Morphological, physiological and reproductive conditions of rafting bull kelp (Durvillaea antarctica) in northern-central Chile (30°S)

The bull kelp Durvillaea antarctica is a common floating alga in the southern hemisphere, but despite the ecological and biogeographic importance of kelp rafts, little is known about the responses of detached kelps to the conditions at the sea surface. The morphological, physiological and reproductive performances of D. antarctica rafts in the Coastal System of Coquimbo, Chile (CSC, ~30°S), were examined during winter and summer of two successive years (2010/11 and 2011/12). Epibionts (Lepas spp.) that only attach to floating objects were used as indicator for the floating time of kelp rafts. Photosynthetic efficiency and reproductive maturity of both benthic and floating algae varied seasonally, with a stronger decrease in summer than in winter. Blade size (measured as weight proportion of kelp individuals), phlorotannin concentrations and antioxidant activities were lower in floating than in benthic algae. Environmental conditions and floating time affected the blade tissues, with stronger negative effects during summer. These results confirm that floating persistence of D. antarctica in the CSC is suppressed during the summer months, which indicates that the dispersal potential of this (and other) floating algae varies seasonally.     

Long-distance dispersal of a subantarctic brooding bivalve (Gaimardia trapesina) by kelp-rafting

The probability of successful dispersal by sessile benthic invertebrates is thought to strongly influence their geographic distribution and population genetics. Generally, species with long-lived planktonic larvae are expected to exhibit wider distribution patterns than those species which brood their young, due to their presumably greater potential for dispersal. In some cases, however, brooding species exhibit broad distributions and show evidence of genetic exchange with geographically distant populations. One potential factor that has been invoked as an expianation is dispersal by floating and rafting of adults and egg masses. Several studies have shown that it is possible for sessile adults to disperse on the order of several to many thousand kilometers by rafting on debris in ocean currents. With very few exceptions, however, direct evidence of rafting in the open ocean has been lacking. We present evidence of long-distance (1300 to 2000 km) dispersal of a brooding pelecypod,Gaimardia trapesina (Lamarck, 1819), in the Southern Ocean in the vicinity of Cape Horn, the Falkland Islands, and the antarctic island South Georgia (54°S; 37°W). Data on survival and fecundity rates ofG. trapesina and the prevalence of kelp rafts collected during the austral winter of 1993 indicate that dispersal by rafting can occur over ecologically relevant time scales and could potentially serve as a significant means of genetic exchange between populations.     

Dispersal of faunal and floral propagules associated with drifting Macrocystis pyrifera plants

The potential of drifting Macrocystis pyrifera kelp for transporting associated animals and plants long distances around the southern oceans was assessed by anchoring kelp holdfasts off the Tasmanian coast in 1985, monitoring the turnover of organisms, and relating species survival to water-transport times and species geographic distributions. Although most of the common animal species and approximately half of the plant species associated with Tasmanian M. pyrifera holdfasts were still present on kelp holdfasts after 191 d at sea, very few of these species have been recorded from New Zealand. It therefore seems unlikely that M. pyrifera plants with intact holdfasts are presently drifting to New Zealand. Drifting kelps probably become negatively buoyant in the Tasman Sea because dissolved nitrate concentrations are insufficient for normal plant growth. Moreover, even if some kelp plants do drift to New Zealand it is possible that their holdfasts rapidly disintegrate in the open ocean because of the abundance of the boring isopods Phycolimnoria spp. in Tasmanian holdfasts. In contrast to the restricted distributions of Tasmanian holdfast-inhabiting species, most of the identified species collected from M. pyrifera holdfasts at subantarctic Macquarie Island also occurred 5 000 km west at Kerguelen Island. Because of the extensive ranges of many subantarctic species, the good probability of survival of epifaunal species on drifting kelps, and the high surface-water nitrate concentrations and low holdfast-densities of Phycolimnoria spp. in the higher latitudes, it is likely that M. pyrifera-mediated transport of faunal and floral propagules has recently occurred, and is probably presently occurring, in subantarctic waters.     

