Patterns of organizations of intertidal and shallow subtidal vegetation in wave exposed habitats of central Chile

Wave-exposed rocky intertidal habitats of central Chile exhibit zonation of algal morphologies rather than strict patterns of species zonation. In low shore areas, there is a vertical sequence of perennial belts of calcareous crusts, kelp-like forms and expanded cushions or non-calcareous crusts. The calcareous crusts are represented by species of Mesophyllum, the kelp-like forms include Lessonia nigrescens and Durvillaea antarctica, while the cushions are represented by Gelidium chilense and G. lingulatum and the noncalcareous, expanded crusts by Codium dimorphum. Thin and thick blades, represented by Iridaea laminarioides, Ulva rigida and Porphyra columbia and filamentous forms including Ceramium rubrum, Centroceras clavulatum and Polysiphonia spp. are more patchy than the lower, perennial belts. They may, however, form distinct temporal monocultures at upper intertidal levels. Upper and lower limits of the various zones are set by interactions of several factors, the relative importance of which can change seasonally. When some of the factors restricting species distribution are experimentally removed, other interactions among factors become limiting.Within each zone, species are morphologically similar, with the abundance of species being regulated by symmetric competitive interactions. Competition is often asymmetric at the boundaries of zones except when adults of small-sized forms interact with morphologically similar juveniles of larger forms. Irrespective of their extremely different morphologies, the permanent, zone-forming algal species generally combine escape from grazers or defensive adaptations with clear competitive abilities. Nevertheless, there is a clear competitive hierarchy which is expressed in vertical displacements and zonation. The lowershore habitats could potentially be occupied by any of the different types of algae. Fast growth and large size allow the kelps to occupy this zone pushing the calcareous crust dominated-zone down into shallow subtidal areas and displacing the cushions and fleshy crusts into the low and middle intertidal regions. In turn, these last forms can displace thick and thin foliose forms and filaments to upper levels on the shore. Displaced forms may exist as patches at various levels of the shore.     

Disturbance and reef topography maintain high local diversity in Ecklonia radiata kelp forests

Disturbance of competitive‐dominant plant and algae canopies often lead to increased diversity of the assemblage. Kelp forests, particularly those of temperate Western Australia, are habitats with high alpha diversity. This study investigated the roles of broad‐scale canopy loss and local scale reef topography on structuring the kelp‐dominated macroalgal forests in Western Australia. Eighteen 314 m² circular areas were cleared of their Ecklonia radiata canopy and eighteen controls were established across three locations. The patterns of macroalgal recolonisation in replicate clearances were observed over a 34 month period. Macroalgal species richness initially increased after canopy removal with a turf of filamentous and foliose macroalgae dominating cleared areas for up to seven months. A dense Sargassum canopy dominated cleared areas from 11 to 22 months. By 34 months, partial recovery of the kelp canopy into cleared areas had occurred. Some cleared areas did not follow this trajectory but remained dominated by turfing, foliose and filamentous algae. As kelp canopies developed, the initial high species diversity declined but still remained elevated relative to undisturbed controls, even after 34 months. More complex reef topography was associated with greater variability in the algal assemblage between replicate quadrats suggesting colonising algae had a greater choice of microhabitats available to them on topographically complex reefs. Shading by canopies of either Sargassum spp. and E. radiata are proposed to highly influence the abundance of algae through competitive exclusion that is relaxed by disturbance of the canopy. Disturbance of the canopy in E. radiata kelp forests created a mosaic of different patch types (turf, Sargassum‐dominated, kelp‐dominated). These patch types were both transient and stable over the 34 months of this study, and are a potential contemporary process that maintains high species diversity in temperate kelp‐dominated reefs.     

