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.     

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.     

PRESENCE OF SPOROPHYLLS IN FLOATING KELP RAFTS OF MACROCYSTIS SPP. (PHAEOPHYCEAE) ALONG THE CHILEAN PACIFIC COAST1

Some species of macroalgae continue to live for extended periods of time after detachment and may even maintain reproductive structures, yet very little is known about this process. Here, we describe the presence of sporophylls (with sporogenous tissues) on floating kelp rafts of Macrocystis spp. along the coast of Chile. Surveys were conducted at nine sites (18–50° S) during austral summer 2002, and floating kelp rafts were seen and collected at seven of these nine sites (between 22 and 50° S). Fifteen (26.8%) of the 56 samples had sporophylls, indicating maintenance of sporophylls after detachment. Some of the kelp sporophytes with reproductive blades showed signs of having been afloat for long periods (indicated by the large size of attached stalked barnacles). Additionally, experiments showed that floating kelps released viable zoospores. To understand the reproductive dynamics of floating kelps, we compared these results with information from attached populations of Macrocystis spp. at nearby coastal sites. In general, attached kelp had higher proportions of sporophylls than floating rafts, suggesting that detachment may negatively affect reproductive status. Nevertheless, floating kelps remained functionally reproductive, suggesting that zoospores may be dispersed via floating rafts. Published reports on other macroalgae indicate that some species (Lessoniaceae, Fucaceae, and Sargassaceae) are fertile and probably release zoospores or zygotes while floating or drifting in ocean currents. Because dispersal distances achieved by spores of most macroalgae are relatively short, release of spores from floating algae may be an alternative mechanism of long‐distance dispersal.     

Production, use and fate of Chilean brown seaweeds: re-sources for a sustainable fishery

Chile is an important producer of brown seaweeds representing 10% of world supply. Landings of Chilean kelp fluctuated between 40,000 t.year⁻¹ in the early ‘80s to 250,000 t.year⁻¹ more recently. Commercialized algae come from natural populations and no mass-cultures of involved species have been established. Four species of brown algae are commercially collected in the country: Lessonia trabeculata, L. nigrescens, Macrocystis pyrifera and M. integrifolia. Since 2000, the demand of alginate sources and food for cultivated abalones dramatically raised the harvesting of these species. Direct evaluations of Lessonia spp. and Macrocystis standing-stocks were made along 700 km of coast in northern Chile. Lessonia spp. estimated populations exceed 900,000 t, whereas M. integrifolia does not exceed 300 t. The insights provided indicate that sustainability of Chilean kelp subjected to intense harvesting would require management programs including the following bio-ecological recommendations: (1) to harvest the entire plant including the holdfast; (2) to harvest plants larger than 20 cm in diameter; (3) to harvest plants sparsely, selecting mayor specimens; (4) rotation of harvesting areas; and (5) for Macrocystis, to cut the canopy 1–2 m from the surface. They must be implemented in a National Program of Kelp Management, elaborated by government, scientists, fisherman, and industry.     

EFFECT OF TEMPERATURE AND GRAZING ON GROWTH AND REPRODUCTION OF FLOATING MACROCYSTIS SPP. (PHAEOPHYCEAE) ALONG A LATITUDINAL GRADIENT1

Macroalgal rafts frequently occur floating in coastal waters of temperate regions of the world’s oceans. These rafts are considered important dispersal vehicles for associated organisms with direct development. However, environmental factors may limit the floating potential of kelp and thereby the dispersal of associated organisms. To examine the effect of water temperature and grazing on growth, reproductive output, and survival of floating Macrocystis spp., experiments were conducted in outdoor tanks during austral summer 2006/2007 at three sites along the Chilean Pacific coast (20° S, 30° S, 40° S). At each site, Macrocystis spp. was maintained individually at three different water temperatures (ambient, ambient − 4°C, ambient + 4°C) and in the presence or absence of the amphipod grazer Peramphithoe femorata for 14 d. High water temperatures (>20°C) provoked rapid degradation of Macrocystis spp. rafts. At moderate temperatures (15°C–20°C), algal survival depended on the presence of associated grazers. In the absence of grazers, algal rafts gained in biomass while grazing caused considerable losses of algal biomass. Algal survival was the highest under cooler conditions (<15°C), where raft degradation was slow and grazer-induced biomass losses were compensated by continuing algal growth. Our results indicate that floating kelp rafts can survive for long time periods at the sea surface, but survival depends on the interaction between temperature and grazing. We suggest that these processes limiting the survival of kelp rafts in warmer temperatures may act as a dispersal barrier for kelp and its associated passengers.     

