Spatial distribution patterns of the soft corals Alcyonium acaule and Alcyonium palmatum in coastal bottoms (Cap de Creus, northwestern Mediterranean Sea)

Current knowledge on the abundance and distribution patterns of different soft coral species is relatively limited when compared to other benthic suspension feeders such as gorgonians and hard coral species. To overcome this scarcity of information, the distribution patterns of the soft corals Alcyonium acaule and Alcyonium palmatum were investigated in northwestern Mediterranean benthic communities over a wide geographical (60 km of coastline) and bathymetrical (0–70 m depth) extent using a remotely operated vehicle. A. acaule was the most abundant species in the study area with highest recorded density of 18 col m^?2 found at depths of 35–45 m in areas that are directly exposed to strong near-bottom currents. Conversely, A. palmatum was only found as scattered solitary colonies at greater depths in soft bottoms, with maximum density of 2.4 col m^?2. Medium and large colonies of A. acaule were preferentially found on sloping and vertical rocky bottoms where they form dense patches. High-density patches of A. acaule were preferentially found on vertical rocky bottom, while isolated colonies were preferentially observed on coralligenous substrata as well as on flat soft and maërl substrates. A. acaule biomass distribution showed highest values between 40 and 45 m depth, and between 60 and 65 m depth. This suggests that deeper populations are formed by colonies that are bigger than the equivalent shallower ones. Although both species are almost genetically identical, ecologically they are very different. For this reason, conservation plans should consider the differential ecological traits shown by these two soft coral species.     

Widespread mass mortalities of the green sea urchin in Nova Scotia,Canada

Mass mortalities of sea urchins exceeding 84 000 t live weight or 2 900 t organic weight occurred during the autumns of 1980 and 1981 along the southern coast of Nova Scotia, Canada. The kill was nearly complete along 160 km of shoreline (straight-line distance) and was inter-mittent along another 280 km. Sea urchins in a variety of rocky habitats were affected, including areas exposed to and sheltered from ocean swell, with both dense and no macrophyte cover, and from 0–13 m depth. The cause of the mortalities was presumed to be a biological agent and was transferable in the laboratory. Colonization of the habitat by subtidal macroalgae and a subsequent large increase in benthic primary production is expected to follow.     

Heat waves,baby booms,and the destruction of kelp beds by sea urchins

Large populations of sea urchins, Strongylocentrotus droebachiensis (Müller), destroyed kelp beds along the Atlantic coast of Nova Scotia in the 1960's and 1970's. The origin of these large sea urchin populations is not understood. We have investigated the potential influence of variable growth and development of the planktonic larvae of sea urchins (in response to temperature and food abundance) on recruitment of benthic juveniles. The adult sea urchins were collected at Sandy Cove, Digby County, Nova Scotia, Canada, in December 1986. Temperature strongly affected larval size and the growth of the echinus rudiment within the range 3° to 9°C, and larvae grew most rapidly at 14°C. Food abundance had a smaller effect on larval growth, and these effects were apparent only at high temperature. Larvae fed the same concentration of two different algal food species grew and developed similarly. Correspondence between spring temperature variation and qualitative variation in sea urchin recruitment, as well as strong temperature effects on larval growth in culture, and the occurrence of a large, positive temperature anomaly in June 1960, all suggest that temperature effects on larval growth and development may have led to intense sea urchin recruitment in 1960 and the appearance of large adult populations 4 to 6 yr later. This result invites further research.     

Stability of sea urchin dominated barren grounds following destructive grazing of kelp in St. Margaret's Bay,Eastern Canada

Kelp regeneration was observed for the first time in St. Margaret's Bay, Nova Scotia, Canada in an area known to have been devoid of macroalgae for several years. The regeneration was destroyed by sea urchins within 10 months. Experimentally induced kelp regeneration met a similar fate under normal grazing pressure. At the lowest sea urchin biomass and density encountered re-establishment of mature kelp stands seems highly unlikely. The sea urchin population required to suppress kelp regeneration is fed by benthic microalgae. Diatoms and other pioneer algal community species were found in the guts of sea urchins. The mean standing crop of benthic microalgae was found to be, 2.2 g C m^-2 and production estimated as ca 15 g C m^-2yr^-1 at 8m depth. Most of the primary production of St. Margaret's Bay has been lost with the disappearing kelp populations.     

