Feeding preferences of gammarid amphipods among four species ofFucus

Each of fourFucus species occurring on the Atlantic coast of Canada was presented to two species of gammarid amphipods in no-choice and multiple-choice laboratory experiments. We also measured concentrations of anti-herbivore phlorotannins in each of theFucus species. The study site was Cranberry Cove in Nova Scotia, Canada, and experiments were conducted in 1988. In a multiple-choice experiment,F. vesiculosus was very heavily grazed. Other species were much less affected. In a no-choice experiment,F. evanescens, F. spiralis andF. vesiculosus were eaten at similar rates, whileF. distichus was barely grazed. These results correlate with measurements of phorotannin concentrations in each of the species.F. distichus had much higher concentrations than did the other species (which were not significantly different among themselves). There is no general clear-cut relationship between grazer deterrence and competitive capability within the fucoid guild studied.     

Effect of diurnal variations in natural irradiance on the apical length growth and light saturation of growth in five species of benthic macroalgae

The diurnal variation in apical length growth rate of five benthic macroalgae from the Swedish West Coast (Fucus spiralis L., F. vesiculosus L., F. serratus L., Ascophyllum nodosum (L.) Le Jolis, and Chondrus crispus Stackhouse) was measured during 1.5 to 2.0-h intervals for 3 to 5 d in September 1984. All species showed a maximum in growth rate in the morning, followed by a continuous decline throughout the rest of the day (most pronounced in F. serratus, A. nodosum and C. crispus). By comparing the growth rates with the average irradiance during the same interval, saturation irradiances for length growth in natural daylight regime were estimated; 60 to 70 Wm^-2 for F. spiralis and F. vesiculosus, 90 to 100 Wm^-2 for A. nodosum, 30 to 40 Wm^-2 for F. serratus and 10 to 20 Wm^-2 for C. crispus. Average growth rates during the dark hours ranged from 33% of average day growth rates in A. nodosum to 63% in C. crispus, respectively.     

Improved survival under heat stress in intertidal embryos (<Emphasis Type="Italic">Fucus</Emphasis> spp.) simultaneously exposed to hypersalinity and the effect of parental thermal history

Intertidal organisms exposed to thermal stress normally experience other stresses simultaneously, but how these combined stresses modify tolerance to heat, especially for embryos, is poorly understood. Tolerance of fucoid algal embryos to heat, with and without acclimation to a sublethal temperature and with simultaneous exposure to hypersaline media, was examined. Embryos of Fucus vesiculosus L. (mid-intertidal zone) were less tolerant than embryos of Fucus spiralis L. (upper intertidal zone); without acclimation and with a growth temperature of 14°C, about half of the embryos survived 3 h exposure to 33°C in F. vesiculosus and of 35°C in F. spiralis. Conditions experienced by parental thalli (4°C versus 14°C storage) significantly affected the heat tolerance of embryos grown for 24 h post-fertilization at 14°C in F. vesiculosus, a result that is important for biologists using fucoid algae as model systems. Acclimation to a sublethal temperature (29°C) or exposure to the LT₅₀ (33°C, F. vesiculosus; 35°C, F. spiralis) in 100 psu seawater (2850 mmol kg^–1 osmolality) resulted in 30–50% higher levels of embryonic survival. Higher levels of HSP60s were found in embryos exposed to 29–33°C than to 14°C; lower levels of HSP60s were present in embryos exposed to the LT₅₀ under hypersaline conditions than in normal seawater. Contemporaneous studies in 1995–1996 of substratum temperature and desiccation levels were made at Schoodic Point, Maine (USA) underneath F. spiralis and F. vesiculosus canopies and in Semibalanus balanoides patches. This study extends the bioindicator utility of heat-shock proteins in studies of intertidal organisms and demonstrates the importance of integrated stress responses in survival of a single stress factor (e.g. temperature).Electronic Supplementary Material Supplementary material is available in the online version of this article at     

Selfing in <Emphasis Type="Italic">Parvulastra exigua</Emphasis>: an asterinid sea star with benthic development