Latitudinal variation in intertidal algal community structure: the influence of grazing and vegetative propagation

Summary The hypothesis that sea urchin grazing and interactions with turf-forming red algae prevent large brown algae from forming an extensive canopy in the low intertidal zone of southern California was tested with field experiments at two study sites. Experimental removal of sea urchins resulted in rapid algal recruitment. Crustose coralline algae which typically dominate the substratum in areas with dense urchin populations were quickly overgrown by several species of short-lived green, brown and red algae. The removal of urchins also significantly increased the recruitment of two long-lived species of large brown algae (Egregia laevigata and Cystoseira osmundacea at one study site and E. laevigata and Halidrys dioica at the other). The experimental plots at both sites were eventually dominated by perennial red algae.A two-factorial experiment demonstrated that sea urchin grazing and preemption of space by red algae in areas where urchins are less abundant are responsible for the rarity of large brown algae in the low intertidal of southern California. The three dominant perennial red algae, Gigartina canaliculata, Laurencia pacifica and Gastroclonium coulteri, recruit seasonally from settled spores but can rapidly fill open space with vigorous vegetative growth throughout the year. These species encroach laterally into space created by the deaths of large brown algae or by other disturbances. Once extensive turfs of these red algae are established further invasion is inhibited. This interaction of algae which proliferate vegetatively with algae which recruit only from settled spores is analogous to those which occur between solitary and colonial marine invertebrates and between solitary and cloning terrestrial plants.It is suggested that a north-south gradient in the abundance of vegetatively propagating species, in grazing intensity and in the frequency of space-clearing disturbances, may account for latitudinal variation in intertidal algal community structure along the Pacific coast of North America.     

Phenology of the rhodomelarian algae Neorhodomela aculeata and Ceramium kondoi and their survival strategies against herbivorous snails

The seasonal growth and reproductive phenology of Neorhodomela aculeata (Perestenko) Masuda and Ceramium kondoi Yendo, and the food preferences of herbivorous snails were examined to elucidate (i) why snails select the fronds of N. aculeata for their habitat; and (ii) the survival strategies of the two red algae under grazing pressures. The maximal lengths and weights of both algal species were recorded for each season over a 12‐month period beginning with the spring of 2003. C. kondoi grew in length at a faster rate than N. aculeate, whereas the turf alga N. aculeata produced new branches from the tips of broken branches. The reproductive period of C. kondoi was between the spring and summer but the reproductive organs of N. aculeata were observed throughout the year. The algal loss rate of fresh N. aculeata to snails was low but snails had a food preference for N. aculeata when compared to C. kondoi in an artificial food experiment. These results indicate that snails may adapt to chemical compounds characteristic of N. aculeata and that the alga further reduces predation damage by its structural resistance. In conclusion, the survival strategies of C. kondoi appear to be rapid growth, seasonal sexual reproduction, and a delicately branched frond morphology that reduces stable feeding patterns of its predators plus high tissue nitrogen content, whereas the survival strategy of N. aculeata includes regenerative growth responses, structural toughness and chemical defenses while under the grazing pressure of herbivorous snails.     