Marine macroalgae as foods for fishes: an evaluation of potential food quality

Synopsis A revitalized view of feeding by herbivorous marine fishes is sought through two questions. First, What characteristics of major taxa of algae identify them as predictably high or low quality foods? Second, are marine algae valuable foods for fishes which do not mechanically disrupt cell walls and do not harbor specialized enzymes or microbes capable of lysing cell walls? Energy, ash and nutrient content of 16 species of marine algae were employed to assess food quality of fleshy red, green, brown and calcareous red algae. On the basis of ash, calories, total protein and total lipid content, fleshy algae should be superior to calcareous algae as foods for fishes; in addition, green algae should be superior to brown algae and brown algae superior to red algae. When the probable digestibility of storage and extracellular carbohydrates is considered, green and red algae are predicted superior to brown algae as food. Two species of damselfishes (Pomacentridae) from the Gulf of California,Eupomacentrus rectifraenum andMicrospathodon dorsalis, eat red and green algae and ignore brown and calcareous algae. They feed, therefore, in a fashion consistent with predictions based only on algal chemistry. These fishes absorb at least 20–24% of the biomass, 57–67% of the protein, 46–56% of the lipid and 37–44% of the carbohydrate contained in algae eaten in the wild. Since these damselfishes do not masticate their food, it appears that herbivorous fishes can digest major fractions of algal nutrients without mechanical destruction of algal cells.     

Biogenic habitat structure and characteristics of temperate reef fish assemblages

Assemblages of non-cryptic, substrate-oriented species of fish were compared on a series of reefs in Southern California, USA. Reefs were grouped according to algal cover: dense beds of giant kelp (Macrocystis pyrifera) with turf understorey; sparse beds of giant kelp with foliose algae understorey: foliose algae < 1 m in height; and open barrens. Despite affinities to particular algal substrates by many individual species, we detected no differences in fish species richness and only weak differences in species composition among reefs of different habitat types. Planktivores and species that consume macro-invertebrates were less likely to occur on reefs that supported giant kelp; the frequencies of occurrence of three other trophic groups (piscivores, herbivores and micro-carnivores) were unaffected by giant kelp. Algal composition on reefs in Southern California is temporally highly dynamic. Changes in macro-algal composition of reefs influenced population dynamics of two fish species (black surfperch and striped surfperch) examined. Overall, the weak spatial variation in fish assemblages on reefs in Southern California appears to result from relatively unspecialized ecological requirements of many species combined with temporal changes in algal structure on reefs that are rapid relative to generation times of the fish. We hypothesize that the degree of spatial differentiation in assemblages of substrate-associated species of fish may be inversely related to the temporal constancy of biogenic reef structure.     

Comparative feeding ecology of two herbivorous damselfishes (Pomacentridae: Teleostei) from the Gulf of California,Mexico

Two species of benthic damselfishes from the Gulf of California, Mexico, use contrasting behaviors when feeding on benthic algal communities. The small (±70 g) Cortez damselfish, Eupomacentrus rectifraenum (Gill, 1862), feeds selectively from a multi-species algal mat, eats fleshy red and green algae and ignores brown and calcareous algae. The giant blue damselfish, Microspathodon dorsalis (Gill, 1862), is a large (±450 g), lethargic, nonselective feeder which grazes on a near monoculture of a fleshy red alga, Polysiphonia sp. Feeding activity for both species is low in the morning peaks during late afternoon, and drops sharply as night approaches. Based on feeding rates, gut-filling times, and weights of gut contents, Cortez damselfish process six to eight full guts of food and giant blue damselfish three full guts of food per day. The algal mat exhibits high standing crops (291–618 g dry wt · m^?2) and low productivity, but the preferred food of the Cortez damselfish (Ulva) appears to colonize the mat frequently and grow rapidly. The Polysiphonia dominated community on giant blue damselfish territories exhibits low standing crops (23 g · m^?2) and high productivity (34–47 times that of the mat per gram algae). Even though the feeding behaviors and resources used by the two damselfishes differ, both species eat similar food (delicate red and green fleshy algae, and depend on rapid colonization and/or high productivity to maintain their primary foods in the grazed algal community.     

