Ecotypic differentiation in the kelpLaminaria saccharina: Phase-specific adaptation in a complex life cycle

Comparison of cultured sporophytes and gametophytes in common-garden experiments confirmed the existence of ecotypic differences in light-related traits among populations ofLaminaria saccharina (L.) Lamour. Cultured sporophytes from the turbid habitat in Long Island Sound, New York, USA, grew faster under both limiting and saturating daily irradiances than sporophytes from shallow and deep habitats along the Atlantic coast of Maine. Rapid growth of turbid plants was attributable to several factors, including high photosynthetic capacity and efficiency [due to differences in photosynthetic unit (PSU) number and size], low respiration rates, and high surface area:weight ratios. In contrast to sporophytes, microscopic gametophytes from the three kelp populations grew at similar rates under limiting and saturating daily irradiances. Biomass-specific photosynthesis vs irradiance (PI) parameters were similar for gametophytes from the shallow, deep, and turbid sites, despite population differences in chlorophyll-specific PI parameters and PSU characteristics. However, turbid gametophytes produced microscopic sporophytes more rapidly than gametophytes from the shallow and deep sites, apparently due to a lower blue-light requirement for gametogenesis. Ecotypic differences in sporophytes and gametophytes ofL. saccharina from shallow, deep, and turbid habitats can be understood as phase-specific adaptations.     

Temperature acclimation of respiration and photosynthesis in the brown algaLaminaria saccharina

Sporophytes of the brown algaLaminaria saccharina (L.) Lamour grown at 15°C contained significantly more chlorophylla (chla) than did similar plants grown at 5°C. The increase in chla in 15°C plants was due to increased numbers of photosystem II reaction centes, and possibly to increased photosynthetic unit size, compared with 5°C plants. These changes were associated with increased values (photosynthetic efficiencies) in 15°C-grownL. saccharina relative to 5°C-grown plants. The changes in together with reduced respiration rates allowed 15°C-grownL. saccharina to achieve net photosynthesis and light-saturated photosynthesis at a lower photon fluence rate (PFR) than 5°C plants when both groups were assayed at the same temperature (15°C). The photon fluence rates necessary to reach the compensation point and achieve light-saturated photosynthesis (I _c andI _k , respectively) increased with increasing incubation temperature inL. saccharina grown at both 5 and 15°C. However, acclimation responses to growth temperature compensated for the short-term effect of temperature onI _c andI _k . Consequently, plants grown at 5 and 15°C were able to achieve similar rates of light-limited photosynthesis, and similarI _c andI _k values at their respective growth temperatures. These responses are undoubtedly important for perennial seaweeds such asL. saccharina, which frequently grow in light-limited habitats and experience pronounced seasonal changes in water temperature.Please address all correspondence and requests for reprints to I.R. Davison     

Changes in morphology and physiology of an East Mediterranean sponge in different habitats

The sponge Tetilla sp. (Tetractinomorpha: Tetillidae) is a common species in the eastern Mediterranean. This sponge inhabits four different habitat types differing in wave impact and irradiance levels. Two of these habitats (a shallow cave and deep water) are characterized by relatively calm water, whereas the other two (shallow exposed site and tide pools) are in turbulent water with high energy flow. The present study examined the influence of physical (depth, illumination and water motion) and biotic factors on morphology, skeletal plasticity and reproductive traits among the four spatially separated populations. Sponges from tidal pools had significantly larger body volume than sponges from deep water and from shallow caves (ANOVA: tidal-deep P<0.0001; tidal-shallow caves P<0.05). Sponges from exposed habitats were significantly larger than deep-water sponges (ANOVA: P=0.01). In addition, individuals from tide pools and from the exposed habitat had a significantly higher proportion of structural silica than sponges from the calmer deep water and from the cave sites. Oxea spicules in sponges from the calm habitats were significantly shorter than in those from the tidal pools and the exposed habitats. The percentage of spicules out of a sponges dry weight in individuals transplanted from deep (calm) to shallow (turbulent) water significantly increased by 21.9±12.9%. The new spicule percentage did not differ significantly from that of sponges originally from shallow water. Oocyte diameter differed significantly between habitats. The maximal size of mature eggs was found in deep-water sponges in June (97±5 m). In the shallow habitats, a smaller maximal oocyte diameter was found in the cave, in May (56.5±3 m). Furthermore, oocyte density in shallow-water sponges was highest in May and decreased in June (with 88.2±9 and 19.3±9 oocytes mm^–2, respectively). At the same time (June), oocyte density of deep-water sponges had just reached its maximum (155±33.7 oocytes mm^–2). The difference in oocyte size and density between deep- and shallow-water individuals indicates an earlier gamete release in the shallow sponge population. The results suggest that plasticity in skeletal design of this sponge indicates a trade off between spicule production and investment in reproduction.Communicated by O. Kinne, Oldendorf/Luhe     

