Detached kelps from distant sources are a food subsidy for sea urchins

Trophic subsidies link habitats and can determine community structure in the subsidised habitats. Knowledge of the spatial extents of trophic interactions is important for understanding food webs, and for making spatial management practices more efficient. We demonstrate trophic linkages between detached (drift) fragments of the kelp Ecklonia radiata and the purple sea urchin Heliocidaris erythrogramma among discrete rocky reefs separated by kilometres. Sea urchins were abundant at one inshore reef, where the biomass of drift was usually high. There, sea urchins trapped detached kelp at high rates, although local kelp abundance was low. Most detached kelp present on the reef was retained by sea urchins. Detached seagrass, which was abundant on the reef, was not retained by sea urchins in large quantities. Experiments with tethered pieces of kelp showed that sea urchins only consumed detached fragments, and did not consume attached kelps. Comparisons of the morphology of detached fragments of kelp collected from the inshore reef to attached kelps from reefs further offshore showed that a large proportion (30-95%, varying among dates) of the fragments originated at distant reefs (>/=2 km away). At the inshore reef, the sea urchin H. erythrogramma is subsidised by detached kelps, and detached kelp fragments have been transported across landscapes. Cross-habitat resource subsidies therefore link discrete reef habitats separated by kilometres of non-reef habitat.     

Dispersion and phagokinesis in the echinoid Evechinus chloroticus (Val.)

Movement in the echinometrid sea urchin Evechinus chloroticus (Val.) was investigated in relation to the availability of a preferred food species, the laminarian alga Ecklonia radiata (C. Ag.). Patterns of microhabitat occupancy, dispersion, and movement were described in two habitats: Coralline Flats, dominated by encrusting coralline algae; and Sediment Flats, occupied by turfing corallines, Ecklonia and the mussel Modiolus areolatus (Gould). In both habitats, sea urchins were positively associated with encrusting coralline algae, despite the fact that encrusting corallines occupied a minor proportion of the substratum in the Sediment Flat habitat. Sea urchins were significantly clumped in both habitat types, and more so in sites within the Sediment Flats habitat. Movement was not directional in either habitat, but the magnitude of movement was almost twice as high in sites from the Coralline Flats habitat, where kelp was absent, as in the Sediment Flats sites, where kelp was present. Behaviour in response to the presence of Ecklonia in both habitats was investigated experimentally. In both habitats, urchins exposed to a “drift” kelp plant at a distance of 1–2 m exhibited net movement approximately twice as great as that shown by control urchins, not exposed to a kelp plant. Despite the relatively small distances the urchins were from the kelp, there was no evidence of directionality in movement. We conclude from the present study that further understanding of the feeding behaviour of urchins requires that a distinction be made between attached and drift algae, and between directionality and magnitude of movement, and that the interaction of these and other factors are best investigated under field conditions.     

An investigation of dispersal of the introduced brown alga Undaria pinnatifida (Harvey) Suringar and its competition with some species on the man-made structures of Torquay Marina (Devon, UK)

As part of a study into the ecology of the recently introduced macroalga Undaria pinnatifida in the British Isles, the competitive ability of this adventive kelp with native species was investigated in Torquay Marina (Devon, UK). Sites of introduction were surveyed for the presence of native species filling a similar ecological niche and long-term competition experiments set up over 6 years. Experiments involving clearance, removal and monitoring showed that Undaria has a low competitive ability in the infralittoral and sublittoral of a rocky breakwater. Undaria did not manage to colonise the nearby (200 m) shoreline for at least 7 years from the date of its original introduction to the floating pontoons of Torquay Marina. Undaria did out-compete native species in a marina environment: two kelp species, Laminaria digitata and Laminaria saccharina, and a sea squirt, Styela clava.     

Guluronate content of alginate in Laminaria japonica and Laminaria angustata fronds in laboratory culture

The guluronate (G) content of alginate in the fronds of Laminaria japonica and Laminaria angustata, cultured in the laboratory from zoospore via gametophyte using PESI medium, was determined by the ¹H‐nuclear magnetic resonance spectroscopic method. The G content of alginates in young fronds cultured in various conditions were shown to exceed 55%. These values were remarkably higher than those in field kelp. The G content increased with extending culture period and at low temperature.     

