Recruitment tolerance to increased temperature present across multiple kelp clades.

Kelp systems dominate nearshore marine environments in upwelling zones characterized by cold temperatures and high nutrients. Worldwide, kelp population persistence and recruitment success generally decreases with rising water temperatures coupled with low nutrients, making kelp populations vulnerable to impending warming of the oceans. This response to climate change at a global scale, however, may vary due to regional differences in temperature variability, acclimation, and differential responses of kelp species to changing conditions. Culture experiments were conducted on 12 eastern Pacific kelp taxa across geographic regions (British Columbia, central California, and southern California) under three nitrate levels (1, 5, and 10 µmol/L) and two temperatures (12°C and 18°C) to determine sporophyte production (i.e., recruitment success). For all taxa from all locations, sporophytes were always present in the 12°C treatment and when recruitment failure was observed, it always occurred at 18°C, regardless of nitrate level, indicating that temperature is the driving factor limiting recruitment, not nitrate. Rising ocean temperatures will undoubtedly cause recruitment failure for many kelp species; however, the ability of species to acclimatize or adapt to increased temperatures at the warmer edge of their species range may promote a resiliency of kelp systems to climate change at a global scale.

Plant-animal diversity relationships in a rocky intertidal system depend on invertebrate body size and algal cover.

Considerable research has examined the influence of herbivores on the maintenance of plant diversity, but fewer studies have examined the reciprocal effect of plant diversity on the animals that use the plant community for food and shelter, particularly in marine systems. Several mechanisms could underlie such effects. Animal diversity and abundance could be increased by complementary use of different plants by different animals, or by an indirect effect of plant diversity on plant production that results in more total plant biomass in high plant-diversity communities. Alternatively, plant species identity could play a dominant role leading to sampling effects or no effect of diversity at all. We conducted a six-year field manipulation of the richness of rocky shore seaweeds in northern California and measured the effects of algal richness and identity on the invertebrate community, from meiofauna to macrofauna. We found that diverse algal communities hosted more species of both large and small invertebrates than the average algal monoculture but that the mechanisms underlying this pattern differed substantially for organisms of different size. More species of macrofauna occurred in the polycultures than in any of the monocultures, likely due to the greater total cover of algae produced in polycultures. Rare and common macrofaunal taxa responded to host plant species richness in opposite ways, with more occurrences of rare taxa and lower abundance of very common taxa in the polycultures. In contrast, meiofaunal richness in polycultures was no different than that of monocultures of finely branched species, leading to strong effects of algal identity. Our findings are similar to those from terrestrial systems in that the effects of plant diversity we observed were most related to the greater amount of habitat in polycultures as a result of overyielding in algal biomass. However, our findings differ from those in terrestrial systems in that the primary mechanisms for both richness and identity effects appear related to the value of plants as shelter from harsh abiotic conditions or predation rather than food, and in that animal body size altered the mechanisms underlying diversity effects.

EFFECTS OF CLIMATE CHANGE ON GLOBAL SEAWEED COMMUNITIES.

Seaweeds are ecologically important primary producers, competitors, and ecosystem engineers that play a central role in coastal habitats ranging from kelp forests to coral reefs. Although seaweeds are known to be vulnerable to physical and chemical changes in the marine environment, the impacts of ongoing and future anthropogenic climate change in seaweed-dominated ecosystems remain poorly understood. In this review, we describe the ways in which changes in the environment directly affect seaweeds in terms of their physiology, growth, reproduction, and survival. We consider the extent to which seaweed species may be able to respond to these changes via adaptation or migration. We also examine the extensive reshuffling of communities that is occurring as the ecological balance between competing species changes, and as top-down control by herbivores becomes stronger or weaker. Finally, we delve into some of the ecosystem-level responses to these changes, including changes in primary productivity, diversity, and resilience. Although there are several key areas in which ecological insight is lacking, we suggest that reasonable climate-related hypotheses can be developed and tested based on current information. By strategically prioritizing research in the areas of complex environmental variation, multiple stressor effects, evolutionary adaptation, and population, community, and ecosystem-level responses, we can rapidly build upon our current understanding of seaweed biology and climate change ecology to more effectively conserve and manage coastal ecosystems.

