The effects of sedimentation and light on recruitment and development of a temperate, subtidal, epifaunal community

Sedimentation has often been implicated as a factor in structuring epifaunal assemblages on hard substrata yet there are few studies that empirically test this relationship. This study investigated the effects of light and sedimentation conditions on the initial recruitment and subsequent development of an encrusting community at Lough Hyne, S.W. Ireland. Slate settlement panels were subjected to four different conditions of sedimentation and light. The five replicates of each treatment were photographed at approximately 6-week intervals over 13 months. Values for Brillouin diversity (HB) and number of recruits were greatest for shaded panels sheltered from sediment. Number of species was much higher on downward facing panels relative to upward facing panels, regardless of conditions of sedimentation and light. Sedimentation resulted in reduced cover for some species, such as Lithothamnion spp., whilst others, such as Anomia ephippium, were less affected by this variable. Total space occupied was significantly higher for inverted panels, possibly indicating geotaxic behaviour of some larvae. Number of species on each panel condition was more similar for sets of panels exposed for shorter periods relative to the 13-month experiment, implying that post-settlement mortality may differ between treatments.     

Feasibility of assessing the community composition of prasinophytes at the Helgoland Roads sampling site with a DNA microarray

The microalgal class Prasinophyceae (Chlorophyta) contains several picoeukaryotic species, which are known to be common in temperate and cold waters and have been observed to constitute major fractions of marine picoplankton. However, reliable detection and classification of prasinophytes are mainly hampered by their small size and few morphological markers. Consequently, very little is known about the abundance and ecology of the members of this class. In order to facilitate the assessment of the abundance of the Prasinophyceae, we have designed and evaluated an 18S rRNA gene-targeted oligonucleotide microarray consisting of 21 probes targeting different taxonomic levels of prasinophytes. The microarray contains both previously published probes from other hybridization methods and new probes, which were designed for novel prasinophyte groups. The evaluation of the probe set was done under stringent conditions with 18S PCR fragments from 20 unialgal reference cultures used as positive targets. This microarray has been applied to assess the community composition of prasinophytes at Helgoland, an island in the North Sea where time series data are collected and analyzed daily but only for the nano- and microplankton-size fractions. There is no identification of prasinophytes other than to record them numerically in the flagellate fraction. The samples were collected every 2 weeks between February 2004 and December 2006. The study here demonstrates the potential of DNA microarrays to be applied as a tool for quick general monitoring of this important picoplanktonic algal group.     

Predation and space utilization patterns in a marine epifaunal community

The relative abundance patterns of several sessile epifaunal species occurring subtidally on large artificial substrata (pilings) were examined under experimental conditions involving the manipulation of densities of the echinoid Arbacia punctulata (Lamarck). The foraging activities of this predator could denude the substratum of most species with notable exceptions including the colonial hydroid Hydractinia echinata Fleming and the sponge Xestospongia halichondroides (Wilson). Moreover, these were the only two of the twenty most common species which did not significantly change in relative abundance over the experimental period. Both species had low recruitment rates and were commonly associated with substrata which had been submerged for several years. Neither species aggressively interacted with adjacent spatial competitors but instead, appeared to employ a defensive space utilization ‘strategy’. Provision of unoccupied substrata by Arbacia was apparently the major factor favoring both recruitment and growth of Hydractinia, which covered up to 30% of the area on the oldest pilings. More recently submerged substrata were covered by species such as Schizoporella errata (Waters) which had a much higher recruitment rate but was commonly overgrown by several other species. Recruitment rate, competitive ability, and vegetative growth are discussed in terms of the size of the substratum and the possibility of biased sampling in fouling studies. The widespread introduction of large artificial substrata into the natural environment has considerably altered the structure of the natural habitat and constitutes a potentially important selective force for changes in settlement preferences, especially among species such as Hydractinia which persist and become abundant on these substrata.     

