaternary changes in kelp forest distribution and productivity probably occurred in coastal upwelling systems along active continental margins worldwide, which would have resulted in complex shifts in the relative productivity of terrestrial and marine components of coastal ecosystems.     

Detrital carbon production and export in high latitude kelp forests

Oecologia - The production and fate of seaweed detritus is a major unknown in the global C-budget. Knowing the quantity of detritus produced, the form it takes (size) and its timing of delivery are...     

Fauna associated with detached kelp in different types of subtidal habitats of the White Sea

The fauna, associated with Laminaria and other largebrown macroalgae was studied by using SCUBA anddredging in two different types of underwater habitatsof the White Sea.In shallow water fjords and bays, with a depth of nomore than 30–40 m, detached kelp (mainly Laminaria saccharina, L. digitata and Alaria esculenta) formed large accumulations. One ofthese benthic accumulations, which has existed morethan 20 years, was studied. It covers about2000 m², and is about 2 m thick. The upper layerof the accumulation of fronds is characterized by highturbulence and is well aerated. The lower layer ischaracterized by anoxic conditions. Mats of sulphurbacteria were not observed, although fronds in themiddle layer were covered by layers of cyanobacteria.About 50 species of macroinvertebrates were found,mainly species that are normally associated withliving kelp, such as the detritivorous species Ophiura robusta and Gammarus oceanicus, and fewspecies that are specific inhabitants of organic-richbiotopes in the White Sea such as Capitellacapitata, Ophryotrocha irinae and Nebaliabipes. It was remarkable that in the shallow waterbasins of the White Sea, the process of decompositionof brown algae in the sublittoral takes place withoutsea urchins, and no other macrofaunal form plays anecological role in the mechanical breakdown of theplant substratum, even not in the large accumulationsof detached kelp.Along the open rocky shoreline, communities associatedwith dead detached kelp were situated at a depth of60–90 m, 40–50 m below the belt of living kelp. Inthis deep zone, no macroinvertebrates typical of thekelp community in the photic zone were found. Duringthe passage from the shoreline to the deeper benthiccommunity, where sea urchins were dominant, all plantdebris became fragmented. These deeper benthiccommunities appeared to be the zone for decompositionof the detached kelp.     

55Ice age kelp forests: climate-driven changes in kelp forest distribution since the last glacial maximum

The University of Connecticut presents the Benthic Marine Algal Herbarium of Long Island Sound Digital Collection. The collection is available on the web for teaching and identification purposes. Phase One of this project will be finished later by early next year. Currently, the collection has approximately 60% of all Long Island Sound macroalgae species. The taxonomy is part of a collaboration between the University of Connecticut and the Northeast Algal Society (NEAS). Thus far, the specimens have come from Dr. Charles Yarish's herbarium and from the Environmental Laboratory at Millstone Power Station on Long Island Sound. The botanical illustrations are from Illustrated Key to the Seaweeds of New England by Martine Villalard-Bohnsack. Database features include the ability to create searches and generate sets based on subjects, division, class, order, family, genus, habitat, species, keyword, and location. Data retrieved includes JPG image, cataloged data, map regions, synonyms, and illustrated microscopic, cross section and surface view for some species. The web site is available for use at: http://www.algae.uconn.edu     

Habitat characteristics can influence fish assemblages in high latitude kelp forests

Kelp forests are patchy fish-associated habitats, which can vary greatly in their size, foundation species, and several physical habitat attributes. The structure of fish assemblages can vary with these characteristics and with the location of the assemblage within the forest, i.e. edges versus interiors. This study quantified the biological and physical heterogeneity within different sized kelp forests and identified which factors are important in structuring the associated fish assemblages. Fish and habitat surveys were conducted at the edge and interiors of ten kelp forests of varying sizes in Kachemak Bay, south-central Alaska. Fish assemblage structure was not correlated with the species composition of surface canopy forming foundation kelps (Nereocystis leutkeana and Eualaria fistulosa) or with kelp forest size. Instead, it correlated with the abundances of two understory foundation kelps (Agarum clathratum and Saccharina latissima), substratum rugosity, and water depth. Together these benthic attributes explained 53.6 % of the fish assemblage variability. Additionally, significantly different fish assemblages were found at edge compared to interior locations with the relative abundance of seven species (Artedius fenestralis, Ammodytes hexapterus, Blepsias cirrhosis, Gadus macrocephalus, Hexagrammos stelleri, Pholis laeta, and Sebastes melanops) explaining 91.4 % of the variability. This study highlights the importance of habitat characteristics such as understory foundation species, substratum rugosity, water depth and location within a patch on the variability of fish assemblages in high latitude kelp forests.     