BIOGEOGRAPHY AND ECOLOGY OF THE KELP/RED ALGAL SYMBIOSIS

A kelp/red algal symbiosis is described from nature based on extensive collections from the San Juan Islands, Washington. Kelp gametophytes were found as endophytes in the cell walls of seventeen species of red algae in three different kelp communities. Host red algae were mostly filamentous (e.g., Pleonosporium vancouverianum) or polysiphonous (e.g. Polysiphonia paniculata). The kelp gametophytes completed vegetative and reproductive development in the hosts with gametangia formed at the host surface and with sporophytes up to several mm in height being produced while still attached to the host. To date, none of the kelp gametophytes from nature have been identified to genus or species, although the gametophyte of Nereocystis luetkeana is a potential candidate for the symbiosis. Preliminary observations from Nova Scotia and the Isle of Man have not found the association in the Atlantic Ocean. Laboratory studies in Korea successfully reconstructed the symbiosis in the red alga Aglaothamnion oosumiense using zoospores of Undaria pinnatifida but not Laminaria religiosa. Here we outline the development of the symbiosis and discuss the potential adaptive significance of the kelp/red algal interaction.     

FORECASTED CO2 MODIFIES THE INFLUENCE OF LIGHT IN SHAPING SUBTIDAL HABITAT1

Some abiotic conditions are well known to play disproportionately large roles in shaping contemporary assemblages, yet their roles may not continue to have similar magnitudes of effect into the future. We tested whether forecasted levels of CO₂ could alter the strength of influence of an abiotic factor (i.e., light intensity) well known for its strength of influence on the subtidal ecology of photosynthetic organisms. We investigated these dynamics in two subtidal algal species that form contrasting associations with kelp forests, one negatively associated with kelp canopies (turf‐forming brown algae, Feldmannia spp.) and the other positively associated with kelp as understory (calcifying red crustose algae, Lithophyllum sp.). Using an experimental approach, we assessed the independent and combined effects of [CO₂] (control and elevated) and light (shade, low ultraviolet B [UVB], full light) on growth, recruitment, and relative electron transport rate (rETR). Under control [CO₂], the effects of light corresponded to the relative light environments of the two groups of algae. The influence of light on the percentage cover and biomass of understory crusts was substantially reduced under elevated [CO₂], which caused crusts to grow less. While elevated [CO₂] had the opposite effect of positively influencing turf cover and biomass, it had the same effect of reducing the structuring effects of light and UVB. Hence, if we are to predict the ecological consequences of future CO₂ conditions, the role of contemporary processes cannot be assumed to produce similar effects relative to other processes, which will change with a changing climate.     

Seasonal changes in subarctic sea urchin populations from different habitats

This paper documents seasonal variation in certain sea urchin (Strongylocentrotus polyacanthus) characteristics in habitats of varying environmental conditions. At Shemya Island, Alaska, three habitat types [dense kelp beds, intermediate kelp beds, and algal barrens (low to no foliose algal cover)] were monitored seasonally from September 1995 to August 1996, for live and drift foliose algae. In general, drift algal abundance was greater in areas with more attached kelp, but this varied with season. Along with drift algae, sea urchin density, test size diameter, gonad and nutrition indices, and mobility were seasonally sampled within each habitat. Densities were highest in the algal barrens and lowest in the kelp beds. Seasonally, densities varied between summer/fall, and winter/spring, with lower numbers in the winter/spring. Test size was largest in the kelp habitats when compared to the intermediate or barren sites. Test size was seasonally consistent in the kelp habitats but not in the intermediate or barren sites. Here, test size did vary depending on season (larger urchins were found in winter). The gonad index showed much seasonal variation at the kelp and intermediate kelp sites, but was relatively more stable over time in the barren habitats. Between habitats, gonad and nutrition indices were larger in areas with kelp. Urchin movement varied seasonally between habitats, with more overall movement and variation in barren habitats. These results illustrate the importance of small-scale temporal and spatial variation. Monitoring for 1 year demonstrated that certain parameters varied more in areas of higher foliose algal cover (gonad indices), while other parameters varied more in low kelp areas (test size and movement). These results suggest that studies involving urchins should consider both time of year and overall algal community composition when conducting any type of experimental or monitoring work.     

