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.     

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.     

The seasonal selection by temperature of heterotrophic bacteria in an intertidal sediment

The seasonal selection by temperature of bacteria in an intertidal sediment was investigated, and a simplified method of demonstrating the temperature adaption of a mixed heterotrophic bacterial population was suggested. The method relied upon counting the bacteria which grew at only two separate incubation-temperatures, and compared favourably with more tedions methods which utilise replicated cultures grown at a large number of incubation temperatures. Using this technique, a temperature adaptation index was calculated for the heterotrophic bacterial population and changes in the value of this index were shown to be correlated with seasonal changes of environmental temperature.     

Short- and long-term effects of eutrophication on the secondary production of an intertidal macrobenthic community

Secondary production of a macrobenthic community at an intertidal mudflat was estimated for 33 successive months. Sampling was carried out along a eutrophication gradient, including non-eutrophied Zostera meadows, an intermediate muddy area, and a strongly eutrophied sand-muddy flat, where macroalgal blooms of Enteromorpha spp. usually occur. The Zostera meadows were always the most productive habitat (145–225 g ash-free dry weight m^–2 year^–1). In the short term, the macroalgal bloom benefited the total estuarine production by enhancing the annual production in the eutrophied area. Nevertheless, our results show that this increase was short lived and in no way sufficient to match the production in the Zostera meadows. In the long term, the present study provides evidence that the disappearance of macrophyte beds, as a result of ongoing eutrophication, constitutes a major threat to the sustainability of the estuarine ecosystem.Communicated by S.A. Poulet, Roscoff     

Coefficient of variation of sea surface temperature (SST) as an indicator of coral bleaching

Coral bleaching has become a major problem on reefs around the world in recent decades. It is believed that mean temperature alone is the primary force driving this ecological phenomenon. We propose that variance in temperature in the short term is just as important as the mean. Thirty years of daily sea surface temperature (SST) data have been collected by the University of Puerto Rico at Mayaguez Marine Laboratory in La Parguera, PR. These data were collated and analyzed initially (by Amos Winter) for their relationship to coral bleaching in this area. We found that the data fell into three categories: high mean temperatures associated with severe bleaching, cooler mean temperatures associated with no bleaching, and years of high SSTs but with no coral bleaching. Here, we examined the relationship between mean temperature during those months in which bleaching occurred, temperature variance (as measured by standard deviation), and coefficient of variation (CV; i.e., SD standardized by the mean). We also derived a critical threshold temperature and level of resolution in time for calculating these statistics to clearly describe the circumstances of bleaching versus non-bleaching events, particularly at marginal bleaching temperatures. These characteristics were compared for the four warmest months of the year (July–October) for four warm bleaching years (1969, 1987, 1990, and 1995), four cool non-bleaching years (1984, 1985, 1986, and 1988), and two warm non-bleaching years (1994 and 2000). No relationship was found between the mean SST and SD in terms of predicting bleaching. The two primary statistics which, in concert, did indicate bleaching, however, were the short-term, biweekly mean temperature and its the associated CV. Bleaching occurs in association with both high temperatures and a high CV. The CV becomes a critical determinant of bleaching only when temperatures are ∼29.1–29.8°C. The warm, non-bleaching years were generally characterized by a CV of < 1.9 and a temperature range between 28.5 and 29.9°C. We conclude that increased mean SSTs alone are not sufficient to induce coral bleaching; a high variance in SST at marginal, lower bleaching temperatures can induce bleaching, and likewise, a low variance of such will not induce bleaching. This variance is most clearly described by the CV.     

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.     

Seasonal variation in ecophysiological patterns in macroalgae from an Arctic fjord. I. Sensitivity of photosynthesis to ultraviolet radiation

