The importance of competitor identity,morphology and ranking methodology to outcomes in interference competition between sponges

The nature or structure of competition within communities has been a dynamic area of practical and theoretical research for a number of decades. Certain components of some communities have proved hard to incorporate within such studies; in marine benthic work this has included the sponges. In this study, intra- and inter-phyletic interactions involving sponges (7,817 in total) were studied on 125 rocks (ranging in surface area from 10 to 2,438 cm ²) at a site experiencing low disturbance levels at Lough Hyne Marine Nature Reserve, Co. Cork, Ireland. The outcomes of sponge-sponge (intra-phyletic) interactions were significantly different in both outcome and structure to those involving sponges and representatives of other phyla. Typically sponge-sponge competition resulted in a much higher number of tied outcomes than did inter-phyletic encounters. Sponges over-grew all groups of organisms with the exception of some cnidarians and ascidians. The morphology of any sponge species (thickness in most cases) was important in determining the outcome of intra-phyletic, but not inter-phyletic, interactions. Sponges which exhibited thick (>2 mm) crusts were, in the majority of interactions, superior competitors compared to the thin (<1 mm) crusts. However, the method used to rank species (i.e. wins:loss or wins:total ratio) made a significant difference to the ranking of sponge competitors. The transitivity of the sponge assemblage investigated was calculated using the index formulated by Tanaka and Nandakumar (1994) as 0.23, indicating the assemblage to be organised as a network rather than a hierarchical system. We suggest this network of principally tied outcomes (due to standoffs) may be maintained in part by chemical interaction. However, in the absence of disturbance, the network is more likely to be mediated through processes of growth and regression of tissues during summer and winter months respectively.     

Functional morphology and zonation of three species of sea anemones from rocky shores in southern Chile

Three species of sea anemones, clearly exhibiting zonation on the shore, are grouped into intertidal [Phymactis clematis (Drayton, 1846), Anthothoe chilensis (Lesson, 1830)] and infralittoral [Antholoba achates (Drayton, 1846)] forms. A series of observations revealed that intertidal forms have the ability to retain water in the coelenteron during exposure to air,/while the infralittoral form cannot do so. This different behaviour is attributed to morphological characteristics, such as the specific body shape and the structure of the sphincter and retractor muscles. These features, in combination with specific abiotic factors of the various habitats under consideration, determine the upper limits of distribution of each species, while zonation below these limits seems to be affected in addition by biological interactions.     

Relationship between depth, sediment, latitude, and the structure of benthic infaunal assemblages on the mainland shelf of southern California

A regional benthic survey was conducted in 1994, and the data were used to assess the relationship among three habitat factors (depth, sediment grain size, and latitude) and the distribution of benthic infaunal assemblages on the southern California coastal shelf. Benthic samples were collected with a 0.1 m² Van Veen grab from 251 sites on the continental shelf (10–200 m deep) from Point Conception, California, to the United States–Mexico international border. The relationship between habitat and assemblages was investigated by conducting a Q-mode cluster analysis to define groups of stations with similar species composition and then examining whether differences were present in physical habitat attributes among those groups of stations. Analysis of data from 175 uncontaminated sites yielded four habitat-related benthic infaunal assemblages along the southern California coastal shelf: a shallow-water assemblage from 10–32 m, a mid-depth assemblage between 32 and 115 m, and two deep-water (115–200 m) assemblages, one in fine and one in coarse sediments. These empirically defined points in the depth and sediment grain size gradients can be used to define reference habitats for the development of biocriteria. Benthic abundance and diversity were greatest in the mid-depth assemblage, conforming to predictions for benthic assemblages in regions of upwelling. Within the 500 km of coastline examined, latitude was not an important factor in defining assemblages. Received: 3 December 1999 / Accepted: 9 October 2000     

Competitive interactions among sessile organisms in Tomioka Bay,south Japan: importance of light conditions on the panel surface

Interspecific competitive interactions among sessile epibenthos were studied by suspending PVC panels at Tomioka Bay, south Japan, for a maximum period of 16 mo. Interactions were monitored from photographs of a fixed area of the panels. Four panels were suspended during two different months in autumn 1991, and the development of the community was followed until December 1992. Altogether, 6511 interspecific overgrowth interactions were recorded, of which 37 resulted in standoffs and the rest in overgrowths. The competitive relationship observed in this sessile assemblage followed the pattern of a hierarchy with numerous backloops. Among the 36 species, belonging to the seven taxonomic groups encountered during the study, the colonial ascidian Didemnum moseleyi was recorded as the dominant species (with respect to competitive ability) while the barnacle Balanus trigonus was the weakest species. The month of panel exposure and whether or not the panel surface was shaded had a significant influence on the competitive ability of the sessile organisms. The order of hierarchy of the most dominant species changed with the month of panel submersion and its light conditions. Among the several abundant species tested, longer residence times were recorded for serpulid worms than for the colonial species. A significant, positive relationship was obtained between the areal cover of competitively dominant sessile organisms and the number of their interspecific interactions. From the short residence time of sessile organisms and the significant relationship between their areal cover and number of interspecific interactions, it is concluded that the interspecific interactions played important role in the species succession.     