Nutrient-limited productivity of calcareous versus fleshy macroalgae in a eutrophic,carbonate-rich tropical marine environment

The results of a study of nutrient enrichment with nitrogen (N) and phosphorus (P) on productivity and calcification of fleshy and calcareous algae are reported in this study. Plants were collected from a nearshore eutrophic site in the Florida Keys (USA) and experimentally pulsed during the night with combinations of N and P. After several days of pulsing (7–10 days), net productivity, calcification, and alkaline phosphatase activity (APA), were measured. Productivity of fleshy algae were frequently enhanced by N, P, and N+P, during both summer and winter. Phosphorus limited the productivity of Hydroclathrus clathratus during winter and Ulva spp. during summer, whereas nitrogen limited the productivity of Laurencia intricata during both seasons. During summer, Dictyota cervicornis productivity was not enhanced by any nutrient enrichment. Nitrogen limited the productivity of the three calcareous species Penicillus capitatus, Penicillus dumetosus and Halimeda opuntia during winter and that of H. opuntia during summer. Neither N nor P enrichment increased calcification of calcareous species, and P enrichment greatly inhibited calcification of P. dumetosus during winter. Nutrient enrichment enhanced the productivity of the fleshy species to a greater extent than that of calcareous algae. The seawater DIN:SRP molar ratio was low at our eutrophic study site (molar ratio average of 3:1 during winter and 9:1 during summer) compared to more oligotrophic sites in the Florida Keys, suggesting that in carbonate-rich environments, eutrophication shifts nutrient regulation of productivity from P to N. APA activities of fleshy macroalage were higher than calcareous algae, and rates of all macro algae were 2- to 7-fold higher in summer compared to winter. Productivity was also about 3-fold higher in fleshy compared to calcareous species and about 2-fold higher in summer compared to winter. These results suggest that nutrient enrichment enhances productivity of fleshy algae to a greater extent than that of calcareous algae. Thus, overgrowth of calcareous algae by more opportunistic fleshy forms could reduce carbonate accretion in tropical coastlines experiencing increased eutrophication.     

Relationships between macroalgal functional form groups and substrata stability in a subtropical rocky-intertidal system

The general hypothesis that morphological, physiological, and ecological adaptations of macro algal functional-form groups can be related to the level of disturbance encountered in a natural environment was examined. Two articulated calcareous coralline algae (Amphiroa van-bosseae Lemoine, 24% cover and Corallina frondescens Post. & Rupr. 20%) and one non-articulated coralline alga (Lithophyllum sp., 16%), all late-successional predation-tolerant strategists, comprise most of the community cover on stable bedrock substrata at Punta Las Cuevitas, Sonora, Mexico. Conversely, Ulva rigida C. ag. (26% cover) and a ralfsioid crust (23%), shows to be early-successional opportunistic strategists, cover more of the disturbed boulder habitat. Porolithon sonorense Daws., a stress-tolerant strategist, is uniquely abundant on both substratum types (13% cover on boulders, 10% on bedrock). The sheet-like and filamentous algae, prevalent in the temporally unstable habitat, generally show greater productivity (>2×) than the thicker and calcareous forms conspicuous in the more constant environment. It appears that selection for delicate thalli with high productivities, as well as selection for tougher morphologies having lower photosynthetic rates due to greater proportions of structural tissues, are widespread, divergent evolutionary forces among marine algae. Experiments with captive sea urchins (Echinometra vanbrunti Agassiz), in conjunction with fish-preference data published for some of the same algae studied here, offer strong support for the functional-form model. Parrotfishes, rudderfishes, surgeonfishes, damselfishes and E. vanbrunti, in the Gulf of California, preferentially feed on delicate, early-successional, sheet-like, and filamentous algae, while rejecting or ignoring the more structured, late-successional and calcareous algae. There is no significant (P > 0.05) gradation in calorific content between the first four of the six functional groups (i.e., Sheet-, Filamentous-, Coarsely Branched- and Thick Leathery-Groups), but the mean value for these fleshy forms (2.6 kcal · g ash-free dry wt^?1) is significantly greater than that for the last two groups (0.3 kcal, Jointed Calcareous- and Crustose-Groups). The approach used in this study demonstrates a realistic technique for predicting macrophyte community composition from knowledge of the disturbance levels in a given habitat or the reverse. The form group-disturbance relationship has important implications for future biological monitoring of rocky-inter-tidal and subtidal systems.     