Contrasting effects of giant kelp on dynamics of surfperch populations

Summary The effect of giant kelp, Macrocystis pyrifera, on the population dynamics of two temperate reef fishes, striped surfperch (Embiotoca lateralis) and black surfperch (E. jacksoni), was examined. Based on an understanding of how particular reef resources influence abundances of the surfperch and of the effect of giant kelp on those resources, we anticipated that Macrocystis would adversely affect populations of striped surfperch but would enhance those of black surfperch. The natural establishment of giant kelp at sites at Santa Cruz Island, California, resulted in the predicted dynamical responses of surfperch. Abundances of striped surfperch declined rapidly when and where dense forests of giant kelp appeared, but showed little change where Macrocystis was continuously absent over the 8 y period of study. Abundances of adult black surperch, which increased following the appearance of giant kelp, were lagged by >1 y because the dynamical response involved enhanced local recruitment. No change in abundance of black surfperch populations was evident at areas without giant kelp.The mechanism by which giant kelp altered the dynamics of the surfperch involved modification of the assemblage of understory algae used by surfperch as foraging microhabitat. Foliose algae (including Gelidium robustum) were much reduced and turf was greatly enhanced following the appearance of Macrocystis; these two benthic substrata are the favored foraging microhabitat for striped surfperch and black surfperch respectively. Populations of both surfperch species tracked temporal changes in the local availability of their favored foraging microhabitat. Thus, while neither species used Macrocystis directly, temporal and spatial variation in giant kelp indirectly influenced the dynamics of these fishes by altering their foraging base. These results indicate that the dynamics of striped surfperch and black surfperch were governed to a large degree by density-dependent consumer-resource interactions. The present work underscores the predictive value that arises from a knowledge of the mechanisms by which processes operate.     

Scales of dispersal among hosts in a herbivorous marine amphipod

Abstract The spatial arrangement of plants may constrain the expression of herbivore preferences due to interaction between the scales of herbivore mobility and vegetation structure. I tested the hypothesis that the presence of the herbivorous marine amphipod Peramphithoe parmerong (Ampithoidae) on a low‐preference, poor‐quality host (the brown alga Padina crassa) was due to limited dispersal and the clumped nature of algal distributions. In contrast to limited dispersal in laboratory assays, amphipods rapidly colonized vacant algae in the field with natural densities being achieved within 1 day. These high rates of colonization did not decline with increasing distance from existing algal beds over the spatial scales separating Pa. crassa from the higher‐quality host algae (Sargassum spp.). Thus, the spatial arrangement of host algae did not interact with herbivore mobility to constrain host choices.     

The introduced alga Undaria pinnatifida (Laminariales,Alariaceae) in the lagoon of Venice

Since its appearance in 1992 in the lagoon of Venice, the brown algae Undaria pinnatifida (kelp) has gradually expanded along the banks of canals both at Chioggia and Venice, becoming the dominant species in the local algal community chiefly from February to July. In Chioggia Island, where another brown seaweed (Sargassum muticum) is present since 1992, the spreading of Undaria reaches a plateau. In Venice Island, instead, the colonisation process is still in progress. During 1999, the kelp has colonised the main canals (e.g. The Grand Canal) and, subsequently, the small inner ones. In order to evaluate the dynamics of substrate re-colonisation by Undaria, two mechanical eradications were carried out during (March) and after the fertile period (July) of the algae. Such experimental manipulations provided evidence of the fast re-colonisation potential of the algae mainly attributable to its efficient reproductive system. Eradication made during the fertile period, in fact, permitted the kelp development during the following year, while re-colonisation has started 2 years later where eradication was performed after the reproductive period. A significant decrease in the surface covered by other species has been observed both in shallow (Ulva rigida, Enteromorpha spp., Antithamnion pectinatum, Chondracanthus acicularis) and deeper areas (Rhodymenia ardissonei) during the period of maximum development of U. pinnatifida. On the basis of the results of this study, the following conclusions can be drawn: (1) the alga U. pinnatifida is continuously expanding in lagoon environments of Venice; (2) in order to limit its spreading, mechanical eradications would be done on a large spatial scale and before the zoospores release; (3) there is suggestive evidence of competition between Undaria and the remaining indigenous algae.     

Feeding preferences of the sandhopper Orchestoidea tuberculata: the importance of algal traits

