Modelling nearshore intertidal habitat use of young-of-the-year Pacific sand lance (Ammodytes hexapterus) in Barkley Sound, British Columbia, Canada

The Pacific sand lance (Ammodytes hexapterus) is an ecologically important prey species for many vertebrate marine predators in the Pacific Northwest. In this study, we examined the use of intertidal habitat by young of the year (YOY) sand lance in Barkley Sound, British Columbia. We selected 14 of the original 29 independent environmental variables for modelling based on univariate analysis for variable selection. These were then used to model sand lance presence-absence using a classification tree approach. Based on our models we found that sand lance avoided mud and intertidal eelgrass. For sites that had very little mud and no intertidal eelgrass, sand lance preferred sediment size mean ≥1,918 μm or sites with sediment size mean ≤1,918 μm but with relatively well sorted sediment (sorting values ≤2.56 SDs, used as a heterogeneity index of the substrate grain size). Adjacent subtidal characteristics were not found to be important at this scale of study. This suggests intertidal substrate characteristics and presence-absence of intertidal eelgrass are the main influences on occurrence of YOY in the intertidal. Our results support the hypothesis that sand lance are associated with particular sediment types, however intertidal sediment types used by sand lance in our study differ from known preferences of sand lance for subtidal sediment types. This difference may be due to unique habitat constraints for intertidal versus subtidal regions. Although it is unknown if the results of this study are widely applicable, the results begin to identify intertidal habitat features that are important for Pacific sand lance.     

A unique algal ridge system in the Exuma Cays, Bahamas

 An algal ridge system discovered along the Exuma Cays, Bahamas constructs bioherms to a thickness of at least 1.5 m and is associated with modern intertidal stromatolites. These algal ridges are unique because they grow in atypical environments characterized by relatively low wave energy, high rates of sedimentation and low rates of herbivory. They also are composed primarily of the branching crustose coralline alga, Neogoniolithon strictum, which heretofore was not known to form algal ridges. Lateral growth rates of crusts, vertical growth rates of branches and survivorship of transplanted N. strictum were greatest in the shallow fore reef zone of the algal ridge. The alga is also capable of surviving and growing when covered with sediment for at least 100 days. Under such conditions it transforms from a branched to an unbranched morphology. Parrotfish grazing, which is said to limit the abundance of branched corallines and algal ridges, was two orders of magnitude lower than in published accounts from other reef systems of the Caribbean and one order of magnitude less than that found on nearby coral reefs of the west Exuma Sound. Neogoniolithon strictum, a delicate and open-branched coralline, persisted for over a year without grazing damage when transplanted to a depth of 2.3 m. This algal ridge-building coralline becomes a well-indurated limestone following submarine lithification of sediment that infills the open branch framework. As a result, N. strictum ridges are comparable to the dense frameworks associated with most algal ridges. Observations of N. strictum -associated bioherms along Central America suggest this ridge system may exist elsewhere under conditions similar to those described for the Bahamas. Accepted: 23 March 1996     

Shark Bay stromatolites: Microfabrics and reinterpretation of origins

Summary Detailed analysis of microfabrics in Hamelin Pool stromatolites leads to reinterpretation of the origins of these structures. Previous studies have concluded that Shark Bay stromatolites form primarily as a result of sediment trapping and binding by microorganisms. Our results suggest that microbial precipitation of microcrystalline carbonate (micrite), as both framework and cement in these stromatolites, is also a fundamental, heretofore unrecognized, process in their formation. Microbial trapping and binding is the primary mechanism of stromatolite accretion in the intertidal zone, forming grainy, calcarenite structures. Microbial precipitation is the primary accretionary mechanism in the subtidal zone, forming muddy, micritic stromatolites. Microbial precipitation also lithifies trapped and bound sediment in the calcarenite stromatolites. Recognition of microbially precipitated micrite in Shark Bay stromatolites is important, as many ancient stromatolites are micritic.     