Development of a subtidal epibenthic community: factors affecting species composition and the mechanisms of succession

Summary Macroinvertebrate grazers and temporal variability were found to strongly influence species composition of communities that developed subtidally on plexiglas panels. On panels exposed to the naturally high densities of sea urchins and sea stars, only grazer-resistant algal crusts, a diatom/blue-green algal film and short-lived filamentous algae became abundant. On those panels protected from grazers, however, other algae and sessile invertebrates were also common. Both the effects of grazing and the abundance of individual taxa differed on panels immersed at different times of the year.Resident species also affected subsequent recruitment. Some colonists were found more frequently on panels with established communities than on recently immersed plates. Others became more abundant on younger than on older panels. Considerable small-scale spatial variation in the abundance of species was also found among panels within treatments and appeared to persist throughout the 13 months of the study.I suggest that since the interactions that determine which mechanisms are important in succession occur between individuals (generalized here to species), not between successional stages, factors such as those examined that can determine which species will interact, indirectly determine the mechnaisms that are important in the development of a community. Models that deal with interactions between successional stages may lack the detail neccessary to predict or explain changes in species composition in diverse communities.     

Markov chain analysis of succession in a rocky subtidal community

We present a Markov chain model of succession in a rocky subtidal community based on a long-term (1986-1994) study of subtidal invertebrates (14 species) at Ammen Rock Pinnacle in the Gulf of Maine. The model describes successional processes (disturbance, colonization, species persistence, and replacement), the equilibrium (stationary) community, and the rate of convergence. We described successional dynamics by species turnover rates, recurrence times, and the entropy of the transition matrix. We used perturbation analysis to quantify the response of diversity to successional rates and species removals. The equilibrium community was dominated by an encrusting sponge (Hymedesmia) and a bryozoan (Crisia eburnea). The equilibrium structure explained 98% of the variance in observed species frequencies. Dominant species have low probabilities of disturbance and high rates of colonization and persistence. On average, species turn over every 3.4 years. Recurrence times varied among species (7-268 years); rare species had the longest recurrence times. The community converged to equilibrium quickly (9.5 years), as measured by Dobrushin's coefficient of ergodicity. The largest changes in evenness would result from removal of the dominant sponge Hymedesmia. Subdominant species appear to increase evenness by slowing the dominance of Hymedesmia. Comparison of the subtidal community with intertidal and coral reef communities revealed that disturbance rates are an order of magnitude higher in coral reef than in rocky intertidal and subtidal communities. Colonization rates and turnover times, however, are lowest and longest in coral reefs, highest and shortest in intertidal communities, and intermediate in subtidal communities.     

Helgoland (Germany): hemogenetic study of an island population

24 haemogenetic markers (5 erythrocyte antigenes, 6 polymorphisms of serum proteins, 12 polymorphisms of red cell enzymes) had been studied in up to 80 individuals from the island of Helgoland (Germany). The cluster analysis separates clearly the Helgoland sample from the neighbouring populations as well as from European standard data. This special position is interpreted partly by genetic peculiarities developed in the course of time, partly as a consequence of genetic drift.     

Regional-scale analysis of subtidal rocky shore community

The French monitoring network, REseau BENThique (REBENT), was launched by the Ministry of the Environment in 2003 following the 1999 Erika oil spill. REBENT aimed to acquire baseline knowledge of coastal benthic habitat distributions with a special focus on biological diversity. This study analyzed data from 38 subtidal rocky reef sites collected by a single diving team of marine biologists along the coast of Brittany from 2004 to 2010. At each site, the depth limits of the algal belts were determined between 0 and ?40 m Chart Datum (CD); the flora and fauna compositions and abundances were sampled at ?3 and ?8 m CD. A total of 364 taxa (156 flora and 208 fauna), belonging to 12 phyla, were identified. The results showed that the depth limit and density of kelp beds increased as water turbidity decreased; moreover, several changes in community structure could be related to water turbidity and temperature. Thus, northern and southern Brittany showed strong differences in diversity and structure of the dominant kelp species (Laminaria hyperborea and Saccorhiza polyschides). The results from this kelp habitat composition survey (dominant kelp species and indicator species) provided important information for local pressure assessments, like increases in turbidity. The data also provided a reference that could be useful for detecting changes in coastal water temperatures due to global warming.     