Factors such as nutritional quality and the secondary metabolite content of food resources have been shown to influence the feeding behavior of herbivores in many marine habitats. For intertidal macroalgae consumers on sandy beaches, the influence of these factors on feeding behavior and the consequences on their performance is poorly understood. In this study, we evaluated the relationships of nutritional quality, chemical defenses (phlorotannins), and the structure of three macroalgal species that form the bulk of imported wrack subsidies to beaches in southern Chile, with the feeding behavior, absorption efficiency, and growth rate of the talitrid amphipod Orchestoidea tuberculata, one of the most abundant organisms in this environment. The amphipods preferred Durvillaea antarctica over Lessonia nigrescens and Macrocystis pyrifera when simultaneously offered fresh pieces of each alga. Similar results were observed when artificial food made of dry powdered algae of each species was provided, suggesting that the structure of these three algae did not influence preference. The performance of amphipods when reared on a diet of a single algal species matched feeding preferences; higher growth rates were observed in treatments with the preferred alga, D. antarctica. These results imply that D. antarctica is a superior food item for O. tuberculata when compared to L. nigrescens or M. pyrifera, and also that the alga’s intrinsic quality (i.e., not structure) may influence dietary preference in these consumers. The higher content of proteins and carbohydrates found in D. antarctica may explain why this macroalga represents better quality food for O. tuberculata. Phlorotannin content did not have obvious negative effects on diet choice or growth, as D. antarctica, the alga with greater content of these secondary metabolites, was preferred and associated with higher growth rates of O. tuberculata. However, it is necessary to emphasize that the low phlorotannins concentrations registered in the three macroalgae species examined in this study, may not have been sufficient to deter O. tuberculata. When the amphipods were fed with each alga individually, they consumed significantly higher quantities of D. antarctica, which suggests that O. tuberculata did not eat more to compensate for the lower nutritional quality of the other algal species in order to maintain growth. Nor was compensation for lower food quality achieved by increasing absorption efficiency. Our results imply that the composition of the macroalgae arriving on the beach can significantly affect the performance and subsequent life history traits of O. tuberculata and by extension other amphipod species.     

Seasonal patterns of food availability: influences on the reproductive output and body condition of the herbivorous crab Grapsus albolineatus

 Rocky shores in Hong Kong experience marked seasonal differences in climate resulting in seasonal changes in macroalgal assemblages. The tropical rocky shore crab, Grapsus albolineatus, feeds selectively on filamentous algae through the year but the abundance of these algae and foliose algae is greatly reduced during the summer when encrusting algae dominate the shores and the crab’s diet. This switch in diet may have implications for the reproductive output of this crab. Standing crop of algae varied greatly through the year, peaking in March and April, when the nutritional quality of the algae was also highest. On the shore, available algal protein and energy were both lowest in July. The crab selected an algal diet rich in nutrients than that available to it on the shore for all months of the year except September to December for protein, and July and August for energy. The input of animal matter considerably increased the protein content of the diet, but made little difference to the energy content. Growth and body condition were greatest during March and June, coinciding with the peak in algal biomass. Storage of nutrients in the hepatopancreas of G. albolineatus commenced in November, coinciding with the increase in biomass and quality of algae on the shores, and then, peaking in May and June, these nutrients were utilised by the reproductive organs during the reproductive season which ran from April to November. The middle of the reproductive season coincided with the period when the standing crop of algae on the shore was at its lowest levels and poorest quality. Cycles of growth, reproduction and storage in G. albolineatus appear to be directly influenced by seasonal patterns of algal food availability. Nutrient storage in the hepatopancreas, during periods when food is nutritionally rich and is most abundant, does not guarantee reproductive success of the crab, but appears to be a prerequisite. Selection of an optimal diet (energy- and protein-rich) in a seasonal environment by ingesting abundant nutrient-rich algal species and the opportunistic consumption of animal matter strongly influences growth and reproductive output of G. albolineatus. Received: 18 March 1996 / Accepted: 17 July 1996     

PHYLOGENETIC AND BIOGEOGRAPHIC AFFINITIES OF THE HALYMENIALES AND RHODYMENIALES (RHODOPHYTA) FROM THE GULF OF MEXICO

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.     

MORPHOGENESIS OF MONOSTROMA OXYSPERMUM (KÜTZ.) DOTY (CHLOROPHYCEAE) IN AXENIC CULTURE,ESPECIALLY IN BIALGAL CULTURE1

The typical morphology of Monostroma oxyspermum (Kütz.) Doty is lost in axenic culture. In synthetic media of the ASP type, it grows as a colony-like mass composed of round cells with numerous rhizoids. Such a mass is a fragile structure which falls apart upon shaking, or slight touch, into small cell-groups and single cells or cells with a long rhizoid. Only temporary saccate or monostromatic fronds appear and reach 1–2 mm in length when grown in enriched seawater media, but disintegrate and become a colony-like mass. The typical morphology is easily restored by adding at specific intervals filtrates of bacterial cultures and supernatant medium from axenic brown and red algal cultures to the basal medium (ASP7), or by reinfecting the Monostroma with an appropriate bacterial flora. Furthermore, the typical morphology in also maintained by bialgal cultures between Monostroma and other axenic strains of various species of seaweeds except the species belonging to the Chlorophyceae. Monostroma thus appears to utilize some substances released by most species of brown and red algae for its typical growth. Active substances released by bacteria, brown and red algae have not yet been identified and purified. However, it is demonstrated that in axenic cultures many species of seaweeds produce active extracellular substances which play an important role in growth and Morphogenesis of other species of seaweeds.     