Photoinhibition and recovery of the kelp Laminaria saccharina at optimal and superoptimal temperatures

Effects of high irradiance on photosynthetic characteristics were examined in sporophytes of the kelp Laminaria saccharina Lamour. from 1992 to 1994. Exposure to high irradiance (700 mol photons m^-2s^-1) for 1 h at optimal temperature (12°C) caused a 40 to 60% decline in photosynthetic efficiency (alpha), quantum yield, and the ratio of variable to maximum chlorophyll fluorescence (F_v/F_m), an indicator of Photosystem II efficiency. Although the photoinhibition effects were partly attributable to protective mechanisms, a concurrent increase in minimal fluorescence (F_o) indicated damage to Photosystem II reaction centers. The magnitude of photoinhibition was proportional to irradiance and duration; however, F_v/F_m was significantly reduced after exposure to irradiances as low as 40 to 50 mol photons m^-2s^-1 for 1 h, or to 700 mol photons m^-2s^-1 for only 5 min. In contrast, photosynthetic capacity (P_max) was affected only at much higher irradiance. Superoptimal temperatures up to 24°C did not exacerbate high-light effects. At 25°C, however, alpha and P_max were more susceptible to photoinhibition than at lower temperatures. Recovery from photoinhibition was examined by following F_v/F_m and F_o for 24 h after exposure to high light. Recovery of F_v/F_m was fastest during the first 1 to 3 h, and slowed or ceased after 6 to 12 h, while recovery of F_o was relatively constant over 12 h. Dithiothreitol, which blocks formation of energy-dissipating xanthophylls, reduced both the initial rate and extent of recovery. Chloramphenicol, which blocks chloroplast-encoded protein synthesis, had little effect on initial rates of recovery, but stopped recovery after 3 h. Thus, L. saccharina appears to rely on the xanthophyll cycle to protect the photosynthetic apparatus, and reversal of this protective mechanism causes the rapid initial recovery in F_v/F_m. Longterm recovery depends on repair of damaged reaction centers. Both the rate and extent of recovery were temperature-dependent. The initial rate was higher at 18 to 22°C than at 12°C, but the extent of recovery over 24 h declined with increasing temperature. High temperatures, therefore, appear to enhance protective mechanisms, but disrupt repair processes. L. saccharina from Long Island Sound, an ecotype adapted to low light and high temperature, showed slightly but consistently greater effects of photoinhibition than plants from the Atlantic coast of Maine, but exhibited faster recovery at superoptimal temperatures.     

Variation in colony geometry modulates internal light levels in branching corals, <Emphasis Type="Italic">Acropora humilis</Emphasis> and <Emphasis Type="Italic">Stylophora pistillata</Emphasis>

Colonial photosynthetic marine organisms often exhibit morphological phenotypic plasticity. Where such plasticity leads to an improved balance between rates of photosynthesis and maintenance costs, it is likely to have adaptive significance. To explore whether such phenotypic plasticity leads to more favourable within-colony irradiance for reef-building branching corals, this relationship was investigated for two coral species Acropora humilis and Stylophora pistillata, along a depth gradient representing light habitats ranging from 500 to 25 μmol photons m⁻² s⁻¹, during 2006 at Heron Island, Great Barrier Reef (23.44°S, 151.91°E). In the present study changes in flow-modulated mass transfer co-varied with light as a function of depth. In low-light (deep) habitats, branch spacing (colony openness) in A. humilis and S. pistillata was 40–50% greater than for conspecifics in high-light environments. Also, branches of A. humilis in deep water were 40–60% shorter than in shallow water. Phenotypic changes in these two variables lead to steeper within-colony light attenuation resulting in 38% higher mean internal irradiance (at the tissue surface) in deep colonies compared to shallow colonies. The pattern of branch spacing was similar for S. pistillata, but this species displayed an alternate strategy with respect to branch length: shade adapted deep and cave colonies developed longer and thinner branches, allowing access to higher mass transfer and irradiance. Corals in cave habitats allowed 20% more irradiance compared to colonies found in the deep, and had a 47% greater proportion of irradiance compared to colonies in the shallow high-light environment. Such phenotypic regulation of internal light levels on branch surfaces partly explains the broad light niches of many branching coral species. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Adaptation strategies of the corallimorpharian <Emphasis Type="Italic">Rhodactis rhodostoma</Emphasis> to irradiance and temperature

