Biomass, diversity and production of rocky shore macroalgae at two nutrient enrichment and wave action levels

The littoral zone of temperate rocky shores is normally dominated by perennial macroalgae (e.g. Fucus, Ascophyllum, Laminaria), but nutrient enrichment and/or permanently decreased wave action may lead to structural community changes from dominance of perennials to increased amounts of annual opportunistic species (mainly green algae). Macroalgal biomass, diversity and production as well as relationships between the two latter were studied using Solbergstrand’s rocky shore mesocosms in SE Norway in connection with a long-term experimental manipulation of nutrient addition and wave action (high and low levels of both factors applied in a crossed way to eight outdoor basins). After more than 2 years of experimental treatment, the total standing stock of macroalgae was larger in low nutrient than in high nutrient treatments as well as in high wave compared to low wave treatments (in autumn only). For macroalgal functional groups, bushy and filamentous brown and filamentous red algae were generally favoured by low nutrient concentrations, while annual filamentous and sheet-like green algae were stimulated by the nutrient enrichment. There was only one significant interaction between nutrient enrichment and wave action (for brown filamentous algae in autumn) and also only one significant main effect of the wave treatment (for bushy brown algae in autumn). Surprisingly, the high nutrient treatments supported a higher diversity of macroalgae, whereas the low nutrient treatments generally showed higher production rates. Moreover, significantly negative correlations were found between macroalgal diversity and primary productivity in both summer and autumn. This study shows that it is the biological components of the communities subjected to external forcing (nutrient addition or decreased wave action) that regulate production and this contradicts the common misperception that resource production in natural systems simply can be fast-forwarded by fertilization. The negative relationships between diversity and productivity, although a consequence of unexpected results for diversity and production, are also novel and hint towards species identities having more important functional consequences than general species dominance patterns and the amount of species per se. These results also emphasise the context dependency of findings within the field of biodiversity and ecosystem functioning.     

Effects of detrital non-native and native macroalgae on the nitrogen and carbon cycling in intertidal sediments

The decay of non-native and native seaweed mixing may modify sediment biogeochemistry and organic matter transfers within benthic food webs according to their composition and biomass. The non-native species Sargassum muticum was deliberately added to the sediment of an intertidal sandflat at different biomass and mixed to the native species Ulva sp. and Fucus vesiculosus. The sediment porewater was then ¹³C and ¹⁵N enriched to test whether both detrital diversity and biomass influenced the transfer of porewater carbon and nitrogen to the sediment and to the macrofauna consumers. More ¹⁵N-nitrogen was mobilized to sediments and macrofauna when the 3-species detrital mixing was buried, probably because this mixing provided species-specific compounds such as polyphenols due to the presence of S. muticum and F. vesiculosus, as well as large amounts of nitrogen due to the presence of Ulva. Our study revealed the importance of detrital diversity and non-native seaweeds for the nitrogen cycling in the benthic food web.     

Estimating variation in surface emissivities of intertidal macroalgae using an infrared thermometer and the effects on temperature measurements

Accurate measurements of surface temperatures with an infrared (IR) thermometer require input of the emissivities of the surfaces being measured; however, few determinations of the emissivities of intertidal organisms’ surfaces have been made. Emissivities of intertidal macroalgae were measured to determine whether algal species, measurement angle, hydration, and layering affected them. Emissivities were similar and averaged 0.94 among 11 of 13 species. The species with lower and more variable emissivities (Chondracanthus exasperatus and Desmarestia viridis) differed in morphology from the other species, which were relatively flat thin blades with little surface texture. Measurement angle caused emissivities to decrease significantly in Mazzaella splendens but not in three other species. Hydration and layering of Ulva lactuca also had no effect. At 22 °C, measured temperatures were within 1 °C of actual temperatures when thermometer emissivity settings ranged from 0.75 to 1.00. When emissivities were set lower than actual values, measured temperatures were lower than actual temperatures at 15 °C and higher than actual temperatures at 60 °C. When the IR thermometer was used to measure surface temperatures of nine species of intertidal algae immediately before they were inundated by the incoming tide, temperatures were higher in mid intertidal than low intertidal individuals and higher on a sunnier day than an overcast day. Temperatures of U. lactuca increased with increasing height on the shore, but temperatures of Ulvaria obscura did not. Temperatures were also higher in Fucus distichus blades than receptacles, and lower in U. lactuca and M. splendens occurring in the lower layers of stacks of algae.     

