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     

Scale-dependent changes in the importance of larval supply and habitat to abundance of a reef fish

While there is great interest in the degree to which local interactions "scale-up" to predict regional patterns of abundance, few studies in marine systems have simultaneously examined patterns of abundance at both the large scale (tens of kilometers) typical of larval movement and the small scale (meters) typical of post-settlement interactions. We addressed this gap by monitoring larval supply, adult survivorship, and giant kelp (Macrocystis pyrifera, a primary habitat-forming species) abundance for 13 populations of kelp bass (Paralabrax clathratus) spread over approximately 200 km in the Santa Barbara Channel, California, USA. At the small, within-site scale, both recruitment and adult survivorship of kelp bass were density-dependent and positively related to kelp abundance. At the larger, among-site scale, the spatial pattern of adult kelp bass abundance was predicted well by the pattern of kelp bass larval supply, but there was a consistent negative spatial relationship between kelp abundance and kelp bass larval supply despite the positive effects of kelp on kelp bass at the smaller spatial scale. This large-scale negative relationship was likely a product of a channel-wide spatial mismatch between oceanographic conditions that favor kelp survival and those that concentrate and distribute fish larvae. These results generally support the recruit-adult hypothesis: kelp bass populations are limited by recruitment at low recruit densities but by density-dependent competition for food resources and/or predator refuges at high recruit densities. At the same time, spatial variation in kelp abundance produced substantial spatiotemporal heterogeneity in kelp bass demographics, which argues for a multispecies, metacommunity approach to predicting kelp bass dynamics.     

Temporal and spatial variability in settlement of the sea urchin<Emphasis Type="Italic"> Paracentrotus lividus</Emphasis> in the NW Mediterranean

Settlement into the benthic habitat may be an important process in regulating sea urchin abundance, which potentially modifies the structure of benthic communities. Strong settlement events may increase sea urchin abundance beyond a certain threshold, leading to the formation of coralline barrens (overgrazed communities with a dominance of encrusting coralline algae). To understand the role of settlement in regulating sea urchin populations we first need to determine settlement variability. Temporal variation in settlement of the sea urchin Paracentrotus lividus was monitored at three sites in the Medes Islands, NW Mediterranean, during three settlement seasons (March 1998 through October 2000). Spatial variation in settlement was studied in 1999 at 50 sites along a gradient of exposures to waves and currents, inside and outside the archipelago, and separated by distances from tens to thousands of meters. Bathymetric distribution of settlement was also studied in 2000 at six sites at 5, 10, 15 and 20 m depths. Settlement of P. lividus occurred in a single annual peak within 3 weeks in May–June. Differences in settlement between years were more than two orders of magnitude. Spatial variability was found at all scales investigated, showing strong patchiness at the smallest spatial scales (tens of meters). Sea urchins settled preferentially at depths between 5 and 10 m. Substratum type, level of protection, and adult population densities were not significant in determining settlement. However, settlement was found to be related to the degree of exposure to waves and currents, indicating that physical processes are very important at the spatial scales investigated. This greatly variable settlement is a necessary, although not sufficient, condition to create gradients of adult P. lividus abundance. Further studies should be designed to investigate the interaction between settlement strength and post-settlement mortality.Communicated by O. Kinne, Oldendorf/Luhe     

Flowering of the seagrass Posidonia oceanica in a north-western Mediterranean coastal area: temporal and spatial variations