Corallimorpharians may dominate some habitats on coral reefs and compete with stony corals for access to light, yet little is known concerning their photosynthetic traits. At Eilat in the northern Red Sea, we observed that the abundance of individuals of the corallimorpharian Rhodactis rhodostoma decreased significantly with depth on the reef slope. Field and laboratory experiments revealed that they employ several mechanisms of photoadaptation to high irradiance on the shallow reef flat. Their endosymbiotic microalgae (zooxanthellae) varied significantly in both abundance and chlorophyll content with level of irradiance. Use of a diving pulse amplitude modulated fluorometer revealed that the zooxanthellae of R. rhodostoma effectively disperse excess light energy by expressing significantly higher values of non-photochemical quenching and maximum excitation pressure on photosystem II when experimentally exposed to high light (HL) versus low light (LL). Host corallimorpharian tissues mediated this response by shielding the algal symbionts from high irradiance. The endoderm of host tentacles thickened significantly and microalgal cells were located further from the mesoglea in HL than in LL. The clades of zooxanthellae hosted by the corallimorpharians also varied with depth. In shallow water, all sampled individuals hosted clade C zooxanthellae, while in deep water the majority hosted clade D. The photosynthetic output of individuals of R. rhodostoma was less affected by HL than was that of a stony coral examined. When exposed to both high temperature (HT) and HL, individuals of R. rhodostoma reduced their maximum quantum yield, but not when exposed to HL at low temperature (LT). In contrast, colonies of the scleractinian coral Favia favus reduced their photosynthetic output when exposed to HL in both temperature regimes. After 2 weeks of HT stress, R. rhodostoma polyps appeared to bleach completely but re-established their zooxanthella populations upon return to ambient temperature. We conclude that mechanisms of photoadaptation to high irradiance employed by both the endosymbiotic zooxanthellae and host corallimorpharians may explain in part the abundance of R. rhodostoma on some shallow reef flats. The ability to survive for weeks at HT while bleached also may allow corallimorpharians to repopulate shallow reef areas where scleractinians have been killed by thermal stress. B. Kuguru and G. Winters contributed equally to this work.     

Effects of four environmental variables on photosynthesis-irradiance relationships in Antarctic sea-ice microalgae

The effects of temperature, salinity, growth irradiance and diel periodicity of incident irradiance on photosynthesis-irradiance (P-I) relationships were examined in natural populations of sea-ice microalgae from McMurdo Sound in the austral spring of late 1984. Both P _m ^b (photosynthetic rate at optimum irradiance) and ^b (initial slope or P-I curve) were temperature-dependent reaching optimal rates at approximately +6° and +2°C, respectively. P-I relationships showed little difference at 20 and 33 S; however, no measurable photosynthesis by sea-ice microalgae was detected in a 60 S solution of brine collected from the upper layers of congelation ice. Although diel periodicity characteristic of the under-ice light field appeared to have little effect on P-I relationships, changes in growth irradiance had a profound effect. An increase in growth irradiance from 7 E m^-2 s^-1 (ambient) to 35 or 160 E m^-2 s^-1 resulted in a transient three-fold increase in P _m ^b and I _k (index of photoadaptation) during the first four days, followed by a sharp decline. The effects of these environmental factors on ice algal photosynthesis may influence the distribution of microalgae in sea-ice environments.     