Effects of sea urchin disease on coastal marine ecosystems

Outbreaks of disease in herbivorous sea urchins have led to ecosystem phase shifts from urchin barrens to kelp beds (forests) on temperate rocky reefs, and from coral to macroalgal-dominated reefs in the tropics. We analyzed temporal patterns in epizootics that cause mass mortality of sea urchins, and consequent phase shifts, based on published records over a 42-year period (1970–2012). We found no evidence for a general increase in disease outbreaks among seven species of ecologically important and intensively studied sea urchins. Periodic waves of recurrent amoebic disease of Strongylocentrotus droebachiensis in Nova Scotia coincide with periods when the system was in a barrens state and appear to have increased in frequency. In contrast, following a major epizootic that decimated Diadema antillarum throughout the Caribbean in 1983, subsequent outbreaks of disease were highly localized and none have been reported since 1991. Epizootics of Strongylocentrotus in the NW Atlantic and NE Pacific, and Paracentrotus and Diadema in the eastern Atlantic, have been linked to climate change and overfishing of sea urchin predators. The spatial extent of recurrent disease outbreaks in these species, and the frequency of phase shifts associated with these epizootics, has decreased over time due to the expansion of the macroalgal state and its stabilization through positive feedback mechanisms. Longitudinal studies to monitor disease outbreaks in sea urchin populations and improved techniques to identify causative agents are needed to assess changes in the frequency and extent of epizootics, which can profoundly affect the structure and functioning of coastal marine ecosystems.     

Changing lobster abundance and the destruction of kelp beds by sea urchins

In a study area in Nova Scotia, Canada, abundance of the lobster Homarus americanus decreased by nearly 50% in 14 years. The lobster is a major predator of sea urchins, and during the past 6 years the sea urchin Strongylocentrotus droebachiensis has destroyed 70% of the beds of Laminaria spp. in the area. Implications for management are discussed.     

Comparing community structure on shells of the abalone Haliotis midae and adjacent rock: implications for biodiversity

This paper concerns the effects on biodiversity of depletion of the South African abalone Haliotis midae, which is a long-lived species with a large corrugated shell that provides a habitat for diverse benthic organisms. We compared community structure on H. midae shells with that on adjacent rock at three sites (Cape Point and Danger Point sites A and B) and at two different times of the year at one of these sites. Shells of H. midae consistently supported communities that were distinctly different from those on rock. In particular, three species of non-geniculate (encrusting) corallines, Titanoderma polycephalum, Mesophyllum engelhartii and Spongites discoideus, were all found either exclusively or predominantly on shells, whereas another non-geniculate coralline, Heydrichia woelkerlingii, occurred almost exclusively on adjacent rock. The primary rocky substratum, however, supported a higher number of species than abalone shells. Possible reasons for the differences between the two substrata include the relative age, microtopography and hardness of the substrata; the abundance of grazers on them; and the relative age of different zones of the abalone shell, which support communities at different stages of succession. Diversity on shells was lowest in zones that were either very young or very old, in keeping with the intermediate disturbance hypothesis. The distinctiveness of shell epibiota will increase β diversity despite having a lower α diversity than that of adjacent rock. Decimation of H. midae by overfishing therefore has implications for biodiversity conservation.     

Trophic ecology and fecundity of Fundulus heteroclitus in Chezzetcook Inlet,Nova Scotia

The ontogenetic and seasonal dietary shifts of the mummichog Fundulus heteroclitus (L.), from eelgrass (Zostera marina L.) beds off the shore of Halifax County, Nova Scotia, Canada were examined over a 5 mo period in 1980. Switches to abundant, stationary, or high-calorie prey types were observed in the foraging period. Smaller fish generally took less diverse prey. The indices of foregut fullness varied according to body length and season. The successive 10 mm total-length classes of the mummichog (14 to 101 mm TL) consumed whole prey of increasing size. In comparison with populations living further south, the Nova Scotia fish had a shorter spawning period, lower fecundity, and smaller eggs.     