Physical disturbance and community organization on a subtidal cobble bed

This study examines the role of physical disturbance in regulating the development and maintenance of macroalgal community structure in a subtidal cobble bed on the Atlantic coast of Nova Scotia, Canada, after a mass mortality of sea urchins. Displacement by hydrodynamic forces of cobbles with and without attached macroalgae was measured in mark-recovery experiments. Cobbles without macroalgae experienced low levels of overturning or displacement by waves. In contrast, cobbles with large fleshy algae (such as kelps and fucoids) were displaced by drag forces along the seabed during periods of moderately strong wave action. Displacement of these “drifters” occurred once the attached alga reached a critical mass (relative to the weight of the anchoring cobble), which varied among species of differing morphology. The rate of displacement of drifters was an exponential function of the ratio of attached algal biomass to rock weight. The loss of larger erect forms from the cobble bed results in a patchwork of low-lying turfs. Clusters of small boulders, experimentally transplanted to the cobble bed, accumulated a greater biomass of large fleshy algae than surrounding cobbles after one year, likely because of the greater stability of boulders. Algal-mediated hydrodynamic displacement of cobbles appears to be an important form of physical disturbance in the cobble habitat, restricting the types and sizes of algae that can grow on cobbles following an urchin die-off.     

Development of a marine subtidal epibiotic community in Hong Kong: implications for deployment of artificial reefs

A 2-year study was conducted in Hong Kong to examine the effects of substratum, season and length of submersion on the development of a subtidal epibiotic community using four types of settlement panels (concrete, steel, wood and tyre). The season and length of submersion had a strong influence on the total biomass and on community structure while the type of substratum had very little impact on the total biomass or the structure of the epibiotic community. The season of submersion determined the species composition of the newly submerged surfaces. In the spring and summer, tubeworms were the most abundant. In the autumn and winter, barnacles and tunicates dominated. Community succession was not obvious in the first year of submersion as it was intermingled with strong seasonal settlement, growth and death of barnacles and tunicates. In the second year of submersion, green mussels and tunicates settled and grew to occupy most of the panel surfaces, forming an assemblage that was characteristic of climax communities in the local subtidal waters. The results suggest that the type of construction material has limited impact on the development of epibiotic communities on artificial reefs deployed in Hong Kong; the season of submersion may affect community structure in the early successional stage, but not the characteristics of the climax communities. This study indicates that the type of substratum should not be of concern when deploying artificial reefs in the subtidal waters in this region. The design of artificial reefs should focus more on other physical and economical aspects such as durability, flow dynamics, stability, cost, and effects on the ambient environment.     

Scale-dependant control of motile epifaunal community structure along a coral reef fishing gradient

Large-scale fishing is mostly conducted using towed gears that reduce the biomass and diversity of benthic invertebrates. However, it is impossible to differentiate between the physical disturbance effect of towed gears from the effect of fish predator removal upon benthic invertebrate communities. Here we explore the impact of fish removal alone on the community structure of small motile coral reef invertebrates (epifauna) along a subsistence fishing intensity gradient in the Lau group, Fiji. We deployed settlement plates at three areas in each of six fishing grounds and examined the density and class richness of the motile epifaunal communities and the associated algal communities in relation to the structure of fish and benthic communities. Motile epifaunal density was unrelated to fishing intensity. However, at smaller inter-area scale (0.5-10 km) motile epifaunal density was negatively related to plate algal biomass, whereas at the larger inter-fishing-ground scale (4-180 km) motile epifaunal density was positively related to the rugosity (substrate complexity) of the surrounding benthos. The class richness and diversity (Margalef's d) of motile epifaunal communities were negatively related to fishing intensity, but unrelated to grazing intensity, rugosity or algal biomass at either scale. Benthic community structure varied significantly with fishing intensity; hard-coral cover was lower and turf-algal cover was higher at high fishing pressure. The variation in benthic community structure was associated with variation in fish community structure, which in turn varied with fishing intensity. Motile epifaunal community structure upon plates was linked to the structure of the surrounding benthic community, but was not directly linked to the plate algal community. We suggest the decline in richness of the motile epifauna community along the fishing gradient is attributable to either to exploiter-mediated coexistence or the reduction in ‘habitat quality’ of the surrounding benthos. At the large spatial scale substrate complexity is the key determinant of motile epifaunal density, suggesting predation by fishes plays an important structuring role at this scale. Assuming that rugosity is inversely related to predation risk then this study represents the first evidence for spatial-dependence on the top-down (predation) vs. bottom-up (algal biomass) control of community structure. We argue fisheries exploitation, in the absence of a physical disturbance can negatively influence motile epifaunal community structure at large spatial scales.     