Effect of feeding regime and irradiance on the photophysiology of the symbiotic sea anemone Aiptasia pulchella

To determine how the animal and algal components of the symbiotic sea anemone Aiptasia pulchella respond to changes in food availability and culture irradiance, sea anemones from a single clone were maintained at four irradiance levels (320, 185, 115, and 45 E m^-2 s^-1) and either starved or fed for 5 wk. Changes in protein biomass of sea anemones maintained under these conditions were not related to the productivity of zooxanthellae, since the protein biomass of fed A. pulchella decreased with increase in irradiance and there was no difference in protein biomass among starved sea anemones at the four irradiance levels. Except for the starved high-light sea anemones, the density of symbiotic zooxanthellae was independent of culture irradiance within both starved and fed. A. pulchella. Starved sea anemones contained over twice the density of zooxanthellae as fed sea anemones. Within both starved and fed individuals, chlorophyll per zooxanthella increased with decreasing culture irradiance while algal size remained constant (in fed sea anemones) at about 8.80 m diameter. Chlorophyll a: c ₂ ratios of zooxanthellae increased with decreasing culture irradiance in zooxanthellae from starved sea anemones but remained constant in zooxanthellae from fed sea anemones. As estimated from mitotic index data, the in situ growth rates of zooxanthellae averaged 0.007 d^-1 and did not vary with irradiance or feeding regime. Photosynthesis-irradiance (P-I) responses of fed A. pulchella indicated an increase in photosynthetic efficiency with decreasing culture irradiance. But there was no consistent pattern in photosynthetic capacity with culture irradiance. Respiration rates of fed sea anemones also did not vary in relation to culture irradiance. The parameter I _k , defined as the irradiance at which light-saturated rates of photosynthesis are first attained, was the only parameter from the P-I curves which increased linearly with increasing culture irradiance. The daily ratio of net photosynthesis to respiration for A. pulchella ranged from 1.6 to 2.8 for sea anemones maintained at the three higher irradiances, but was negative for those maintained at 45 E m^-2 s^-1. Since the final protein biomass was greatest for sea anemones maintained at the lowest irradiance, these results indicate that sea anemone growth cannot be directly related to productivity of zooxanthellae in this symbiotic association.     

Biomass, photosynthesis and growth of Laminaria saccharina in a high-arctic fjord, NE Greenland

Biomass, photosynthesis and growth of the large, perennial brown alga Laminaria saccharina (L.) Lamour. were examined along a depth gradient in a high-arctic fjord, Young Sound, NE Greenland (74°18'N; 20°14'W), in order to evaluate how well the species is adapted to the extreme climatic conditions. The area is covered by up to 1.6-m-thick ice during 10 months of the year, and bottom water temperature is <0°C all year round. L. saccharina occurred from 2.5 m depth to a lower depth limit of about 20 m receiving 0.7% of surface irradiance. Specimen density and biomass were low, likely, because of heavy ice scouring in shallow water and intensive feeding activity from walruses in deeper areas. The largest specimens were >4 m long and older than 4 years. In contrast to temperate stands of L. saccharina, old leaf blades (2-3 years old) remained attached to the new blades. The old tissues maintained their photosynthetic capacity thereby contributing importantly to algal carbon balance. The photosynthetic characteristics of new tissues reflected a high capacity for adaptation to different light regimes. At low light under ice, or in deep water, the chlorophyll a content and photosynthetic efficiency (!) were high, while light compensation (E_c) and saturation (E_k) points were low. An E_c of 2.0 µmol photons m^-2 s^-1 under ice allowed photosynthesis to almost balance, and sometimes exceed, respiratory costs during the period with thick ice cover but high surface irradiance, from April through July. Rates of respiration were lower than usually found for macroalgae. Annual elongation rates of leaf blades (70-90 cm) were only slightly lower than for temperate L. saccharina, but specific growth rates (0.48-0.58 year^-1) were substantially lower, because the old blades remained attached. L. saccharina comprised between 5% and 10% of total macroalgal biomass in the area, and the annual contribution to primary production was only between 0.1 and 1.6 g C m^-2 year^-1.     