GRAZING ON GIANT KELP MICROSCOPIC PHASES AND THE RECRUITMENT SUCCESS OF ANNUAL POPULATIONS OF MACROCYSTIS PYRIFERA (LAMINARIALES, PHAEOPHYTA) IN SOUTHERN CHILE(1).

The giant kelp Macrocystis pyrifera (L.) C. Agardh is widely distributed in the Northern Hemisphere and Southern Hemisphere, yet it exhibits distinct population dynamics at local to regional spatial scales. Giant kelp populations are typically perennial with the potential for year-round reproduction and recruitment. In southern Chile, however, annual giant kelp populations exist and often persist entirely on secondary substrata (e.g., shells of the slipper limpet Crepipatella fecunda [Gastropoda, Calyptraeidae]) that can cover up to 90% of the rocky bottom. In these populations, the macroscopic sporophyte phase disappears annually during winter and early spring, leaving a 3-4 month period in which a persistent microscopic phase remains to support the subsequent year's recruitment. We tested the effects of a suite of grazers on the recruitment success of this critical microscopic phase at two sites in southern Chile. Field experiments indicated that the snail Tegula atra negatively impacted M. pyrifera sporophyte recruitment, but that recruitment was highest in the presence of sessile female limpets, C. fecunda. Conversely, small male C. fecunda (biofilm grazers) did not regulate kelp recruitment. Laboratory observations showed that C. fecunda males only grazed on microscopic kelp gametophytes and small (<250 µm) sporophytes, rejecting larger sporophytes, whereas T. atra grazed on all the kelp stages. Recruitment to the C. fecunda treatments far exceeded that to bare rock in the absence of grazers but was not due to the physical presence of C. fecunda shells. We concluded that the key to M. pyrifera recruitment success in southern Chile is its capacity to colonize secondary substrates provided by the slipper limpet C. fecunda.

ABIOTIC REGULATION OF INVESTMENT IN SEXUAL VERSUS VEGETATIVE REPRODUCTION IN THE CLONAL KELP LAMINARIA SINCLAIRII (LAMINARIALES, PHAEOPHYCEAE)1.

Clonal kelp taxa may reproduce both sexually and vegetatively resulting in a potential trade-off in the allocation of acquired carbon and nitrogen resources. Such trade-offs may dictate a different response of clonal kelps to varying environmental conditions relative to aclonal kelp taxa. Laboratory temperature and nutrient manipulation experiments demonstrated that investment in sexual and vegetative reproduction in Laminaria sinclairii (Harv. ex Hook. f. et Harv.) Farl., C. L. Anderson et D. C. Eaton was regulated by different abiotic factors. Sorus production (investment in sexual reproduction) and blade growth were significantly higher at 12°C compared to 17°C, regardless of nutrient concentration. Net carbon storage and depletion in rhizomes were observed in the low- and high-temperature treatments, respectively, suggesting that carbon stores were not responsible for increased growth. Rhizome elongation (investment in vegetative reproduction), on the other hand, was significantly higher in 12 µM NO3- than in 2 µM NO3- , irrespective of temperature. This increase in rhizome growth was concurrent with elevated rhizome percent tissue nitrogen levels also observed in treatments with higher nutrients, again indicating a growth response to treatment independent of previous nutrient stores. These results suggest that regulation of growth and investment in sexual reproduction in L. sinclairii is similar to that in aclonal kelps (i.e., warmer temperatures result in decreased reproductive output). Additionally, depletion of carbon and nitrogen from rhizomes in suboptimal conditions confirms the role of clonal kelp rhizomes in carbon and nutrient storage.

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.

PHENOTYPIC PLASTICITY RECONCILES INCONGRUOUS MOLECULAR AND MORPHOLOGICAL TAXONOMIES: THE GIANT KELP, MACROCYSTIS (LAMINARIALES, PHAEOPHYCEAE), IS A MONOSPECIFIC GENUS(1).