Diet,food preference,and algal availability for fishes and crabs on intertidal reef communities in southern California

Synopsis Herbivory by wide-ranging fishes is common over tropical reefs, but rare in temperate latitudes where the effects of herbivorous fishes are thought to be minimal. Along the west coast of North America, herbivory by fishes on nearshore reefs is largely restricted to a few members of the Kyphosidae, distributed south of Pt. Conception. This paper presents information on natural diets and results from feeding choice experiments for two abundant kyphosids from intertidal habitats in San Diego, California —Girella nigricans andHermosilla azurea, and similar data for the lined shore crab,Pachygrapsus crassipes, which also forages over intertidal reefs. These results are compared with the availability of algae in intertidal habitats measured during summer and winter, on both disturbed and undisturbed habitats. The diets of juveniles ofG. nigricans andH. azurea collected from nearshore habitats were dominated by animal prey (mainly amphipods), but adults of these fishes, andP. crassipes, consumed algae nearly exclusively, with 26, 10, and 14 taxa of algae identified fromG. nigricans, H. azurea, andP. crassipes, respectively. Algae with sheet-like morphologies (e.g.Ulva sp.,Enteromorpha sp., members of the Delesseriaceae) were the principal algae in the diets of the fishes, and calcareous algae (e.g.Corallina sp.,Lithothrix aspergillum) and sheet-like algae (Enteromorpha sp.) comprised the greatest identifiable portion of the shore crab's diet. Feeding choice experiments indicated that the fishes preferred filamentous algae (e.g.Centroceras clavulatum, Polysiphonia sp.,Chondria californica) and sheet-like algae (e.g.Enteromorpha sp.,Ulva sp.,Cryptopleura crispa) over other algal morphologies, whereas the shore crab chose jointed calcareous algae (e.g.Lithothrix aspergillum, Corallina vancouveriensis, Jania sp.) most frequently. The diets and preferences for algae by the fishes were generally most similar to the assemblage of algae available in early successional (disturbed) habitats during summer when sheet-like and filamentous algae are abundant. The shore crab exhibited the opposite trend with a diet more similar to late successional (undisturbed) habitats.     

Harmful algae on tropical coral reefs: Bottom-up eutrophication and top-down herbivory