Parvulastra exigua, a species that lacks a dispersive larva, is paradoxically one of the most widely distributed and abundant sea stars in Australia. The potential that self-fertilization and successful development of self-generated progeny might contribute to population maintenance in this species was investigated through induction of spawning in isolated individuals collected from five populations near Sydney, New South Wales (34°S, 151°E), during the spawning period (August–October 2010, 2011). Evidence of selfing differed significantly among the five populations but the level of selfing within individual egg masses did not. Overall, there was a mean of 8.9% fertilization in egg masses with embryos. Some clutches developed to the juvenile stage while others arrested development before hatching. The presence of miniscule testis tissue in gonads that appeared to be ovaries indicated that hermaphroditism may be more common in some populations than indicated by anatomy. Mixed mating (selfing + outcrossing), pseudocopulation during egg laying and developmental success of self-fertilized embryos may contribute to the ecological success of P. exigua. These might influence population genetic structure and would facilitate establishment of founder populations by a small number of migrants.     

Feeding preferences of periwinkles among four species ofFucus

A widely accepted view of intertidal community organizatiton in the NW Atlantic proposes that fucoid vegetation is maintained by the actions of predators which remove species competitively superior toFucus species. Herbivory is an important component of these predatory interactions, but has been studied largely with reference to the interaction betweenF. vesiculosus andLittorina littorea. There are many species of fucoids and herbivorous invertebrates on the shores of the NW Atlantic and this paper reports field and laboratory experiments performed in 1987 (in or near Halifax, Canada) on the effects of grazing by three species ofLittorina on adults and juveniles of four species ofFucus. In laboratory experiments, portions ofFucus species were presented singly (no choice) or together (multiple choice) to single species ofLittorina. AdultF. distichus was grazed to only a small extent.F. evanescens andF. vesiculosus adult tissues were heavily grazed in most laboratory experiments.F. spiralis adults were heavily grazed in no choice experiments, but were grazed only slightly in multiple choice tests. When adult fucoid tissues were exposed to a mixed suite of grazers in the field, onlyF. vesiculosus was grazed heavily. Tests done on adult tissues showed clearly thatFucus species are highly variable in their vunerability to grazers. Moreover, we found variability among periwinkle species in grazing rates on adultFucus. We also found a great disparity in the feeding rates of different periwinkle species on juveniles ofFucus. Therefore, it is not possible to extrapolate from a single periwinkle species/single fucoid species interaction when attempting to identify vegetation structuring processes.     

Phosphate uptake by intertidal algae in relation to zonation and season

The removal of phosphate from ambient seawater by whole plants of five species of fucoid algae, collected from the east coast of N. Ireland in 1988 and 1989, was followed over 6-h periods. A transient uptake pattern was observed forPelvetia canaliculata (L.) Dcne. et Thuret,Fucus spiralis L.,F. vesiculosus L. andF. serratus L., consisting of an initial period of high uptake, followed by a phase of zero uptake and then a period at an intermediate rate.Ascophyllum nodosum (L.) Le Jolis had a constant slow rate of uptake over 6 h. The initial uptake rate ofF. spiralis was significantly greater than that of any other species. Phosphate uptake over a 2-h period was measured at concentrations ranging from that of ambient seawater to 25µg-at. l^–1 for whole plants ofF. spiralis andF. serratus, using a large scale batch method. A small scale batch method was used for whole plants ofP. canaliculata and sections of the other four species investigated. Uptake abilities of the algae at low concentrations of phosphate were compared using the parameterV ₁ (the uptake rate at 1µg-at. l^–1) and at high concentrations usingV _max, the maximum uptake rate. These kinetic parameters of uptake were calculated using a method that avoids bias and permits statistical evaluation of the results. The fucoid algae studied could be divided into two distinct groups on the basis of their abilities to take up phosphate from seawater.P. canaliculata andA. nodosum had low values ofV ₁ in winter, which were also correlated with their positions on the shore and did not vary between winter and summer. TheFucus species had higher values ofV ₁ in winter, which were also correlated with their positions on the shore. In summer, however,V ₁-values for these species decreased and no longer correlated with their shore heights. TheV _max-value forF. spiralis was higher in winter than in summer but was signifcantly greater than that of any other species at all times of year. The ecological significance ofV _max is discussed in relation to nutrient limitation and the possible occurrence of patches of high nutrient concentration in the intertidal environment.     

