Effect of canopy removal on a rocky shore community metabolism and structure

The effects of the permanent removal of the canopy-forming alga Fucus serratus was studied in terms of both functioning and diversity on a mid–low rocky shore ecosystem in the south-west English Channel (48°N 43.686′, 3°W 59.282′). Ecosystem functioning was examined as net or gross primary productivity (NPP or GPP) and respiration (Resp) measured through CO₂ fluxes. Diversity was examined as number and composition of species. Measurements were performed in situ, during emersion times, without altering target assemblages. The experiment was designed with two treatments [control (C) or canopy removed (CR)] and five replicates, and was conducted over an 18-month period (from February 2006 to August 2007) to integrate the seasonal variability. The mean GPP and Resp were severely reduced in CR treatment compared to control throughout the survey. The mean NPP was not affected at first, due to the development of opportunistic green algae, but was drastically reduced after 9 months of experiment. The canopy removal affected neither the number of species nor their distribution among trophic groups, and the algal community was only slightly affected. The abundance and biomass of mobile invertebrates, however, were greatly reduced in the absence of canopy. This indicates an important effect of the dominant alga on the higher trophic levels of the community. At this tidal level, the canopy did not seem to affect the community by dampening the environmental stress but by providing food, habitat or both.     

A seasonal stable isotope survey of the food web associated to a peri-urban rocky shore

We characterized the structure and functioning of the food web associated to an intertidal rocky shore in the harbour of Brest (Western Brittany, France), through a seasonal δ¹³C and δ¹⁵N survey. The dominance of benthic-derived organic matter, and particularly resuspended sediment, in the local pool of suspended organic matter, is highlighted. This benthic-derived organic matter dominates the diet of filter-feeders, including a certain degree of selectivity displayed by various taxa belonging to this trophic group. The food web structure appeared very stable temporally, which suggested the dominance of mixed diets, rather than the consumption of pure sources. It was constituted of 4 trophic levels, including a large diversity of predators, likely favoured by the diversity of microhabitats that characterize this intertidal habitat. From our results, the trophic functioning of this peri-urban rocky shore community does not appear functionally simplified as previously reported for other anthropized and/or artificial rocky ecosystems. We suggest that the topological complexity of rocky habitats, including a diversity of microhabitats, might be responsible of the important animal diversity, in spite of the anthropogenic disturbances characterizing this peri-urban area.     

Polychaete assemblages along a rocky shore on the South Adriatic coast (Mediterranean Sea): patterns of spatial distribution

This study reports on the distribution of polychaete assemblages along a depth gradient from 5 to 25 m, along a stretch of rocky coast near Otranto (South Adriatic sea, Italy). Three depths were sampled in May and November 2000. At each depth three sites (about 100 m apart) were sampled by scraping off three replicate quadrats of 20×20 cm. The experimental design enabled identification of across- and along-shore spatial patterns of variation of polychaete assemblages. A total of 4,168 specimens, belonging to 152 taxa were collected. Multivariate analysis showed that the polychaete assemblages differed significantly among depths with the clearest differences between the shallowest sites (5 m) and the deeper ones (15–25 m). A considerable source of variation among sites at each level of the shore was also exhibited by the analyses, with the greatest differences among sites within depths recorded at 5 m. The species most contributing to the differentiation of assemblages among depths and sites within each depth were identified. Some potential causes of the observed differences are hypothesized and discussed. The importance of quantitative observation to provide the context for studying the underlying ecological processes is also stressed.Communicated by R. Cattaneo-Vietti, Genova     

Biomass, diversity and production of rocky shore macroalgae at two nutrient enrichment and wave action levels