Assimilation efficiency of a temperate-zone intertidal fish (Cebidichthys violaceus) fed diets of macroalgae

We studied assimilation efficiencies of the temperate-zone intertidal fish Cebidichthys violaceus (Girard, 1854) fed in the laboratory on each of the following species of macroalgae: Spongomorpha coalita (Chlorophyta), Ulva lobata (Chlorophyta), Iridaea flaccida (Rhodophyta) and Porphyra perforata (Rhodophyta). Together, these 4 algae make up over 75% of the natural summer diet of C. violaceus. Assimilation efficiency was calculated by proximate organic analysis of food and feces; the amount of ash in food and feces was used as a standard. Depending on the algal species, the fish assimilated 43 to 81% of the protein, 21 to 44% of the lipid, 45 to 62% of the carbohydrate and 31 to 52% of all three classes of organic material combined. These data are the first results showing that a temperate-zone marine fish can assimilate macroalgal constituents. Protein, carbohydrate and total organic material were absorbed more efficiently from rhodophytes than from chlorophytes. Conversely, lipid was absorbed more efficiently from chlorophytes than from rhodophytes. These results are compared with previous work showing that C. violaceus in nature eats more chlorophytes than rhodophytes, but in laboratory preference tests prefers rhodophytes to chlorophytes.     

Effects of wave-induced lightflecks in the intertidal zone on photosynthesis in the macroalgae Postelsia palmaeformis and Hedophyllum sessile (Phaeophyceae)

Intertidal algae are exposed to a highly variable photic regime because of crashing waves. We measured photosynthetic rate of whole fronds of the seaweeds Postelsia palmaeformis (Ruprecht) (at Bodega Marine Laboratory, 1991) and Hedophyllum sessile (Setchell) (Phaeophyceae) (at Friday Harbor Laboratories, 1990) in flashing and steady-state photosynthetically active radiation (PAR) of equal irradiance, using a recirculating metabolism chamber designed to minimize the thickness of diffusional boundary layers on the surfaces of algal tissues. The dimensionless ratio of photosynthetic rate under flecking PAR (P_fleck) and the sum of photosynthetic rate under steady state PAR (P_s) of high and low irradiance was computed for lightfleck periods from 0.2 to 100 s. P_fleck:P_s is a measure of the light-flash utilization-efficiency in flashing light, and was greater than unity at periods between lighflecks of 2 to 30 s, with a peak at 10 s. We used a novel optical fiber irradiance meter to measure PAR incident on fronds of P. palmaeformis as they were washed about by waves breaking in the intertidal zone, and compared the light records to that obtained by a stationary sensor under the canopy. PAR flashing period was closely correlated with the period of breaking waves in stands of P. palmaeformis. We estimated the seasonal spectrum of period between light flashes in stands of this species by examining the spectral density of ocean waves at Bodega Marine Laboratory, Bodega Bay, California. The wave spectrum peaks at a period (10 s) where light-flashes may theoretically enhance the light-flash utilization-efficiency of seaweeds. We calculate that the enhanced light-flash utilization-efficiency wrought by wave-induced light-flashes may contribute to significant gains in primary productivity of these macroalgae under some conditions.     

Effect of freezing on photosynthesis of intertidal macroalgae: relative tolerance of Chondrus crispus and Mastocarpus stellatus (Rhodophyta)

The effect of freezing on photosynthetic metabolism was studied in the red algae, Chondrus crispus and Mastocarpus stellatus. Plants of both species were collected from the intertidal at Chamberlain or Kresge Point, Maine, USA (43°56N, 69°54W) between February and March 1987. Photosynthetic rates were measured immediately after freezing at-20°C and following recovery periods in seawater. Photosynthesis in C. crispus declined rapidly following freezing, falling to 70% of control values within 1 h and 30% after 3 h exposure. Minimum photosynthetic rates (7 to 9% of controls) occurred following freezing exposures of 12 h or more. Full photosynthetic recovery in C. crispus after 3 h at-20°C required 48 h. Photosynthesis in C. crispus did not fully recover in plants frozen for 6 h or more. In contrast, photosynthesis in M. stellatus was relatively unaffected by freezing exposures of <12 h. Twelve hours or more at-20°C reduced photosynthesis to 55% of controls. Photosynthesis in M. stellatus fully recovered from 24 h at-20°C within 24 h. In both species the reduction of photosynthesis by freezing was associated with damage to the plasma membrane and reduced efficiency of energy transfer from phycobilisomes to chlorophyll a, but did not appear to involve ribulose-1,5-bisphosphate carboxylase oxygenase activity. The freezing tolerance of C. crispus and M. stellatus positively correlates with their respective intertidal distributions, suggesting that freezing may be involved in controlling the distributions of these species on the shore.     