Alloimmune responses between Acropora hemprichi conspecifics: nontransitive patterns of overgrowth and delayed cytotoxicity

Allogeneic interactions between Acropora hemprichi (Ehrenberg) colonies were followed in situ up to 18 mo between 1990 and 1992 in Eilat, Red Sea. We established all 21 possible branch pair combinations among seven large colonies in three to eight replicates each (n=92 pairs), in addition in 14 pairs of grafts within-colonies. All isografts resulted in fusion. Allogeneic interactions revealed a variety of effector mechanisms which involved the appearance of dark blue-purple color in the tissues of interacting branch tips (<1 cm), unilateral overgrowths (1.0 to 1.5 mm mo^-1 for 1 to 8 mo), rejections, healings, continuous overgrowths, reversals, repeated rejections and more. Direction of overgrowth was usually specific to a given combination of colonies. A hierarchial network of overgrowths was obtained by analyzing overgrowth directions of all pairwise combinations, which included 30 transitive and five nontransitive three-way combinations. These outcomes, similarly to responses of allogeneic encounters recorded from other coral species, show that colonies can respond selectively to different types of allogeneic challenges. It is suggested that corals may have the capacity to distinguish self from nonself by recognizing nonself attributes presented on conspecifics.     

Variability of competition at scales of 10<Superscript>1</Superscript>, 10<Superscript>3</Superscript>, 10<Superscript>5</Superscript>, and 10<Superscript>6</Superscript> m: encrusting arctic community patterns

Variability in interference competition was studied in benthic marine communities of the arctic and subarctic Atlantic intertidal and shallow subtidal zones. We sampled multiple square-metre quadrats at distances of 10¹, 10³, and 10⁵ m apart around the high polar island of Spitsbergen (Svalbard Archipelago). We also took some similar samples in Iceland and in the Faeroe Islands (10⁶ m apart from Spitsbergen samples). Encrusting fauna were present on high arctic intertidal rocks but we only found competitive interactions on subtidal substrata. On subarctic Icelandic and Faeroese shores, in contrast, spatial competition was common even in the intertidal zone. Analysis of variance of competition intensity data (numbers of interactions per area) revealed multiple factors to be significant influences explaining variability. Amongst the 10¹-, 10³-, and 10⁵-m spatial scales, only the largest emerged as a significant term. Whether intra- or interspecific competition dominated the types of interactions varied greatly between sites: 21–97% of competition was intraspecific. The proportion of competitive encounters resulting in a decided outcome (i.e. a win for one competitor and a loss for the other, rather than a tie or standoff between them) showed little variability at any scale. All the values of competition transitivity (how hierarchical a pecking order is) were very high compared to values reported in the literature from any other (polar or non-polar) locality. Variability in this measure was generally <10% across scales. We conclude from our data that great care must be taken in interpreting patterns of competition between similar taxa in large-scale space or time. Not only did most aspects of competition in our study communities vary significantly at the 10⁵-m scale but different aspects of competition varied at different scales and by hugely different amounts.Communicated by J.P. Thorpe, Port Erin     

Comparisons of genetic population structures in four intertidal brachyuran species of contrasting habitat characteristics

Genetic population structures along the Japanese coast, analyzed by sequence data from the mitochondrial DNA COI region, were determined for four intertidal brachyuran species in the superfamily Thoracotremata (Ocypode ceratophthalma, Gaetice depressus, Chiromantes dehaani and Deiratonotus japonicus), which were characterized by different habitat requirements. O. ceratophthalma (seashore; supratidal sand) and C. dehaani (estuarine; supratidal marsh) showed no significant genetic differentiation among Japanese populations. The Japanese populations of O. ceratophthalma, however, were found to genetically differentiated from the Philippine population. G. depressus (seashore; intertidal cobbles) exhibited significant genetic differentiation between the Amami-Ohshima population and other local populations. D. japonicus (estuarine; intertidal cobbles) showed significant genetic differentiation among many local populations separated by about 30–1,200 km. The different patterns of genetic population structure recorded for the four species, thus, do not simply correspond to habitat type.     