The benthic invertebrates of the Salton Sea: distribution and seasonal dynamics

The Salton Sea, California's largest inland water body, is an athalassic saline lake with an invertebrate fauna dominated by marine species. The distribution and seasonal dynamics of the benthic macroinvertebrate populations of the Salton Sea were investigated during 1999 in the first survey of the benthos since 1956. Invertebrates were sampled from sediments at depths of 2–12 m, shallow water rocky substrates, and littoral barnacle shell substrates. The macroinvertebrates of the Salton Sea consist of a few invasive, euryhaline species, several of which thrive on different substrates. The principal infaunal organisms are the polychaetes Neanthes succinea Frey & Leuckart and Streblospio benedicti Webster, and the oligochaetes Thalassodrilides gurwitschi Cook, T. belli Hrabe, and an enchytraeid. All but Neanthes are new records for the Sea. Benthic crustacean species are the amphipods Gammarus mucronatus Say, Corophium louisianum Shoemaker, and the barnacle Balanus amphitrite Darwin. Neanthes succinea is the dominant infaunal species on the Sea bottom at depths of 2–12 m. Area-weighted estimates of N. succinea standing stock in September and November 1999 were two orders of magnitude lower than biomass estimated in the same months in 1956. During 1999, population density varied spatially and temporally. Abundance declined greatly in offshore sediments at depths >2 m during spring and summer due to decreasing oxygen levels at the sediment surface, eventually resulting in the absence of Neanthes from all offshore sites >2 m between July and November. In contrast, on shoreline rocky substrate, Neanthes persisted year round, and biomass density increased nearly one order of magnitude between January and November. The rocky shoreline had the highest numbers of invertebrates per unit area, exceeding those reported by other published sources for Neanthes, Gammarus mucronatus, Corophium louisianum, and Balanus amphitrite in marine coastal habitats. The rocky shoreline habitat is highly productive, and is an important refuge during periods of seasonal anoxia for Neanthes and for the other invertebrates that also serve as prey for fish and birds.     

Utilization of purple and red sea urchins (Strongylocentrotus purpuratus Stimpson and S. franciscanus Agassiz) as food by the white sea urchin (Lytechinus anamesus Clark) in the field and laboratory

White sea urchins (Lytechinus anamesus Clark) attacked purple (Strongylocentrotus purpuratus Stimpson) and red (S. franciscanus Agassiz) sea urchins at Anacapa Island, California. Densities of white urchins were highest in the deep algal crust-dominated community where up to 6% of purple and 25% of red urchins were being attacked by white urchins. Up to 9% of Lytechinus anamesus in an area were actively eating stronglylocentrotids and usually, more than one white urchin was involved in the attack. In areas with low densities of white urchins, no strongylocentrotids were being attacked.After 36 h in the laboratory, there was no difference in the number of white urchins attacking injured or healthy purple urchins in each of the three experimental densities of white urchins. However, both injured and healthy urchins were attacked by more white urchins in high density. When given a choice between injured purple urchins or fresh kelp, white urchins overwhelmingly chose kelp. Data suggest that white urchins utilize other urchin species as an alternative source of food when more preferred food is absent, but will switch to preferred food should it become available.     

Changes in growth and reproduction of green sea urchins, Strongylocentrotus droebachiensis (Müller), during repopulation of the shallow subtidal zone after mass mortality

Following a disease outbreak that caused mass mortality of green sea urchins, Strongylocentrotus droebachiensis, along the Atlantic coast of Nova Scotia in September 1999, changes in growth and reproduction were monitored over 3.75 years as surviving individuals migrated from deep water to repopulate the shallow subtidal zone at a wave-exposed site. Urchins were sampled at 4 depth strata: at 24 m on a boulder field where the population was unaffected by the disease, at 12 and 16 m on a steeply sloping bedrock ramp, and at 8-10 m along the lower margin of a kelp bed (Laminaria digitata) where urchins formed a grazing front by January 2002. Urchins migrating shoreward from the deep-water refuge responded rapidly to increased algal productivity in the shallows through increased growth and reproduction. Measures of annual increments of skeletal elements (rotules) from urchins across the depth gradient indicated that the fastest growing individuals from the source population formed the grazing front. Urchins in the front reached a larger asymptotic size and produced larger gonads than urchins lower on the ramp. The annual cycle in gonad index showed a pronounced spring spawning period across all depths; a secondary fall spawning was evident at the front and 12 m. The presence of mature, fertilizable ova and short response time to spawning induction in both spring and fall supported the occurrence of two spawning periods.     