Fractal properties of spatial distribution of intertidal benthic communities

We studied the spatial distribution of intertidal macrozoobenthos, microphytobenthos (diatom algae) and sediments at scales from decimeters to kilometers using an index of spatial homogeneity, D _I. Sediments were found to be randomly distributed, making up a mosaic of silty and sandy sites. On the contrary, the estimated spatial variability of macrofauna within all the scales up to 5500 m depended upon neither extent (total area covered) nor grain (finest spatial resolution) but only their ratio. We treat this as evidence of statistical self-similarity (fractal property) of the pattern. For diatoms, spatial heterogeneity of community structure was also self-similar in the range from 0.25 to 75 m (within a single bight). At larger scales, microalgae showed a combination of patchy structure with pronounced gradient along the shoreline from brackish-water to marine flora. Thus, fractal properties of both groups became manifested at scales corresponding to their mean body size. The ranges of fractal patterns were approximately equal to 10³–10⁵ if measured in body size units. We suggest that fractal-like spatial structures may be a general feature of communities, and speculate on the nature of such patterns. Received: 16 February 1999 / Accepted: 30 December 1999     

The distribution of intertidal phytopsammon in an Oregon estuary

The sediments of two tidal flats in Yaquina Bay, Oregon, USA, were studied to determine the distribution and abundance of the interstitial microalgal communities. The hydrography of the bay, as well as fluetuations in various physical and chemical parameters appear to regulate the biomass and the vertical and intertidal distribution of these organisms.     

Impact of sewage on the distribution,abundance and community structure of rocky intertidal macro-organisms

The biological effects of a low-volume domestic sewage discharge were studied near Wilson Cove, San Clemente Island, California (USA), from February to June, 1972. There were fewer species and less cover near the outfall (7 macro-invertebrates, 17.6%; 13 macrophytes, 91.7%) than in nearby “unpolluted” control areas (9 macroinvertebrates, 9.2%; 30 macrophytes, 103.4%). The outfall biota was less diverse than that of the controls, as shown by 5 different diversity indices. A great reduction in community stratification (spatial heterogeneity) and, hence, community complexity occurred near the outfall; this reduction in stratification was primarilly due to the absence of Egregia laevigata, Halidrys dioica, Sargassum agardhianum and Phyllospadix torreyi. These were replaced in the mid-intertidal near the outfall by a low turf of blue-green algae, Ulva californica, Gelidium pusillum and small Pterocladia capillacea, and in the lower intertidal by Serpulorbis squamigerus covered with Corallina officinalis Var. chilensis. A statistically-based determination of assemblages or groups of organisms (i.e., cluster analysis) revealed 3 discrete outfall and 3 discrete control area groups; 3 assemblages contained samples from both areas. The distributional patterns of these groups indicate that near the outfall the degree of dilution of discharged sewage is more important in regulating zonation than is tidal height. The enhancement of the suspension feeder Serpulorbis squamigerus and the omnivores Ligia occidentalis, Pachygrapsus crassipes and Anthopleura elegantissima in the outer fringe of the outfall plume hypothetically is due to their ability to utilize sewage as a food source. A critical effect of the outfall may be to decrease environmental stability thereby favoring rapid-colonizers and more sewage-tolerant organisms. The outfall macrophytes were characterized by relatively higher net primary productivities, smaller growth forms, simpler and shorter life histories, and most were components of early successional stages.     

Vertical stratification of mobile epiphytal arthropod assemblages between the canopy and understorey of subtidal macroalgae

This study examined patterns of faunal stratification between vertical strata by partitioning subtidal macroalgal communities into vertically contiguous (and hence paired) canopy and understorey habitats. Such floristic strata are often clearly defined in temperate subtidal communities and have been the focus of a range of other, primarily botanical, studies. Previous studies that have touched upon this subject have used statistical tests that require spatially independent samples, yet have collected samples that are often spatially contiguous and therefore not statistically independent. Here an analytical approach for examining faunal stratification in subtidal macroalgal assemblages using split-plot ANOVA for univariate data and a novel multivariate test suitable for analysing assemblage-level paired-sampled data is proposed. Differences in faunal assemblage structure between habitats were primarily attributed to the distribution of two functional feeding groups: filter-feeding and scavenging arthropods. Filter-feeders were more abundant amongst the canopy, whereas scavengers were more abundant in the understorey. No pattern was detected for herbivorous arthropods. These patterns are consistent with the feeding ecology of these functional groups and reflect expected differences in the relative availability of trophic resources between habitats, commensurate with the physical structure of the environments. Contrary to expectation, understorey algal habitats supported faunal assemblages that were as diverse, or more diverse than the canopy habitats, despite comprising smaller habitable spaces (in terms of algal biomass) and generally supporting fewer faunal individuals. Trophic stratification of epifaunal assemblages is suggested as a mechanism whereby faunal diversity is enhanced in these environments.     