Photosynthetic performance of Laminaria solidungula measured in situ in the Alaskan High Arctic

Photosynthetic performance in the kelp Laminaria solidungula J. Agardh was examined from photosynthesis irradiance (P-I) parameters calculated from in situ ¹⁴C uptake experiments, using whole plants in the Stefansson Sound Boulder Patch, Alaskan Beaufort Sea, in August 1986. Rates of carbon fixation were determined from meristematic, basal blade, and second blade tissue in young and adult sporophytes. Differences in saturating irradiance (I _k, measured as photosynthetically active radiation, PAR), photosynthetic capacity (P _max), and relative quantum efficiency () were observed both between young and adult plants and between different tissue types. I _k was lowest in meristematic tissue (20 to 30 E m^–2 s^–1) for both young and adult plants, but consistently 8 to 10 E m^–2 s^–1 higher in young plants compared to adults in all three tissues. Average I _k for non-meristematic tissue in adult plants was 38 E m^–2 s^–1. Under saturating irradiances, young and adult plants exhibited similar rates of carbon fixation on an area basis, but under light limitation, fixation rates were highest in adult plants for all tissues. P _max was generally highest in the basal blade and lowest in meristematic tissue. Photosynthetic efficiency () ranged between 0.016 and 0.027 mol C cm^–2 h^–1/E m^–2 s^–1, and was highest in meristematic tissue. The relatively lower I _k and higher exhibited by L. solidungula in comparison to other kelp species are distinct adaptations to the near absence of light during the eight-month ice-covered period and in summer when water turbidity is high. Continuous measurement of in situ quantum irradiance made in summer showed that maximum PAR can be less than 12 E m^–2 s^–1 for several days when high wind velocities increase water turbulence and decrease water transparency.The Univeristy of Texas Marine Science Institute Contribution No. 695     

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.     

Photosynthetic characteristics of Prochloron sp./ascidian symbioses

The prokaryotic green alga Prochloron sp. (Prochlorophyta) is found in symbiotic association with colonial didemnid ascidians that inhabit warm tropical waters in a broad range of light environments. We sought to determine the light-adaptation features of this alga in relation to the natural light environments in which the symbioses are found, and to characterize the temperature sensitivity of photosynthesis and respiration of Prochloron sp. in order to assess its physiological role in the productivity and distribution of the symbiosis. Colonies of the host ascidian Lissoclinum patella were collected from exposed and shaded habitats in a shallow lagoon in Palau, West Caroline Islands, during February and March, 1983. Some colonies from the two light habitats were maintained under conditions of high light (2 200 E m^–2 s^–1) and low light (400 E m^–2 s^–1) in running seawater tanks. The environments were characterized in terms of daily light quantum fluxes, daily periods of light-saturated photosynthesis (H_sat), and photon flux density levels. Prochloron sp. cells were isolated from the hosts and examined for their photosynthesis vs irradiance relationships, respiration, pigment content and photosynthetic unit features. In addition, daily P:R ratios, photosynthetic quotients, carbon balances and photosynthetic carbon release were also characterized. It was found that Prochloron sp. cells from low-light colonies possessed lower chlorophyll a/b ratios, larger photosynthetic units sizes based on both reaction I and reaction II, similar numbers of reaction center I and reaction center II per cell, lower respiration levels, and lower P_max values than cells from high-light colonies. Cells isolated from low-light colonies showed photoinhibition of P_max at photon flux densities above 800 E m^–2 s^–1. However, because the host tissue attenuates about 60 to 80% of the incident irradiance, it is unlikely that these cells are normally photoinhibited in hospite. Collectively, the light-adaptation features of Prochloron sp. were more similar to those of eukaryotic algae and vascular plant chloroplasts than to those of cyanobacteria, and the responses were more sensitive to the daily flux of photosynthetic quantum than to photon flux density per se. Calculation of daily minimum carbon balances indicated that, though high-light cells had daily P:R ratios of 1.0 compared to 4.6 for low-light cells, the cells from the two different light environments showed nearly identical daily carbon gains. Cells isolated from high-light colonies released between 15 and 20% of their photosynthetically-fixed carbon, levels sufficient to be important in the nutrition of the host. Q₁₀ responses of photosynthesis and respiration in Prochloron sp. cells exposed briefly (15–45 min) to temperatures between 15° and 45°C revealed a discontinuity in the photosynthetic response at the ambient growth temperatures. The photosynthetic rates were found to be more than twice as sensitive to temperatures below ambient (Q₁₀=3.47) than to temperatures above ambient (Q₁₀=1.47). The Q₁₀ for respiration was constant (Q₁₀=1.66) over the temperature range examined. It appears that the photosynthetic temperature sensitivity of Prochloron sp. may restrict its distribution to warmer tropical waters. The ecological implications of these findings are discussed in relation to published data on other symbiotic systems and free-living algae.     