The giant kelp genus Macrocystis C. Agardh (Laminariales, Phaeophyceae) is one of the world's most ecologically and economically important seaweed taxa, yet its taxonomy remains uncertain. Although the genus currently contains four accepted species based on variable holdfast and blade morphology [M. pyrifera (L.) C. Agardh, M. integrifolia Bory, M. angustifolia Bory, and M. laevis C. H. Hay], numerous recent studies on Macrocystis interfertility, genetic relatedness, and morphological plasticity all suggest that the genus is monospecific. We reviewed this evidence and present an explanation for the extreme phenotypic plasticity that results in morphological variability within Macrocystis, driven by the effects of environmental factors on early development of macroscopic sporophytes. We propose that the genus be collapsed back to a single species, with nomenclatural priority given to M. pyrifera.

Complementarity in marine biodiversity manipulations: reconciling divergent evidence from field and mesocosm experiments.

Mounting concern over the loss of marine biodiversity has increased the urgency of understanding its consequences. This urgency spurred the publication of many short-term studies, which often report weak effects of diversity (species richness) driven by the presence of key species (the sampling effect). Longer-term field experiments are slowly accumulating, and they more often report strong diversity effects driven by species complementarity, calling into question the generality of earlier findings. However, differences among study systems in which short- and long-term studies are conducted currently limit our ability to assess whether these differences are simply due to biological or environmental differences among systems. In this paper, we compared the effect of intertidal seaweed species richness on biomass accumulation in mesocosms and field experiments using the same pool of species. We found that seaweed species richness increased biomass accumulation in field experiments in both short (2-month) and long (3-year) experiments, although effects were stronger in the long-term experiment. In contrast, richness had no effect in mesocosm experiments, where biomass accumulation was completely a function of species identity. We argue that the short-term experiments, like many published experiments on the topic, detect only a subset of possible mechanisms that operate in the field over the longer term because they lack sufficient environmental heterogeneity to allow expression of niche differences, and they are of insufficient length to capture population-level responses, such as recruitment. Many published experiments, therefore, likely underestimate the strength of diversity on ecosystem processes in natural ecosystems.

Deep-water kelp refugia as potential hotspots of tropical marine diversity and productivity.

Classic marine ecological paradigms view kelp forests as inherently temperate-boreal phenomena replaced by coral reefs in tropical waters. These paradigms hinge on the notion that tropical surface waters are too warm and nutrient-depleted to support kelp productivity and survival. We present a synthetic oceanographic and ecophysiological model that accurately identifies all known kelp populations and, by using the same criteria, predicts the existence of >23,500 km(2) unexplored submerged (30- to 200-m depth) tropical kelp habitats. Predicted tropical kelp habitats were most probable in regions where bathymetry and upwelling resulted in mixed-layer shoaling above the depth of minimum annual irradiance dose for kelp survival. Using model predictions, we discovered extensive new deep-water Eisenia galapagensis populations in the Galápagos that increased in abundance with increasing depth to >60 m, complete with cold-water flora and fauna of temperate affinities. The predictability of deep-water kelp habitat and the discovery of expansive deep-water Galápagos kelp forests validate the extent of deep-water tropical kelp refugia, with potential implications for regional productivity and biodiversity, tropical food web ecology, and understanding of the resilience of tropical marine systems to climate change.

Community-wide distribution of predator-prey interaction strength in kelp forests.

The strength of interactions between predators and their prey (interaction strength) varies enormously among species within ecological communities. Understanding the community-wide distribution of interaction strengths is vital, given that communities dominated by weak interactions may be more stable and resistant to invasion. In the oceans, previous studies have reported log-normal distributions of per capita interaction strength. We estimated the distribution of predator-prey interaction strengths within a subtidal speciose herbivore community (45 species). Laboratory experiments were used to determine maximum per capita interaction strengths for eight species of herbivores (including amphipods, isopods, gastropods, and sea urchins) that graze on giant kelp (Macrocystis pyrifera) microscopic stages. We found that maximum per capita interaction strength saturated as a function of individual herbivore biomass, likely caused by predator/prey size thresholds. Incorporating this nonlinearity, we predicted maximum per capita interaction strength for the remaining herbivore species. The resulting distribution of per capita interaction strengths was bimodal, in striking contrast to previous reports from other communities. Although small herbivores often had per capita interaction strengths similar to larger herbivores, their tendency to have greater densities in the field increased their potential impact as grazers. These results indicate that previous conclusions about the distributions of interaction strength in natural communities are not general, and that intermediate-sized predators can under realistic circumstances represent the most effective consumers in natural communities.