Long-term studies of macrozoobenthos in intertidal and shallow subtidal habitats near the island of Norderney (East Frisian coast,Germany)

Near the East Frisian island of Norderney two sites are investigated permanently in order to study long-term fluctuations of macrozoobenthos: one transect (since 1977) at the northern side in shallow subtidal waters and another one (since 1976) at the sheltered southern side in the intertidal area of the Wadden Sea. Since 1980 the investigations have been continued in the frame of COST-47, sedimentary intertidal programme (including the shallow subtidal habitats colonized by the Macoma balthica community).The results up to 1984, respectively to 1985, are presented and the changes of abundance of the dominant species are discussed. Discussed are also the influences of water temperatures and sediment disturbances caused by wave action. There is evidence that the intertidal variety of the Macoma balthica community shows a greater stability than the subtidal variety.     

Wave and sediment dynamics along a shallow subtidal sandy beach inhabited by modern stromatolites

To help define the habitat of modern marine stromatolites, wave-dominated flow and sediment transport were studied in the shallow subtidal region (1-2 m depth) along the slightly concave, windward face of Highborne Cay, Exuma, Bahamas - the only face of the cay that includes a population of stromatolites concentrated near the region of highest curvature of the beach. Wave energy impacting this island's most exposed beach was driven by local wind forcing which increases largely in response to the passage of atmospheric disturbances that typically affect the region for periods of a few days. Although some wave energy is almost always noted (maximum horizontal orbital speeds at the bottom are rarely <10 cm s(-1)), wave conditions remain comparatively calm until local winds increase above speeds of approximately 3-4 m s(-1) at which point maximum wave speeds rapidly increase to 50-80 cm s(-1). Stromatolites, which are largely restricted to the shoreward side of a shallow platform reef, are sheltered by the reef beyond which wave speeds are one to four times higher (depending on tidal stage). Moreover, stromatolite populations are predominantly found along a region of this wave-exposed beach that experiences comparatively reduced wave energy because of the curved morphology of the island's face. Maximum wave speeds are 1.4 to 2 times higher along more northern sections of the beach just beyond the locus of stromatolite populations. A quantitative model of sediment transport was developed that accurately predicted accumulation of suspended sediment in sediment traps deployed in the shallow subtidal zone along this beach. This model, coupled with in situ wave records, indicates that gross rates of suspended sediment deposition should be two to three times higher northward of the main stromatolite populations. Regions of the beach containing stromatolites nevertheless should experience significant rates of gross suspended sediment deposition averaging 7-10 g cm(-2) day(-1) ( approximately 4-6 cm day(-1)). Results suggest that one axis of the habitat of modern marine stromatolites may be defined by a comparatively narrow range of flow energy and sediment transport conditions.     

Stromatolite biostromes and associated facies in the late precambrian porsanger dolomite formation of Finnmark,Arctic Norway

The Late Precambrian Porsanger Dolomite Formation, occurring beneath the Varanger tillite in Arctic Norway, consists of various dolomitic lithofacies of subtidal, intertidal and supratidal environments. The lithofacies belong to three facies associations, A, B and C, which are repeated several times in the sequence. Facies association A comprises cryptalgal laminites, dolomicrites and thin-bedded grainstones and flakestones. The environment represented by this facies is broadly intertidal (locally supratidal) flat, with the interbedded carbonate sands representing storm deposits. Facies association B, of shallow subtidal to low intertidal origin, comprises cross-bedded carbonate sands (flakestones, grainstones and oolites) forming units up to 10 m thick. Small stromatolite bioherms (5 m wide, 2 m high) are locally developed within these “high-energy” deposits. Facies association C formed in a subtidal environment consists of laterally extensive (over 20 km) uniformly developed stromatolite biostromes, up to 16 m thick. The biostromes, locally divided by channels filled with grainstones and intraformational conglomerates, are composed of cylindrical and turbinate columnar (SH-V and SH-C) and digitate stromatolites (Gymnosolen, Inseria and Tungussia) in their lower parts. Larger, bulbous (SH-C and LLH-C) and conical (Conophyton) stromatolites occur in the upper parts, as well as the branching conophyte, Jacutophyton.All of the biostromes are always developed above cross-bedded carbonate sands (facies association B). A broadly symmetrical cyclic pattern, A B C B A, of tidal flat deposits (facies association A) passing up into carbonate sands (B), into biostrome (C), overlain by carbonate sands (B) and then tidal flat deposits (A), is repeated four times in the Porsanger Dolomite sequence. The pattern is interpreted in terms of two controls on sedimentation: (1) a slow transgressive phase followed by (2) depositional regression. The former (1) took place either through eustatic sea-level rise or more likely through accelerated subsidence because of tectonic instability and compaction of underlying sediments. This resulted in the sequence: tidal flat sediments, low intertidal/shallow subtidal carbonate sands, subtidal biostrome (A, B, C). Depositional regression through prograding tidal flats, generated the shoaling upward part of the cycle: biostrome, carbonate sands, tidal flat sediments (C, B, A).     