UV-radiation versus grazing pressure: long-term floating of kelp rafts (<Emphasis Type="Italic">Macrocystis pyrifera</Emphasis>) is facilitated by efficient photoacclimation but undermined by grazing losses

Large quantities of floating macroalgae are traveling in coastal waters of the SE Pacific and in other temperate climate zones. While afloat, these algae are potentially exposed to full solar radiation, including UVA and UVB, which can have profound effects on their physiological and growth performance. Latitudinal variations in UV-radiation (UVR) are hypothesized to affect floating algae differently with higher impacts at low latitudes than at high latitudes. In addition, UVR together with grazing might accelerate the demise of floating kelps. This hypothesis was tested with outdoor laboratory experiments in which sporophytes of the giant kelp Macrocystis pyrifera (L.) C. Agardh were exposed to a combination of different UVR regimes (PAR only, PAR + UV) and grazing at three sites along the Chilean coast (20°S, 30°S, and 40°S). A latitudinal trend in irradiance was detected with increasing values from 40°S to 20°S. Surprisingly, floating M. pyrifera responded with a high acclimation potential within this latitudinal UVR gradient. At 20°S, floating kelps were slightly sensitive to UVR, which was reflected in reduced blade growth. At 30°S, physiological responses were hardly affected by the prevailing irradiance but sporophyte growth and thus persistence mainly depended on the presence or absence of amphipod grazers. At high latitudes, grazing had only minor impacts on algal biomass and blade growth, and kelps thrived well under all tested environmental conditions. Overall, our results reveal that floating M. pyrifera was only slightly affected by UVR and that sporophytes can efficiently acclimate over a latitudinal UVR gradient that spans from 20°S to 40°S. Given this high acclimation potential, we suggest that these (and possibly other) positively buoyant algae are important dispersal agents over a wide range of temperate latitude conditions.     

Population structure,northern range limit,and recruitment variation in the intertidal limpet Lottia scabra

The northern range limit of the intertidal limpet Lottia scabra is Cape Arago, Oregon (43°N), where adult survival is excellent, the population is small (<300), and recruitment is low; the range limit may be set by limited recruitment. Between June 2012 and March 2013, 25 sites from the middle of the species range (33°N) to Cape Arago were sampled and population size frequency distributions, densities, and nearest neighbor distances were compared to the amount of rocky and sandy shore and kelp bed size. North and south of 37°N, the densities of new recruits averaged 22 and 86 m^?2, respectively. This shift was associated with the range limit of Macrocystis pyrifera kelp beds; we hypothesize that slower currents in M. pyrifera beds may limit larval dispersal leading to higher recruitment. North and south of 40°N, adult density averaged <1 and 458 m^?2, respectively, with the species absent from many sites to the north. This shift was associated with a sharp drop in the amount of rocky shoreline and an increase in uninhabitable sandy shore. Near the northern range limit, >80 % of the individuals were solitary and may be unable to spawn successfully. Recruitment at Cape Arago was infrequent and likely due to self-recruitment. This study suggests that the range limit was set by the absence of M. pyrifera and too little rocky shore leading to high larval wastage, low settlement, low population densities, and, due to an Allee effect, very small effective population sizes.     

Herbivorous amphipods inhabit protective microhabitats within thalli of giant kelp <Emphasis Type="Italic">Macrocystis pyrifera</Emphasis>

Many small marine herbivores utilize specific algal hosts, but the ultimate factors that shape host selection are not well understood. For example, the use of particular microhabitats within algal hosts and the functional role of these microhabitats have received little attention, especially in large algae such as kelps. We studied microhabitat use of the herbivorous amphipod Peramphithoe femorata that inhabits nest-like domiciles on the blades of giant kelp Macrocystis pyrifera. The vertical position of nest-bearing blades along the stipe of the algal thallus and the position of the nests within the lateral blades of M. pyrifera were surveyed in two kelp forests in northern-central Chile. Additionally, we conducted laboratory and field experiments to unravel the mechanisms driving the observed distributions. Peramphithoe femorata nests were predominantly built on the distal blade tips in apical sections of the stipes. Within-blade and within-stipe feeding preferences of P. femorata did not explain the amphipod distribution. Amphipods did not consistently select distal over proximal blade sections in habitat choice experiments. Mortality of tethered amphipods without nests was higher at the seafloor than at the sea surface in the field. Nests mitigated mortality of tethered amphipods, especially at the seafloor. Thus, protective microhabitats within thalli of large kelp species can substantially enhance survival of small marine herbivores. Our results suggest that differential survival from predation might be more important than food preferences in determining the microhabitat distribution of these herbivores.     

