Patterns of spatial variation of nutrient content,epiphyte load and shoot size of Posidonia oceanica seagrass meadows (Mediterranean Sea)

Knowledge of patterns of spatial variability of vegetative development, epiphyte load and nutrient availability in seagrass meadows is essential for the adequate design of research and environmental monitoring programmes. Differences in shoot size, epiphyte load and nutrient content of leaves and epiphytes of the Mediterranean endemic seagrass Posidonia oceanica at spatial scales ranging from metres to hundreds of metres are evaluated using a hierarchical nested sampling design. The size and epiphyte load of P. oceanica shoots and the nitrogen and phosphorus content of leaves and epiphytes were different in most of the spatial scales considered. Sampling efforts concentrated at the metre scale incorporated most of the variability in size, epiphyte load and nutrient content of the leaves and epiphytes of P. oceanica shoots. Epiphyte load showed no correlation with nutrient content in the epiphytes or in the leaves. However, epiphyte load and shoot size were negatively correlated, which suggests that light penetration in the canopy may be a main determinant of epiphyte load.     

Spatio-temporal variation in the structure of a deep water Posidonia oceanica meadow assessed using non-destructive techniques

The Malta‐Comino Channel (Maltese islands, central Mediterranean), supports extensive meadows of the seagrass Posidonia oceanica that in some places extend to a depth of around 43 m, which is rare for this seagrass. To assess spatial and temporal variation in the state of the deeper parts of the P. oceanica meadow with time, data on the structural characteristics of the seagrass meadow at its lower bathymetric limit were collected during the summers of 2001, 2003 and 2004 from four stations (two stations within each of two sites) located at a similar depth, over a spatial extent of 500 m. Shoot density was estimated in situ, while data on plant architecture (number of leaves, mean leaf length, and epiphyte load) were successfully obtained using an underwater photographic technique that was specifically designed to avoid destructive sampling of the seagrass. Results indicated that P. oceanica shoot density was lower than that recorded from the same meadow during a study undertaken in 1995; the observed decrease was attributed to the activities of an offshore aquaculture farm that operated during the period 1995–2000 in the vicinity of the meadow. ANOVA indicated significant spatial and temporal variations in meadow structural attributes at both sites during the 3‐year study; for example, shoot density values increased overall with time at site A; a indication of potential recovery of the meadow following cessation of the aquaculture operations. Lower shoot density values recorded from site B (compared with site A) were attributed to higher epiphyte loads on the seagrass, relative to those at site A. The findings, which include new data on the structural characteristics of P. oceanica occurring at depths >40 m, are discussed with reference to the use of the non‐destructive photographic technique to monitor the state of health of deep water seagrass meadows.     

Architectural characteristics of two bed types of the seagrass Posidonia oceanica over different spatial scales

Seagrass beds occur in various morphological forms, ranging from small patches to continuous meadows. The endemic Mediterranean seagrass Posidonia oceanica forms dense and extensive stands that occur in several different morphotypes, including reticulate (seagrass interspersed with a different habitat type, such as bare sand) and continuous beds. This study, undertaken in the Maltese Islands, examined whether reticulate and continuous P. oceanica beds, located adjacent to each other and at similar depths, had different within-bed architectural characteristics. Five commonly used architectural measures (shoot density, number of leaves per shoot, mean leaf length, mean leaf width and shoot biomass) were measured from P. oceanica shoots collected from the two bed types at three different spatial scales: (1) tens of metres (‘small’ scale); (2) hundreds of metres (‘medium’ scale); and (3) kilometres (‘large’ scale). Results of 2-factor ANOVA (factor 1=bed type; factor 2=sampling locality) carried out at the three spatial scales indicated significant differences between the two bed types in shoot density (P<0.01) and leaf length (P<0.05) at the small scale, and in leaf number (P<0.05) at the large scale. Significant interactions were also apparent for shoot density (at the large scale) and for shoot biomass (at the medium scale). However, the results obtained did not indicate consistent architectural differences between the two P. oceanica bed types over the spatial scales considered. Spatial variations in within-bed architectural characteristics observed were therefore thought to be attributable mainly to the influence of local environmental factors. The findings are discussed with reference to the conservation and management of P. oceanica habitat.     