Effects of removing sea urchins (Strongylocentrotus droebachiensis): Stability of the barren state and succession of kelp forest recovery in the east Atlantic

Stability properties of the barren state of a kelp forest-sea urchin system were studied in northern Norway. The ability of the sea urchin Strongylocentrotus droebachiensis to maintain high population densities and recover from perturbations, and the succession of kelp forest revegetation, were studied experimentally by reducing the sea urchin density on a barren skerry. Additional information was obtained from community changes following a natural, but patchy, sea urchin mortality that varied between sites. On the barren grounds, high sea urchin densities (30 50 per m²) is maintained by annual recruitment. Severe reductions of sea urchin densities initiated luxuriant kelp growth, while more moderate reductions allowed establishment of opportunistic algae (during spring and early summer), but no kelps. Succession of algal growth, after the severe decline in sea urchin densities, followed a predictable pattern. At first the substrate was colonized by filamentous algae, but within few weeks they were outcompeted by the fast growing kelp Laminaria saccharina. After 3–4 years of the removal experiment, the slower-growing, long-lived kelp L. hyperborea became increasingly dominant. Increased food availability after reduction in sea urchin density led to increased individual growth of the remaining sea urchins. However, the population density did not increase, neither from recruitment nor immigration from adjacent areas with high sea urchin densities. Possibly, early establishment of a dense kelp stand, may represent a breakpoint in the ability of sea urchins to reestablish a barren state. The ability of L. saccharina quickly to invade and monopolize an area may have both positive and negative effects on the succession towards the climax L. hyperborea kelp forest. Competitive interactions may slow the process, but development of a dense stand of L. saccharina will also reduce grazing risk on scattered recruits of the more slowly growing L. hyperborea.     

Production and standing stocks of the kelp Macrocystis laevis Hay at the Prince Edward Islands,Subantarctic

Summary The recently described species Macrocystis laevis Hay is endemic to the Prince Edward Islands. Aerial photographs of Marion Island were used to outline the distribution of the kelp and to assess its cover. M. laevis occurs along the lee shore of the island, between the 5 and 20 m isobaths. Plant densities and gross plant morphology were measured by divers during April/May 1988. Net production was estimated from growth measurements taken in April/May 1988 and 1989 and again during August 1989. The mean biomass of kelp was 0.67 kgC·m^–2 within the kelp beds. Net production was estimated at 7.7 gC·m^–2·d^–1 and 11.5 gC·m^–2d^–1 during the months of April and August respectively. M. laevis had a uniform frond-length frequency distribution, which suggests that only the oldest fronds are lost by wave action or senescence. Based on calculations for M. laevis and Durvillaea antarctica (the two species making up most of the macrophyte biomass) macrophytes are more productive per unit area than the phytoplankton but contribute less to the seas around the Prince Edward Islands by virtue of their small spatial coverage. Neither of the kelps lose much material as particulate or dissolved organic carbon through fragmentation. The extent of grazing on live M. laevis fronds is unknown, and only D. antarctica contributes to a macrofaunal detrital community. The contribution of M. laevis production to the nearshore ecology of the islands seems limited, as we suspect that almost all of its production is exported to the open ocean pelagic system.     

A quantitative scuba-diving survey of the sublittoral macrobenthos at subantarctic Marion Island