High intertidal community organization on a rocky headland in Maine,USA

A mosaic patchwork of the barnacle Balanus balanoides L., the mussel Mytilus edulis L., and the alga Fucus vesiculosus L. was found in the transitional region between the mid and high intertidal zones on a rocky headland on Mount Desert Island, Maine, USA. The development of the mosaic was observed by following recruitment and survival of B. balanoides in denuded patches located at the same tidal level along a 60 m stretch of shore. Barnacle recruitment was least under canopies of F. vesiculosus and greatest in open areas kept moist at low tide by surf. Barnacle survival after settlement was least under the F. vesiculosus canopy due to the whiplash effect of the algal fronds in the surf and greatest in open areas free from competition from mussels. In open areas, early mortality was correlated with settlement density. In areas of dense settlement (60 spat cm^-2) up to 90% mortality resulted within 5 months from crowding associated with growth. In older individuals crowding produced hummocks of elongated, weakly attached barnacles which were more prone to removal by surf than uncrowded barnacles. Mussels exerted competitive dominance over barnacles for space and the presence of mussel beds prevented further barnacle recruitment. Mussels suffered extensive mortality during winter storms when surf removed dense mats of weakly attached mussels. The patchy distribution of mussels and barnacles results from irregular rock substrata producing numerous environmental patches with respect to wave exposure and drainage at low tide, and from densitydependent mortality of both mussels and barnacles which creates patches of new colonizable space within each environmental patch.     

Synergistic effects of ultraviolet radiation and conditions at low tide on egg masses of limpets (<Emphasis Type="Italic">Benhamina obliquata</Emphasis> and <Emphasis Type="Italic">Siphonaria australis</Emphasis>) in New Zealand

The effects of ultraviolet radiation (UVR), desiccation and conditions in tidal pools on embryonic survival were examined for two common pulmonate limpets that lay intertidal benthic egg masses on rocky shores in New Zealand: Benhamina obliquata and Siphonaria australis. Field surveys and manipulative experiments were conducted between December 2006 and September 2007 in the Wellington region of New Zealand (41°17′S, 174°47′E). Egg mass deposition sites in the field were species-specific: B. obliquata deposited eggs primarily in shaded crevices, whereas S. australis predominantly deposited egg masses in the sun and in tidal pools. For both species, however, embryonic mortality was greater in egg masses that had been in full sun compared to shade. For S. australis, there was also high mortality in egg masses in tidal pools or desiccated compared to those that remained submerged in flowing seawater at low tide. In outdoor experiments, embryonic mortality was also always greatest for egg masses exposed to full sun, and lowest for those in shaded treatments. Mortality was also higher if egg masses were in simulated tidal pools, and for S. australis, if desiccated, compared to those submerged in flowing seawater. Periods of particularly sunny conditions with high temperatures also resulted in higher overall mortality. Finally, egg masses of both species that were initially deposited in the shade had greater mortality in response to subsequent UV exposure compared to egg masses initially deposited in full sun. Results from this study suggest that the egg masses of these two species are highly vulnerable to UVR, as well as other intertidal stressors. Embryos of both of these species may be at risk of high mortality particularly during summer when extreme conditions of UV intensity and high temperature coincide with low tide cycles. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Temporal patterns of larval settlement and survivorship of two broadcast-spawning acroporid corals

Acroporid corals are the main reef-building corals that provide three-dimensional habitats for other reef organisms, but are decreasing on many reefs worldwide due to natural and anthropogenic disturbances. In this study, temporal patterns of larval settlement and survivorship of two broadcast-spawning acroporid coral species, Acropora muricata and A. valida, were examined through laboratory rearing experiments to better understand the potential for larval dispersal of this important coral group. Many larvae were attached (but not metamorphosed) to settlement tiles on the first examination 3–4 days after spawning (AS). The first permanent larval settlement (i.e. metamorphosed and permanently settled juvenile polyps) occurred at 5–6 days AS, and most larval settlement (85–97% of total) occurred within 9–10 days AS. Larval survivorship decreased substantially to around 50% by the first week of the experiment and to approximately 10% by the second to third week. The rates of larval attachment, settlement, and the initial drop in survivorship of larvae suggest that effective dispersal of some acroporid species may largely be completed within the first few weeks AS.     