The littoral zone of temperate rocky shores is normally dominated by perennial macroalgae (e.g. Fucus, Ascophyllum, Laminaria), but nutrient enrichment and/or permanently decreased wave action may lead to structural community changes from dominance of perennials to increased amounts of annual opportunistic species (mainly green algae). Macroalgal biomass, diversity and production as well as relationships between the two latter were studied using Solbergstrand’s rocky shore mesocosms in SE Norway in connection with a long-term experimental manipulation of nutrient addition and wave action (high and low levels of both factors applied in a crossed way to eight outdoor basins). After more than 2 years of experimental treatment, the total standing stock of macroalgae was larger in low nutrient than in high nutrient treatments as well as in high wave compared to low wave treatments (in autumn only). For macroalgal functional groups, bushy and filamentous brown and filamentous red algae were generally favoured by low nutrient concentrations, while annual filamentous and sheet-like green algae were stimulated by the nutrient enrichment. There was only one significant interaction between nutrient enrichment and wave action (for brown filamentous algae in autumn) and also only one significant main effect of the wave treatment (for bushy brown algae in autumn). Surprisingly, the high nutrient treatments supported a higher diversity of macroalgae, whereas the low nutrient treatments generally showed higher production rates. Moreover, significantly negative correlations were found between macroalgal diversity and primary productivity in both summer and autumn. This study shows that it is the biological components of the communities subjected to external forcing (nutrient addition or decreased wave action) that regulate production and this contradicts the common misperception that resource production in natural systems simply can be fast-forwarded by fertilization. The negative relationships between diversity and productivity, although a consequence of unexpected results for diversity and production, are also novel and hint towards species identities having more important functional consequences than general species dominance patterns and the amount of species per se. These results also emphasise the context dependency of findings within the field of biodiversity and ecosystem functioning.     

Community succession following massive ice-scour on a rocky intertidal shore: recruitment,competition and predation during early,primary succession

The importance of species interactions and recruitment variability was examined during the first year and a half of primary succession (1988–1989) on an exposed rocky seashore near Halifax, Nova Scotia. Previous work suggested that emergent rock on these shores is normally dominated by fucoid rockweeds because predatory whelks control the sessile animal competitors, mussels and barnacles, and because herbivorous littorinids control ephemeral algal competitors. Abundances of all species except seasonal ephemeral algae were very small throughout this experiment and we found no significant effects of carnivory, herbivory, plant-animal competition or plant-plant competition. A slight facilitation of Fucus recruitment is attributed to a thin mat of ephemeral, blue-green algae. Very few other studies have directly manipulated intertidal ephemeral algae. As primary succession may be very rare in this assemblage, these results may be specific to these circumstances, but they highlight the varying importance of species interactions with variable recruitment. In particular, it appears that variations in recruitment success may be important to community structure, even when recruitment is not limited by propagule supply. The scale of the study also provides insight into successional processes occurring after the recent, extensive ice-scour of exposed seashores in this region.     

Carbon emission associated with respiration and calcification of nine gastropod species from the intertidal rocky shore of Western Europe

Intertidal rocky shores are characterized by vertical zonation that results from the interplay between environmental conditions, organism physiology, and species interactions. Metabolism of intertidal organisms is highly variable between species and it changes with vertical position along the intertidal gradient. The present study aimed to quantify the carbon metabolism of nine intertidal rocky shore gastropods, in order to clarify their respective roles in carbon production during emersion and immersion. The influences of monthly temperature variation and tidal level were tested for each species. Analyses were performed in the laboratory using the infrared gas analyzer method for measuring aerial respiration rates, and the dissolved inorganic carbon and total alkalinity technique for measuring aquatic respiration rate and calcification. Hourly carbon fluxes were calculated for the mean annual temperature of 13 °C measured in both air and underwater in the study area. Respiration rates were similar for emersion (8–25 μmol CO₂ g AFDW^?1 h^?1) and immersion (10–23 μmol DIC g AFDW^?1 h^?1). For all species, underwater respiration fluxes were more influenced by monthly temperature variation than by air fluxes, probably as an adaptation to the rapid changes occurring during emersion. Calcification was an important factor influencing annual carbon fluxes for all studied species; every species showed different calcification rates according to its size and position on the intertidal zone. Annual carbon emissions were calculated using the mean immersion/emersion time of each species. Intertidal gastropod carbon emission was primarily influenced by body biomass and their vertical position within the intertidal zone.     