Inshore-offshore and vertical patterns of phytoplankton biomass and community composition in Santa Monica Bay, CA (USA)

Spatial gradients in biomass and community composition have important consequences for ecosystem structure and function. In this study, small-scale inshore-offshore (1-10 km) and vertical (1-50 m) patterns of microphytoplankton biomass and community composition are described, and the environmental controls of microphytoplankton biomass are evaluated in a coastal ecosystem of the Southern California Bight (SCB). During a two-year period, persistent inshore-offshore gradients in phytoplankton biomass and occasional inshore-offshore gradients in community composition, coincident with regional precipitation, were found, although the strength of the gradients varied between sampling periods. The chlorophyll a maximum was generally present between 15 and 45 m, the cell abundance maximum occurred in surface waters, and there was little evidence of vertical gradients in community composition. Variability in chlorophyll a concentrations was linked to variability in environmental parameters only after some rain and upwelling events. This study demonstrates that inshore-offshore patterns in phytoplankton biomass previously documented at large spatial scales (100-700 km) in the SCB can also persist at smaller scales (1-10 km), although the mechanisms for the gradients are likely to be different at the different spatial scales. The results provide a baseline data set that can be used to focus monitoring and management efforts in the SCB. In particular, this work shows that a limited number of sampling stations are sufficient for phytoplankton monitoring in Santa Monica Bay.     

Harpacticoid copepod response to epiphyte load variations in Posidonia oceanica (L.) Delile meadows

We conducted a field experiment to assess the response of phytal harpacticoids to nutrient‐driven increases of epiphyte load in Posidonia oceanica meadows. First, we evaluated differences in species richness, diversity and assemblage structure of phytal harpacticoids in P. oceanica meadows with differing epiphyte loads. Secondly, we conducted a field experiment where epiphyte load was increased through an in situ addition of nutrients to the water column and evaluated the responses of the harpacticoid assemblages. We predicted that there would be changes in the harpacticoid assemblages as a result of nutrient‐driven increases of epiphyte load, and that these changes would be of a larger magnitude in meadows of low epiphyte load. Our results show that the harpacticoid fauna (>500 μm) present in P. oceanica meadows in the Bay of Palma comprised taxa which are considered phytal and other less abundant ones previously described as sediment dwellers or commensal on other invertebrate species. Nutrient addition had an overall significant effect on epiphyte biomass and on harpacticoid abundance, diversity and assemblage structure, possibly as a response to the increased resources and habitat complexity provided by epiphytes. The abundance of dominant species at each location was favoured by nutrient addition and in some cases correlated with epiphytic biomass, although never strongly. This may indicate that structural complexity or diversity of the epiphytic cover might be more important than the actual epiphytic biomass for the harpacticoid species investigated. More species‐specific studies are necessary to ascertain this and clarify the relationships between harpacticoids and epiphytes in seagrass meadows. To our knowledge, this is the first account of harpacticoid species associated with P. oceanica leaves and the epiphytic community they harbour in the Mediterranean Sea.     

Variability in the structure of epiphytic assemblages of the Mediterranean seagrass Posidonia oceanica in relation to depth

The aim of the present study was to evaluate whether the variability in the structure of the epiphytic assemblages of leaves and rhizomes of the Mediterranean seagrass Posidonia oceanica differed between depths at a large spatial scale. A hierarchical sampling design was used to compare epiphytic assemblages at two different depths (10 and 20 m) in terms of both species composition and abundance and patterns of spatial variability in the Tuscan Archipelago (North Western Mediterranean Sea, Italy). Results showed significant differences in the structure of assemblages on rhizomes and leaves at different depths. These differences were related to species composition and abundance; differences were not significant for total biomass, total percentage cover and percentage cover of animals and algae. Whereas the higher variability was observed among shoots in all the studied systems, patterns of spatial variability at the other spatial scales investigated differed between the two studied depths. Moreover, in the present study, analogous patterns between depths resulted for both the assemblages of leaves and rhizomes, suggesting that factors that change with depth can be responsible for the spatial variability of both the assemblages (leaves and rhizomes), and operate regardless of the microclimatic conditions and the structure of assemblages.     

Composition of epiphytic leaf community of Posidonia oceanica as a tool for environmental biomonitoring