Selective grazing of the isopod Idotea baltica between Fucus evanescens and F. vesiculosus from Kiel Fjord (western Baltic)

Grazing rates of the isopod Idotea baltica on Fucus evanescens and F. vesiculosus (Phaeophyta) were quantified in laboratory feeding preference experiments. Fucus species were offered alone (no-choice) or simultaneously (choice). In three of four no-choice experiments and in all four choice experiments, I. baltica significantly preferred F. vesiculosus to F. evanescens. F. evanescens recently immigrated into Kiel Fjord and has increased in abundance since 1990. One possible reason for the competitive success of this species may be that, compared to F. vesiculosus, it is less preferred by I. baltica, the most abundant mesograzer at the study site.     

Distribution and maximum growth depth of Fucus vesiculosus along the Gulf of Finland

 A survey of the distribution and maximum depth of a continuous Fucus vesiculosus belt was carried out in the Gulf of Finland in 1991. F. vesiculosus is widely distributed throughout the Gulf of Finland, including the vicinity of Vyborg Bay, Russia in the east. The maximum growth depth of F. vesiculosus in the Gulf of Finland reflects two different patterns according to the exposure to wave action. The most robust and continuous F. vesiculosus belt is observed on exposed shores, where the maximum growth depth is 5 to 6 m, with the optimum at 2 to 3 m. On moderately exposed shores the maximum growth depth is 3 m, with an optimum growth depth of <2 m. The maximum growth depth also varies geographically, with a decreasing trend towards the east. Maximum growth depth of F. vesiculosus correlates with light intensity. The compensation point for F. vesiculosus photosynthesis is about 25 μmol m⁻² s⁻¹, and photosynthesis is saturated at a light intensity of 300 μmol m⁻² s⁻¹. Vertical irradiance attenuation measurements in situ in summer revealed that for F. vesiculosus photosynthesis the quantity of light is optimal (200 to 300 μmol m⁻² s⁻¹) at <3 m depth. At depths >5 m the quantity of light is near or below the photosynthesis compensation point and insufficient for growth. These depth limits of light penetration coincide with measured growth depths of F. vesiculosus in the Gulf of Finland. Received: 7 May 1999 / Accepted: 18 November 1999     

Functional divergence in heat shock response following rapid speciation of Fucus spp. in the Baltic Sea

In the Baltic Sea, the broadly distributed brown alga Fucus vesiculosus coexists in sympatry over part of its range (south west Gulf of Bothnia) with the Baltic endemic F. radicans sp. nov, while further north in colder and lower-salinity areas of the Baltic F. radicans occurs alone (north west Gulf of Bothnia). F. radicans appears to have arisen via rapid speciation from F. vesiculosus within the recent history of the Baltic (ca. 7500 BP). Possible functional divergence between the two species was investigated by comparing stress-responsive gene expression in a common-garden experiment. The experiment used two allopatric populations of Fucus vesiculosus from the Skagerrak (North Sea) and Central Baltic, as well as F. radicans from the same Central Baltic site. The two species in sympatry displayed divergent heat shock responses, while F. vesiculosus populations from allopatric sites did not. F. radicans was more sensitive to heat shock at 25°C, either alone or together with high irradiance and desiccation, than Baltic or Skagerrak F. vesiculosus. The results indicate that rapid functional divergence in the inducible heat shock response has occurred between sympatric species on a timescale of thousands of years.     