Summary A SCUBA-diving survey of the macrobenthos of hard substrata in the sublittoral zone at subantarctic Marion Island was conducted during March and April 1988. Dense beds (12 kg m^–2) of the kelp Macrocystis laevis occur in depths > 5 m. Durvillaea antarctica is found along the infralittoral fringe and Desmarestia rossi and Durvillaea sp. occur in a narrow zone from 3 m–6 m. Under-storey algae (chiefly rhodophytes) tend to decrease in biomass with depth, with mean values of 1.57 kg m^–2 at 5m, 0.75 kg m^–2 at 10m and 0.49 kg m^–2 at 15 m. Encrusting coralline algae are particularly abundant in shallow areas (¯x = 0.92 kg m^–2) but are insignificant in deeper areas. Total biomass of macrozoobenthos increased with depth with mean values of 0.12 kg m^–2 at 5 m, 0.34 kg m^–2 at 10 m and 0.46 kg m^–2 at 15 m. Polychaetes, crustaceans, echinoderms, molluscs, sponges and bryozoans dominated the macrozoobenthos in terms of biomass. Approximately 200 species of macrobenthic animals were recorded and numerically, polychaetes, crustaceans, molluscs, nematodes and echinoderms dominated. The sublittoral benthos at Marion Island is compared with that occurring at other subantarctic and Antarctic islands, in particular, the Kerguelen Island group. Zoogeographic trends and the possible effects of nutrient input from seabird guano are briefly discussed.     

Factors influencing depth distribution of soft bottom inhabiting Laminaria saccharina (L.) Lamour. in Kiel Bay,Western Baltic

The kelp Laminaria saccharina dominates soft bottoms in 4–10 m depth in Kiel Bay. Experimental sporophytes transplanted to 2 and 5 m depth showed the typical annual growth pattern of Laminaria species. Surprisingly, 2 m plants died after the first resting phase, whereas 5 m plants survived and showed outgrowth of a new blade generation. Thalli at both depths were infected with the brown algal endophyte Streblonema aecidioides, with host deformations being significantly stronger in 2 m plants. Growth rates of infected sporophytes were reduced. Exclusion of UV light in 2 m depth resulted in less infected thalli. Discs excised from L. saccharina and cultivated in different photon fluence rates from 10–600 µmol m^–2 s^–1 did not differ in growth rate, photosynthesis or dark respiration. Hence, an exclusion of L. saccharina from shallow depths caused by high light cannot be concluded. We suggest the biological interaction with the endophyte S. aecidioides, amplified by UV light, to be most important for the exclusion of L. saccharina from shallow depths in the western Baltic.     

Overgrazing of Kelp Beds Along the Coast of Norway

The aim of this study was to better understand the down-grazing of kelp beds by sea urchins (Strongylocentrotus droebachiensis) along the coast of Norway. Barren grounds were first observed in sheltered areas along the coast of the counties of Trø ndelag, Nordland and Troms in 1974. In the 1980s, the barren grounds spread to areas more heavily exposed to waves. In the 1990s, the kelp beds were re-established in some localities in southern Trø ndelag, initially in wave-exposed areas. In the northernmost parts of Norway, i.e. the counties of Troms and Finnmark, the barren ground areas may still be increasing. Crabs (Cancer pagurus) and common eiders are the most common predators on urchins. Predation on sea urchins in kelp beds is probably not among the factors that limit the sea urchin populations. Along the coast of Nordland and further north, sea urchins are infected by nematodes, resulting in a low, but significant increase in their mortality. No re-growth of kelp beds has been found in the most infected areas. In the late 1960s and the early 1970s, a high occurrence of echinoderm larvae was observed in deeper waters. This was a period with cold water, which may have caused high recruitment of sea urchins. The bet-hedging life strategy of sea urchins may account for the sudden increase in the size of the populations. In the present paper I propose the hypothesis that higher individual growth rates and higher mortality rates in the south than in the north may explain the decrease in the populations, which may in turn account for the re-growth of kelp in the southern areas.     

Urchin grazing of kelp gametophytes in warming oceans

Sea urchins can cause extensive damage to kelp forests, and their overgrazing can create extensive barren areas, leading to a loss of biodiversity. Barrens may persist when the recruitment of kelp, which occurs through the microscopic haploid gametophyte stage, is suppressed. However, the ecology of kelp gametophytes is poorly understood, and here we investigate if grazing by juvenile urchins on kelp gametophytes can suppress kelp recruitment and if this is exacerbated by climate change. We compared grazing of Ecklonia radiata gametophytes by two species of juvenile urchins, the tropical Tripneustes gratilla and the temperate Centrostephanus rodgersii, at winter (19°C), summer (23°C), and ocean warming (26°C) temperatures for the low-latitude range edge of E. radiata, which is vulnerable to ocean warming. We examined the rate of recovery of gametophytes following grazing and determined whether they survived and formed sporophytes after ingestion by sea urchins. Both T. gratilla and C. rodgersii grazed E. radiata gametophytes, reducing their abundance compared to no grazing controls. Surprisingly, temperature did not influence grazing rates, but gametophytes did not recover from grazing in the ocean warming (26°C) treatment. Gametophytes survived ingestion by both species of sea urchin and formed sporophytes after ingestion by T. gratilla, but not C. rodgersii. These results suggest complex grazer–gametophyte interactions, in which both negative (reduced abundance and poor recovery with warming) and positive (facilitated recruitment) effects are possible. Small grazers may play a more important role in kelp ecosystem function than previously thought and should be considered in our understanding of alternate stable states.     