INDUCTION AND REDUCTION OF ANTI‐HERBIVORE DEFENSES IN BROWN AND RED MACROALGAE OFF THE KENYAN COAST1

Herbivory is particularly intense in tropical benthic communities, suggesting preference of constitutive, rather than inducible, anti‐herbivory defense. The objective of the study was to examine whether anti‐herbivore defenses in the red alga Hypnea pannosa J. Agardh and the brown algae Sargassum asperifolium Hering and G. Martens ex J. Agardh and Cystoseira myrica (S.G. Gmelin) C. Agardh could be induced and subsequently reduced in response to grazing by the amphipod Cymadusa filosa Savigny. During a 14‐day treatment phase, algae were exposed to amphipod grazing or were left ungrazed (control). Subsequently, one subset of algae was used in feeding assays, whereas another was cultivated for additional 14 days without consumers (recovery phase). At the end of each phase, bioassays were conducted to detect defensive traits in terms of differences in consumption rates of grazed and control pieces of live algae and agar‐based food containing nonpolar algal extracts. Consumption of grazed live S. asperifolium and H. pannosa specimens was lower than of control algae. Furthermore, nonpolar extracts of grazed S. asperifolium and C. myrica were less preferred than those from control algae. Defensive responses were exclusively detected after the treatment phase, although strong preference of ungrazed H. pannosa and C. myrica over grazed conspecifics continued throughout the recovery phase. These findings suggest that phenotypic plasticity in anti‐herbivory defense of marine macroalgae 1) might be more common than previously shown, 2) could be switched on and off within 2 weeks, and 3) can be found in nonpolar algal extracts.     

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.     

The food of Ancylus fluviatilis (Müll.), a littoral stone-dwelling,herbivore

Summary A. fluviatilis is a herbivore which usually ingests periphyton. Evidence derived from both field observation and laboratory experiments suggests that the limpet prefers diatoms and that within this group Gomphonema is apparently the most attractive genus. These food preferences were most clearly defined in satiated individuals.Snails had no apparent effect on algal community structures in the field situation and it is suggested that this may either result from adaptability of the periphyton with respect to grazing disturbances, or from low snail density and thus low grazing intensity.The food preferences of A. fluviatilis in particular, and other algal exploiters in general, are discussed in terms of plant defense mechanisms. It is argued that an animal's ability to cope with algal cell walls (either in a mechanical or chemical sense) may be one of the major determinants in the food preference behaviour shown by herbivores feeding on algae.     

Herbivory and patch dynamics on rocky reefs in temperate Australasia: The roles of fish and sea urchins

Sea urchins are widely considered to be the major grazers in temperate subtidal systems, with herbivorous fish being browsers of minor importance. This paper reviews spatial and temporal patterns in these herbivores on rocky reels in temperate Australasia, with the aim of assessing their relative impacts on patch structure and dynamics. Herbivorous fishes are widespread and make up a significant numerical component the reel fish fauna. Sea urchins are also abundant, but not all geographic locations support actively grazing species. Both fish and sea urchins exhibit distinct patterns of distribution among depth strata. Within depth strata, all herbivores are restricted to (sea urchins) or forage preferentially in (fish) particular habitat patches, causing a mosaic of different feeding activities. These patches are either related to specific features of the habitat (e.g. Kelp patches, topography) or behavioural interactions. Foraging by sea urchins and demersal-nesting damselfishes is intense and persistent, whereas in the kelp-feeding fish Odax cyanomelas, foraging reaches greatest intensity at predictable locations during a few months of every year. Many fish and sea urchins consume some algae in preference to others. However, feeding preferences may determine the nature of the impact only in fishes. For sea urchins, preference may occasionally determine the order in which algae are consumed, but at high densities they consume all available macroalgae. Impacts of both types of herbivore on the abundance of algae have been recorded. Some sea urchins (e.g. Evechinus chloroticus, Centrostephanus rodgersii) appear to severely modify biogenic habitat structure by maintaining ‘barrens’ (areas devoid of macroalgae) over long periods. In contrast to this, the effects of fishes may be more transitory (e.g. seasonal impact of Odax cyanomelas on brown algae) or occur at smaller spatial scales (e.g. nest sites maintained by male Parma victoriae) Herbivorous and other fishes appear to respond to spatial patterns in algal distributions, rallier than having it major impact upon them. The relative effects of fish and sea urchins on the long-term dynamics of kelp forests are unknown, hut temporal patterns in herbivore abundance and behaviour, and algal demography arc urgent targets for research.     