Drastic decline of an extensive eelgrass bed in Nova Scotia due to the activity of the invasive green crab (Carcinus maenas)

In 2001–2002, there were severe declines of eelgrass (Zostera marina) in estuaries along the Nova Scotia coast of the southern Gulf of St. Lawrence. We examined the relationship between the recent invasion of European green crabs (Carcinus maenas) and the eelgrass decline in Benoit Cove in Tracadie Harbour. The abnormally abundant eelgrass wrack consisted mainly of entire shoots, not the usual blades. Three separate methods yielded similar estimates of the rate of shoot removal from the eelgrass beds (direct quadrat counts, “mark–recapture” of tagged shoots in the eelgrass bed, and rate of shoot appearance in the shore wrack). From 14 July to 8 September 2002, the shoot density in the bed fell by about 75 %. Similar rates of decline occurred in crab enclosures (4.4 crabs m^?2) placed in the eelgrass bed. Green crab foraging, involving the tearing of shoots and the digging of large pits, was the reason for the drastic decline of the eelgrass bed in Benoit Cove. It is clear that an invasion of green crabs to a region where they had not previously existed can both destroy eelgrass beds and restrict their recovery.     

Feeding ecology of the green turtle (Chelonia mydas) at rocky reefs in western South Atlantic

Feeding ecology of green turtles was investigated between January 2005 and April 2008 at Arvoredo Reserve, Brazil (27°17′S, 48°18′W). Data were obtained through the performance of observational sessions, geo-referenced counts, benthic surveys, capture and recapture of individuals, and oesophageal lavages. This protected area was identified as an important green turtle feeding ground, used year-round by juveniles (curved carapace length = 32–83 cm). Turtles fed close to the rocky shores of the area and selected grazing sites commonly at hard-to-reach, near-vertical portions of the rocks. They were less active in cold months, and more abundant at shallow areas of the reef (0–5 m), where their preferred food items occurred. Their diet was dominated by macroalgae species but invertebrates were also present. Their main food item was the red algae Pterocladiella capillacea, which seems to be eaten through periodical cropping of its tips. Observational methods such as the ones applied here could be incorporated to other research programs aiming to understand the relationships between Chelonia mydas feeding populations and their environment.     

Spatial distribution patterns of the gorgonians <Emphasis Type="Italic">Eunicella singularis</Emphasis>, <Emphasis Type="Italic">Paramuricea clavata</Emphasis>, and <Emphasis Type="Italic">Leptogorgia sarmentosa</Emphasis> (Cape of Creus,Northwestern Mediterranean Sea)

Most of the current knowledge on Mediterranean gorgonians is restricted to investigations of those populations found within shallow sublittoral waters, and only limited data are available for populations located below scuba depth. To overcome this lack of information, the occurrence and abundance of the gorgonians Eunicella singularis, Paramuricea clavata, and Leptogorgia sarmentosa were investigated in northwestern Mediterranean benthic communities over a wide geographical (60 km of coastline) and bathymetrical (0–70 m deep) extent using a remotely operated vehicle (ROV). The greatest occurrence and abundances of E. singularis and P. clavata were concentrated in areas that are directly exposed to strong near-bottom currents. E. singularis was the most common and abundant species and displayed great plasticity and amplitude in its environmental preferences. Conversely, P. clavata showed a very patchy distribution that was associated with vertical rocky walls. Only isolated colonies of L. sarmentosa were observed in the study area. Hot spots of abundance of E. singularis and P. clavata were identified below a depth of 40 m, which demonstrates the importance of studying the distribution of benthic species over a wide geographical and bathymetrical extent.     