Disturbance and reef topography maintain high local diversity in Ecklonia radiata kelp forests

Disturbance of competitive‐dominant plant and algae canopies often lead to increased diversity of the assemblage. Kelp forests, particularly those of temperate Western Australia, are habitats with high alpha diversity. This study investigated the roles of broad‐scale canopy loss and local scale reef topography on structuring the kelp‐dominated macroalgal forests in Western Australia. Eighteen 314 m² circular areas were cleared of their Ecklonia radiata canopy and eighteen controls were established across three locations. The patterns of macroalgal recolonisation in replicate clearances were observed over a 34 month period. Macroalgal species richness initially increased after canopy removal with a turf of filamentous and foliose macroalgae dominating cleared areas for up to seven months. A dense Sargassum canopy dominated cleared areas from 11 to 22 months. By 34 months, partial recovery of the kelp canopy into cleared areas had occurred. Some cleared areas did not follow this trajectory but remained dominated by turfing, foliose and filamentous algae. As kelp canopies developed, the initial high species diversity declined but still remained elevated relative to undisturbed controls, even after 34 months. More complex reef topography was associated with greater variability in the algal assemblage between replicate quadrats suggesting colonising algae had a greater choice of microhabitats available to them on topographically complex reefs. Shading by canopies of either Sargassum spp. and E. radiata are proposed to highly influence the abundance of algae through competitive exclusion that is relaxed by disturbance of the canopy. Disturbance of the canopy in E. radiata kelp forests created a mosaic of different patch types (turf, Sargassum‐dominated, kelp‐dominated). These patch types were both transient and stable over the 34 months of this study, and are a potential contemporary process that maintains high species diversity in temperate kelp‐dominated reefs.     

Status,trends and drivers of kelp forests in Europe: an expert assessment

A comprehensive expert consultation was conducted in order to assess the status, trends and the most important drivers of change in the abundance and geographical distribution of kelp forests in European waters. This consultation included an on-line questionnaire, results from a workshop and data provided by a selected group of experts working on kelp forest mapping and eco-evolutionary research. Differences in status and trends according to geographical areas, species identity and small-scale variations within the same habitat where shown by assembling and mapping kelp distribution and trend data. Significant data gaps for some geographical regions, like the Mediterranean and the southern Iberian Peninsula, were also identified. The data used for this study confirmed a general trend with decreasing abundance of some native kelp species at their southern distributional range limits and increasing abundance in other parts of their distribution (Saccharina latissima and Saccorhiza polyschides). The expansion of the introduced species Undaria pinnatifida was also registered. Drivers of observed changes in kelp forests distribution and abundance were assessed using experts’ opinions. Multiple possible drivers were identified, including global warming, sea urchin grazing, harvesting, pollution and fishing pressure, and their impact varied between geographical areas. Overall, the results highlight major threats for these ecosystems but also opportunities for conservation. Major requirements to ensure adequate protection of coastal kelp ecosystems along European coastlines are discussed, based on the local to regional gaps detected in the study.     

Spatial and short-term variability of larval,post-larval and macrobenthic assemblages associated with subtidal kelp forest ecosystems in Central Chile

Identifying patterns of spatial and temporal variability in the composition of communities associated with kelp forests is critical to understand the functioning of this productive, yet vulnerable ecosystem. We used a suite of sampling methods (light attraction and airlift devices) to evaluate the variability of larval, post-larval and macrobenthic assemblages associated with kelp forests (Lessonia trabeculata) in Central Chile (30° to 33°S). Pelagic collections identified two assemblages: early-life stages and emerging macrobenthos, with the later contributing three quarters to the total abundance regardless of the source of illumination (permanent or flashing). Field experiments showed that moon phases affected the structure and composition of the samples. Surveys carried out during new moon showed the highest abundances and taxonomic richness of emergent assemblages. However, species composition varied in both assemblages depending on the moon phase. Although the pelagic assemblages collected at sites with contrasting upwelling intensity did not show differences in community structure, differences in composition were evident for early-life stages. The relationship between pelagic and benthic collections indicated that four decapod crustaceans were represented at both larval and early juvenile stages; however, only the high abundances and densities of Paraxanthus barbiger allowed for estimations of benthic-pelagic coupling. For this species, larval abundances and benthic juvenile densities demonstrated contrasting local and regional patterns, suggesting a decoupling between pelagic and benthic environments. These findings highlight the differential variability in smaller components of kelp forests, but also suggest that post-settlement processes may be driving biological interactions through these highly productive and complex environments.     