“Free space-photosynthesis” in the algae Fucus virsoides and Laminaria saccharina

This paper deals with the role of bicarbonates in the apparent free space (AFS) and their importance for the photosynthetic process. Both the littoral alga Fucus virsoides (Mediterranean Sea) and the sublittoral alga Laminaria saccharina (Baltic Sea) show the same response. Unloading of Clions and subsequent reloading of HCO₃-ions result in a positive photosynthetic rate in distilled water (free of CO₂). Time of exposure of an algal thallus to bicarbonate solutions, as well as the concentration of these solutions, influences AFS-assimilation of the ions.     

Influence of habitat on abundance and size structure of a large temperate-reef fish, Achoerodus viridis (Pisces: Labridae)

The relationship between densities of Achoerodus viridis (Pisces: Labridae) and reef habitats at various localities within New South Wales (NSW), Australia was examined. Types of habitats were quantified at inner, mid and outer estuarine locations in each of two estuaries (Botany Bay and Port Jackson) to determine whether purported patterns of movement from estuaries could be related to differences in habitat. Although the same types of habitat were generally found at all locations, differences in the proportion of habitat types were found between shallow and deep reefs and among inner, mid and outer estuarine locations for both estuaries. Shallow habitats were usually dominated by Ecklonia radiata, turf and/or fringe habitat in Botany Bay, whereas deep sites were generally dominated by urchin-grazed barrens habitat and, sometimes, sponge- and ascidian-dominated deep reef. Shallow sites in Port Jackson were dominated by a mixture of habitats, as were deep reefs at mid-estuarine locations. Other deep reefs in Port Jackson were dominated by E. radiata (inner estuarine) or barrens (outer estuarine) habitat. Thus, patterns of habitat cover were not consistent between estuaries and numbers of fish could not be related to proportional representation of habitat on reefs along estuarine gradients. Univariate and multivariate analyses showed that there was little evidence that any size class of fish was correlated with the proportional representation of a particular habitat or group of habitats. Counts of fish that focused on barrens and E. radiata forest habitats over a period of 10 yr showed that similar numbers and all sizes of fish were found in the two types of habitat. Greater numbers of small fish were, however, found in the E. radiata forest habitat than in the barrens habitat. Estimates of abundance along the coast of NSW (100s to 1000 km) in a range of habitats (e.g. ascidian-dominated reef, kelp forest, urchin-grazed barrens) showed that there was no indication that a particular habitat consistently had greater numbers of A. viridis than other habitats. Therefore, A. viridis of a range of sizes appears to be flexible in its use of habitats on reefs. Received: 24 December 1997 / Accepted: 23 June 1998     

Ecological implications of the presence of the tube-building polychaete <Emphasis Type="Italic">Lanice conchilega</Emphasis> on soft-bottom benthic ecosystems

The common tube-building polychaete Lanice conchilega is known as a habitat structuring species and can form dense aggregations. The effects of L. conchilega on the surrounding benthic community have received little attention, especially in subtidal areas. Therefore, the presence of L. conchilega in different habitats in the North Sea and its effect on the abundance, species richness, diversity and community structure in these habitats are evaluated in the present paper, based on data from the ICES North Sea Benthos Survey of 2000. Lanice conchilega has a wide geographical distribution and a low habitat specialization, but optimally occurs in shallow fine sands. In the present study, the presence of L. conchilega resulted in a density increase and a significant (positive) correlation of the benthos density with the density of L. conchilega. Furthermore, the species richness (number of species) increased with increasing density of L. conchilega. This trend was, however, not consistent: the number of species reached more or less an asymptotic value or even decreased after reaching a critical density of L. conchilega (>500–1,000 ind/m2), as observed in shallow fine sands. The same overall pattern was detected concerning the expected number of species. The N ₁-diversity index showed similar or slightly higher values in L. conchilega patches compared to patches without L. conchilega. From the results of the community analysis, it can be concluded that the species, which were responsible for the increase of the diversity, belonged to the overall species-pool of that habitat. The effects on density and diversity differed between the four discerned habitats (shallow muddy sand, shallow fine sand, shallow medium sand and deep fine sand), and were most pronounced in shallow fine sands. These patterns can be attributed to the habitat structuring capacity of L. conchilega. The mechanisms responsible for the increase of the habitat quality in patches of L. conchilega can be summarized as (1) changes in the hydrodynamics, (2) increases of the habitat stability and oxygen supply, and (3) a creation of habitat heterogeneity in a uniform environment. In this way, L. conchilega alters the habitat characteristics and affects other organisms, and can therefore even be considered as an ecosystem engineer. In other words, L. conchilega patches are responsible for an increased habitat quality in an otherwise uniform habitat, which results in a higher survival of the surrounding benthic species.     