Ontogenetic shift in foraging habit of ocean sunfish Mola mola from dietary and behavioral studies

The ocean sunfish (Mola mola) is typically considered to feed on gelatinous zooplankton, but reports in the literature describe various benthic organisms being found in their stomachs. This might reflect ontogenetic dietary shift, as little was known about the foraging habit of this species. We examined their foraging habits using dietary analyses in combination with a behavioral study in Iwate, Japan (39°22′N, 141°58′E) from 2009 to 2010. Our stomach content analyses (n = 17, 31–250 cm total length) suggested that small sunfish (<50 cm) feed on benthic crustaceans, but large sunfish (>200 cm) feed on jellyfish. Larger sunfish showed higher values of both carbon and nitrogen stable isotope ratios. Deployment of accelerometers and animal-borne cameras on small sunfish in July (49–58 cm, n = 5) suggested their possibility of feeding, while they stayed near the seabed. This indicates that small sunfish might feed on benthic preys. Deployment of accelero-magnetometers on large sunfish in July (84–164 cm, n = 4) clarified that the large sunfish in July swam back and forth between the surface and deep water (>100 m). Temporary decelerations, which were considered to be associated with feeding of planktonic prey, were observed in deep water. Whereas deployment of accelero-magnetometers on large sunfish in November (105 cm, n = 3) showed several bursts, they swam within the mixed layer (0–100 m), which might be associated with chasing of rapid prey. These results suggest that ocean sunfish have heterogeneous diets depending on their body size and possibly season.     

Temporal changes in the trophic ecology of the asymbiotic gorgonian Leptogorgia virgulata

Gorgonians are widespread sub-littoral benthic suspension feeders in the world oceans. However, data on their trophic ecology and role in benthic–pelagic coupling and biogeochemical cycles remain limited. This study assesses the trophic ecology of Leptogorgia virgulata in the Skidaway River estuary, USA (31.9896N, 81.0242W) by analysing carbon and nitrogen isotopic signatures of its soft tissue and different-sized fractions of particulate organic matter (POM) in its environment. Samples were taken in 5 April, 2 August, and 15 October 2012 and 11 January 2013. Results support a distinct temporal shift in the diet of L. virgulata from the POM fraction <10 µm (i.e. pico- and nanoplankton) to the 10–63-µm fraction (i.e. microplankton). This trophic regime is likely associated with the natural abundance of prey items within these size classes, thus suggesting that L. virgulata may be an opportunistic feeder, and seasonal shifts in food availability in the water column affect its diet. As such small prey items affect the bioenergetics of L. virgulata, it is important to understand the implications of changes in food availability associated with environmental drivers on the physiology and population dynamics of this dominant species in the western Atlantic Ocean.     

Continuous plankton records: Geographical variations in numerical abundance,biomass and production of euphausiids in the North Atlantic Ocean and the North Sea

Geographical variations in the numbers, biomass and production of euphausiids and the contribution of common species to the total are described from samples taken during 1966 and 1967 in the North Atlantic Ocean and the North Sea by the Continuous Plankton Recorder at 10 m depth. Euphausiids were most abundant in the central and western North Atlantic Ocean and the Norwegian Sea. Thysanoessa longicaudata (Krøyer) was numerically dominant. Biomass was greatest in the Norwegian Sea and the north-eastern North Sea where Meganyctiphanes norvegica (M. Sars) accounted for 81 and 59%, respectively, of the total biomass. Production was highest off Nova Scotia and in Iberian coastal waters; the dominant species were T. raschi (M. Sars) in the former area and Nyctiphanes couchi (Bell) in the latter. The mean P:B ratios were correlated with temperature.     

Changes in sea urchin populations after the destruction of kelp beds

In St. Margaret's Bay, Nova Scotia, Canada, there are large areas in which sea urchins (Strongylocentrotus droebachiensis) have eliminated beds of kelp (Laminaria spp.). Sites were identified where destruction of kelp beds had taken place 1, 2, 3, 3.5 and 4 years ago. With increase of time since kelp disappearance, the sea urchins showed decreased growth rate, reduced gonad size, but an increase in numbers resulting from high recruitment rates in the first two years after kelp bed destruction. These sea urchin populations, by their browsing, effectively prevent the regeneration of kelp. There is, as yet, no evidence of the sea urchin populations being starved out to allow the kelp to return.     