Modelling seasonal growth and composition of the kelp Saccharina latissima

A dynamical model for simulating growth of the brown macroalga Saccharina latissima is described. In addition to wet and dry weights, the model simulates carbon and nitrogen reserves, with variable C/N ratio. In effect, the model can be used to emulate seasonal changes in growth and composition of the alga. Simulation results based on published, environmental field data are presented and compared with corresponding data on growth and composition. The model resolves seasonal growth, carbon and nitrogen content well, and may contribute to the understanding of how seasonal growth in S. latissima depends simultaneously on a combination of several environmental factors: light, nutrients, temperature and water motion. The model is applied to aquaculture problems such as estimating the nutrient scavenging potential of S. latissima and estimating the potential of this kelp species as a raw material for bioenergy production.     

Ecological studies on macroinvertebrate populations associated with polluted kelp forests in the North Sea

Summary 1. The English Counties of Midlothian and Durham are situated on the North Western Edge of the North Sea. Both counties have a high population density and a variety of industries located in them.2. The coastal seas adjacent to these areas are heavily polluted.3. In one of these industrial areas, County Durham, the coastal waters have been polluted for 150 years — from the beginning of the Industrial Revolution.4. Two gradients of pollution, one estuarine and one off the open coast, are described. The intervening sea coast has little or no pollution.5. A comparative method of pollution surveying is presented which makes use of the epifauna in the holdfast or hapteron of the large brown algaLaminaria hyperborea (Gunn.)Fosl.6. The size of the habitat is measured in cubic millilitres and called ecospace.7. The rate of invertebrate colonisation is measured in ecoperiods.8. Statistical investigation reveals similar habitats in clean and polluted water.9. The normal sequence of habitat colonisation by invertebrates is described for clean waters. It contains species groups commonly found on rocky sublittoral kelp forests elsewhere in the North Atlantic.10. Ecological comparison is made of the community development described for clean and polluted stations.11. Results indicate that in polluted water, invertebrate succession is retarded; the species composition of the community is simplified, 43 % of the species are lost, rates of habitat colonisation by individuals of pollution tolerant species are speeded up; the polluted community undergoes neotenous development i.e. retaining juvenile characteristics throughout development to community maturity.12. Functional aspects of new (pollution present) and old (pollution absent) communities are discussed.13. The marginal presence of some species in polluted water is also discussed.15. Two ecological barriers to normal community development in the polluted environment are postulated.

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Ökologische Studien an Populationen von Makro-Invertebraten auf verunreinigten Seetang-Wäldern in der Nordsee

Kurzfassung Die Arbeit berichtet über erste Ergebnisse einer ökologischen Untersuchung an verschmutzten Küstengewässern im Bereich der Grafschaften Durham und Midlothian (im Nordosten Englands). Die untersuchten Küstenstreifen enthalten sowohl städtische als auch industrielle Gebiete, während die dazwischenliegende Landmasse weitgehend landwirtschaftlich genutzt wird. Die Küstengewässer in Durham sind seit über 100 Jahren verschmutzt, d. h. seit dem Beginn der industriellen Revolution. Das Ausmaß der Verschmutzung wird kurz umrissen. Es wird eine Methode beschrieben, die einen Vergleich der in den verschmutzten Seetang-Gebieten gewöhnlich anzutreffenden Invertebraten-Epifauna ermöglicht. Repräsentative Proben der auf dem Haftorgan der BraunalgeLaminaria hyperborea lebenden Fauna wurden gesammelt und mit Proben aus unverseuchten Gewässern verglichen. Die Ergebnisse deuten an, daß in verschmutzten Gewässern die Artenfolge, die Geschwindigkeit der Aufeinanderfolge, die Häufigkeit der Individuen und die trophische Struktur der Lebensgemeinschaft verändert sind. In verschmutzten Gewässern ist die Entwicklung zu komplexen Lebensgemeinschaften verlangsamt; es entstehen neotene Gemeinschaften. Gründe, welche einer Entwicklung zu einer normalen Lebensgemeinschaft im Wege stehen, werden herausgestellt.