The demand for sensitive biological tools to assess the environmental quality of coastal waters at broad spatial scales is increasing. Many of the tools used are based on the taxonomic composition of biotic assemblages. They usually require a valuable taxonomic expertise while are unique reflecting the overall ecosystem integrity. Here, we evaluate the potential indicator value of several features of the epiphytic community (overall assemblage composition, species richness, and proportion of the main taxonomic groups) growing on the seagrass Posidonia oceanica leaves. We do so by empirically examining their changes along a disturbance gradient where multiple human activities have interactive and cumulative impacts, sampling at different spatial scales and at two different depths (5 and 15 m). Our results show that the specific composition of the epiphytic assemblages (i.e. species composition) closely reflects, in the deep meadows, the combined effects of different anthropogenic stressors along the gradient, showing an integrative and non-specific response. Similarly, an increase in the proportion of hydrozoans, and a decrease in the proportion of rhodophytes and chlorophytes are observed in deep meadows along the gradient. In shallow meadows, grazing and biotic features of the seagrass seem the main forcing factors determining species composition, and therefore masking the response of epiphytes to the deterioration gradient. After address the effect of natural sources of variability (water depth, within- and between-meadow heterogeneity), changes in epiphyte assemblages and in the proportion of hydrozoans, rhodophytes and chlorophytes in relatively deep meadows seem promising monitoring tools for detecting coastal environmental deterioration.     

Temporal and bathymetric variation of epiphyte cover and leaf biomass in a southern Posidonia oceanica (L.) meadow: the case of Mahdia coast,Tunisia

The aim of this research was to study spatial and temporal variation in epiphyte cover and leaf biomass of Posidonia oceanica in Eastern Tunisia. Sampling was conducted at four stations on the Mahdia coast during October and December 2010, and April and August 2011, which correspond respectively to autumn, winter, spring and summer in this area. Posidonia oceanica shoots were collected at two depths (5 and 10 m). Cover of macroinvertebrates and macroalgae was estimated on adult leaves. The results showed that leaf and epiphyte biomasses vary significantly according to sampling date, with the highest values recorded in August. We found a high diversity of epiphytic assemblages on the leaves of P. oceanica with clear qualitative and quantitative dominance of Rhodophyceae compared to other groups, followed by Phaeophyceae. Most epiphyte species on the leaves of P. oceanica in Eastern Tunisia are the same as those in other parts of the Mediterranean Sea. No bathymetrical variation in the epiphytic community was found in our study area, which can be explained by the high levels of water clarity in Mahdia.     

Habitat characteristics and spatial arrangement affecting the diversity of fish and decapod assemblages of seagrass (Zostera marina) beds around the coast of Jersey (English Channel)

Recent research and management plans for seagrass habitats have called for landscape level approaches. The present study examines the spatial utilisation of subtidal seagrass beds by fish and decapods around the coast of Jersey (49°N 02° W). A hierarchical scale of landscape configuration and the plant characteristics of eight seagrass beds were measured and the contributions of these variables as predictors of the properties of the fish and decapod assemblages were evaluated using multiple linear regression models. The results indicated that total diversity had a negative relationship with transect heterogeneity and total species number had a weak negative association with increasing fragmentation. Both total diversity and total species number showed a positive relationship with depth. In fact, in all models of species number and densities, values were higher in deeper seagrass beds. Total decapod density increased with aggregation of seagrass patches within a landscape. In addition to landscape configuration, smaller-scale structural changes in both canopy height and epiphyte load appeared to influence densities of decapod crustaceans. At night, fewer patterns could be explained by the independent variables in the model.     

Photo-library method for mapping seagrass biomass

The creation of seagrass biomass maps by diving/snorkelling is time-consuming and expensive. This paper presents a method for estimating seagrass dry weight using a photo-library of classes of differing seagrass biomass. Field data were collected at seagrass beds in Ngederrak Reef, Palau, in 2006. Photos of 25 × 25 cm quadrats were taken prior to the collection the above-ground biomass for determination of biomass dry weight. Fifteen classes of seagrass biomass and substrate type were identified. The dry weight for each class of seagrass was measured in laboratory. A photo-library was created for biomass classification where each in situ quadrat photo is accompanied with seagrass dry weight of the sample and a photo of the sorted sample taken in laboratory. The photo-library of quadrats was then used to estimate seagrass biomass on photos gathered along 100 m long transects at 2 m intervals. This procedure was conducted by three different observers. The seagrass dry weight estimates were consistent between interpreters even if one of the interpreters had no experience in seagrass research. This approach allows quick collection of seagrass dry weight data over large areas. The method can be used for creating seagrass biomass maps by snorkelling/diving and/or for calibrating and validating biomass maps created by remote sensing.     