<Emphasis Type="Italic">Lacuna vincta</Emphasis> (Mollusca,Neotaenioglossa) herbivory on juvenile and adult <Emphasis Type="Italic">Nereocystis luetkeana</Emphasis> (Heterokontophyta,Laminariales)

Herbivory can be an important factor structuring coastal algal communities. Herbivores may preferentially graze particular algal species or tissue types. Mesograzers, despite their small size, can critically weaken kelp thalli and impact entire kelp beds. We propose that when kelp beds are composed of several kelp cohorts, mesograzers will selectively choose to inhabit younger plants and grazing activities will have a greater impact on younger plants. This study investigated the effects of grazing by the littorinid gastropod, Lacuna vincta, on different age classes of the bull kelp, Nereocystis luetkeana by (1) testing food preference of L. vincta on juvenile, first-year adult, and second-year adult Nereocystis blades in the laboratory, (2) determining substrate (blades of different ages) preference of L. vincta in the laboratory, and by (3) estimating in-situ herbivore abundances and densities on juvenile and adult Nereocystis. Results demonstrated that grazing by L. vincta produced greater damage on juvenile than older Nereocystis tissues. Although L. vincta did not select juvenile versus older kelps as substrate in the laboratory, in situ surveys showed that differences existed between age classes with higher L. vincta densities on juvenile than adult kelp. We conclude that at a local scale, L. vincta can be an important structuring factor in Nereocystis populations due to its high density and grazing ability. Handling editor: K. Martens     

Reproduction and adult longevity of five species of planktonic cyclopoid copepods reared on different diets: a comparative study

1. The nutritional value of a pure algal food, the phytoflagellate Chlamydomonas reinhardtii, and a mixed diet was tested for five planktonic cyclopoid copepods (Acanthocyclops robustus, Cyclops vicinus, Cyclops abyssorum, Mesocyclops leuckarti and Thermocyclops crassus). The algae were offered at high density (4.5 × 10⁵ cells ml^–1; 22.5 mg C l^–1) in a flow-through system. The mixed diet consisted of concentrated natural plankton (rotifers, copepod nauplii, small copepodites and large phytoplankton forms) in the size range 50–150 μm and with a dry mass > 20 mg l^–1. Reproductive parameters (clutch size, interclutch periods, number of clutches produced) and adult longevity were monitored as indicators of nutritive value. 2. All species had a significantly lower reproductive output and a shorter or unchanged adult lifespan on the algal compared with the mixed diet. 3. The species differed considerably in their ability to use algae. Mesocyclops leuckarti produced no clutches with algae, and females died earlier than with the mixed food. Acanthocyclops robustus and C. vicinus produced smaller and fewer clutches, displayed a longer interclutch period and shorter (A. robustus) or similar (C. vicinus) lifespan on the algal food than on the mixed food. Thermocyclops crassus and C. abyssorum produced smaller clutches with the algal food, but interclutch period was shorter with the algal than with the mixed diet (T. crassus) or of the same length with both diets (C. abyssorum). Adult lifespan was the same under both food regimes. Unfed females produced no eggs. 4. The ability to utilize algae, the reproductive output and the reproductive allocation were not related to body size. Acanthocyclops robustus, a species of intermediate size, produced by far the largest clutches and the most eggs per lifetime under both food regimes and invested more in reproduction than the other species. 5. Reproduction was costly. Unfed and non-reproducing females of C. vicinus and C. abyssorum reached the same age or lived longer than fed and reproducing ones. 6. The adult cyclopoids tested are primarily omnivorous, but utilize algae which are frequently sufficient for egg production.