Zooplankton standing stock and community size structure within the epipelagic zone: a comparison between the central Red Sea and the Gulf of Aden

Standing stock and size composition of the zooplankton comunity (>100 m) were studied in four depth strata of the upper 200 m of the water column during a Meteor cruise to the central Red Sea and Gulf of Aden in spring 1987. The central Red Sea was divided into a northern area of higher salinity and a less saline southern part. Both areas exhibited significant differences in zooplankton abundance and standing stock. The latter increased by the ratio 1:2:3 from the northern central Red Sea to its southern part and further south to the Gulf of Aden. For size structure analysis samples were fractionated into three size classes (100 to 300, 300 to 500, 500 to 5000 m). In the central Red Sea the smallest size was dominant whereas in the Gulf of Aden the largest size fraction played a greater relative role than in the central Red Sea. This shift in size structure of the zooplankton community from the Red Sea to the Gulf of Aden is apparently primarily related to ecosystem difference between both areas, leading to a change in species composition. In addition, size reduction of individual species common to both seas may be of some significance in the extreme environment of the Red Sea.     

Dynamic zonation of Nerita plicata,N. undata and N. textilis (Prosobranchia: Neritacea) populations on a rocky shore in Kenya

Vertical zonation of Nerita plicata Linnaeus, N. undata Linnaeus, and N. textilis Dillwyn, inhabiting the rocky cliffs of Nyali, Kenya, was studied through several spring-neap tidal cycles from November to December (northeastern monsoon) 1981 and May to July (southeastern monsoon) 1982. For the first time, a monsoon-related cycle of migration in Nerita plicata and N. undata, superimposed on the shorter term spring-neap cycle is demonstrated. Both species rest predominantly in the upper eulittoral zone and the littoral fringe. N. plicata and N. undata rest at significantly higher levels during the southeastern monsoon (SEM) than during the northeastern monsoon (NEM) (P<0.001). On and around neap-tide days N. plicata, N. undata, and N. textilis occupy significantly distinct mean resting positions (P<0.001) in both seasons; N. undata being the highest in the SEM, followed by N. plicata, with N. textilis always in the lowest level. In the NEM, N. plicata and N. undata sometimes change order. On and around springtide days in both seasons, the upper two species, N. plicata and N. undata, rest close together with no significant difference in their mean resting positions, but with both significantly higher than N. textilis (P<0.001). Both the spring-neap and the monsoon migrations of N. plicata and N. undata are interpreted as either direct or indirect responses to wave action.     

Avian herbivory: an experiment, a field test, and an allometric comparison with mammals

Mechanistic studies on herbivore functional responses have largely taken place in mammals; very little has been done in herbivorous birds so far. Here we aim to fill that gap by experimentally quantifying the (short-term) functional response of a large avian herbivore, the Bewick's Swan (Cygnus columbianus bewickii). We explicitly distinguish between encounter-limited and handling-limited foraging by analyzing the results in the framework of the models of D. E. Spalinger and N. T. Hobbs, originally developed for mammalian herbivory. Bite size in captive swans was experimentally manipulated by varying sward height. The time interval between two bites increased with bite size, which supports the handling-limited model (process 3) and rejects the encounter-limited models (processes 1 and 2). Subsequently, we took the obtained functional response parameters into the field in order to predict, from measurements of sward height, (1) bite sizes, (2) handling times, and (3) short-term intake rates in free-ranging swans. Indeed, for all three variables, the observed values closely matched the experimentally based predictions. Finally, we review functional response parameters available in the literature on avian herbivores and scale them allometrically in relation to mammals. This analysis revealed that maximum bite sizes, and therefore maximum intake rates, in herbivorous birds are smaller than in herbivorous mammals. We hypothesize and provide evidence that birds compensate by longer daily foraging times.     