Petrography and biosedimentology of the Rosso Ammonitico Veronese (middle-upper Jurassic, north-eastern Italy)

A petrographic and biosedimentological study of the Rosso Ammonitico Veronese from the Trento Plateau (north-eastern Italy) shows that diagenetic (neomorphism, recrystallization) and biological processes (microbial content and pigmentation) influenced the formation and alteration of the carbonate matrix. The subject of this article is the interaction of early diagenetic processes and an attempt to explain the different colors of the matrix (red, pink, grey). Nearly 200 samples derived from 14 sections (Callovian to Tithonian) located in the Verona area have been studied by means of classical, cathodoluminescence and SEM microscopy. Calcite and ferruginous microfilaments of different shapes and sizes are present and tentatively attributed to fungi and iron bacteria. These micro-organisms precipitated iron oxy-hydroxides at poorly dysoxic-anoxic sediment–water interfaces. Further liberation of submicronic hydroxides (now hematite) was responsible for the red pigmentation of the carbonate matrices, originally composed of less than 1 μm-sized micrite. Controversial smaller nanograins (0.1–0.5 μm) attributed to nanobacteria or planktonic picoeukaryotes have been observed in the reddish samples. Recrystallization of the micrite leads to the formation of new micritic crystals, between 2 and 4 μm in size, then to microspar crystals. Micritic textures are linked to the different colours of the samples. The intensity of the red colour is correlated with the presence of hematite (former iron hydroxides) and the presence of planar subhedral micritic grains. In contrast, pink and greyish samples are linked to the increasingly coalescent structure of anhedral micritic and microsparitic crystals.     

Sedimentation on Rasdhoo and Ari Atolls, Maldives, Indian Ocean

A first systematic study of composition, texture, and distribution of modern sediments in two Maldivian atolls reveals the predominance of skeletal carbonates. Fragments of corals, calcareous algae, mollusks, benthic foraminifera, and echinoderms are identified in the grain-size fraction >125 μm. Non-skeletal grains such as cemented fecal pellets and aggregate grains only occur in small percentages. Fragments of skeletal grains, aragonite needles, and nanograins (<1 μm) are found in the grain-size fraction <125 μm. Needles and nanograins are interpreted to be largely of skeletal origin. Five sedimentary facies are distinguished (1–5), for which the Dunham-classification is applied. Fore reef, reef, back reef, as well as lagoonal patch reef and faro areas in both atolls are characterized by the occurrence of coral grainstones (1), which also contain fragments of red coralline algae, the codiacean alga Halimeda, and mollusks. On reef islands, coral-rich sediment is cemented to form intertidal beachrock and supratidal cayrock. Skeletal grains in atoll-interior lagoons are mainly mollusks and foraminifera. The lagoon of Rasdhoo Atoll is covered in the west by mudstones (2), in the center by mollusk packstones (3) and mollusk wackestones (4), and by hard bottoms with corals in the east adjacent to channels through the atoll reef margin. The interior lagoon of Ari Atoll contains mollusk wackestones (4) in the center and mollusk-foraminifer packstones (5). Marginal lagoon areas are characterized by hard bottoms with corals. Facies distribution appears to be an expression of depositional energy, which decreases from the atoll margin towards the center in Ari Atoll, and towards the west in Rasdhoo Atoll. Predominant sediment mineralogies include aragonite and high-magnesium calcite. Mean aragonite content decreases from 90% in coral grainstone to 70–80% in mollusk packstone, mollusk wackestone, and mudstone, and to 50% in mollusk-foraminifer packstone. Stable isotopes of oxygen and carbon in bulk samples range from −3 to −1.5 (δ¹⁸O) and from +0.4 to +3.2 (δ¹³C). It is not possible to delineate facies based on O- and C-isotopes.     