Marine macroalgae inhabiting Arctic coastal ecosystems are exposed to pronounced seasonal variations in the radiation regime, including harmful UVB radiation. This study presents the first data on the seasonal changes in the sensitivity of macroalgal photosynthesis towards UV exposure by comparing under-ice, clear-water and turbid-water conditions characteristic for late winter, spring and summer. Various brown (Laminaria saccharina, L. digitata, L. solidungula, Saccorhiza dermatodea, Desmarestia aculeata), red (Palmaria palmata, Devaleraea ramentacea) and one green macroalgal species (Monostroma aff. arcticum) were collected at the same water depth throughout the seasons in the Kongsfjord (Spitsbergen, Svalbard, Norway). Maximum quantum yield (F_v/F_m) and maximum photosynthetic electron transport rates (ETR_max) were determined immediately after collection, after 2 h exposure to artificial UV radiation and after 18 h recovery in dim white light. Photosynthesis of the studied species showed different responses depending on their morpho-functional and physiological characteristics, their life strategies, phenology and depth distribution. Within the genus Laminaria, maximum quantum yield of adult specimens of the deep-water species L. solidungula was most strongly UV sensitive. Adult L. saccharina exhibited a lower UV sensitivity than a 6-month-old specimen. Inhibition of photosynthesis after UV exposure remained at the same level throughout the study period, both in adult L. saccharina and S. dermatodea. However, adult specimens of L. saccharina collected in May showed partial recovery only, whereas photosynthesis of specimens from both species collected later recovered fully. D. aculeata exhibited a remarkable decrease of UV sensitivity during the study period. Photosynthesis of specimens collected under the ice was strongly inhibited by UV, but the degree of inhibition decreased during spring and summer. Concomitantly ETR_max values were low after UV exposure in specimens collected in June, but increased later in the season. P. palmata exhibited a relatively flexible response. Photosynthesis in specimens collected under the ice in June or in turbid water in July/August was relatively strongly inhibited; specimens collected during sunny periods and in clear water in spring showed a much lower degree of photoinhibition after UV exposure. The seasonal pattern of low/high ETR_max values in spring/summer is probably a characteristic of the life strategy of this species. The UV sensitivity of D. ramentacea exhibited a similar seasonal pattern. In M. aff. arcticum, UV sensitivity increased and ETR_max values decreased during the study period, reflecting the life strategy of this annual late winter/spring species. The physiological basis for the seasonal changes in UV sensitivity of photosynthesis is presented in a companion paper (this issue).     

Resource acquisition in two intertidal fucoid seaweeds, Fucus serratus and Himanthalia elongata : seasonal variation and effects of reproductive development

Seasonal variations and the effect of reproductive development on resource acquisition by two intertidal fucoid species, the iteroparous Fucus serratus L. and the semelparous Himanthalia elongata (L.) S. F. Gray were examined. The oxygen-exchange characteristics of vegetative apical tissue of both non-fertile and fertile plants and receptacle tissue were compared at monthly intervals throughout reproductive development. Respiratory rates in non-fertile F. serratus varied seasonally between 1.5 and 8.0 μmol g⁻¹ fresh wt h⁻¹; in fertile plants the receptacle had a significantly lower respiratory rate than the vegetative tissue. The respiratory rate of the vegetative button of fertile H. elongata displayed less seasonal variation and was lower than that of the receptacle, which varied from a maximum of 9.5 μmol g⁻¹ fresh wt h⁻¹ at receptacle initiation in October to a minimum of 2.0 μmol g⁻¹ fresh wt h⁻¹ in February. The maximum photosynthetic rate (P _max) of non-fertile plants of both species did not vary in a distinct seasonal manner (∼60 μmol g⁻¹ fresh wt h⁻¹ for F. serratus and ∼12 μmol g⁻¹ fresh wt h⁻¹ for H. elongata). In fertile plants, the P _max of the receptacle tissue was (∼50% lower in F. serratus, and at its peak three times higher in H. elongata, than that of vegetative tissue. The stable carbon-isotope ratio (δ¹³C) did not differ between different tissue types in F. serratus, but values did vary seasonally, being less negative in the summer than in the winter (−13.5‰ compared to −18‰). The receptacle tissue of H. elongata also displayed a distinct seasonal variation in δ¹³C values (−12‰ in summer, −16‰ in winter), whilst the δ¹³C of the vegetative button did not vary seasonally. The rate of uptake of inorganic nitrogen by the vegetative thallus was lower in H. elongata than in F. serratus. The receptacle tissue of F. serratus had lower uptake rates than the vegetative tissue, whilst the uptake rate by H. elongata receptacle tissue was higher than that of the vegetative button. Received: 14 March 1997 / Accepted: 22 April 1997     

Seasonal changes in the structure of an intertidal community

Quantitative (0.25 m²) samples of macrofaunal (>1.0 mm) invertebrates were taken in each season from one habitat of an intertidal sandbar in the North Inlet estuary near Georgetown, South Carolina, USA. During all seasons the community inhabiting the sample site was numerically dominated by two species of haustoriid amphipods (Acanthohaustorius millsi and Pseudohaustorius caroliniensis). Seasonal changes at the community level were clearly controlled by the population dynamics of the numerically dominant species, and qualitative information on life histories was important to the interpretation of analyses' results.This work was supported by the Environmental Technology Center of Martin Marietta Corporation and the Belle W. Baruch Institute for Marine Biology and Coastal Research. It is Contribution No. 138 of the Belle W. Baruch Institute for Marine Biology and Coastal research.     