     

Economic valuation of kelp forests in northern Chile: values of goods and services of the ecosystem

Kelp beds, besides being one of the most important benthic resources in northern Chile, provide a variety of environmental goods and services. In order to evaluate economically the wild kelp populations in northern Chile (26° to 32° S) more than simply their commercial value as a source of raw materials for alginate extraction, we used several economic indicators to holistically assess the value of a group of brown seaweeds of economic importance, Lessonia spp. and Macrocystis pyrifera: (1) market value of biomass as a source of raw material for extraction of alginic acid, (2) market value of associated species of economic importance, (3) value as a source of scientific information, (4) value as a climate buffer (CO₂ capture and release of O₂), (5) value of associated biodiversity (non-commercial species), (6) value as cultural heritage and (7) value as a reservoir of biodiversity. Existence values of kelp beds which estimate the willingness of citizens to pay and work without payment to preserve the ecosystem were calculated using the contingent valuation technique. The results indicate that kelp beds in northern Chile have a total value of US $540 million. Of this total, kelp fishery accounts for 75 % and associated-species fisheries account for 15 %. In this context, the economic value of Chilean kelp beds is mainly associated with the industry of alginate extraction. By contrast, existence value as a source of scientific information or environmental buffer for CO₂ capture or O₂ production represents only 9 % of the total value, representing a very low relative importance to society. The economic valuation of coastal resources and marine ecosystems is a complementary tool for decision making and implementation of public policies related to the conservation and sustainable exploitation of renewable resources and their ecosystems.     

Landscape epidemiology and control of pathogens with cryptic and long-distance dispersal: sudden oak death in northern Californian forests

Exotic pathogens and pests threaten ecosystem service, biodiversity, and crop security globally. If an invasive agent can disperse asymptomatically over long distances, multiple spatial and temporal scales interplay, making identification of effective strategies to regulate, monitor, and control disease extremely difficult. The management of outbreaks is also challenged by limited data on the actual area infested and the dynamics of spatial spread, due to financial, technological, or social constraints. We examine principles of landscape epidemiology important in designing policy to prevent or slow invasion by such organisms, and use Phytophthora ramorum, the cause of sudden oak death, to illustrate how shortfalls in their understanding can render management applications inappropriate. This pathogen has invaded forests in coastal California, USA, and an isolated but fast-growing epidemic focus in northern California (Humboldt County) has the potential for extensive spread. The risk of spread is enhanced by the pathogen's generalist nature and survival. Additionally, the extent of cryptic infection is unknown due to limited surveying resources and access to private land. Here, we use an epidemiological model for transmission in heterogeneous landscapes and Bayesian Markov-chain-Monte-Carlo inference to estimate dispersal and life-cycle parameters of P. ramorum and forecast the distribution of infection and speed of the epidemic front in Humboldt County. We assess the viability of management options for containing the pathogen's northern spread and local impacts. Implementing a stand-alone host-free "barrier" had limited efficacy due to long-distance dispersal, but combining curative with preventive treatments ahead of the front reduced local damage and contained spread. While the large size of this focus makes effective control expensive, early synchronous treatment in newly-identified disease foci should be more cost-effective. We show how the successful management of forest ecosystems depends on estimating the spatial scales of invasion and treatment of pathogens and pests with cryptic long-distance dispersal.     