Cypris Habitat Selection Facilitated by Microbial Films Influences the Vertical Distribution of Subtidal Barnacle Balanus trigonus

The potential driving force(s) of the vertical distribution of subtidal barnacle Balanus trigonus Darwin were investigated using both field and laboratory experiments. Early juveniles (∼24 h old) placed in intertidal [∼0.5 m above mean low water level (MLWL)] and subtidal (∼3 m below MLWL) habitats survived equally well, indicating that the intertidal absence of B. trigonus in Hong Kong waters was not determined by differential mortality. However, enhanced attachment of cyprids in subtidal habitats indicated the importance of differential larval choice in determining their vertical distribution. In the laboratory, cyprids preferred to attach in response to subtidal microbial films, which may implicate microbial films as a primary cue in driving the adult vertical distribution. Microbial films developed in these two habitats differed in their biomass (=total organic carbon), abundance of bacteria and diatoms (determined by fluorescence microscopy), and bacterial diversity (determined by DNA fingerprinting analysis). For example, 6-day films in subtidal habitat had a significantly higher biomass than in films from intertidal habitat (P<0.05). There was no difference in the biomass of films from these two habitats in 9-day films (P>0.05); however, bacterial abundance was greater in subtidal films than in intertidal films, irrespective of the age of the film, although there was no difference in diatom abundance in films from these two habitats. Neither the abundance of bacteria and diatoms nor the biomass correlated with the attachment preferences of cyprids. This study has not provided any data to prove the existence of inductive and inhibitive (to cyprid attachment) bacterial species in subtidal and intertidal films, respectively; however, results indicate that bacterial community provided qualitative information that might explain the preferential attachment of B. trigonus cyprids in subtidal habitat.     

Deep-water stromatolites andFrutexites Maslov from the early and Middle Jurassic of S-Germany and Austria