Larvae of a colonial ascidian use a non-contact mode of substratum selection on a coral reef

The rate at which larvae successfully recruit into communities of marine benthic invertebrates is partially dependent upon how well larvae avoid benthic predators and settle on appropriate substrata. Therefore, to be able to predict recruitment success, information is needed on how larvae search for settlement sites, whether larvae preferentially settle on certain substrata, and the extent to which there are adequate cues for larvae to find these substrata. This article describes how larvae of the colonial ascidian Diplosoma similis find settlement sites on a coral reef. Direct field observations of larval settlement were made on a fringing reef in Kaneohe Bay, Oahu, Hawaii, between September 1985 and April 1986. A comparison of the substrata that larvae contacted prior to settlement relative to the percentage cover of these substrata on the study reef suggests that larvae are using a non-contact mode of substratum identification to locate suitable settlement sites. This mode of substratum identification allowed 74% of larvae to evade predation by benthic organisms who would otherwise have eaten larvae if they had been contacted. Of those larvae that evaded predation, 88% subsequently settled on the same two substrata upon which most adults are found (dead coral or the green alga Dictyosphaeria cavernosa). This pattern of settlement was probably a result of active selection, since the two substrata cover only 14.4% of the reef's surface and currents had little effect on the direction in which larvae swam. An important contributing factor to the high success rate of larval settlement on suitable substrata was the lack of any temporal decay in substratum preference. It is concluded that for Diplosoma similis larval supply is a sufficient predictor of larval settlement rate. However, for marine invertebrates whose larvae are passively dispersed and exhibit a greater temporal decay in substratum preference, larval settlement should generally have a greater dependency on spatial variation in the abundance of benthic predators and suitable substrata.     

Population structure,northern range limit,and recruitment variation in the intertidal limpet Lottia scabra

The northern range limit of the intertidal limpet Lottia scabra is Cape Arago, Oregon (43°N), where adult survival is excellent, the population is small (<300), and recruitment is low; the range limit may be set by limited recruitment. Between June 2012 and March 2013, 25 sites from the middle of the species range (33°N) to Cape Arago were sampled and population size frequency distributions, densities, and nearest neighbor distances were compared to the amount of rocky and sandy shore and kelp bed size. North and south of 37°N, the densities of new recruits averaged 22 and 86 m^?2, respectively. This shift was associated with the range limit of Macrocystis pyrifera kelp beds; we hypothesize that slower currents in M. pyrifera beds may limit larval dispersal leading to higher recruitment. North and south of 40°N, adult density averaged <1 and 458 m^?2, respectively, with the species absent from many sites to the north. This shift was associated with a sharp drop in the amount of rocky shoreline and an increase in uninhabitable sandy shore. Near the northern range limit, >80 % of the individuals were solitary and may be unable to spawn successfully. Recruitment at Cape Arago was infrequent and likely due to self-recruitment. This study suggests that the range limit was set by the absence of M. pyrifera and too little rocky shore leading to high larval wastage, low settlement, low population densities, and, due to an Allee effect, very small effective population sizes.     

Structure of a Mediterranean cryptobenthic fish community and its relationships with habitat characteristics

In the last 10 years, several studies have been carried out on the fish fauna of the Ustica Island marine reserve, yet no investigation was specifically addressed to the cryptobenthic fish assemblage. The first task of this study, conducted along the shallow rocky reefs of Ustica, was to determine the species composition, diversity and relative density of the resident cryptobenthic fishes. Furthermore, we aimed to assess the effects of some macro- and microscale habitat characteristics on the distribution pattern of fishes. In particular, the effect of predator density was indirectly evaluated by comparing density data collected within and outside the integral reserve zone. Overall, 20 species belonging to Blenniidae, Gobiidae, Tripterygiidae, Scorpaenidae and Gobiesocidae were recorded. Gobius bucchichi, Scorpaena maderensis, Tripterygion delaisi, T. melanurus and T. tripteronotus were the numerically dominant and most common species. The effects of zone (i.e. of predator density), bottom type and depth on species richness, diversity and evenness were not significant. A greater total fish density was observed on stones compared with rocky cliff and plateau, but only in the shallowest depth range. At level of single species, G. bucchichi was more abundant inside than outside the integral reserve, but only on stones and at 0–2 m depth range. Density of G. bucchichi was generally higher on stones than on rocky cliffs or plateau and between 0 and 5 m depth, although these differences were not always significant. T. delaisi was conversely more abundant in the deepest stratum (7–10 m). Canonical analyses demonstrated that bottom type and depth influenced significantly the fish assemblage structure. The observed differences in the assemblage structure relied mainly upon the dominant species. T. tripteronotus was mainly associated with rocky plateau and the intermediate depth range (3–5 m), whereas S. maderensis, T. melanurus and Lipophrys trigloides inhabited preferentially the rocky cliffs. At microscale level, the habitat choice of the investigated species was almost entirely based on whether the substrate was either vegetated or composed of bare rock. T. delaisi and T. tripteronotus were associated with substrata covered by algae, whilst G. bucchichi, S. maderensis and T. melanurus preferred bare rock bottoms. In some species, the electivity indices for the less abundant type of cover, measured at different spatial scale, changed accordingly. For instance, the smaller the size of the sampled area, the higher was the intensity of the association between G. bucchichi and Anemonia viridis.     