Impact of sewage on the distribution,abundance and community structure of rocky intertidal macro-organisms

The biological effects of a low-volume domestic sewage discharge were studied near Wilson Cove, San Clemente Island, California (USA), from February to June, 1972. There were fewer species and less cover near the outfall (7 macro-invertebrates, 17.6%; 13 macrophytes, 91.7%) than in nearby “unpolluted” control areas (9 macroinvertebrates, 9.2%; 30 macrophytes, 103.4%). The outfall biota was less diverse than that of the controls, as shown by 5 different diversity indices. A great reduction in community stratification (spatial heterogeneity) and, hence, community complexity occurred near the outfall; this reduction in stratification was primarilly due to the absence of Egregia laevigata, Halidrys dioica, Sargassum agardhianum and Phyllospadix torreyi. These were replaced in the mid-intertidal near the outfall by a low turf of blue-green algae, Ulva californica, Gelidium pusillum and small Pterocladia capillacea, and in the lower intertidal by Serpulorbis squamigerus covered with Corallina officinalis Var. chilensis. A statistically-based determination of assemblages or groups of organisms (i.e., cluster analysis) revealed 3 discrete outfall and 3 discrete control area groups; 3 assemblages contained samples from both areas. The distributional patterns of these groups indicate that near the outfall the degree of dilution of discharged sewage is more important in regulating zonation than is tidal height. The enhancement of the suspension feeder Serpulorbis squamigerus and the omnivores Ligia occidentalis, Pachygrapsus crassipes and Anthopleura elegantissima in the outer fringe of the outfall plume hypothetically is due to their ability to utilize sewage as a food source. A critical effect of the outfall may be to decrease environmental stability thereby favoring rapid-colonizers and more sewage-tolerant organisms. The outfall macrophytes were characterized by relatively higher net primary productivities, smaller growth forms, simpler and shorter life histories, and most were components of early successional stages.     

Allelic esterase isozymes,their variation with season,position on the shore and stage of development in the cirripede Balanus balanoides

Samples of Balanus balanoides (L.) in the Menai Straits were taken from their upper limit of distribution on the shore, from mean low water (MLW) and at three different times of the year, January, May and August. Individual prosomae were homogenised, electrophoresed on polyacrylamide gels and esterase isozymes determined. Group BbEI esterase isozymes fitted the Hardy-Weinberg predicted values for 4 co-dominant alleles in all samples. A comparison of BbEI phenotype frequency distribution of the upper shore and MLW samples showed no significant difference at the 5% level. The phenotype frequency of samples taken at different times of the year were significantly different at the 5% level, due to an excess of allele A in winter and an excess of allele C in summer. It is argued that the seasonal variation of phenotype frequency is probably not due to selection operating on a balanced polymorphism, but that the expression of the alleles through the synthesis of esterase isozymes is dependent on some environmental agent, postibly temperature or day length.     

Patterns of micro- and meiofaunal abundance in marine sediments,measured with the adenosine triphosphate assay

Adenosine triphosphate (ATP) measurements are used to determine vertical and seasonal distributions of microorganisms and meiofauna in sediments from a 14 m-deep mud bottom in central Long Island Sound on 12 sampling dates from April 1975 to October 1976. Below the topmost 1 cm of sediment, ATP measurements can be useful in estimating and comparing standing stocks of microorganisms and meiofauna. In the top 1 cm, however, large quantities of newly settled bivalves (Yoldia limatula, Nucula annulata, and Mulinia lateralis) and juvenile polychaetes (Owenia fusiformis) in summer and fall months account for total ATP concentrations. The ATP content of individual meiofauna ranges from 1.97 ng individual copepod nauplius^-1 to 190.7±60 ng individual M. lateralis ^-1. In general, the total ATP content of individual polychaetes and bivalves is much higher than that of individuals of other groups. However, on a mg ATP per g wet or dry tissue basis, the ATP content of micro- and meiofaunal taxa are not significantly different. In addition to providing a means for comparing micro- and meiofaunal standing stocks, ATP measurements permit examination of the relative contribution of different meiofauna to the total living biomass of meiofauna in sediments. Total sediment ATP concentrations are greatest in the top 1 cm at all seasons, and decrease with increasing depth in the sediment. Annual concentrations in the topmost centimeter average 4.22 g g dry sediment^-1 and range seasonally from 1.09 to 7.64 g g dry sediment^-1. At a depth of 10 cm, values average 0.16 and range from 0.019 to 0.35 g g dry sediment^-1. High ATP concentrations in surface sediment reflect high concentrations of microorganisms and meiofauna at the sediment-water interface. The top 2 cm of sediment contain 71% of all meiofauna, with 41% occurring in the topmost cm. In general, densities are lowest in the winter and highest during the spring and summer, averaging 490 individuals 10 cm^-2, and varying from 87 to 1366 individuals 10 cm^-2. Because of wide variation in recruitment patterns of the benthos in Long Island Sound, the extremes of the range in meiofaunal densities can be observed in the same month in two different years. In order to monitor and compare standing stocks of organisms less than 1 mm in size in sediments, the ATP assay can save hours of processing time compared with alternate methods such as direct counts.     