Comparisons of genetic population structures in four intertidal brachyuran species of contrasting habitat characteristics

Genetic population structures along the Japanese coast, analyzed by sequence data from the mitochondrial DNA COI region, were determined for four intertidal brachyuran species in the superfamily Thoracotremata (Ocypode ceratophthalma, Gaetice depressus, Chiromantes dehaani and Deiratonotus japonicus), which were characterized by different habitat requirements. O. ceratophthalma (seashore; supratidal sand) and C. dehaani (estuarine; supratidal marsh) showed no significant genetic differentiation among Japanese populations. The Japanese populations of O. ceratophthalma, however, were found to genetically differentiated from the Philippine population. G. depressus (seashore; intertidal cobbles) exhibited significant genetic differentiation between the Amami-Ohshima population and other local populations. D. japonicus (estuarine; intertidal cobbles) showed significant genetic differentiation among many local populations separated by about 30–1,200 km. The different patterns of genetic population structure recorded for the four species, thus, do not simply correspond to habitat type. An erratum to this article can be found at     

Mesozooplankton community structure in offshore and coastal waters of the Ionian Sea (eastern Mediterranean) during mixed and stratified conditions

Depth-stratified samples, collected during a period where the water column was vertically mixed (March 2000) and a period of thermal stratification (September 2000), were analyzed in order to investigate the horizontal and vertical distribution patterns and composition of mesozooplankton, especially copepod species assemblages, in a pelagic (Ionian Sea) and a coastal area (Patraikos Gulf) of the eastern Mediterranean. Total mesozooplankton abundance and biomass were significantly lower in the highly oligotrophic offshore waters of the Ionian Sea when compared to the semi-enclosed Patraikos Gulf during both seasons. Small-sized copepods dominated the mesozooplankton community. An ‘offshore’ and a ‘coastal’ copepod assemblage were defined in the surface layer (0–50 m) only during March when differences in environmental conditions (i.e., temperature, salinity and fluorescence) were strong between the two areas. Copepod vertical community structure in offshore waters differed between sampling months. In March one assemblage (0–200 m) was mainly identified, while in September three distinct assemblages (0–50, 50–100 and 100–200 m) were observed, related to different vertical distribution patterns of the various copepod species. A pronounced seasonal change of the dominant copepods was evident in the surface layer, where strong differences in hydrological properties were observed from March to September. Below this layer, the copepod community was relatively stable showing decreasing seasonal differences with increasing depth.     

Invasion patterns inferred from cytochrome oxidase I sequences in three bryozoans, Bugula neritina, Watersipora subtorquata, and Watersipora arcuata

Nucleotide variation in cytochrome c oxidase subunit I (COI) was used to examine population structure in three invasive bryozoans: Bugula neritina (Linnaeus, 1758), Watersipora subtorquata (d’Orbigny, 1852), and W. arcuata (Banta, 1969). These species are found on ship hulls and have a short (≤2 days) larval phase. Samples were collected from 1998–2001 at multiple sites in Australia, and in Hong Kong, New Zealand, Hawaii, California, Curaçao, and England. B. neritina is known to include three cryptic species, including species Type S (Davidson and Haygood in Biol Bull 196:273–280, 1999) which occurs on the east and west coasts of the USA. One haplotype recorded previously in the USA, S1, was found to be widespread, occurring throughout Australia and in Hong Kong, Curaçao, Hawaii, and England. W. subtorquata, a Caribbean–Atlantic species which has invaded southern Australia, New Zealand, and California, had low nucleotide diversity in these areas (π=0.0016±0.0014), consisting of three haplotypes connected by one or two nucleotide mutations. W. arcuata, an Eastern-Pacific native, had comparatively high diversity (π=0.0221±0.0115) in introduced populations from Australia and Hawaii. In each species, identical haplotypes were identified on separate coastlines providing evidence of widespread, rather than genetically independent, introductions. The major contrast in nucleotide diversity suggests that different propagule-source models explain introductions.     

Abundance,diversity and temporal variability in a California intertidal nudibranch assemblage

A 40-month study of a nudibranch assemblage within a 250 m² intertidal area at Asilomar State Beach, California, USA, indicated that dominance was shared among 9 species which comprised 87% of the total number of individuals enumerated. These 9 species also showed high frequency of occurrence in the area. The number of nudibranch species counted per month was shown to be independent of weather and tide conditions, but the numbers of individuals and diversity values were correlated with wave action, lower values and numbers being tabulated during times of storms. Cumulative plots of diversity and species number based upon 15 min enumeration times indicated adequate assessment of diversity and species number after 60 min of sampling. The number of species and individuals and the diversity varied from month to month over the 40 months, but these variations were shown to be statistically insignificant or to be correlated with weather. Hence, it was concluded that the assemblage was a stable one, structured primarily by the 9 dominant species. No statistically significant seasonal or annual changes were observed in the assemblage. Diversity was found to correlate most highly with species number and less with number of individuals. The average number of species found per month was 13, the average number of individuals 104, and the average monthly diversity (HH}) was 1.82. Significant positive and negative correlations between abundances of certain species were found. These were attributed to occurrence of prey items or to synchrony of reproduction. There was no evidence of migration, and resident status for one dominant species was established. Comparison of the present study with earlier work suggested that a stable nudibranch assemblage was not merely a local phenomenon.     