A conceptual paradigm, the “Relative Dominance Model”, provides the perspective to assess the interactive external forcing-mechanisms controlling phase shifts among the dominant benthic functional groups on tropical coral reefs [i.e., microalgal turfs and frondose macroalgae (often harmful) versus reef-building corals and calcareous coralline algae (mostly beneficial due to accretion of calcareous reef framework)]. Manipulative experiments, analyses of existing communities and bioassays tested hypotheses that the relative dominances of these functional groups are mediated by two principal controlling factors: nutrients (i.e., bottom-up control) and herbivory (i.e., top-down control). The results show that reduced nutrients alone do not preclude fleshy algal growth when herbivory is low, and high herbivory alone does not prevent fleshy algal growth when nutrients are elevated. However, reduced nutrients in combination with high herbivory virtually eliminate all forms of fleshy micro- and macro-algae. The findings reveal considerable complexity in that increases in bottom-up nutrient controls and their interactions stimulate harmful fleshy algal blooms (that can alter the abundance patterns among functional groups, even under intense herbivory); conversely, elevated nutrients inhibit the growth of ecologically beneficial reef-building corals. The results show even further complexity in that nutrients also act directly as either limiting factors (e.g., physiological stresses) or as stimulatory mechanisms (e.g., growth enhancing factors), as well as functioning indirectly by influencing competitive outcomes. Herbivory directly reduces fleshy-algal biomass, which indirectly (via competitive release) favors the expansion of grazer-resistant reef-building corals and coralline algae. Because of the sensitive nature of direct/indirect and stimulating/limiting interacting factors, coral reefs are particularly vulnerable to anthropogenic reversal effects that decrease top-down controls and, concomitantly, increase bottom-up controls, dramatically altering ecosystem resiliencies.     

Antarctic patterns of shallow subtidal habitat and inhabitants in Wilke’s Land

Studies of east Antarctic marine assemblages on hard substrata are rare. In relation to sea-ice breakout, we assessed benthic patterns of habitat and inhabitants between islands and bays at each of two depths (6 and 12 m) across the Windmill Islands coast. Island sites experience sea-ice breakout in the austral spring, while bay sites typically retain sea-ice cover into the summer and in some places the cover is virtually permanent. Composition of assemblages differed between sheltered bays and exposed islands. Islands were dominated by macroalgae, which also varied with depth. Immediately below the ice–foot zone at 6 m, substratum space were monopolised by foliose red (Palmaria decipiens) and foliose brown (Desmarestia sp.) algae, whereas at 12 m large canopies of Himantothallus grandifolius was abundant. The understorey consisted of a mixture of turfs and encrusting red algae at 6 m, and coralline algae at 12 m. Sheltered bays had large areas of sediment/algal complex and no canopy-forming macroalgae. We found more sponges and hydroids in bays, and more brittle stars around islands. Experiments testing factors that covary with exposure and depth in Antarctica, such as light, sedimentation and ice scour are necessary to determine processes that maintain these striking patterns.     

Interactions between herbivorous fish guilds and their influence on algal succession on a coastal coral reef

Herbivory is an important mechanism affecting algal succession, particularly on coral reefs where the relationship between algae and corals is largely controlled by herbivores. However, different functional groups of herbivores may have contrasting effects on succession, which may explain different trajectories of coral reef recovery after disturbance. Here, the effects of different herbivore groups (roving herbivores = foragers and territorial damselfish = farmers) were isolated by a multi-factorial experiment carried out on a coastal coral reef with high macroalgal cover, high farmer densities and relatively low forager abundance. The effects of foragers and farmers were distinguished by monitoring algal succession on settlement tiles placed inside and outside exclusion cages, with orthogonal treatments established inside and outside damselfish territories (with appropriate cage controls). Within 12 months, algal assemblages on ungrazed tiles inside exclusion cages proceeded rapidly from fine filamentous turfs, to corticated algae, to tough erect (e.g. Amphiroa spp.) and foliose (e.g. Peyssonnellidae) calcified algae. Farmers had a dramatic impact on succession, essentially arresting the development of the algal community at a point where it was dominated by palatable filamentous algae of the genus Polysiphonia. Fleshy macroalgae such as Sargassum spp. were excluded from farmer territories. In contrast, foragers did not suppress fleshy macroalgae, but rather, appeared to decelerate succession and promote a relatively diverse assemblage. In contrast to forager-dominated reefs, farmer territories did not appear to function solely as forager exclusion areas or promote algal diversity as a result of intermediate grazing pressure. The relatively strong effects of farmers observed here may represent a future scenario for coral reefs that are increasingly subject to overfishing of large grazing fishes.     