Sexual reproduction in Posidonia oceanica (L.) Delile is likely to be highly variable both in space and time. However, there has been little effort to identify at what spatial scale variability exists in the distribution and abundance of reproductive shoots or the magnitude of these variations. Such information is important for elucidating several aspects of the reproductive strategy of this species. Here, by using a reconstructing technique, I estimated the rate at which flowering events occurred over the last decades in three P. oceanica meadows along the coastline of the north-western Mediterranean (Italy). Then, variation in inflorescence frequency (i.e. the percentage of shoots that flowered in a particular year of the total number of shoots present in the sample in that year) was examined on a hierarchy of spatial scales, ranging from a few metres to meadows separated by tens of kilometres over a 3-year period. Separate analyses were also conducted to examine the patterns of spatial variability of several plant characteristics (i.e. shoot age and density of vegetative shoots) that may influence the timing of flowering. In total, ten reconstructed flowering events were recorded during the period 1983–1999. Despite the similar climatic conditions experienced in the three meadows surveyed, no inflorescence scars were found in one meadow. In the other two meadows, the flowering rate fluctuated considerably from year to year (0–26%), and simultaneous flowerings were detected only in 3 years (1993, 1994 and 1995). The pattern of spatial variability in the inflorescence frequency varied among years. An investigation of the flowering events recorded along the north-western Mediterranean coast reveals that in the studied period flowering also occurred across a broad range of meadows. Therefore, flowering could be a far more widespread phenomenon in this sector of the Mediterranean basin than previously thought. Analysis of variance showed significant variation in 1993 at the mean inflorescence frequency on very small scales (metres to tens of metres), whereas variation was not found at any of the scales investigated in 1994 and 1995. Overall, variation in the inflorescence frequency within individual meadows (65–93%) was at least an order of magnitude greater than that among separate meadows (0.5–19%). Patterns of spatial variation in the plant characteristics examined did not reflect those of flowering. It may be hypothesised that internal rhythms of populations, in conjunction with changes in environmental conditions, which are consistent over relatively large spatial (kilometres), are involved in the control of flowering induction. Environmental heterogeneity (in terms of nutrient availability) on a very local scale (metres to tens of metres) and intrinsic factors might affect the distribution of reproductive shoots within P. oceanica meadows.Communicated by R. Cattaneo-Vietti, Genova     

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.     

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.     

Lack of phlorotannin induction in the kelp Laminaria hyperborea in response to grazing by two gastropod herbivores

Phlorotannins (polyphenolic compounds) are found exclusively in brown algae and are commonly regarded as an important component of algal defense against herbivores. The variation in phlorotannin concentration within and among individual seaweed specimens of the same species can be substantial, and seaweeds may respond to herbivore damage by an increased production of phlorotannins in order to prevent further herbivory. In the present study, we investigated the natural variation of phlorotannins in different parts of the kelp Laminaria hyperborea in an observational field survey. We also performed a manipulative induction experiment in which we studied the effect of natural grazing by the patellid limpet Ansates (=Helcion) pellucida and the littorinid snail Lacuna vincta, as well as artificial damage on the production of phlorotannins in L. hyperborea. There was a large variation in phlorotannin content both among and within L. hyperborea individuals and populations. The control plants in the induction experiment had a higher phlorotannin content compared to plants in natural populations, but grazing by A. pellucida or L. vincta, or artificial damage did not induce a higher phlorotannin concentration in the kelps. Instead, the phlorotannin content in wounded kelps was significantly lower than in the control plants, possibly due to removal of phlorotannin-rich outer layers of the kelp thallus, or to an induced compensatory growth response in the L. hyperborea plants. These results imply that L. hyperborea phlorotannins probably do not function as an inducible chemical defense against A. pellucida and L. vincta, instead grazing by these herbivores decreases the phlorotannin content of the kelps.     

Floating kelps in Patagonian Fjords: an important vehicle for rafting invertebrates and its relevance for biogeography