Habitat use and group size of pied cormorants (<Emphasis Type="Italic">Phalacrocorax varius</Emphasis>) in a seagrass ecosystem: possible effects of food abundance and predation risk

Both food abundance and predation risk may influence habitat use decisions. However, studies of habitat use by birds in marine environments have focused only on food abundance. I investigated the possible influences of food abundance and predation risk from tiger sharks (Galeocerdo cuvier) on habitat use by pied cormorants (Phalacrocorax varius) over two spatial scales and on cormorant group size. Cormorants were usually solitary, but group size was highest in shallow habitats during months when shark density was low. Regardless of season, cormorant density within shallow habitats was higher over seagrass than sand, and cormorants were distributed between these two microhabitats proportional to prey density. Therefore, cormorants appear to respond to prey abundance at a relatively narrow spatial scale (i.e., tens of meters). At the habitat-patch scale (~1 km), the density of cormorants and their prey (teleosts) was higher in shallow habitats than in deep ones, but the density of cormorants was influenced by an interaction between water temperature (i.e., season) and habitat. There was decreased use of shallow habitats as water temperature, and the density of tiger sharks, increased. When shark density was low, cormorants were distributed across habitats roughly in proportion to the abundance of fish, suggesting that cormorants respond to food abundance at the scale of habitat patches. However, as shark abundance increased, the relative density of cormorants dropped in the dangerous shallow habitats such that there was a greater density of cormorants relative to their food in deep habitats when sharks were abundant. This suggests that pied cormorants trade-off food and risk by accepting lower energetic returns to forage in safer habitats. This study provides the first evidence that marine habitat selection by birds may be influenced by such a trade-off, and provides further evidence that tiger sharks are important in determining habitat use of their prey and mediating indirect interactions within Shark Bay.Communicated by P. W. Sammarco, Chauvin     

Habitat modification affects recruitment of abalone in central New Zealand

The habitat experienced during early life-history stages can determine the number and quality of individuals that recruit to adult populations. In a field experiment, biogenic habitat complexity was manipulated (presence or absence of foliose macroalgae) at two depths (2–3 m and 5–6 m) and the habitat-dependent effects on recruitment of the black foot abalone (Haliotis iris) were examined at three field sites along the south coast of Wellington, New Zealand (41°20′S, 174°47′E), between July and November 2005. Recruit density (<5 weeks post-settlement) was measured on cobbles covered with crustose coralline algae. Habitats of low complexity (barrens treatments) had consistently greater densities of recruits than habitats of high complexity (algae treatments). However, recruits in algae habitats were larger, and for deep habitats, there was greater survival in algae habitats compared with barrens habitats. While depth had no significant effect on early recruit (<2 weeks post-settlement) density, late recruit (<5 weeks post-settlement) density was greater in shallow habitats, and so it seems recruit survival was greater in shallow habitats. In this experiment, algal habitat complexity had strong effects on early recruit abundance, but habitat-dependent variations in recruit growth and survival may modify initial patterns of abundance and determine recruitment to adult abalone populations.     