Severe storm disturbances and reversal of community structure in a southern California kelp forest

Regular observations made over a period of 5 yr in four permanent transects provided data on plant, sea urchin, and fish densities which indicate that two unusually severe winter storms in 1980 (Storm I) and 1983 (Storm II) had different effects on a southern California kelp-forest community. Storm I removed all canopies of the giant kelp Macrocystis pyrifera, but spared most understory kelps, mainly Pterygophora californica. Hence, the previously large accumulation of detached drift kelp, mostly M. pyrifera, disappeared. Denied their preferred diet of drift kelp, the sea urchins Strongylocentrotus franciscanus and S. purpuratus then emerged from shelters to find alternative food. Without effective predators, they consumed most living plants, including the surviving understory kelps. This weakened the important detritus-based food chain, as indicated indirectly by declining abundances of algal turf and fish (Embiotocidae) that eat small animals living in turf. In 1983, Storm II reversed the process by eliminating exposed urchins, while clearing rock surfaces for widespread kelp settlement and growth. By summer 1984, the kelp grew to maturity to form extensive canopies despite elevated water temperatures during summer and fall of 1983. Thus, severe storms may have vastly different effects on community structure, depending on the state of the community before the disturbance.     

Population structure of the Spanish sardine Sardinella aurita: natural morphological variation in a genetically homogeneous population

The importance of larval dispersal in determining the distribution and abundance of benthic marine organisms is well recognized; however, the contribution of post-larval dispersal has not been measured. I compared the dispersal of swimming larvae with that of rafting colonies in a population of compound ascidians, Botrylloides sp., living attached to leaves of the eelgrass Zostera marina in Tomales Bay, California, USA in 1990–1992. Colonies rafting on broken eelgrass traveled over 200 times farther and had comparable recruitment success relative to swimming larvae. The recruitment of rafting colonies into new habitats was facilitated by the ability of thesecolonial animals to grow asexually onto surrounding substrata. Rafting colonies brooded larvae that were subsequently relased after the colony settled into a new habitat. These results suggest that colonization of new habitats can occur by post-larvae as well as larvae, and that long-range dispersal by species with short-lived larvae may occur by post-larval rafting.     

Relationship of diversity and habitat area in North Pacific plastic-associated rafting communities

Plastic and other anthropogenic debris (e.g., rubber, tar) augment natural floating substrates (e.g., algal rafts, pumice) in the open ocean, allowing “islands” of substrate-associated organisms to persist in an otherwise unsuitable habitat. We examined a total of 242 debris objects collected in the eastern Pacific in 2009 and 2011 (32–39°N, 130–142°W) and the western Pacific in 2012 (19–41°N, 143–156°E). Here, we ask: (a) What taxa are associated with plastic rafts in the North Pacific? and (b) Does the number of taxa associated with plastic debris vary with the size of the debris “island?” We documented 95 rafting taxa from 11 phyla. We identified several potentially invasive plastic-associated rafting taxa, including the coral pathogen Halofolliculina spp. In concordance with classic species–area curves, the number of rafting taxa was positively correlated with the size of the raft. Our findings suggest that diversity patterns on plastic debris are compatible with the concept of island biogeography.     

Effects of grazing by two species of sea urchins (Strongylocentrotus franciscanus and Lytechinus anamesus) on recruitment and survival of two species of kelp (Macrocystis pyrifera and Pterygophora californica)

We studied the effects of grazing by two species of sea urchins on two species of kelp (Macrocystis pyrifera and Pterygophora californica) in the San Onofre kelp bed in southern California from 1978 through 1981. Both red sea urchins, Strongylocentrotus franciscanus, and white sea urchins, Lytechinus anamesus, were abundant and lived in aggregations. The purple sea urchin (S. purpuratus) was rare at the study site and was not studied. The aggregations of red urchins were either relatively small and stationary (for over 3 yr) or relatively large and motile (advancing at about 2 m mo^–1). Both stationary and moving aggregations were observed at the same time, and within 100 m of one another. Stationary aggregations of red urchins probably subsisted mainly on drift kelp and had no effect on kelp recruitment or on adult kelp abundance. In contrast, red sea urchins in large, motile aggregations or fronts ate almost all the macroalgae in their path. The condition of their gonalds indicated that red urchins in fronts were starved relative to red urchins in the small, stationary aggregations. Large, motile aggregations developed after 2 yr of declining kelp abundance (probably due largely to storms). We propose that a scarcity of drift algae for food results in a change in the behavior pattern of the red urchins and thus leads to the formation of large, motile aggregations. The aggregations of white urchins, which occurred along the offshore margin of the kelp bed, were large, but relatively stationary. The white urchins rarely ate adult kelps, but grazed extensively on early developmental stages of kelps and evidently prevented seaward expansion of the bed. The spatial distribution of both types of red urchin aggregations appeared to be unrelated to predation pressure from fishes or lobsters.Please address all requests for reprints to the senior author at his present address.     