Photosynthetic response to temperature and desiccation of the intertidal alga Mastocarpus papillatus

When exposed to air during low tide, intertidal macroalgae experience a terrestrial environment and often encounter extreme levels of heating and desiccation. Two aspects of photosynthesis may be influenced by this increase in temperature and decrease in water content during exposure to air: (1) the rate of aerial photosynthesis itself, and (2) the rate at which aquatic photosynthesis recovers upon immersion in water at high tide. This laboratory study examines the effect of air temperature and desiccation on photosynthesis of the intertidal macroalga Mastocarpus papillatus Kützing. Plants were collected at Hopkins Marine Station, California, USA (36°37N; 121°54W) between July and December 1990. When apical tips were exposed to 15 to 25°C air for 2 h, photosynthesis was rapidly recovered upon reimmersion in seawater. Recovery was delayed, but complete, when tissue was exposed to 30°C air, but did not occur after exposure to 35°C air. Desiccation did not influence either the rate or the ultimate level of recovery upon reimmersion. Photosynthesis in air generally decreased with increasing desication, with no net photosynthesis occurring below 25% relative water content. Net photosynthesis of hydrated thalli increased with air temperature from 15 to 30°C, then decreased at 35°C. Dark respiration of hydrated thalli increased over the entire temperature range. This study indicates that thallus heating and desiccation during periods of exposure to air can potentially influence the total carbon budget of M. papillatus.     

The Wahlund effect and the geographical scale of variation in the intertidal limpet Siphonaria sp.

Samples of Siphonaria sp. were collected between 1978 and 1982 from sites covering its known geographic range, from Kalbarri, Western Australia to Port Robe, South Australia. Geographic variation of 7 polymorphic enzymes was examined in this intertidal pulmonate limpet, and was found to be consistently small, indicating a large-scale influence of gene flow due to planktonic dispersal. Despite this large-scale uniformity, there is fine-scale genetic patchiness, which is repeated, rather than accumulated, on the larger scale. Throughout its geographic range, Siphonaria sp. shows deficits of heterozygotes for all 7 loci. The consistency among loci indicates that the causes of the deficits are populational, rather than locus-specific. A Wahlund effect, the departure from Hardy-Weinberg equilibrium due to mixing of individuals from groups with different allelic frequencies, is the simplest explanation of such deficits. The limited geographic variation of allelic frequencies, however, is grossly inadequate to produce these deficits through a Wahlund effect. Similarly, temporal variation in allelic frequencies in recruits does not explain the deficits. The largest contributor to a Wahlund effect appears to be binomial sampling variance among small local breeding groups. Thus, mixing of larvae on a scale of metres, rather than among geographical areas, apparently produces the deficits of heterozygotes.     

Estimating the biogenic enhancement factor of weathering using an inverse viability method

To determine the influence of the biosphere on weathering we use a dynamic model of the global carbon cycle. It takes into account the most important processes for the long-term evolution of the Earth. The model is solved under a slowly changing environment of increasing solar luminosity and volcanic activity and continental area. By comparing the model results for the global average temperature with data derived from δ¹⁸O values from cherts it is possible to quantify the biogenic enhancement factor of weathering. For this purpose a newly developed inverse viability method is applied, which allows for calculating the range of possible biogenic enhancement factors consistent with the data. We find that in the Precambrian the weathering was 5.4 times lower than in the Phanerozoic era. This supports the hypothesis that the Cambrian explosion was caused by a positive feedback between the spread of biosphere, increased silicate weathering, and a consequent cooling of the climate.     