Stability of strong species interactions resist the synergistic effects of local and global pollution in kelp forests

Foundation species, such as kelp, exert disproportionately strong community effects and persist, in part, by dominating taxa that inhibit their regeneration. Human activities which benefit their competitors, however, may reduce stability of communities, increasing the probability of phase-shifts. We tested whether a foundation species (kelp) would continue to inhibit a key competitor (turf-forming algae) under moderately increased local (nutrient) and near-future forecasted global pollution (CO(2)). Our results reveal that in the absence of kelp, local and global pollutants combined to cause the greatest cover and mass of turfs, a synergistic response whereby turfs increased more than would be predicted by adding the independent effects of treatments (kelp absence, elevated nutrients, forecasted CO(2)). The positive effects of nutrient and CO(2) enrichment on turfs were, however, inhibited by the presence of kelp, indicating the competitive effect of kelp was stronger than synergistic effects of moderate enrichment of local and global pollutants. Quantification of physicochemical parameters within experimental mesocosms suggests turf inhibition was likely due to an effect of kelp on physical (i.e. shading) rather than chemical conditions. Such results indicate that while forecasted climates may increase the probability of phase-shifts, maintenance of intact populations of foundation species could enable the continued strength of interactions and persistence of communities.     

Exploitation and recovery of a sea urchin predator has implications for the resilience of southern California kelp forests

Size-structured predator–prey interactions can be altered by the history of exploitation, if that exploitation is itself size-selective. For example, selective harvesting of larger sized predators can release prey populations in cases where only large individuals are capable of consuming a particular prey species. In this study, we examined how the history of exploitation and recovery (inside marine reserves and due to fisheries management) of California sheephead (Semicossyphus pulcher) has affected size-structured interactions with sea urchin prey in southern California. We show that fishing changes size structure by reducing sizes and alters life histories of sheephead, while management measures that lessen or remove fishing impacts (e.g. marine reserves, effort restrictions) reverse these effects and result in increases in density, size and biomass. We show that predation on sea urchins is size-dependent, such that the diet of larger sheephead is composed of more and larger sized urchins than the diet of smaller fish. These results have implications for kelp forest resilience, because urchins can overgraze kelp in the absence of top-down control. From surveys in a network of marine reserves, we report negative relationships between the abundance of sheephead and urchins and the abundance of urchins and fleshy macroalgae (including giant kelp), indicating the potential for cascading indirect positive effects of top predators on the abundance of primary producers. Management measures such as increased minimum size limits and marine reserves may serve to restore historical trophic roles of key predators and thereby enhance the resilience of marine ecosystems.     

Allelic and Genotypic Composition of Ancestral Spanish and Colonial Californian Gene Pools of Avena Barbata: Evolutionary Implications

Spanish explorers and colonists inadvertently started a massive experiment in evolutionary genetics when they accidentally introduced Avena barbata to California from Spain during the seventeenth and eighteenth centuries. Assays of the Spanish and Californian gene pools of this species for 15 loci show that the present day Spanish gene pool, particularly that of Southwestern Spain, is identical or virtually identical to that of California for five loci and closely similar for nine loci. Despite their similar allelic and single-locus genotypic compositions, the present-day Spanish and Californian gene pools are differently structured on a multilocus genetic basis. Evolutionary implications of these results are discussed.     

The effects of harvesting of the South African kelp (Ecklonia maxima) on kelp population structure, growth rate and recruitment

Ecklonia maxima is an economically important kelp in South Africa. The harvested kelp is used mainly as feed for cultured Haliotis midae (abalone) on farms all along the South African South and West Coast. The effects that different harvesting methods have on the growth of sub-canopy kelps, kelp population structure and kelp recruitment were tested in a kelp bed at Bordjies Rif near Cape Town. Two 30 × 10 m sites were set up, about 100 m apart, in near monoculture stands of E. maxima. Each 30×10 m area was subdivided into three treatments. In treatment 1 (T1) the whole ‘head’ of each kelp sporophyte that reached the surface was cut off between the bulb and the primary blade (‘lethal’ method). In treatment 2 (T2) (‘non-lethal’ method), the secondary fronds of all sporophytes that reached the surface were cut 20–30 cm from their junction with the primary blade, and removed. In the control plot, the kelp plants were not treated. Harvesting treatments were done approximately every four months, at low spring tide, from 3 March 2003 to 3 November 2003 (three treatments). The effects of harvesting on the kelps depend largely on the size of plant and the time the fronds were removed; however, no seasonal pattern could be observed. The different treatments had no effect on the growth rate, population structure or recruitment of the kelp. This means that factors other than light play an important role in the growth, structure and recruitment of the kelp beds in False Bay. Results are discussed in relation to current commercial harvesting practices.