Blue mussels, Mytilus edulis, at the edge of the range: population structure, growth and biomass along a salinity gradient in the north-eastern Baltic Sea

The blue mussel, Mytilus edulis, is the most conspicuous animal species in the northern Baltic rocky sublittoral. In the studied area the species lives at the margin of its salinity tolerance. Although dwarfed by the low-salinity conditions, blue mussels in the northern Baltic are very abundant and have a decisive role in the benthic and pelagic ecosystems. We studied abundance, size distribution, biomass and growth rate of blue mussels along a 270 km salinity gradient in the northern Baltic Sea. Samples (n=317, 1998-1999) from moderately exposed and exposed rocky shores at seven study areas were collected in the southern Archipelago Sea in the west and into the central Gulf of Finland in the east, where the species is becoming increasingly rare. The results show a marked decline in mean mussel size and biomass from the saline west to the less saline east. The growth rate also decreased with lower ambient salinity. However, abundance of small mussels was considerably higher in the central and eastern parts of the study area. Vertically, the highest biomass was recorded at intermediate depths (5 and 8 m), being lower at both shallower (3 m) and deeper bottoms (12 m). It is concluded that salinity is the foremost factor determining size structure and growth rate among populations within the area. The results suggest that predation further influences the population structure of blue mussels living at the edge of their range in the central Gulf of Finland ultimately set by their salinity tolerance.     

Respiration and lipid content of the Arctic copepod<Emphasis Type="Italic"> Calanus hyperboreus</Emphasis> overwintering 1 m above the seafloor at 2,300 m water depth in the Fram Strait

In August 2000 high concentrations of the dominant herbivorous copepod Calanus hyperboreus were detected in the Arctic Fram Strait, west of Spitsbergen, 1 m above the seafloor at 2,290 m water depth. Individuals from that layer were sampled by a hyper-benthic net attached to the frame of an epi-benthic sledge. For comparison, the vertical distribution of C. hyperboreus in the water column was studied simultaneously by a multiple opening/closing net haul from 2,250 m depth to the surface. Maximum abundance was found close to the surface with 6.6 and 10.0 ind. m^?3 at 0–50 m and 50–100 m depth, respectively. However, the major fraction of the population (>40%) occurred between 1,000 and 1,500 m depth. In the deepest layer (2,000–2,250 m) abundance measured 2.2 ind. m^?3 and was twice as high as between 100 and 1,000 m depth. In comparison to individuals from surface waters, copepods from the hyper-benthic layer were torpid and did not react to mechanical stimuli. Stage CV copepodids and females from the deep sample contained 4–10% less lipid and showed significantly reduced respiration rates of 0.24 and 0.26 ml O₂ h^?1 g^?1 dry mass (DM) as compared to surface samples (0.49 and 0.43 ml O₂ h^?1 g^?1 DM). All these observations indicate that the hyper-benthic part of the population had already started a dormant overwintering phase at great depth. Based on the lipid deposits and energy demands, the potential maximum duration of the non-feeding dormant phase was estimated at 76–110 days for females and at 98–137 days for CV copepodids, depending on what indispensable minimum lipid content was assumed. In any case, the estimated times could not meet the necessary requirements for a starvation period of >6 months until the next phytoplankton bloom in the following spring. The ecological implications of these results are discussed with respect to the life cycle and eco-physiological adaptations of C. hyperboreus to its high-Arctic habitat.