Do sulphuric acid and the brown alga <Emphasis Type="Italic">Desmarestia viridis</Emphasis> support community structure in Arctic kelp patches by altering grazing impact,distribution patterns,and behaviour of sea urchins?

Macrobenthic community structure and the distribution of the green sea urchin (Strongylocentrotus droebachiensis) were recorded inside and outside (=barrens) of kelp patches (Alaria esculenta) at Kongsfjordneset, Svalbard between August 2002 and October 2006. In manipulative field experiments, conducted at Kongsfjordneset, Svalbard in August 2002, the effect of the presence of the brown seaweed Desmarestia viridis on sea urchin distribution and kelp grazing was determined. Additionally, we studied the effect of sulphuric acid, which is produced and stored by D. viridis, on sea urchin movements in the laboratory at Ny-Ålesund, Svalbard, in May 2006. Sea urchin densities were two- to threefold lower in kelp patches than on barrens. The macrobenthic community inside kelp patches hosted 39% more species and was of different species composition than on barrens. Anchored pieces of the kelp A. esculenta were less consumed when surrounded by D. viridis than non-surrounded conspecifics. Changes in pH affected the behaviour of sea urchins. Exposing sea urchins to 500 μl seawater at pH 7.5 caused them to stop, while the exposure of as little as 25 μl at pH 1 caused sea urchins to move in the opposite direction. Acid-mediated escape responses in sea urchin behaviour suggest chemical protection by D. viridis as an additional mechanism to mechanical protection in the generation of kelp refuges. These results improve our understanding of how isolated kelp beds can persist over a wide range of environmental conditions, like wave-sheltered sites, and suggest that changes in community structure may be in part attributable to altered trophic interactions.     

Notes on the distribution and biology of the deep-sea crab Bathynectes maravigna (Brachyura: Portunidae) in the Mediterranean Sea

Data on the distribution and biology of the deep-sea portunid crab Bathynectes maravigna are reported for the Mediterranean Sea, based on several fisheries research surveys. Densities are low and, therefore, biological data are scarce. In the western Mediterranean, the species is much commoner in Alborán Sea than in the Catalano-Balearic Sea. Occurrences are also scarce in the southern Adriatic and northwestern Thyrrenian Sea, as well as in the Ionian Sea. The Alborán Sea and the seas surrounding the southern Italian peninsula are the areas where densities are the highest. The occurrence depth range was found to be 245–786 m, but most of the occurrences took place deeper than 500 m. Sizes ranged between 9 and 51 mm carapace length in males and between 12 and 51 mm in females. Ovigerous females have been only reported in October–December and March–May. Eighty three percent of both males and females are right-handed. Sexual dimorphism was present in cheliped length with males having longer chelae than females. The species appears to be much commoner in those areas where Atlantic influence is the highest.     

Disentangling the links between habitat complexity and biodiversity in a kelp‐dominated subantarctic community