Blood pressure in a scottish island community

A survey of mean blood pressure levels in age-and-sex-matched samples of the population of the island of Tiree appears to support the local belief that blood pressure is higher on the island than on the mainland. This conclusion may be related to the observation of a greater mean width of the heart, measured radiologically, in the island population compared with the mainland population. The greater mean body weight observed in the island compared with the mainland may be a factor. Blood pressure in men between the ages of 40 and 59 years is higher in Tiree than in selected samples in England and America.     

The macrofauna and macroflora associated withLaminaria digitata andL. hyperborea at the island of Helgoland (German Bight,North Sea)

This paper describes the macroflora and macrofauna associated with two bull kelp species,Laminaria hyperborea andL. digitata, at the island of Helgoland, North Sea. During a study period of seven months (March–September 1987), 29 macroflora species and 125 macrofauna species were found. The dominant taxonomic groups were Polychaeta (25 species), Bryozoa (17), Amphipoda (14), Hydrozoa (10) and Ascidiae (8). The species maximum was in July. In general,L. hyperborea was preferred as a substrate for settlement toL. digitata. Composition of the communities associated with kelp changed during the season according to exposure to wave action, and according to location on the kelp thallus. The rhizoid community of both kelps bore more species at exposed locations. Wave-exposedL. digitata lacked obvious faunal settlement on both phylloid and cauloid. Phylloid and cauloid ofL. hyperborea were chosen as an attractive substrate at both sheltered and wave-exposed locations, showing an association of encrusting bryozoan and hydrozoan colonies.     

Diversity and abundance of mollusks in the sublittoral epifaunal community of Punta Patilla, Venezuela

The diversity of a sublittoral epifaunal mollusk community of Punta Patilla, Sucre State, Venezuela, was studied from September 1990 to September 1991. We identified 25 species (14 bivalves and 11 gastropods) of mollusks that inhabit gravel, soft sand and bottoms covered by Thalassia testudinum. Total diversity indices were H' = 3.42. J' = 0.74 and 1-D = 0.85. Monthly diversity reached its maximum in March 1991 (3.12 bits/ ind.), June 1991 (2.88 bits/ind.) and September 1991 (2.95 bits/ind.); minimum diversity was recorded in August 1991 (1.20 bits/ind.). A log series model showed a diversity index alpha = 4.56 for species abundance data and alpha = 3.11 for biomass data. The more abundant species were Chione cancellata, Anigona listeri, Chione granulata and Area zebra among the bivalves, and Chicoreus brevifrons, Turritella variegata and Phllonotus pomum among the gastropods (which present maximum biomass). The average total biomass (56.80 g/m2) is low when compared to reports from other tropical zones.     

Long-term changes in the macrozoobenthos around the rocky island of Helgoland (German Bight,North Sea)

The paper briefly summarizes what is known about long-term changes (facts, causes, consequences) in the macrozoobenthos of intertidal and subtidal hard-bottom communities around the island of Helgoland (German Bight, North Sea). There is increasing observational evidence that these communities (spectrum and abundances of species) are changing on a long-term temporal scale. The reasons are diverse and mainly anthropogenic. A shift in North Sea climate towards more oceanic conditions may be among the most important factors driving the recent changes in species spectrum. Many of the species which have been recorded as new to the Helgoland area during the past decade are southern (oceanic) species which may be considered as indicators of a warming trend.Communicated by K. Wiltshire     

Foraging behavior and activity budgets of sea stars on a subtidal sediment bottom community