Floating macroalgae are common dispersal vehicles for associated benthic invertebrates. In order to investigate the importance of kelp rafts for species dispersal in the Patagonian Fjord Region (PFR), the abundance and distribution pattern of floating kelps (Macrocystis pyrifera, Durvillaea antarctica) and of the invertebrate fauna associated with M. pyrifera were evaluated during austral spring of 2002–2005, 2008 and 2010. In the southernmost Magellan Region (MR), benthic M. pyrifera were additionally sampled to compare the community structures in both conditions. Floating kelps were abundant throughout the entire PFR, harbouring a diverse and abundant invertebrate fauna. The density of floating kelps increased towards the south. In the MR, a loss of species was observed between benthic and floating condition (e.g. decapods, echinoderms, several peracarid species), but a high diversity of organisms from all major phyla were observed on rafts. Throughout the PFR, the predominant rafting species belonged to the peracarids, molluscs and annelids, but the community composition differed between floating samples from the northern and southern zones of the PFR. Relative abundances of peracarids were higher in northern zones, whereas molluscs and annelids dominated in the southern areas. Species of the peracarid genera Peramphithoe, Gondogeneia, Bircenna and Limnoria were shared between all areas. The results suggest that kelp rafts not only contribute to local population connectivity in the PFR, but could also be an important dispersal vehicle for rafting species along the PFR in north–south direction, crossing the biogeographic boundary around Taitao Peninsula. Furthermore, the MR appears to be an important stepping stone for species dispersal via kelp rafting in the subantarctic region.     

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     

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.     

Feeding responses of the bivalves Crassostrea gigas and Mytilus trossulus to chemical composition of fresh and aged kelp detritus

The chemical composition of kelps (e.g. polyphenolics) deters grazing by herbivores, but kelp detritus is potentially a source of nutrition for suspension feeders. The effects of kelp detritus derived from two species [Agarum fimbriatum Harvey and Costaria costata (Turner) Saunders] on feeding of oysters, Crassostrea gigas Thunberg, and mussels, Mytilus trossulus Gould, were examined in feeding experiments. Fresh and aged kelp particles were sequentially presented in combination with the microalga Rhodomonas lens at an initial total concentration of 5᎒^-4 ml^-1. Aging of kelp particles for 4 days in seawater significantly reduced the concentration of polyphenolics without changing the total carbon or nitrogen content. Clearance rates of both mussels and oysters were significantly lower in the presence of fresh versus aged kelp particles, and clearance rates declined overall with declining polyphenolic concentrations. Video endoscopy was used to examine feeding selectivity at the level of the gill in oysters in the same food treatments used in the clearance rate experiments. Comparison of particle composition in the water versus the pseudofeces in both oysters and mussels was also used as a measure of feeding selectivity. When presented with R. lens in combination with fresh and aged kelp particles selectivity for R. lens tended to be greater against fresh than aged particles, and there was some indication that this was stronger for A. fimbriatum than for C. costata particles. The ability to select was lower at very high polyphenolic concentrations, which may reflect poisoning of sensory binding sites. These data suggest that bivalves distinguish among particles of varying chemical composition and respond by changing their clearance rates and their selectivity.     

Interactions between an invasive and a native bryozoan (<Emphasis Type="Italic">Membranipora membranacea</Emphasis> and <Emphasis Type="Italic">Electra pilosa</Emphasis>) species on kelp and <Emphasis Type="Italic">Fucus</Emphasis> substrates in Nova Scotia,Canada

Most research on biological invasions to date has focused on the population dynamics of very successful and disruptive introduced species; however, additional knowledge of the biology of the native species is essential for understanding interactions between the two and may reveal factors that limit invasion success. The invasive bryozoan Membranipora membranacea interacts with native Electra pilosa on two substrates in northwest Atlantic subtidal habitats: highly dynamic and fast-growing kelps; and smaller, more stable, and slow-growing fucoid algae. We quantified the relative abundance and evaluated encounter outcomes in different seasons of these two bryozoans on kelp and Fucus at four sites in Nova Scotia. We also examined the effects of substrate (kelp, Fucus), temperature (7, 10, 13°C), and food (limited, unlimited) on growth rates of E. pilosa in laboratory experiments and using field manipulations. We compared our findings on factors affecting the growth of E. pilosa to those on M. membranacea obtained in similar and thus directly comparable experiments from a previous study. The proportional abundance of M. membranacea was greater than that of E. pilosa on kelps, but the opposite was observed on Fucus. Competitive standoffs between the two bryozoans were more frequent than expected, with no differences recorded between substrates; most encounters were won by M. membranacea. Growth of E. pilosa was faster on Fucus than kelp, decreased with increasing temperature only on Fucus, and was not affected by food. Growth rate of E. pilosa in all treatments was slower than that previously measured for M. membranacea. Faster growth and strong overgrowth abilities likely interact on kelps to ensure success of the invasive bryozoan. Success can be limited by low space availability, which in turn restricts growth rate, and consequently, colony size, such as on fucoid substrates. The incorporation of alternative contexts into invasion research can reveal factors involved in the resilience of native communities.     