Reproductive traits of pioneer gastropod species colonizing deep-sea hydrothermal vents after an eruption

The colonization dynamics and life histories of pioneer species determine early succession at nascent hydrothermal vents, and their reproductive ecology may provide insight into their dispersal and population connectivity. Studies on the reproductive traits of two pioneer gastropod species, Ctenopelta porifera and Lepetodrilus tevnianus, began within a year after an eruption on the East Pacific Rise (EPR) that eliminated vent communities near 9°50′N from late 2005/early 2006. Standard histology was used to examine gamete release, instantaneous female fecundity, and time to maturation. Both species exhibited two-component oocyte size–frequency distributions indicating quasi-continuous reproduction with high fecundity. In samples collected in December 2006, both C. porifera and L. tevnianus individuals were reproductively mature. The smallest reproducing C. porifera were 4.2 mm (males) and 5.4 mm (females) in shell length, whereas reproductive L. tevnianus were smaller (2.3 and 2.4 mm in males and females, respectively). Most C. porifera were large (>6.0 mm) compared to their size at metamorphosis and reproductively mature. In contrast, most L. tevnianus were small (<1.0 mm) and immature. Reproductive traits of the two species are consistent with opportunistic colonization, but are also similar to those of other Lepetodrilus species and peltospirids at vents and do not fully explain why these particular species were the dominant pioneers. Their larvae were probably in high supply immediately after the eruption, due to oceanographic transport processes from remote source populations.     

Self-fertilization and maternal enzymes in the planulae of the coral Goniastrea favulus

Electrophoretic analysis of planulae obtained from controlled breeding experiments with the coral Goniastrea favulus during spawning events in 1986 and 1987 on the Great Barrier Reef, Australia, shows that offspring are produced equally frequently by outcrossing or self-fertilization, leading to a genetically determined rate of selfing of around 50%. Field rates of selfing may be greater than this as a consequence of the limited dispersal of gametes within this species. However, the level of heterozygote depression found in one population is well below that which would be predicted for such levels of selfing. In one enzyme system, glucosephosphate isomerase, the maternal phenotype continues to be expressed in the planula larva for between 11 and 17 d after fertilization. This effect is postulated to result from a residual of maternal enzyme and RNA derived from a relatively large egg.     

Food and habitat choice in floating seaweed clumps: the obligate opportunistic nature of the associated macrofauna

The species composition of macrofauna associated with floating seaweed rafts is highly variable and influenced by many factors like spatial and temporal variation, period since detachment and probably also the seaweed species. The presence of seaweed preferences was assessed by a combination of in situ seaweed samplings and multiple-choice aquarium experiments in a controlled environment, using the seaweed-associated grazing organisms Idotea baltica and Gammarus crinicornis. Results from the sampling data confirm that the seaweed composition influences macrofaunal species composition and abundance: samples dominated by Sargassum muticum displayed higher densities but lower diversities compared to samples dominated by Ascophyllum nodosum and Fucus vesiculosus. Seaweed preference was also apparent from the multiple-choice experiments, but did not exactly match the results of the community analysis: (1) I. baltica had high densities in seaweed samples (SWS) dominated by F. vesiculosus and A. nodosum, while in the experiments, this isopod was most frequently associated with Enteromorpha sp. and F. vesiculosus, and fed mostly on S. muticum, A. nodosum and Enteromorpha sp.; (2) G. crinicornis had high densities in SWS dominated by F. vesiculosus, while in the experiments, this amphipod was most frequently associated with S. muticum, but fed most on A. nodosum and F. vesiculosus. It is clear from the laboratory experiments that preference for habitat (shelter) and food can differ among seaweed species. However, food and habitat preferences are hard to assess because grazer preference may change if choices are increased or decreased, if different sizes of grazers are used, or if predators or other grazers are added to the experiments. The effects of seaweed composition may also be blurred due to the obligate opportunistic nature of a lot of the associated macrofaunal species.     