The consequences of scale: assessing the distribution of benthic populations in a complex estuarine fjord

Evidence suggests that patterns of benthic community structure are functionally linked to estuarine processes and physical characteristics of the benthos. To assess these linkages for coarse-sediment shorelines, we used a spatially nested sampling design to quantify patterns of distribution and abundance of both macroinfauna and macroepibiota. We examined replicate beach segments within a site (∼1 km), sites within areas of relatively uniform salinity and temperature (∼10 km), and areas (∼100 km) in the two major basins of Puget Sound, Washington. Because slight variations in physical characteristics of a beach can lead to significant alterations in biota, we minimized confounding physical influences by working only in the predominant shoreline habitat type in Puget Sound, a mixture of sand, pebbles and cobbles. Species richness decreased steadily from north to south along gradients of declining wave energy, increasing temperature and decreasing salinity. A few taxa were confined to the South Basin, but many more were found in the North. Most of the variability in population abundance was captured at the smaller spatial scales. Physical conditions tend to become increasingly different with distance among sites. Communities became more different from north to south as species intolerant of more estuarine conditions dropped out. There was significant spatial autocorrelation among populations on neighboring beach segments for 73 of the 172 species sampled. Populations of these benthic species may be connected via dispersal on scales of at least km in Puget Sound. Our results strengthen prior conclusions about the strong linkages between the biota and physical patterns and processes in estuaries. It is important for monitoring and impact-detection studies to account for natural variation of physical gradients across the sampling scales used. Nested, replicated sampling designs can facilitate the detection of environmental change at spatial scales ranging from global (e.g., warming or El Niño), to regional (e.g., estuary-wide changes in salinity patterns), to local (e.g., from development at a site).     

Spatial organization of a sedimentary macrobenthic community located on the West African Equatorial margin

Multiscale variability of a macrobenthic community inhabiting fine sediments on the West African Equatorial margin was assessed during three cruises as part of the BIOZAIRE research program. Spatial scales ranged from 15 to 550 m (within station) and from 1000 to 7000 m (between stations). Principal Coordinates of Neighbour Matrices (PCNM) allowed the detection of multiscale patterns of variability within a location, while a binary coding of the sampling stations was used at broad spatial scales. Significant patterns were observed at 170 and >4000 m. The two spatial scales displayed similar biotic responses, with the Scaphopoda and Bivalvia having patterns that were opposite to the Pholoidae. A spatio-temporal patchwork of environmental variables might explain the observed patterns. Practically, these results emphasis the need to develop balanced and crossed designs among the sources of variation (geography, time, depths).     

Factors influencing landscape pattern of the seagrass Halophila decipiens in an oceanic setting

The scale of landscape pattern formation of an ecological community may provide clues as to the processes influencing its spatial and temporal dynamics. We conducted an examination of the spatial organization of an annual seagrass (Halophila decipiens) in an open ocean setting at two spatial scales and growing seasons to identify the relative influence of external (hurricanes) versus internal (clonal growth) factors. Visual surveys of seagrass cover were conducted over 2 years within three replicate 1 km² study areas each separated by ∼25 km in an inshore–offshore transect along the southwest coast of Florida at depths between ∼10 and 30 m. A towed video sled allowed observations of seagrass cover of 1 m² areas approximately every 6 m over thousands of meters of evenly spaced transects within the study areas (coarse scale). The towed video revealed that 17.5% of the seafloor was disturbed irrespective of location or sample time. Randomly selected 10 × 10 m quadrats within the larger, 1 km² study areas were completely surveyed for seagrass cover by divers at 0.625 m² resolution (fine scale). The coarse-scale observations were tested using both conventional geostatistics and an application of a time-series technique (Runs test) for scale of seagrass cover contiguity. Fine-scale observations were examined using conventional geostatistics and a least squares approach (cumulative logistic).     

Patterns of variation of intertidal species of commercial interest in the Parque Litoral Norte (north Portugal) MPA: Comparison with three reference shores

Marine Protected Areas (MPAs) are world-wide established with the aim of conserving biodiversity and preventing overexploitation of marine organisms. Evaluating the effectiveness of MPAs is needed in order to support and implement their management, but it is complicated by the large natural variability in space and time of distribution and abundance of natural populations. Here, we tested the hypothesis that patterns of total abundance and size-frequency distribution of two intensively harvested intertidal species (the sea urchin Paracentrotus lividus and the mussel Mytilus galloprovincialis) differed between a protected and three reference shores along the rocky coast of north Portugal. Response variables were in terms of mean values and measures of variance at different spatial scales (from centimetres to metres) and over time (along a period of about 12 months). A further comparison involved the estimation of the reproductive potential of sea urchins, quantified as variations of Gonad Index (GI = gonad dry weight/body dry weight × 100) at the scale of shore. Results did not generally support a predictable direct effect of protection, as the total abundance and the abundance of larger individuals of both species and GI did not differ between the MPA and reference shores. However, a considerable temporal and spatial variability at smaller scales was detected for several response variables. Such findings have implications for management of MPAs, highlighting the need for sampling designs properly replicated in space and time, in order to examine their effectiveness, and for considering spatial and temporal heterogeneity of target populations and driving processes as a criterion for their implementation and design.     