Summary Despite extensive discussions during the last 20 years stromatolites are still used by many geologists as unequivocal indicators of very shallow-water conditions. We investigated four stratigraphic units from the Lower and Middle Jurassic of southern Germany (Posidonien-Schiefer, Amaltheen-Ton) and of the Northern Calcareous Alps (Adneter Kalk, Klauskalk), which were formerly interpreted as shallow marine sediments by some authors due to the occurrence of stromatolites. Our interpretations of the macro-, micro- and ultrafacies of these sediments are not compatible with shallow-water settings. We therefore propose a deep-marine, aphotic origin of these stromatolites. Former interpretations of the Posidonien-Schiefer as a shallow-water deposit are mainly based on the occurrence of stromatolites. We favour the model of a temporarily stagnant, deep, aphotic basin for these planktonrich sediments. Particles resembling ooids, but lying within mudstones cannot be taken as evidence for shallow agitated water. They either formed within the mud or are allochthonous. The deep-water setting of the red limestone of the Alpine Early and Middle Jurassic is indicated by a lack of platform-typical components like coated grains and phototrophic benthos and by shells of plankton and nekton forming a major part of the sediment. Stromatolites occur on the steep slope of a drowned Rhaetian reef with an estimated relief of 50–100 m and immediately below and within radiolarian limestones, deposited below the aragonite compensation depth (ACD). The aphotic stromatolites show some morphological differences to their shallow water counterparts. In all of our sections they occurred during intervals of reduced sedimentation. They form only thin horizons and probably grew very slowly. Mineralizations by Fe−Mn oxides and phosphate are very common. The presence of a microbial film is evident from binding of sedimentary particles, but the nature of the microbes is not known. Growth habits within the very distinct environments of red limestone and black shales show some common features, but also clear differences. The microproblematicumFrutexites Maslov is a very common component in deep-water stromatolites, but may also itself form small crusts or dendrolites. It occurs in two different forms. Opaque, slender forms with indistinct outlines probably grew within the weakly lithified sediment. Thicker, transparent forms with well defined outlines are found in cavities and probably also grew on the seafloor. Well preserved specimens display an internal fabric of radially arranged fibres of Fe−Mn oxides and calcite. It is suggested that calcite or aragonite were one original mineralogy ofFrutexites, which was later replaced by Fe−Mn oxides or phosphate. It is not certain whetherFrutexites is an organic, biomineralized structure or an inorganic mineralization, but the variable mineralogy and growth forms in different environments point to an organic origin. But even if organic, the occurrence in cryptic habitats and negative phototactic growth-directions make it clear thatFrutexites was not phototrophic.     

Evaluation of ammonium and phosphate release from intertidal and subtidal sediments of a shallow coastal lagoon (Ria Formosa – Portugal): a modelling approach

During an annual cycle, overlying water and sediment cores were collected simultaneously at three sites (Tavira, Culatra and Ramalhete) of Ria Formosa’s intertidal muddy and subtidal sandy sediments to determine ammonium, nitrates plus nitrites and phosphate. Organic carbon, nitrogen and phosphorus were also determined in superficial sediments. Ammonium and phosphate dissolved in porewater were positively correlated with temperature (P < 0.01) in muddy and sandy sediments, while the nitrogen-oxidized forms had a negative correlation (P < 0.02) in muddy sediments probably because mineralization and nitrification/denitrification processes vary seasonally. Porewater ammonium profiles evidenced a peak in the top-most muddy sediment (380 μM) suggesting higher mineralization rate when oxygen is more available, while maximum phosphate concentration (113 μM) occurred in the sub-oxic layer probably due to phosphorus desorption under reduced conditions. In organically poor subtidal sandy sediments, nutrient porewater concentrations were always lower than in intertidal muddy sediments, ranging annually from 20 μM to 100 μM for ammonium and from 0.05 μM to 16 μM for phosphate. Nutrient diffusive fluxes predicted by a mathematical model were higher during summer, in both muddy (104 nmol cm⁻² d⁻¹––NH₄⁺; 8 nmol cm⁻² d⁻¹––HPO₄⁻²) and sandy sediments (26 nmol cm⁻² d⁻¹––NH₄⁺; 1 nmol cm⁻² d⁻¹––HPO₄⁻²), while during lower temperature periods these fluxes were 3–4 times lower. Based on simulated nutrient effluxes, the estimated annual amount of ammonium and phosphate exported from intertidal areas was three times higher than that released from subtidal areas (22 ton year⁻¹––NH₄⁺; 2 ton year⁻¹––HPO₄⁻²), emphasizing the importance of tidal flats to maintain the high productivity of the lagoon. Global warming scenarios simulated with the model, revealed that an increase in lagoon water temperature only produces significant variations (P < 0.05) for NH₄⁺ in porewater and consequent diffusive fluxes, what will probably affect the system productivity due to a N/P ratio unbalance.     