Drought stress,rain and recovery of the intertidal seaweed <Emphasis Type="Italic">Fucus spiralis</Emphasis>

Non-motile organisms of intertidal shores such as seaweeds have to cope with a great variability of environmental factors. In this survey, we studied whether different morphotypes of the intertidal seaweed Fucus spiralis L. are also reflected in a characteristic performance. Desiccation and recovery of this Phaeophyceae were investigated in field experiments near Aljezur, Portugal. Fucus spiralis is exposed to serious desiccation during periods of falling tide, resulting in a tissue water loss of about 90%. Due to large semidiurnal tidal ranges in this area, two morphotypes can be distinguished: F. spiralis growing in the lower intertidal (LZ) is thicker and fleshier compared with plants in the upper intertidal (HZ), and this is reflected in a significant difference in fresh and dry mass. During sunny days and at low tide, effective quantum yields (Φ_PSII) decreased significantly after 2 h desiccation. This continued until re-submersion. The photosynthetic performances of HZ and LZ plants also differed significantly after LZ plants were already submerged and photosynthetisizing, but the HZ specimens still exposed to air. Recovery experiments after desiccation treatments showed fast recovery within 6 min after re-submersion in both morphotypes. HZ specimens showed a slower recovery, which indicates a protection measure to the adverse conditions in the upper intertidal. In 24 h desiccation treatments, however, HZ specimens expressed a significantly higher maximum fluorescence yield F _v /F _m recovery. Simulated rainfalls during low tides caused photosynthetic activity to drop to 50% of initial F _v /F _m , independent of the length of the rain period. Treated plants also fully recovered after 6 min re-submersion in seawater. A comparison of single fronds and tufts clearly indicated advantages of the tuft growth strategy: tufts showed higher Φ_PSII at prolonged emersion times. Our study indicated a clear relationship between size and drought resistance, which was primarily due to the smaller and hardy HZ plants that withstand longer desiccation times without damage.     

A comparison of two cartographic exposure methods using Fucus vesiculosus as an indicator

Morphological variation and vertical distribution of Fucus vesiculosus were quantified at several sites in the Finnish archipelago (Baltic Sea). F. vesiculosus samples were obtained from skerries at geographical distances of 1 km or more (large scale) and at intervals of ca 100 m around a single island (small scale). The results were examined in relation to wave exposure, calculated by Baardseth and effective fetch cartographic methods. Despite the fact that the exposure indices were calculated differently they correlated strongly. Vegetative morphological characteristics of F. vesiculosus illustrate the morphological differences both within and between exposure gradients. The tallest and widest F. vesiculosus plants were found at the sheltered end of the large-scale exposure gradient. Those from equally sheltered sites of the island were smaller in all respects. Thus, the trend from small narrow plants to large wide sheltered plants was expressed differently over the different geographical scales. Consequently localities with similar exposure indices may have morphologically different F. vesiculosus populations. Shores with similar cartographic exposure indices can be different in nature. Underwater topography and shore locations, either close to the mainland or at the outermost sites of the archipelago, affect the exposure. Although a sheltered shore is indicated, the sublittoral zone may be quite exposed to the movements of water. In contrast, in an open shore environment underwater rocks, boulders and shallow water areas can provide sheltered habitats. The depth range of the F. vesiculosus belt exhibited two distinctive patterns. At sheltered sites, around islands in the outermost reaches of the archipelago F. vesiculosus can grow to a maximum depth of 5 m. In exposed habitats the belt becomes narrower, reaching a maximum depth of 3 m. Closer to the mainland F. vesiculosus is found at exposed sites to a maximum depth of 5 m; the depth range at sheltered sites is narrower, only reaching depths of 2 m or less. In conclusion, the changes in plant morphology and in the vertical belt distribution are similar to each other along both gradients at the exposed ends of the wave action spectrum; however, the two gradients diverge at the sheltered ends of the spectrum. Received: 10 August 1998 / Accepted: 11 January 1999     

Competition and facilitation between germlings of<Emphasis Type="Italic"> Ascophyllum nodosum</Emphasis> and<Emphasis Type="Italic"> Fucus vesiculosus</Emphasis>