Method of estimating the standing stock of Trochus niloticus incorporating Landsat satellite data,with application to the trochus resources of the Bourke Isles,Torres Strait,Australia

The habitat of Trochus niloticus in the Bourke Isles, Torres Strait, was classified into areas of a Landsat image, using high-ratio values of green (Band 2) to red (Band 3) light, along the windward reef margins. These shallow-water (< 15 m) areas have a coral and rubble/algal pavement cover, which constitutes the optimal habitat for this gastropod. The habitat was sampled to estimate the abundance of T. niloticus. The proportion of commercial-sized individuals was estimated by measuring the basal width of all individuals in a sample. A multistage sample design incorporating three spatial scales -100 m² (transect), 1500 m² (site) and 1 km² (reef) — was used to provide variance estimates for sample-design optimisation and to provide data on the spatial variation of abundance. Most variation (68%) in abundance was within reefs and was attributable to differences in reef cover. Variations in abundance and time costs for sampling 2 and 4 m transects were compared; the 2 m transect was more efficient than the 4 m transect. The abundance estimates were combined with habitat-area estimates and the proportion of commerical-sized individuals was estimated at a standing stock of 186000 (24% precision), or 14 t of commerical-sized T. niloticus.     

Thermal tolerance of larvae of Pollicipes elegans, a marine species with an antitropical distribution

For the antitropical gooseneck barnacle Pollicipes elegans, population-specific physiological temperature tolerance of larvae may serve as a barrier to larval dispersal across the warmest regions of the tropical Pacific Ocean. Thermal tolerance ranges of larvae of three different populations of P. elegans sampled in 2011 and 2012 (Mexico [MX], El Salvador [ES], and Peru [PE]) were investigated by measuring three indicators of physiological performance: swimming activity, oxygen consumption, and lethality or LT₅₀. The thermal tolerance profiles, which include measurable optimum (maximum aerobic performance), pejus (“getting worse”) and pessimum (worst aerobic performance) ranges, of larvae from the three populations were consistent with their characteristic environmental temperatures. In MX, larvae live close to the upper border of their optimum during warm months and so have a limited capacity to tolerate higher-than-normal temperatures. Larvae from the ES population likewise appear to live within their optimum temperature range, but these larvae lack a detectable pessimum range, suggesting they would be unable to cope with temperatures above their pejus range. Larvae from PE have a broad optimum but no pejus range. Different thermal tolerance ranges provide strong evidence for population-dependent physiological adaptations in P. elegans. For the southern (PE) and northern (MX) P. elegans populations, high tropical temperatures are likely to be a strong direct physiological barrier to larval survival and dispersal, which is in contrast to the more thermally tolerant ES population.     