Age-dependent zonation of the periwinkle Littorina littorea (L.) in the Wadden Sea

On sedimentary tidal flats near the island of Sylt (German Bight, North Sea) abundance and size distribution of periwinkles, Littorina littorea L., were studied in low intertidal and in shallow and deep subtidal mussel beds (Mytilus edulis L.). In low intertidal mussel beds, surveys revealed that high densities (1,369±571 m^–2) of juvenile snails (≤13 mm) were positively correlated with strong barnacle epigrowth (Semibalanus balanoides L. and Balanus crenatus Bruguière) on mussels. A subsequent field experiment showed that recruitment of L. littorea was restricted to the intertidal zone. Abundances of periwinkles (213±114 m^–2) and barnacles abruptly decreased in the adjacent shallow subtidal zone, which served as a habitat for older snails (>13 mm). L. littorea was completely absent from disjunct deep (5 m) subtidal mussel beds. Snail abundance varied seasonally with maxima of >4,000 m^–2 in low intertidal mussel beds in October and minima in July, just before the onset of new recruitment. I suggest that the presence of cracks and crevices among the dense barnacle overgrowth in intertidal mussel beds favoured recruitment and survival of juvenile snails. Larger (older) specimens are assumed to actively migrate to the less favourable adjacent subtidal. Therefore, intertidal mussel beds are considered as nurseries for the population of L. littorea in the Wadden Sea. Received in revised form: 25 September 2000 Electronic Publication     

Intertidal and Subtidal Fouling Assemblages in a Patagonian Harbour (Argentina, Southwest Atlantic)

The composition of the early stages of intertidal and subtidal fouling assemblages in Comodoro Rivadavia harbour (Argentina, 45°52′ S, 67°28′ W) and the influence of shore level and season on their structure were analysed. At the beginning of each season, stones were glued to the substratum with epoxy putty and distributed along 4 vertical transects at intervals of 20 m, at 3 levels: upper intertidal, middle intertidal, and subtidal. Substrata remained in the field for 84–100 days. A total of 48 samples (4 seasons × 3 levels × 4 replicates) were analysed. Species richness increased with depth, with 6 taxa in the upper intertidal, 23 in the middle intertidal and 31 in the subtidal. Seasonal differences in richness were less distinct. Green, red and brown algae were the dominant groups. Invertebrates were mainly represented by filter-feeding, sessile organisms, such as cheilostome bryozoans, spirorbid polychaetes and acorn barnacles. The barnacle Balanus glandula and the bryozoan Cryptosula pallasiana were the only non-indigenous species found in this study. Highly significant differences in structure among shore levels and seasons were evidenced by a two-way ANOSIM test. The upper intertidal is characterized by the filamentous green algae Urospora penicilliformis and Ulothrix flacca. The barnacle Balanus glandula is the most abundant species in the middle intertidal. The subtidal is defined mainly by the presence of the spirorbid polychaetes Paralaeospira levinseni and Romanchella perrieri, and the keyhole limpet Fissurella radiosa. Ordination of samples by season was less clear than by shore level.     

Habitat linkages: the effect of intertidal saltmarshes and adjacent subtidal habitats on abundance, movement, and growth of an estuarine fish

In this study we used pinfish (Lagodon rhomboides) in field experiments to examine linkages between intertidal saltmarsh and adjacent subtidal habitats. Pinfish are more than twice as abundant in intertidal marshes adjacent to seagrass beds than in those adjacent to the unvegetated subtidal bottom. Movement of pinfish between the marsh edge and the adjacent subtidal habitat was greater for fish captured in areas with both intertidal and subtidal vegetation than in those with intertidal vegetation and adjacent unvegetated mudflats. This movement provides an important link between habitats, allowing transfer of marsh-derived secondary production to subtidal seagrass beds and vice versa. Pinfish held in enclosures with both intertidal and subtidal vegetation were, on average, approximately 90% heavier than fish held in enclosures with intertidal vegetation and unvegetated subtidal bottom. Because saltmarshes and seagrass beds contribute to the production of living marine resources, active measures are being taken to preserve and restore these habitats. The results from this study have direct application to decisions concerning site selection and optimal spatial proximity of saltmarsh and seagrass habitats in the planning of restoration and mitigation projects. To maximize secondary production and utilization of intertidal marshes, managers may opt to restore and/or preserve marshes adjacent to subtidal seagrass beds. Received: 31 May 1996 / Accepted: 23 October 1996     