Capability of dynamic photoinhibition in Arctic macroalgae is related to their depth distribution

The capability of several macroalgal species to protect photosynthesis from excessive irradiance by dynamic photoinhibition was investigated relative to their depth distribution in summer 1995 in the Kongsfjord (79°N; 12°E, Ny Ålesund, Spitsbergen, Norway). Photoinhibition of photosynthesis was induced by exposure of algae from different water depths to a high photon fluence rate of 500?μmol?m^?2?s^?1 for 2?h. Changes in optimal quantum yield (F _v/F _m) were measured during the inhibition phase. Recovery of photosynthesis was subsequently induced by dim white light (10?μmol?m^?2?s^?1) and observed as changes in the variable fluorescence. With a newly developed mathematical model different parameters of the response kinetics of inhibition and recovery were calculated and related to the depth distribution of each algal species. It is shown that two components with slow and fast reaction kinetics, respectively, are involved in photoinhibition and recovery of photosynthesis. Their possible molecular bases are discussed. The half-life time (τ) of the inhibition and recovery phases, i.e. the time necessary to reach half maximal response, is clearly related to the depth distribution of the investigated species. Algae collected close to the water surface show a fast reaction of both photoinhibition and recovery and, hence, have a low τ. With increasing depth the reactions become slower and τ increases. τ was highest in deep water algae. Further analysis of the reaction kinetics in Laminaria saccharina shows that the relative proportion of the two kinetics involved change with the collection depth. In contrast, a significant difference in the reaction rates of both kinetics was not observed.     

Hydrology and grazing jointly control a large-river food web

Inputs of fresh water and grazing both can control aquatic food webs, but little is known about the relative strengths of and interactions between these controls. We use long-term data on the food web of the freshwater Hudson River estuary to investigate the importance of, and interactions between, inputs of fresh water and grazing by the invasive zebra mussel (Dreissena polymorpha). Both freshwater inputs and zebra mussel grazing have strong, pervasive effects on the Hudson River food web. High flow tended to reduce population size in most parts of the food web. High grazing also reduced populations in the planktonic food web, but increased populations in the littoral food web, probably as a result of increases in water clarity. The influences of flow and zebra mussel grazing were roughly equal (i.e., within a factor of 2) for many variables over the period of our study. Zebra mussel grazing made phytoplankton less sensitive to freshwater inputs, but water clarity and the littoral food web more sensitive to freshwater inputs, showing that interactions between these two controlling factors can be strong and varied.     

Lateral or along-shore distribution of the polychaetous annelids of an intertidal,sandy habitat

The lateral or along-shore distribution of the polychaetous annelids of a subtropical, sandy, intertidal habitat in Florida (USA) was studied on three transects (covering 2.5 km of shoreline), sampled quarterly for 2 years. Although the average species number and density of individuals were not significantly different at specific tidal levels, the species composition was highly variable over time. Composition of density dominants was identical along-shore, and down-shore density distribution of selected species was also identical. Analysis by entire transects rather than by individual stations decreased along-shore heterogeneity by 10 to 15%.     

Relationships between intertidal zonation and circatidal rhythmicity in littoral gastropods

The presence and phase of circatidal rhythmicity was correlated with vertical zonation and other ecological factors. Ten species were studied in the field and in aktographs under controlled conditions in the laboratory. Retina plicata, Melanerita atramentosa, Bembicium nanum, Austrocochlea obtusa and Morula marginalba occupied mid- or upper-littoral zonations, and were subjected to regular tidal influence. They were active at high water and for a period after the ebb, possibly as these were the times of least desiccation. Each possessed a circatidal activity rhythm under non-tidal conditions. Midlittoral Amphinerita polita and lower-littoral Theliostyla albicilla were active at low water, possibly because their prefered habitats reduced desiccation and other selective forces, for example wave action and predation, determined the phase of the rhythmicity. Supra- and upper-littoral Nodilittorina pyramidalis and Melarapha unifasciata and lower- and infralittoral specimens of Bembicium auratum displayed no overt circatidal rhythmicity, possibly because they were not subjected to regular tidal action.     