Effects of the El Niño southern oscillation on Turbo torquatus (Gastropoda) and their kelp habitat

Abstract Turbo torquatus (hereafter Turbo) were abundant and patchily distributed, especially in algal dominated habitats in shallow water (less then 10 metres) on rocky reefs in central New South Wales, Australia. Although the assemblage of algae was similar in barrens with and without crevices, Turbo were most abundant in crevices, suggesting that shelter was important. Experimental removal of the kelp canopy resulted in a great decrease in the number of Turbo. This was despite cleared patches containing more filamentous food algae, further highlighting the importance of shelter. The density of Turbo in kelp forests ranged from six to seven per square metre in times of abundance and less then one per square metre at other times over a 12‐year period. Variation in the resource base (i.e. food algae and kelp cover) was strongly linked to the abundance of Turbo. Abundance of Turbo was lowest when the density of adult kelp was low (less than 14 plants per square metre). The condition of kelp was severely affected during the 1997–1998 and 2002 El Niño events and was compromised 2–4 years after each event. These pulse events and related loss of shelter probably contributed to a decline in abundance of Turbo. This model was further supported when Turbo abundance increased with a subsequent increase in the density of kelp.     

A quantitative study of calcareous stream and Tintenstriche algae from the Malham district,Northern England

The algal floras of two limestone streams and two calcareous Tintenstrichen are described and compared. The most important factor governing the distribution and abundance of species was the availability of water. Permanently wet stands, dominated by filamentous blue-green algae (Schizothrix calcicola and Phormidium incrustatum) had a greater species diversity than stands subject to frequent drying which were dominated by coccoid blue-green algae (Gloeocapsa spp.).

Significant correlations were found between S. calcicola and water pH (+ve), total species numbers and pH (+ve), filamentous blue-green algae and aufwuchs thickness (+ve) and Calothrix numbers and rock mass colonized (+ve).

The algal flora of the Tintenstrichen and the streams differed, although both developed upon the same limestone formation. The results are discussed with reference to previous work, substratum stability, aufwuchs structure, water chemistry, light and temperature.     

Modification of the physical environment by an <Emphasis Type="Italic">Ecklonia radiata</Emphasis> (Laminariales) canopy and implications for associated foliose algae

Macroalgal canopies modify their surrounding environment and thereby influence the structure of associated algal assemblages. Canopies can modify many factors that can be hard to separate and, consequently, the importance of individual factors often remains unknown. Experiments were carried out to test the hypotheses that Ecklonia radiata canopies modify light, sediment cover and water motion, and that each of these physical factors separately influence the assemblage of associated foliose algae. We measured light, sediment cover and water motion across six naturally occurring E. radiata densities and found a reduction in light and sediment cover as kelp density increased. The outcome for water motion was inconclusive. We also manipulated each of these three factors, while controlling for the two others, to determine the separate effects of light, sediment cover and water motion on the assemblage of foliose algae. Reduction in light had a strong effect on the foliose assemblage, reducing species richness and biomass. Reduction in sediment cover and water motion did not cause separate effects at the level of the assemblage, but the biomass of individual species of foliose algae indicated both positive and negative effects. We conclude that E. radiata canopies at Marmion, Western Australia, modify at least two factors of their physical environment, light and sediment cover. However, only light is modified to an extent where it has effects at the assemblage-level because, in contrast to the effects of sediment cover and water motion, the direction of responses are consistent among individual species of algae.     

Development of filamentous green algae in the benthic algal community in a littoral sand-beach zone of Lake Biwa

Temporal changes of biomass and dominant species in benthic algal communities were investigated in a littoral sand-beach zone in the north basin of Lake Biwa from December 1999 to September 2000. Chlorophyll-a amounts of benthic algal communities per unit area of the sandy sediments rapidly increased from late April to June. Increases in biomass of the benthic algal communities are considered to result from the propagation of filamentous green algae Oedogonium sp. and Spirogyra sp. The cell numbers of filamentous green algae and chlorophyll-a amounts of benthic algal communities at depths of 30 and 50cm at a station protected by a breakwater in May were significantly higher than those of a station exposed directly to wave activity. Thus, the biomass accumulation of the benthic algal communities seems to be regulated strongly by wave disturbance. The development of filamentous green algae may contribute to the increase in biomass of the benthic algal community and to the changes in seasonal patterns of biomass in the sand-beach zone of Lake Biwa. We consider that the development of the filamentous green algal community in the littoral zone of Lake Biwa is the result of eutrophication.     