Algal succession in a Macrocystis pyrifera forest

Algal succession within a subtidal forest of the giant kelp Macrocystis pyrifera was studied by following colonization and community development on concrete blocks fastened to the bottom. Sets of blocks were placed in the bed at 3-month intervals. Subsequent algal development on each set was followed for over a year. All macroscopic species attached to the substrata were noted, and the number and lenght of basal branches determined every 1 to 3 months. Colonizing plants fell into 3 categories: rapid-growing ephemerals, and rapid and slow-growing perennials. Ephemerals such as Giffordia (Ectocarpus) mitchellae, Colpomenia peregrina, and diatom films generally produced and initial bloom on the blocks but were gradually replaced by perennials (articulated corallines, Rhodymenia spp., Gigartina spp.) characteristic of the mature kelp community. These stages, rather than representing “ecological” succession, seemed to reflect differences in growth rate and success in interspecific competition for space and light. Colonization on the blocks varied with season, indicating that most species have either a spring-summer or fall-winter period of maximum reproduction. M. pyrifera sporophyte colonization was greatest in spring. During community development, algal diversity (H), number of species (s) and evenness (J) all reached a peak within 100 to 200 days regardless of the time the blocks were started. Diversity and number of species then fell as ephemeral species disappeared. These species were apparently unable to compete with perennials and, once gone, did not recolonize. Evenness remained high.     

Geographic variation in polyphenolic levels of Northeastern Pacific kelps and rockweeds

Brown algal polyphenolic compounds are secondary metabolites whose functions may include protecting plants from pathogens or damage by UV radiation, and deterring feeding by herbivores. We present here the first analysis of spatial variation (at scales from tens of meters to hundreds of kilometers) in concentration of these compounds in two orders of brown algae from the northeastern Pacific Ocean. In kelps (order Laminariales), variation among sites was significant in only 25% of species examined and was consistent within families (high in the Alariaceae and low in the Laminariaceae and Lessoniaceae). In rockweeds (order Fucales, family Fucaceae), site variation was high in three of four species examined. Both the proportion of high polyphenolic kelp species and the magnitude of spatial variation within species from both kelps and rockweeds were much higher than would have been predicted from previous studies in other regions. In one kelp (Laminaria groenlandica), significant differences between sites occurred at scales of only tens of meters. No latitudinal clines were observed. Differences in phenolic concentrations of kelps spanned nearly an order of magnitude in one species, Hedophyllum sessile. Phenolic levels were significantly higher in members of the Fucales than the Laminariales, but showed no significant differences between intertidal and subtidal species. Received: 22 July 1996 / Accepted: 26 October 1998     

Hydrology influences population genetic structure and connectivity of the intertidal amphipod Corophium volutator in the northwest Atlantic

The mechanisms driving genetic structure in marine systems are elusive due to the difficulty of identifying temporal and spatial barriers to dispersal. By studying marine invertebrate species with limited dispersal potential, genetic structure can be directly related to physical and biological factors restricting connectivity. In the northwest Atlantic, the benthic brood-rearing amphipod Corophium volutator is distributed across basins with distinct circulation patterns and has the potential to disperse passively during its adult stage. We analyzed spatial genetic variation and migration rates across C. volutator’s North American range with sequence data for mitochondrial DNA and three novel nuclear markers using frequency and coalescent-based methods. We found low genetic differentiation within basins, but strong subdivision within the Bay of Fundy and a striking biogeographic break between the Bay of Fundy and Gulf of Maine, suggesting that genetic drift may act on populations in which connectivity is restricted due to the limitation of passive dispersal by hydrological patterns.     

Trophic significance of kelps in kelp communities in Brittany (France) inferred from isotopic comparisons

The relative contribution of kelps and other food sources to the diet of consumers in shallow coastal communities has been hotly debated in recent years. It has been suggested that considering proxies instead of phytoplankton isotopic signatures can lead to an overestimation of kelp contribution through isotopic mixing models. We analysed spatial patterns in carbon isotopic ratios of the dominant primary sources and consumers at two subtidal sites in Brittany (France) prior to the anticipated 2011 phytoplankton bloom to determine which of kelp and phytoplankton is the dominant food source in Laminaria hyperborea forests. We found simultaneous spatial variations for consumers, kelps and deposited particulate organic matter, suggesting that kelp-derived organic matter is a key component of associated food webs for at least part of the year.     