Sublethal effects of chronic oil exposure on the intertidal clam Macoma balthica

When exposed to Prudhoe Bay crude oil in flowing seawater for 180 days, the small intertidal clam Macoma balthica showed behavioral, physical, physiological and biochemical changes. At a high concentration of oil in seawater (3.0 mg l^-1) burrowing rate decreased, respiration rate increased, growth was inhibited, and very high mortalities resulted. The lowest concentration of oil in seawater (0.03 mg l^-1) inhibited growth and caused reabsorption of gametes. One group of adverse oil effects which was related to sluggishness and disorientation of the clams appeared after a week of exposure to oil; another group related to a negative energy balance was not observed until 60 days. We conclude that chronic exposure of M. balthica to oil-in-seawater concentrations even as low as 0.03 mg l^-1 will, in time, lead to population decreases.Please address requests for reprints to Dr. D. G. Shaw at the Institute of Marine Science     

Energy relations of the bivalve Mercenaria mercenaria on an intertidal mudflat

Measurements of the filtration rate and oxygen consumption of the bivalve Mercenaria mercenaria (L.) have been made using flowing water systems, and then combined with growth and survivorship data to construct an energy budget for a tidal-flat population in Southampton Water. Estimated consumption (1292 kcal m^-2 year^-1) is mainly deposited as faeces and pseudofaeces (759 kcal) or excreted (160 kcal); 29% (374 kcal) is assimilated, of which 241 kcal are used for respiration, 72 kcal for flesh production and 61 kcal for gamete production. M. mercenaria contributes significantly to other trophic groups in Southampton Water; predators remove an estimated 55 kcal m^-2 year^-1, and 829 kcal pass to the scavenger/decomposer chain.     

Long-term effects of ultraviolet radiation on growth and photosynthetic performance of polar and cold-temperate macroalgae

Long-term effects of artificial ultraviolet radiation (UV) and natural solar radiation on growth and photosynthetic activity, as measured by chlorophyll fluorescence, were investigated in 13 different polar and cold-temperate macroalgal species. Isolates of five different species from the Arctic and Antarctic were exposed to different light treatments of photosynthetically active radiation (PAR), PAR+UVA and PAR+UVA+UVB. Eight different species collected on the island of Helgoland, North Sea, Germany, were studied in the laboratory and under natural solar radiation conditions. Increase in fresh weight and changes in photosynthetic performance were monitored over a period of 3-4 weeks. The sublittoral polar species, particularly the Antarctic red algal species Gymnogongrus antarcticus and G. turquetii, the Arctic cold-temperate brown alga Alaria esculenta and, very drastically, the Arctic-endemic Laminaria solidungula, exhibited strong inhibiting effects of artificial UVB radiation on growth. In the cold-temperate sublittoral growth of the red algae Phycodrys rubens and, to a lesser extend, Membranoptera alata was substantially inhibited by UV radiation. In contrast, eulittoral species, e.g. Fucus serratus, did not show any differences in growth with respect to artificial irradiation conditions, with or without UV radiation. In the laboratory, some individuals of the green alga Codium fragile exhibited strong morphological changes of the whole thallus, particularly under UVB exposure. In the experimental outdoor set up, growth of most of the algal species was already inhibited by the full solar UV waveband, but, apart from Polyides rotundus, no additional UVB effect could be detected. Changes of in vivo fluorescence were not always consistent with the measurable changes in growth rate, indicating that physiological processes leading to an inhibition of growth may act independently of changes in photosynthetic activity. For the polar species, a general correlation between the natural vertical distribution in the field and the individual sensitivity towards UV radiation was indicated, while for eulittoral species from Helgoland no clear relationship was found. The obtained results show that measuring growth is a good ecological parameter to monitor long-term effects of UV radiation on single macroalgal species and the possible resulting changes of whole algal communities in coastal ecosystems.     

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.     

Analysis of an intertidal population of the amphipod Gammarus palustris using a modified version of the egg-ratio method

An intertidal population of Gammarus palustris Bousfield was sampled over a 23-month period and analyzed by a modified form of the egg-ratio method. Total numbers and size-frequency distributions revealed small numbers in the early spring which later increased to a peak in the summer months. This was followed by a marked reduction near the end of sumer and a subsequent rise in numbers during the early fall. Finally there was a second decrease to very low numbers in the late fall and early winter. Average brood size was calculated and used as an indication of food limitation. Attempts to correlate this with per capita rate of change (r), birth rate ( \((\hat b)\) and death rate ( \((\hat d)\) ) suggested that the population increase in the early summer was food-rather than predator-limited. The late summer decline appears to be the result of predation pressure as well as food limitation. The fall peak is attributed to juvenile release from ovigerous femals escaping previous predation and to an increase in adult males resulting from a release in predation pressure. The winter decline may be due to adverse environmental conditions.