潮间带土壤盐度与电导率的关系

以5：1的水土质量比浸提潮间带土壤，质量法测定的土壤盐度(质量分数)，与电导法测定的稀释了5倍的土壤浸提液(5：1水土质量比)的电导率值之间有极显著的相关性。质量法实测的土壤盐度与电导率计算的土壤盐度之间的相对误差多在5％以下。因此，用电导法测定潮间带土壤盐度不失为一种简便、快速的方法，尤其是样品数较多时更可节约时间。     

Control of the intertidal distribution of Gastroclonium coulteri in monterey bay,California, USA

Transplantation and selective removal experiments were carried out on Gastroclonium coulteri (Harvey) Kylin at Mussel Point, California, USA, between June, 1975 and May, 1979. Transplanted G. coulteri survived higher in the intetidal than the natural population, except during March and April when extreme low tides, and resultant severe desiccation, occurred during the daytime. Transplants survived, but did not grow, in low intertidal tidepools. Transplants to the shallow subtidal survived and increased in overall length. Transplants to the deeper subtidal zone died within 1 month. Transplants from the subtidal to the intertidal survived, but decreased in size. At the upper boundary of G. coulteri, removal of Gigartina papillata and Rhodoglossum affine resulted in a decrease in the percent cover of Gastroclonium coulteri. Presence of the other algae thus had a beneficial rather than competitive effect on G. coulteri. Removal of G. coulteri resulted in an increase in the percent cover of G. papillata and R. affine, demonstrating that the lower boundary of these two species is set by competition with G. coulteri. At the lower boundary of Gastroclonium coulteri, removal of Phyllospadix spp. or Prionitis lanceolata led to an increase in percent cover of G. coulteri. Removal of G. coulteri did not lead to an increase in percent cover of Phyllospadix spp. Thus, the lower boundary of G. coulteri is set by competition with Phyllospadix spp. or Prionitis lanceolata, but the upper boundary of Phyllospadix spp. is not set by competition with G. coulteri. The results support the hypothesis that upper boundaries tend to be set by physical factors (here, exposure at low tide), and lower boundaries tend to be set by biological factors (here, competition).     

Changes in diatom patch-size distribution and degradation in a spatially self-organized intertidal mudflat ecosystem

Self-organized spatial patterns have been proposed as possible indicators for regime shifts in ecosystems. Until now, this hypothesis has only been tested in drylands. Here, we focus on intertidal mudflats where regular spatial patterns develop in early spring from the interaction between diatom growth and sedimentation but disappear when benthic herbivore abundance increases in early summer, accompanied by a dramatic shift to a bare mudflat. We followed the patch-size distributions of diatom biofilms during this degradation process. As time progressed, we found a temporal change in the spatial configuration occurring simultaneously with the loss of the diatom-sediment feedback. This indicates a gradual failure in time of the self-organization process that underlies regular patterning in this ecosystem. The path to degradation co-occurred with the loss of the larger patches in the ecosystem, which resulted in a decrease of the truncation in the patch-size distribution. Hence, our study in mudflat ecosystems confirms the general hypothesis that spatial patterns can provide important clues about the level of degradation. Nevertheless, our study highlights the need for thorough study about the type of spatial patterns and the nature of the underlying feedbacks before a reliable assessment of ecosystem status can be made, as changes in patch-size distribution differed markedly with those observed in other ecosystems.     

Interactions between two introduced species: Zostera japonica (dwarf eelgrass) facilitates itself and reduces condition of Ruditapes philippinarum (Manila clam) on intertidal flats

Dwarf eelgrass (duckgrass; Zostera japonica) and Manila clams (Ruditapes philippinarum) are two introduced species that co-occur on intertidal flats of the northeast Pacific. Through factorial manipulation of clam (0, 62.5, 125 clams m⁻²) and eelgrass density (present, removed by hand, harrowed), we examined intra- and interspecific effects on performance, as well as modification of the physical environment. The presence of eelgrass reduced water flow by up to 40% and was also observed to retain water at low tide, which may ameliorate desiccation and explain why eelgrass grew faster in the presence of conspecifics (positive feedback). Although shell growth of small (20–50 mm) clams was not consistently affected by either treatment in this 2-month experiment, clam condition improved when eelgrass was removed. Reciprocally, clams at aquaculture densities had no effect on eelgrass growth, clam growth and condition, or porewater nutrients. Overall, only Z. japonica demonstrated strong population-level interactions. Interspecific results support an emerging paradigm that invasive marine ecosystem engineers often negatively affect infauna. Positive feedbacks for Z. japonica may characterize its intraspecific effects particularly at the stressful intertidal elevation of this study (+1 m above mean lower low water).