A simple spatially explicit ideal-free distribution: a model and an experiment

This investigation presents a simple spatially explicit analysis of the ideal-free distribution. The traditional ideal-free distribution assumes discrete sites with definite boundaries, and predicts how many individuals should occupy each site. In contrast, the present analysis assumes that a forager’s gains gradually decline with distance from a site, and asks where in space individuals ought to be. Although many interesting situations may arise, the analysis asks how individuals should position themselves as the distance between two identical sources increases. Nash equilibrium positions should follow a pitchfork pattern as the distance between sites is increased; that is, an individual should maintain a position between two sources when they are close together but should move nearer one of the sources when they are far apart. In addition, the text describes an experimental study that parallels the theoretical analysis. The experiment supports the predicted pitchfork pattern, and provides somewhat weaker support for the predicted differences in ”individual” and ”paired” pitchforks. Received: 14 June 2000 / Revised: 20 September 2000 / Accepted: 7 October 2000     

Interlinked temporal changes in environmental conditions, chemical characteristics of sediments and macrofaunal assemblages in an estuarine intertidal sandflat (Seto Inland Sea, Japan)

Five field surveys were conducted in an estuarine intertidal sandflat of the Seto Inland Sea (Japan) between April 1994 and April 1995. Chlorophyll a, pheopigments, total organic carbon and acid-volatile sulphides (AVS) of surface and subsurface sediments, and macrofaunal assemblages were investigated in parallel at 15 stations. Monthly hydrological data of low-tide creek water adjacent to the flat were used as a complementary environmental characterisation of the study area. Strong temporal changes were found among sampling dates, most remarkably in autumn with a major increase of algal detritus and AVS, a sharp reduction in macrofaunal abundances and species richness, and a massive mortality of the clam Ruditapes philippinarum. This dystrophic event was preceded by a photoautotrophic and hypertrophic spring–summer characterized by abundant fresh (i.e., living) algal material, including microphytobenthos and macroalgae (Ulva sp.). In summer, abundant macrofaunal assemblages reached the highest biomass values (455 g wet weight m⁻² or 60.6 g ash free dry weight m⁻²), with a major contribution of filter-feeding bivalves Musculista senhousia and R. philippinarum. These are among the highest values reported in the literature for sedimentary shores. From autumn, there was a progressive recolonisation of macrofauna, initiated by few opportunistic polychaetes (e.g., Cirriformia tentaculata and Polydora sp.), apparently promoting a fast sediment recovery in winter, and followed by new bivalve recruits in the next spring. This study provides the first evidence of significant and interlinked within-year changes in chemical characteristics of sediments and macrofaunal assemblages in an estuarine intertidal flat at a small spatial scale (i.e., tens of meters). This demonstrates the high temporal variability of species–environment relations in these systems and a close relationship in seasonally driven trophodynamic processes among primary producers and benthic consumers. We conclude that a thorough parallel evaluation of the temporal changes in chemical characteristics of sediments should be taken into account in assessing the year-round distribution and changes of intertidal macrofauna, particularly in eutrophic, estuarine intertidal flats.An erratum to this article can be found at     

Diving behaviour of Little Penguins from four colonies across their whole distribution range: bathymetry affecting diving effort and fledging success

Little Penguins, Eudyptula minor, breed in several small colonies in New Zealand and Australia. In this study, we compare the birds’ diving performances at different sites situated throughout their breeding range. Environmental conditions and breeding success vary drastically amongst colonies, but all birds feed on similar types of prey and face similar limitations on their foraging range. We examined several diving parameters and calculated the proportion of foraging zone available during breeding to examine whether oceanographic and geographic factors in the foraging zone can explain variations in diving behaviour and fledging success among the different colonies. In colonies with high fledging success, Penguin Island and Oamaru, penguins made shallow dives <50 m depth and had lower diving effort. More than 90% of the foraging zone was in waters <50 m depth in these colonies. Motuara Island also has shallow waters with 95% <50 m depth, but the fledging success was low. Phillip Island has only 42% of waters <50 m and comparatively low fledging success. Thus, penguins dived deeper and showed a higher diving effort in colonies with lower fledging success (Motuara Island and Phillip Island), indicating that they were disadvantaged compared to conspecifics from other colonies that dived shallower and with a lesser diving effort. We concluded that bathymetry is an important factor, but not the only one, which influences fledging success.