Across-shelf transport of gastropod larvae in the central Bahamas: rapid responses to local wind conditions

Surveys for veliger larvae of the large gastropods Strombusgigas and Strombus costatus were carried out in duplicate transectsrunning offshore (0.26–5 km) from Lee Stocking Island,Exuma Cays, Bahamas. The across-shelf distribution of the larvaeduring the 1991 spawning season was highly variable over periodsof 3–6 days. Larvae of both species are known to inhabitthe upper few meters of the water column, and analysis of windstress patterns over the study period showed that across-shelfdistribution of the larvae was affected by wind forcing. Seaward(recruitment-unfavorable) transport near Lee Stocking Islandwas most highly correlated with wind stress toward the north-northwest(335–357). The resultant wind stress during the studywas toward 279, suggesting that recruitment of conch to nurserygrounds on the Great Bahama Bank in the Exuma Cays occurs underanomalous summer wind conditions. The correlation between theacross-shelf distribution of conch veligers and wind stresswas highest with extinction coefficients of 0.17–0.23in the exponentially filtered wind stress. This indicates thatlarval distribution over the 5-km-long transects responded rapidlyto changes in wind stress. Over 75% of the effective wind stressoccurred over just 6–8 h preceding collections; therefore,short-term changes in wind stress, such as afternoon strengtheningof the trade winds, may have a significant influence on across-shelflarval transport. ³Present address: Northeast Fisheries Science Center, NationalMarine Fisheries Service NOAA, 74 Magruder Road, Highlands,NJ 07732, USA     

The potential for seaweed resource development in subarctic Canada; Nunavik,Ungava Bay

Ungava Bay is ice covered 6–7 months of the year and evidence of ice scouring of seaweeds is extensive in the intertidal and shallow subtidal. Maximum tidal amplitudes of 16 m, among the highest in Canadian waters, compound this impact. Despite this level of annual perturbation, very extensive and dense beds of fucoids in the intertidal and laminarians in the subtidal are common on the western shores of Ungava Bay. Ground surveys of 24 intertidal stations combined with satellite images delineated 82,000 tons standing crop of Fucus vesiculosus and Fucus evanescens in Payne Bay, of which 36,000 tons were considered harvestable. Subtidally, kelp cover reached peak biomass at 5–10 m consisting of three primary species, Saccharina longicruris, Laminaria digitata, and Laminaria solidungula. In the area of Payne Bay, kelp beds of 100 ha were common, averaging 9–12 kg m⁻² wet weight. The productivity of brown algae at these latitudes has been assumed to be low relative to southern latitudes. Direct measurement of lineal growth indicates productivity is intermediate between arctic and temperate populations. The potential for medium level industrial harvest exists under conservative management strategies within the constraints of subarctic logistics.     

Direct effects of physical stress can be counteracted by indirect benefits: oyster growth on a tidal elevation gradient