Kelp and sea urchin settlement mediated by biotic interactions with benthic coralline algal species

Species interactions can influence key ecological processes that support community assembly and composition. For example, coralline algae encompass extensive diversity and may play a major role in regime shifts from kelp forests to urchin-dominated barrens through their role in inducing invertebrate larval metamorphosis and influencing kelp spore settlement. In a series of laboratory experiments, we tested the hypothesis that different coralline communities facilitate the maintenance of either ecosystem state by either promoting or inhibiting early recruitment of kelps or urchins. Coralline algae significantly increased red urchin metamorphosis compared with a control, while they had varying effects on kelp settlement. Urchin metamorphosis and density of juvenile canopy kelps did not differ significantly across coralline species abundant in both kelp forests and urchin barrens, suggesting that recruitment of urchin and canopy kelps does not depend on specific corallines. Non-calcified fleshy red algal crusts promoted the highest mean urchin metamorphosis percentage and showed some of the lowest canopy kelp settlement. In contrast, settlement of one subcanopy kelp species was reduced on crustose corallines, but elevated on articulated corallines, suggesting that articulated corallines, typically absent in urchin barrens, may need to recover before this subcanopy kelp could return. Coralline species differed in surface bacterial microbiome composition; however, urchin metamorphosis was not significantly different when microbiomes were removed with antibiotics. Our results clarify the role played by coralline algal species in kelp forest community assembly and could have important implications for kelp forest recovery.     

Differentiation of benthic marine primary producers using stable isotopes and fatty acids: Implications to food web studies

A two-dimensional biomarker approach, using stable isotopes (δ¹³C, δ¹⁵N) and fatty acids, was used to evaluate differences both amongst and within benthic primary producer types (seagrass, fleshy red algae, calcareous red algae, brown algae, and seagrass periphyton) that are typical of the nearshore, temperate Australian region. The primary source of variance (as examined by permutational ANOVA) for all biomarkers examined was amongst primary producer types, as opposed to amongst species within type. δ¹³C showed a clear separation (Monte Carlo p < 0.05) between seagrass (range of means = −10.1 to −14.0‰) and macroalgae (−14.6 to −25.2‰), but could not differentiate amongst the algal types examined. Similarly, distinct δ¹⁵N signatures (p < 0.05) were found only for seagrass (range of means = 3.6-4.1‰) versus calcareous red algae (4.6-5.5‰), with all other types overlapping in their mean δ¹⁵N values. In contrast, multivariate analysis of fatty acid data (using Canonical Analysis of Principal coordinates; CAP) distinguished not only between seagrass and macroalgae, but also between red and brown algae (and to a limited extent between the calcareous and fleshy red algal types). The principal unsaturated fatty acids in the samples were C₂₀ polyunsaturates (found primarily in the macroalgae and periphyton), and C₁₈ mono- and polyunsaturates, with high proportions of 18:2n-6 and 18:3n-3 typical of the seagrasses. The C₁₈ monounsaturate 18:1n-7 was one of the most diagnostic compounds for the red algae examined, being present in very low amounts in seagrass and virtually absent in the brown algae. Conversely, brown algae were high in 18:4n-3, with 20:4n-3 particularly diagnostic of the kelp Ecklonia radiata. In contrast to stable isotopes, fatty acids helped distinguish different algal groups, thereby providing support that a two-dimensional approach using stable isotopes and fatty acids is likely to provide the most useful tool to distinguish primary producers in food web structure.