海南岛新村湾营养负荷对海菖蒲的影响研究

对海南岛新村湾海草床的优势种——海菖蒲(Enhalus acoroides)的茎枝特征、茎枝密度、生物量和不同组织营养素含量进行了研究,并探讨了海水和沉积物间隙水营养负荷对其的影响。结果表明,(1)海菖蒲的茎枝特征(叶长、宽和枝重)、茎枝密度和地上生物量存在显著的空间差异,这些因子与海水和沉积物间隙水DIN含量呈负相关;(2)海菖蒲TN、TP质量分数随着样区的变化而产生显著差异,海菖蒲TN质量分数与其所在样区海水和沉积物间隙水DIN浓度呈显著正相关;(3)随着水体N负荷的增加,海菖蒲叶单位面积附着藻类生物量显著增加。本次研究的结果表明,网箱养殖引起的营养负荷是导致海草衰亡的潜在原因之一,引起了新村湾网箱养殖区海草床的退化。     

Effects of fish farming on seagrass (Posidonia oceanica) in a Mediterranean bay: seagrass decline after organic loading cessation

The effects of fish farming on a seagrass (Posidonia oceanica) meadow at Fornells Bay, Minorca (Balearic Islands) were studied. Changes in plant and meadow features (e.g. shoot morphology, shoot density, biomass, rhizome growth, nutrient and soluble sugars concentrations…) in three stations along a transect from a disturbed (organic pollution due to fish cultures) to an undisturbed site were assessed. The fish culture had ceased in 1991; however, seagrass decline, already reported in a previous study for the period 1988–1990, was still taking place at the time of sampling (July 1994). Differences between stations were very clear; the station closest to the fish cages showed reduced shoot density, shoot size, underground biomass, sucrose concentration and photosynthetic capacities. In contrast, shoots from the most polluted station showed higher P-concentration in tissues and higher epiphyte biomass than the other two. Since water conditions had recovered completely by the time of the sampling mission, it is proposed that the persistence of the seagrass decline was due to the excess organic matter remaining in the sediment.     

Does Encope emarginata (Echinodermata: Echinoidea) affect spatial variation patterns of estuarine subtidal meiofauna and microphytobenthos?

Foraging macrofauna, such as the sand dollar Encope emarginata, can modify sediment properties and affect spatial distribution patterns of microphytobenthos and meiobenthos at different spatial scales. We adopted a spatial hierarchical approach composed of five spatial levels (km, 100 s m, 10 s m, 1 s m and cm) to describe variation patterns of microphytobenthos, meiobenthos and sediment variables in shallow subtidal regions in the subtropical Paranaguá Bay (Southern Brazil) with live E. emarginata (LE), dead E. emarginata (only skeletons — (DE), and no E. emarginata (WE). The overall structure of microphytobenthos and meiofauna was always less variable at WE and much of variation at the scale of 100 s m was related to variability within LE and DE, due to foraging activities or to the presence of shell hashes. Likewise, increased variability in chlorophyll-a and phaeopigment contents was observed among locations within LE, although textural parameters of sediment varied mainly at smaller scales. Variations within LE were related to changes on the amount and quality of food as a function of sediment heterogeneity induced by the foraging behavior of sand dollars. We provide strong evidence that top-down effects related to the occurrence of E. emarginata act in synergy with bottom-up structuring related to hydrodynamic processes in determining overall benthic spatial variability. Conversely, species richness is mainly influenced by environmental heterogeneity at small spatial scales (centimeters to meters), which creates a mosaic of microhabitats.     

Staggered nested designs to identify hierarchical scales of variability

Staggered nested sampling designs are virtually unknown in biology, but are widely used in manufacturing because they are highly efficient. I used a staggered nested design to address variation in the abundance of an intertidal clam Austrovenus stutchburyi (Gray) at four spatial scales. The scales were addressed by sampling cells containing five samples spaced at 50, 5, 0.5, and 0.05 m from one another. Estimates of variation from staggered nested designs can be made via ANOVA and principal component methods. ANOVA and principal component methods estimated roughly equal variability among those four spatial scales of abundance, and at all but the smallest scales variances were estimated very imprecisely. Staggered nested designs are highly efficient, applicable to any nested design, and deserve to be investigated more widely.