Habitat complexity is one of the most important factors modulating species diversity. This feature comprises several interrelated attributes, such as number, size, and spatial arrangement of complexity‐forming elements. However, the separate and joint effects of these attributes on diversity and community structure are still not well understood. Here, we assess the relationships between several structural‐complexity attributes of the subantarctic kelp Lessonia flavicans and species richness, total abundance, and structure of kelp‐associated macrobenthic communities. We predicted that longer thalli and larger holdfasts favor greater species richness and total abundance of invertebrate organisms. To test the prediction, an observational sampling program was established in two sites of the Strait of Magellan. Uni‐ and multivariate analyses revealed both positive and negative effects of kelp structural‐complexity attributes on diversity. Holdfast diameter and maximum frond length, followed by thallus wet weight, had the strongest positive fits to species richness and total abundance; the number of stipes, on the other hand, was negatively associated with both response variables. Longer fronds were associated with greater abundances of spirorbid polychaetes. Larger holdfasts supported larger abundances of Nereididae and Terebelidae polychaetes and the limpet Nacella mytilina. Contrarily, kelps with longer fronds and more stipes supported fewer amphipods. In this way, we demonstrate that different dimensions of habitat complexity can have contrasting effects on diversity and community structure, highlighting the fundamental role of multiple dimensions of kelp habitat complexity for local biodiversity.     

Diversity,biogeography and host specificity of kelp endophytes with a focus on the genera Laminarionema and Laminariocolax (Ectocarpales,Phaeophyceae)

Endophytic filamentous brown algae are known to invade stipes and fronds of kelps with potentially negative effects for the hosts. They have simple filamentous thalli and are difficult to identify based on morphology. We investigated the molecular diversity of 56 endophytes isolated from seven different kelp species from Europe, Chile, Korea and New Zealand by sequencing two unlinked molecular markers (5’COI and ITS1). A majority of 49 of the isolated endophytes (88%) belonged to the genera Laminarionema and Laminariocolax. The endophyte Laminarionema elsbetiae was isolated from Saccharina latissima and S. japonica tissues in Europe and Korea, respectively, and showed highly similar sequences in both regions. Three different species of the genus Laminariocolax were identified, the most common of which was L. aecidioides, an endophyte with a worldwide distribution and a broad host range. The other two species, L. tomentosoides and a species described here as Laminariocolax atlanticus sp. nov., were associated with different kelp species in the northern hemisphere and the North Atlantic, respectively. Our results suggest that specific host-endophyte patterns could exist locally, as found in kelps in Brittany where all endophytes isolated from S. latissima were L. elsbetiae, all endophytes isolated from Laminaria digitata were Laminariocolax tomentosoides, and those isolated from Laminaria hyperborea were Laminariocolax atlanticus and L. aecidioides. However, this pattern was not consistent with the results from other places, such as Western Scotland and Helgoland where the same kelp species are present.     

The microcopepod fauna in the Eastern Mediterranean and Arabian Seas: a comparison with the Red Sea fauna

Multiple opening-closing nets of 0.05 mm mesh size were employed to study the community structure and vertical distribution of microcopepods at selected stations in the Red Sea, Arabian Sea and Eastern Mediterranean Sea down to a maximum depth of 1850 m. Calanoids, cyclopoids (Oithona and Paroithona) and poecilostomatoids (mainly Oncaea) were the 3 most abundant orders. In the epipelagic zone (0–100 m), these orders occurred at similar abundance levels, whereas in the meso- and bathypelagic zones the poecilostomatoid genus Oncaea dominated numerically by about 60–80% of all copepodids.The species diversity of Oncaea in the Red Sea is compared with preliminary results from the two adjacent regions. In the deep Eastern Mediterranean Sea, the number of species appears to be similar to that in the deep Red Sea and low as compared to the deep Arabian Sea. In this latter area an extremely speciose Oncaea fauna was found at depth below the oxygen-minimum-zone (900–1850 m). The results are related to the differences in the hydrographic conditions of these 3 areas.     

Frond elongation rates of shallow waterMacrocystis pyrifera (L.) Ag. In northern Baja California,Mexico

Macrocystis pyrifera (L.) Ag.frond elongation rates were measured during autumn-winter, spring and summer in a shallow water (7.5 m depth) kelp bed in Bahía Papalote, northern Baja California. Frond elongation was maximum during spring and minimum during winter. Frond growth was significantly correlated with solar radiation, and was highest in the smallest size fronds (0–2 m). Average frond elongation rate varied between 0.3–11% d^–1 during the study period. The relationship between average frond length and elongation rate followed an exponential curve with a negative slope during autumn-winter and summer, but was best described by a straight line during spring. Standard growth rates were obtained by a graphic method. Standard growth rates had intermediate values between those reported for southern California and southern Baja California.M. pyrifera growth cycle shows a different trend from what has been previously reported.