Long-distance transport of 14C-labelled assimilates in the Fucales: directionality,pathway and velocity

A directed translocation of ¹⁴C-labelled photoassimilates was shown to occur in the thalli of six species of brown seaweeds belonging to the Fucales. The apical growing regions of these algae act as strong sinks in the source-tosink relationship of the long-distance transport. Physical interruptions of the thalli showed that the midrib is the main pathway of the assimilates in Fucus vesiculosus and F. serratus. ¹⁴C-histoautoradiography has shown that translocation takes place through the filamentous cells of the medulla. The translocation velocity of the labelled assimilates, calculated from time source profiles of the tracer, was estimated to be 2 to 4 cm · h^-1. It is suggested that the sieve elements of the Fucales act in a way similar to the conducting sieve elements of the Laminariales.     

The structure of subtidal algal and invertebrate assemblages at the Chatham Islands,New Zealand

We examined the distribution and abundance of organisms on subtidal rocky reefs at nine sites around the Chatham Islands, a remote group 780 km east of southern New Zealand. We sampled five depth strata ranging from 1 to<16 m to identify spatial patterns in the abundance of algae and invertebrates and to assess their variation within and among sites. This information is used to discuss hypotheses concerning community structure at this remote locality. Several patterns were apparent. The immediate subtidal was occupied by the southern bull kelp Durvillaea spp. A suite of 11 fucalean species were dominant to a depth of 10 m with an average abundance of 28 m^-2, while one species, Carpophyllum flexuosum, occurred mostly in deeper water. Only two laminarian species of algae were present at the islands. The indigenous Lessonia tholiformis was abundant at 2.5 to 15 m and was not found in deeper water, while the giant kelp Macrocystis pyrifera was abundant at two sites in 12 to 18 m. The commercially valuable abalone Haliotis iris was extremely abundant in shallow water, with an overall mean of 6 m^-2 at 5 m. The sea urchin Evechinus chloroticus was common, but reached high densities only in small (<25 m²) patches. The characteristic urchin-dominated zones reported in kelp beds world-wide were not seen. There was considerable site-to-site variation in the occurrence and abundance of individual species. Some differences between sites were associated with shelter from swell (e.g. M. pyrifera was found only in sheltered sites) and physical habitat (e.g. juvenile H. iris were found only beneath boulders inshore), but much of the variation could not be explained by physical or depth-related factors alone. We hypothesize that the differences in these kelp bed assemblages compared to mainland New Zealand are partially due to the high degree of endemism at the Chatham Islands. Local variation cannot be explained by herbivory, and is most likely the result of the various life-history characteristics of the major habitat-forming species, the large brown algae.     

Matching spatial distributions of the sea star Echinaster sepositus and crustose coralline algae in shallow rocky Mediterranean communities

Understanding why a species is present in a particular location and the consequences of its presence is complex but necessary to identify the mechanisms that generate and maintain ecological diversity. The common sea star Echinaster sepositus can be either very abundant or non-existing in nearby localities of the western Mediterranean. Yet, the factors that shape its distribution and the impact of the sea star on natural communities remain uninvestigated. Here, we quantified multiple biotic and abiotic factors that may affect the distribution of E. sepositus and tested whether this sea star can shape the organization of the community it inhabits. Our results showed that the distribution of this sea star was highly contagious and positively correlated with the abundance and distribution of crustose coralline algae from tens of meters to tens of kilometers. Despite significant differences in community composition between localities with high or low abundance of the sea star, experimental addition of E. sepositus to natural communities failed to shift the composition of the algal community in 4 months. Overall, our results suggest that within habitat variability in the abundance of crustose coralline algae may explain the abundance of E. sepositus at multiple geographic scales, emphasizing the need to investigate small-scale processes at larger geographic scales.     