Photoadaptation of photosynthesis in Gonyaulax polyedra

Gonyaulax polyedra Stein exhibited a combination of photoadaptive strategies of photosynthesis when only a single environmental variable, the light intensity during growth, was altered. Which of several biochemical/physiological adjustments to the light environment were employed depended on the level of growth irradiance. The photoadaptive strategies employed over any small range of light levels appeared to be those best suited for optimizing photosynthetic performance and not photosynthetic capacity. (Photosynthetic performance, P _i, is defined as the rate of photosynthesis occurring at the level of growth irradiance.) Among all photosynthetic parameters examined, only photosynthetic performance showed a consistent correspondence to growth rates of G. polyedra. Above 3500 to 4000 W cm^-2, where photosynthetic performance was equal to photosynthetic capacity, cells were not considered light-limited in either photosynthesis or growth. At these higher light levels, photosynthetic perfomance, cell volume, growth rates and respiration rates remained maximal; photosynthetic pigment content varied only slightly, while the photosynthetic capacity of the cells declined. At intermediate light levels (3000 to 1500 W cm^-2), photosynthesis, not growth, was light-limited, and photoadaptive strategies were induced which enhance absorption capabilities and energy transfer efficiencies of chlorophyll a to the reaction centers of G. polyedra. Photosynthetic capacity remained constant at about 280 mol O₂ cm^-3 h^-1, while photosynthetic performance ranged from 100 to 130 mol O₂ cm^-3 h^-1. Major increases in photosynthetic pigments, especially peridinin-chlorophyll a-proteins and an unidentified chlorophyll c component, accompanied photoadaptation to low irradiances. Maximal growth rates of 0.3 divisions day^-1 were maintained, as were respiration rates of about-80 mol O₂ cm^-3 h^-1 and cell volumes of about 5.4×10^-8 cm^-3 cell^-1. Below about 1250 W cm^-2, photosynthesis in G. polyedra was so light-limited that photosynthetic performance was unable to support maximal growth rates. Under these conditions, G. polyedra displayed photostress responses rather than photoadaptive strategies. Photostress was manifested as reduced cell volumes, slower growth, and drastic reductions in pigmentation, photosynthetic capacity, and rates of dark respiration.     

Ecotypic differentiation in the marine diatom Skeletonema costatum: influence of light intensity on the photosynthetic apparatus

Three genetically distinct clones of Skeletonema costatum (Grev.) Cleve were grown at 20°C under high (274 E m^-2 s^-1) and low (27 E m^-2 s^-1) light conditions and their photoadaptive photosynthetic responses compared. When all three clones were grown under low light, pigment analyses and fluorescence excitation spectra demonstrated that the accessory pigments, chlorophyll c and fucoxanthin, became more important in light-harvesting compared to chlorophyll a. Photosynthetic unit sizes increased for Photosystems I and II in low light, but photosynthesis vs irradiance characteristics were not reliable predictors of photosynthetic unit features. Fluorescence excitation spectra and photosynthesis vs irradiance (P-I) relationships indicated that changes in energy transfer occurred independent of changes in pigment content. Large increases in accessory pigment content were not accompanied by large increases in excitation from these pigments. Changes in energy transfer properties were as important as changes in PSU size in governing the photoadaptive responses of S. costatum. When the three clones were grown under identical conditions, each had a separate and distinct pattern of photoadaptation. Significant differences among clones were found for pigment ratios, photosynthetic unit sizes for Photosystems I and II and efficiency of energy transfer between pigments. These strikingly different photoadaptive strategies among clones may partially account for the great ecological success of the diatom species. This is the first quantitative investigation of the importance of both chlorophyll c and fucoxanthin to the adaptive responses of diatoms to light intensity, and represents the most complete characterization of the photoadaptive responses of a single species of marine phytoplankter to differences in light environment.     

Population differentiation and taxonomic status of the exploited limpet Patella candei in the Macaronesian islands (Azores,Madeira, Canaries)

There has been considerable confusion in the taxonomy of limpets of the North East Atlantic Ocean and Mediterranean Sea, particularly those from the Macaronesian islands. The present study compared populations of the intertidal limpet Patella candei d'Orbigny from the Azores, Madeira and the Canaries with those of P. caerulea Linnaeus and P. depressa Pennant from the European and African continental coasts. No major differences in radular morphology were detected between the three species. However, electrophoretic analysis of 15 enzyme loci gave overall genetic identity (I) values of 0.5 between the three species, indicating that they cannot be regarded as conspecific as previously thought, and suggesting that P. candei is endemic to the Macaronesian islands. Comparisons of P. candei within these islands showed that, although populations did not differ with respect to radular morphology and soft-body parts, populations from the Azores were distinct from those in Madeira and the Canaries in shell shape and gene frequencies. Individuals from the Azores had, no average, taller shells and longer radulae, while those in Madeira and the Canaries had a shallow, depressed and stellate shell form. This was interpreted as being due to the wider habitat distribution of the species in the Azores compared to Madeira and the Canaries. Electrophoretic results showed that P. candei from the Azores differed from P. candei in Madeira and the Canaries by almost 40% of the loci investigated (I=0.660), suggesting that the former is a separate endemic species. An I value of 0.969 between populations in Madeira and the Canaries was typical of conspecific populations.