The paradigmatic gradient for intertidal marine organisms of increasing physical stress from low to high elevation has long served as the basis for using direct effects of duration of water coverage to predict many biological patterns. Accordingly, changes in potential feeding time may predict the direction and magnitude of differences between elevations in individual growth rates of sessile marine invertebrates. Oysters (triploid Crassostrea ariakensis) experimentally introduced at intertidal (MLW+0.05 m) and subtidal (MLW–0.25 m) elevations in racks provided a test of the ability to use duration of water coverage to predict changes in growth. During early-to-mid winter, a depression of 38–47% in shell growth of intertidal oysters matched the 36% reduction in available feeding time relative to subtidal oysters. In late winter as solar heating of exposed oysters increased, growth differences of 52–55% departed only slightly from the predicted 39%. In spring, however, duration of water coverage failed to predict even the correct direction of growth change with elevation as intertidal oysters grew 34% faster despite 39% less feeding time. Intense seasonal development of shell fouling by other suspension feeders like ascidians, mussels, and barnacles on subtidal (94% incidence) but not on aerially exposed intertidal (21–38% incidence) oysters may explain why duration of water cover failed to predict spring growth differences. Less intense fouling develops on intertidal oysters due to the physiological stress of aerial exposure on settlers, especially during higher temperatures and longer solar exposures of spring. Fouling by suspension feeders is known to reduce growth of the host through localized competition for food and added energetic costs. Thus, in springtime, indirect effects of aerial exposure providing a partial refuge from biological enemies overwhelmed direct effects of reduced duration of water coverage to reverse the expected pattern of slower intertidal growth of a marine invertebrate.     

Amphipods exclude filamentous algae from the Western Antarctic Peninsula benthos: experimental evidence

Hard bottom, subtidal communities along the Western Antarctic Peninsula are dominated by forests of large, chemically defended macroalgae that support a very dense assemblage of amphipods. Free-living filamentous algae are rare in the subtidal, but filamentous algal endophytes are common in many of the larger macroalgae, both likely as the result of amphipod grazing pressure. Filamentous algae are common in the intertidal, but primarily in the upper intertidal and on high-energy shores where amphipods are likely to be excluded much of the time. We tested the hypothesis that free-living, filamentous algae would be rapidly consumed if transplanted from the intertidal to the subtidal, and our results clearly supported this hypothesis. The filamentous, intertidal green alga Cladophora repens was transplanted to the benthos in 6 different macroalgal habitats. Control algae were transplanted in 3 m deeper waters nearby (usually 12 m or less laterally) but suspended 3 m off the bottom where amphipods are absent or rare. Overall consumption during approximately 6 h on the bottom ranged from 22 to 98% of the initial biomass, while significantly less biomass loss occurred in the water column.     

Ecophysiology of stromatolitic microbial mats,Stocking Island,exuma cays,Bahamas

Intertidal stromatolites, covered by cyanobacterial mats, were recently discovered at Stocking Island, Exuma Cays, Bahamas. Ecophysiological responses (CO₂ fixation, N₂ fixation, and photoacclimation) of these cyanobacterial mats to experimental manipulations were examined to identify potential environmental variables controlling community structure and function. The mats exhibit horizontal zonation that shifts from soft to crusty to hard in a seaward direction. Cluster analysis of chemotaxonomic photopigments (chlorophylls and carotenoids) revealed that visually distinct mat types are composed of distinct phototrophic assemblages. Under reduced irradiance, diatoms within the mats photoacclimated by increasing accessory photopigments (diadinoxanthin, fucoxanthin, and chlorophyll c ₁ c ₂) and cyanobacteria reduced the photoprotective carotenoid echinenone. In a 4-day nutrient addition bioassay experiment, nitrate, phosphate, dissolved organic carbon, and trace metal enrichments did not enhance CO₂ fixation, but phosphate enrichments tripled N₂ fixation rates. The addition of DCMU increased N₂ fixation rates relative to nonamended light and dark rates, indicating light (photosystem I) enhanced nitrogenase activity. Soft mats appear to represent the early stages of colonization and stabilization of mat communities. Active growth following stabilization results in the formation of partially-lithified crusty mats, which eventually become highly-lithified and form hard mats. Collectively, our results suggest that Stocking Island stromatolitic mats have low growth rates and consequently exhibit slow responses to increased nutrient availability and changes in ambient irradiance. In general, intertidal stromatolitic mats at Stocking Island appear to exhibit low rates of CO₂ and N₂ fixation relative to nonlithifying temperate cyanobacteral mats. Although production is low, respiration is likewise low, leading to the suggestion that high production to respiration ratios (P:R) may be necessary for lithification of intertidal stromatolitic mats.