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.     

Interhemispheric comparison of recruitment to intertidal communities: pattern persistence and scales of variation

Recruitment variation can be a major source of fluctuation in populations and communities, making it difficult to generalize results. Determining the scales of variation and whether spatial patterns in the supply of individuals are persistent over time can provide insight into spatial generality and the application of conservation and metacommunity models. We examined these issues using eight-year-long data sets of monthly recruitment of intertidal mussels (Mytilus spp., Perumytilus purpuratus, Semimytilus algosus, Brachidontes granulata) and barnacles (Balanus glandula, Chthamalus dalli, Jehlius cirratus, Notochthamalus scabrosus) at sites spanning > 900 km along the coasts of Oregon-northern California (OR-NCA, 45.47-39.43 degrees N) and central Chile (CC, 29.5-34.65 degrees S). We evaluated four general "null" hypotheses: that despite different phylogenies and great spatial separation of these taxa, their similar life history strategies and environmental settings lead to similar patterns of recruitment (1) between hemispheres, (2) in time, (3) in space, and (4) at larger and smaller spatial scales. Hypothesis 1 was rejected: along the OR-NCA coast, rates of recruitment were between two and three orders of magnitude higher, and patterns of seasonality were generally stronger and more coherent across space and time than along CC. Surprisingly, however, further analysis revealed regularities in both time and space for all species, supporting hypotheses 2 and 3. Temporal decorrelation scales were 1-3 months, and characteristic spatial scales of recruitment were approximately 250 km. Contrary to hypothesis 4, for the ecologically dominant species in both hemispheres, recruitment was remarkably persistent at larger mesoscales (kilometers) but was highly stochastic at smaller microscales (meters). Across species, increased recruitment variation at large scales was positively associated with increased persistence. Our results have several implications. Although the two regions span distinct latitudinal ranges, potential forcing processes behind these patterns include similar large-scale climates and topographically locked hydrographic features, such as upwelling. Further, spatial persistence of the recruitment patterns of most species at the mesoscale supports the view that marine protected areas can be powerful conservation and management tools. Finally, persistent and yet contrasting spatial patterns of recruitment among competing species suggest that recent metacommunity models might provide useful representations of the mechanisms involved in species coexistence.     

Differential meiospore size and tolerance of ultraviolet light stress within and among kelp species along a depth gradient

 Kelp are differentially stratified along a gradient of UV exposure (as a function of water depth). The role of ultraviolet light in seaweed zonation has not been fully explored. This study found a significant meiospore size difference within and among the kelp species examined: Pterygophora californica Ruprecht (high to mid-subtidal), Macrocystis integrifolia Bory (high to mid-subtidal), Laminaria groenlandica Rosenvinge (high-subtidal), Alaria marginata Postels and Ruprecht (low-intertidal) and Hedophyllum sessile Setchell (mid-intertidal). This size difference was correlated with the depth distribution of adult plants, with the largest meiospores originating from shallow-dwelling adult kelp exposed to high UV light. Under UV stress in the laboratory, meiospores from adults growing in high-UV environments displayed greater germination and survival rates than the progeny of adult kelp occupying lower-UV environments. This suggested that in Barkley Sound, British Columbia, Canada, differential tolerance to UV (possibly determined by meiospore size) may limit the upper settlement position of kelp species and individuals. Tolerance to UV may be an important determinant of kelp zonation on rocky shores. Received: 16 June 1998 / Accepted: 6 January 2000