We dived at regular intervals (8, 12 or 14 h) over periods of 9 to 23 days to quantify the allocation of time to different activities by large adults of the four common sea stars foraging in a sediment bottom community (8 to 11 in depth) in the northern Gulf of St. Lawrence. The relative time spent at different activities was similar for Leptasterias polaris and Asterias vulgaris, the most frequent activity being moving over the bottom (42-46% of the time), presumably in search of prey, secondly being stationary (26-27%), and thirdly digesting prey (17-20%). Capture of prey was less frequent, 13% of the time for L. polaris (mainly digging for bivalves) and 5% for A. vulgaris. Compared to L. polaris and A. vulgaris, Crossaster papposus spent more time being stationary and periods of moving were of shorter duration. Solaster endeca differed from the other sea stars because of the high proportion of time spent digesting prey (40%) and the small proportion of time spent moving (22%). Capture events were not observed for C. papposus and S. endeca, likely because they exclusively fed on epifaunal species, which were rapidly captured. For all four species, the most frequent changes between activities were between moving and being stationary. The number of days per prey eaten (length of a feeding bout) was about 11 for S. endeca, 6.6 for L. polaris, 3.1 for C. papposus and 2.8 for A. vulgaris. All sea stars appear to be energy maximizers that spend most of their time finding, capturing and digesting prey.     

Movement,activity pattern,and rôle of a hermit crab population in a sublittoral epifaunal community

The movements and activity of hermit crabs in a sublittoral soft-bottom community in the North Adriatic Sea (Gulf of Trieste) were monitored by means of observations by a SCUBA diver and time-lapse photography. Tagging experiments showed that the hermit crabs exhibit a pattern of movement which causes them to remain in a defined area for extended periods. During a two-year period, the radius of activity of a crab is estimated to be under 15 m. This leads to a high recovery rate of tagged shells. The average speed of a hermit crab was 2.1 m/h and the average distance travelled per day was 21.6 m as calculated from the analysis of the time-lapse films. This distance is an order of magnitude larger than that estimated from the tagging experiments (1.1 m/day). The activity of the crabs was restricted to the sediment surface between multi-species clumps. The approximate area of the sediment surface traversed by a crab is 0.4 m²/day. Many inter- and intraspecific encounters were observed, particularly among hermit crabs and the brittle stars Ophiothrix quinquemaculata (D.Ch.) and Ophiura lacertosa (Pennant). The daily activity pattern of the crabs is related to the light and dark phases on the sediment surface. The high density and conspicuous activity of the hermit crabs point to the important rôle of the pagurids in the investigated community.     

Effects of periodic hypoxia on mortality, feeding and predation in an estuarine epifaunal community

The York River Estuary, a tributary of the Chesapeake Bay, USA, experiences periodic low oxygen stress (hypoxia), yet epifaunal species form dense communities there. We studied hypoxia tolerance of common epifaunal species in the York River by exposing sessile and mobile epifauna to high and low oxygen concentrations in laboratory aquaria. Mortality in hypoxia varied among species, ranging from 0% to 100%, with trends of decreased tolerance by mobile species relative to sessile species. While most species tested experienced some mortality after being exposed to hypoxia (at 1 mg O(2)/l or 0.5 mg O(2)/l) for 5 days, many species had a median lethal time (LT(50)) in hypoxia greater than 1 week (3 of 6 species at 1 mg O(2)/l and 6 of 14 species at 0.5 mg O(2)/l), the maximum duration of typical hypoxic episodes in the York River, suggesting that hypoxia may cause little mortality for some species in this system. However, hypoxia had sub-lethal effects on behavior in all species tested. Epifaunal animals responded to hypoxia with behaviors that moved them higher in the water column or by entering resting states until hypoxia passed. Feeding and predation by a variety of taxa (the hydroid Obelia bicuspidata, the mud crab Neopanope sayi, juvenile blue crabs Callinectes sapidus, the flatworm Stylochus ellipticus, and the nudibranch Doridella leucolena) decreased during hypoxia, despite varying mortality responses to low oxygen stress, suggesting that short hypoxic episodes may create predation refuges for prey species. At least one highly tolerant species (O. bicuspidata) showed substantially decreased growth in hypoxia. Although relatively high tolerance of hypoxia by many estuarine epifaunal species limits serious disturbance during brief hypoxic episodes, hypoxia's greatest impact on York River epifaunal communities might be through its indirect effects on behavior and predation.