Competitive interactions between germlings of Ascophyllum nodosum (L) Le Jolis and Fucus vesiculosus L. were studied both in the laboratory and on a shore of the Isle of Man, in the Irish Sea. Both intra- and interspecific competition were investigated by comparing the performance of algal germlings both in monocultures and mixed populations of the two species. The growth of germlings of both species reduced with increasing density. F. vesiculosus always grew faster than Ascophyllum and did best in mixed cultures, whereas Ascophyllum did least well when mixed with Fucus germlings. Clearly the adverse effects of F. vesiculosus on A. nodosum were greater than those of Ascophyllum cohorts. At the same total density, the survival and growth of Ascophyllum declined with an increasing proportion of Fucus germlings, implying that poor recruitment of A. nodosum results from strong competition with F. vesiculosus. However, under desiccation stress on the shore, F. vesiculosus enhanced the survival of A. nodosum at the early germling stage even though competition may occur again at the late stage. Thus, whether interactions between germlings take the form of competition or facilitation depends on the environmental conditions.Communicated by O. Kinne, Oldendorf/Luhe     

Adaptation of light-harvesting pigments to downwelling light and the consequent photosynthetic performance of the eulittoral rockweeds Ascophyllum nodosum and Fucus vesiculosus

We compared the effect of habitat and water depth on the light-harvesting pigment content for Ascophyllum nodosum and Fucus vesiculosus at two near-shore stations in Long Island Sound (USA). Excised pieces of seaweeds were attached at depth intervals to a vertically buoyed line, and left in situ for 7 days. For comparison, fronds were collected from sun and shade habitats in the littoral zone. The three major antenna (light-harvesting) pigments increased in concentration with depth or shade. Chlorophyll c to a ratios remained stable at about 0.2. Fucoxanthin to chlorophyll a ratios decreased by 20 to 30% with depth or shade. Although pigment composition for the two rockweed species was equivalent, the maximum photosynthetic performance of F. vesiculosus exceeded that of A. nodosum by a factor of 2, while the compensation depths for 4 m-adapted A. nodosum and F. vesiculosus under natural limiting light conditions were equivalent. Plants held at 4 m had higher photosynthetic rates compared with plants held at 0 m, no matter the depth of measurement. Indirect evidence indicates that the enhanced photosynthesis of 4 m-adapted plants is due not only to higher concentrations of antenna pigments but to other physiological factors as well. We conclude that the clearly delineated vertical distribution of these two canopy species, the F. vesiculosus zone over the A. nodosum zone, is not determined by light quantity or quality, but by biotic factors as evidenced by the experiments of Menge which are cited herein.     

Settlement of <Emphasis Type="Italic">Macoma balthica</Emphasis> larvae in response to benthic diatom films

The role of multi-species benthic diatom films (BDF) in the settlement of late pediveliger larvae of the bivalve Macoma balthica was investigated in still-water bioassays and multiple choice flume experiments. Axenic diatom cultures that were isolated from a tidal mudflat inhabited by M. balthica were selected to develop BDF sediment treatments characterized by a different community structure, biomass, and amount of extracellular polymeric substances (EPS). Control sediments had no added diatoms. Although all larvae settled and initiated burrowing within the first minute after their addition in still water, regardless of treatment, only 48–52% had completely penetrated the high diatom biomass treatments after 5 min, while on average 80 and 69% of the larvae had settled and burrowed into the control sediments and BDF with a low diatom biomass (<3.5 μg Chl a g⁻¹ dry sediment), respectively. The percentage of larvae settling and burrowing into the sediment was negatively correlated with the concentration of Chl a and EPS of the BDF. This suggests higher physical resistance to bivalve penetration by the BDF with higher diatom biomass and more associated sugar and protein compounds. The larval settlement rate in annular flume experiments at flow velocities of 5 and 15 cm s⁻¹ was distinctly lower compared to the still-water assays. Only 4.6–5.8% of the larvae were recovered from BDF and control sediments after 3 h. Nonetheless, a clear settlement preference was observed for BDF in the flume experiments; i.e., larvae settled significantly more in BDF compared to control sediments irrespective of flow speed. Comparison with the settlement of polystyrene mimics and freeze-killed larvae led to the conclusion that active selection, active secondary dispersal and, at low flow velocities (5 cm s⁻¹), passive adhesion to the sediment are important mechanisms determining the settlement of M. balthica larvae in estuarine biofilms.