Factors influencing food choice by the seaweed-eating marine snail Norrisianorrisi (Trochidae)

Through two-choice gustatory experiments, a␣preference hierarchy was established␣for␣the␣herbivorous gastropod Norrisianorrisi Sowerby, with laminarialean kelps preferred over all other seaweeds. Among the kelps, laminae of Macrocystispyrifera were slightly preferred over Egregiamenziesii, and both were strongly preferred over sporophylls of Eiseniaarborea. E.arborea, the least preferred kelp, was consistently chosen over other algae common in the snail's habitat (Halidrysdioica, Dictyotaflabellata, and Pterocladiacapillacea) and over seaweeds believed to be edible and palatable based on their morphology, structure, and secondary chemistry (Endarachnebinghamiae, Mazzaellaflaccida, and Ulvalobata). The morphologies and structural toughness of tested seaweeds varied significantly as did their nutritional (% carbon, % nitrogen, C:N ratio, and % ash) contents and phlorotannin concentrations; however, snails preferred to feed on kelps regardless of nutritional content, toughness or phlorotannin concentration; and among kelps preferred to feed on the least tough species (based on penetrometer measurements), which also were those containing the lowest phlorotannin concentrations. Preference for kelp was not upheld in experiments using agarose thalli to which freeze-dried powder, of either the kelp Eiseniaarborea or non-kelp Endarachnebinghamiae was added, suggesting the destruction of attractant chemicals during the making of the artificial foods. Our data suggest that the preference of N. norrisi for kelps over other potentially edible and palatable seaweeds may not be related to nutritional content, but instead may have evolved in response to factors such as availability, habitat provision, or refuge from predation. Received: 27 September 1996 / Accepted: 7 October 1997     

Colonisation of kelp imitations by epiphyte and holdfast fauna; a study of mobility patterns

The kelp Laminaria hyperborea provides a habitat for a diverse invertebrate community. Invertebrate abundance is particularly high in the kelp holdfasts and associated with epiphytes on the stipes. We investigated colonisation patterns of this fauna on artificial substrata (holdfast and epiphyte mimics) exposed for short time periods in the kelp forest in two seasons, summer and late autumn. Within 7 days, 99 taxa common on natural kelp substratum colonised the artificial substrata in large numbers, demonstrating high mobility and indicating that there may be a rapid exchange of individuals and species among algae. Gastropods and amphipods were the most abundant colonisers. In summer, a specific “epiphyte fauna” could be distinguished from a “holdfast fauna”, and species associated with epiphytes were significantly more abundant on the artificial substrata. Colonisation was more rapid in summer than in late autumn, which may be related to animal population density. The high dispersal rate can facilitate the interaction with other communities and with higher trophic levels in the kelp-forest food web. Published online: 13 August 2002     

Is kelp detritus a good food for suspension feeders? Effects of kelp species, age and secondary metabolites

The food quality of detrital particles derived from three species of kelps was evaluated in a laboratory feeding experiment utilizing two species of suspension feeders, the serpulid polychaete Pseudochitonopomaoccidentalis and the mussel Mytilustrossulus. Fresh and aged kelp particles were also evaluated, and growth in all treatments was compared to growth on ad libidum phytoplankton rations. Fresh particles from Laminariagroenlandica, aged particles from Agarumfimbriatum and Alariamarginata, and mixed phytoplankton promoted the highest growth rates in both consumers. Growth was inversely related to total polyphenolic concentration in the fresh kelp particles. The increase in quality of both Agarumfimbriatum and Alariamarginata particles with age corresponded with a rapid loss of polyphenolic secondary metabolites and an increase in total nitrogen. Received: 26 November 1996 / Accepted: 9 January 1997     

Methyl iodide (CH3I) production by kelp and associated microbes

Methyl iodide (CH₃I) and methyl bromide (CH₃Br) were produced by an axenic kelp-tissue culture and methyl iodide was produced by microbial cultures derived from decaying kelp tissue, demonstrating that methyl iodide is a product of both kelp and microbial metabolism. Methyl iodide production by blade tissue from five kelp species was not enhanced by desiccation, but production rates were enhanced by drastic cellular disruption. Wounding and tissue age of Macrocystis pyrifera had no effect on methyl iodide production. Undefined microbial populations, obtained from decaying kelp, produced methyl iodide at low rates while growing on suspensions of powdered kelp tissue in seawater and on artificial media. This study (1984–1985 on kelp collected from California, USA) confirms that direct methyl iodide production by kelp is not globally significant, but suggests that production by marine microbes associated with seaweed degradation may be of global importance.Please address all requests for reprints to Dr. S. L. Manley in Santa Cruz