Exaggerated male eye span influences contest outcome in stalk-eyed flies (Diopsidae)

Evolution of male weapons or status signals has been hypothesized to precede evolution of female mating preferences for those traits. We used staged male fights among three species of Malaysian stalk-eyed flies (Diptera: Diopsidae) to determine if elongated eye span, which is preferred by females in two sexually dimorphic species, influences contest outcome. Extreme sexual dimorphism, with large males possessing longer eye span than females, is shared by Cyrtodiopsis whitei and C. dalmanni. In contrast, C. quinqueguttata exhibits a more ancestral condition – short, sexually monomorphic eye stalks. Videotape analysis of 20-min paired contests revealed that males with larger eye span and body size won more fights in the dimorphic, but not monomorphic, species. To determine if males from the dimorphic species use eye span directly to resolve contests, we competed male C. dalmanni from lines that had undergone artificial selection for 30 generations to increase or decrease eye span. We found that eye span, independently of body size, determines contest outcome in selected-line males. Furthermore, in both dimorphic species, the average encounter duration declined as the eye span difference between contestants increased, as expected if males use eye span to assess opponent size. The number of encounters also increased with age in dimorphic, but not monomorphic, species. Selected-line males did not differ from outbred males in either fight duration or number of encounters. We conclude that exaggerated male eye stalks evolved to influence both competitive interactions and female mating preferences in these spectacular flies. Received: 20 July 1998 / Received in revised form: 2 February 1999 / Accepted: 12 March 1999     

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.     

Spatial variability of infaunal nematode and polychaete assemblages in two muddy subtidal habitats

Spatial variation of populations and their assemblages is an important component of many aspects of ecology, including the maintenance of species diversity. In nature, organisms are generally aggregated in patches or form gradients or other spatially related patterns. This investigation quantified the degree of spatial structure in benthic invertebrate assemblages. It examined the distribution of infaunal assemblages of different size, mobility and contrasting life-history strategies (i.e. meiofaunal nematodes and macrofaunal polychaetes) in two offshore muddy habitats in the Celtic Deep and the NW Irish Sea off the west coast of the United Kingdom. The more heterogeneous habitat in the NW Irish Sea was characterised by higher tidal stress and bottom temperature while greater water depth, mean particle diameter and organic carbon content was typical for the comparatively homogeneous environment in the Celtic Deep. In both habitats, the environmental conditions became increasingly dissimilar with separation. A total of 125 nematode and 88 polychaete species were recorded with 69% of the nematode and 49% of polychaete species present at both study sites. Occurrence frequency of species, species diversity and average similarity of assemblage composition was higher in the Celtic Deep than in the NW Irish Sea. Results from correlation analyses revealed statistically robust relationships of community similarity and sample distance. Given their small size and low mobility, nematodes were more susceptible to within-habitat physical variability than larger-sized, more mobile polychaetes. This, coupled with limits to long-distance dispersal and likely restrictions in gene flow, resulted in a significant decrease in community similarity with distance at the spatial scales sampled (i.e. within 0.1–23 km). Polychaetes, in contrast, combined a higher dispersal potential (>23 km) with a relatively high tolerance to within-habitat environmental variability and these were the most likely causes for non-significant relationships between the similarity of their assemblages and sample distance. The potential mechanisms causing the observed variation and implications of results for environmental management strategies are discussed. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Temperature, salinity and prey availability shape the marine migration of Arctic char, Salvelinus alpinus, in a macrotidal estuary

The influence of salinity, temperature and prey availability on the marine migration of anadromous fishes was determined by describing the movements, habitat use and feeding behaviours of Arctic char (Salvelinus alpinus). The objectives were to determine whether char are restricted to the upper water column of the inter-/subtidal zones due to warmer temperatures. Twenty-seven char were tracked with acoustic temperature/pressure (depth) transmitters from June to September, 2008/2009, in inner Frobisher Bay, Canada. Most detections were in surface waters (0–3 m). Inter-/subtidal movements and consecutive repetitive dives (maximum 52.8 m) resulted in extreme body temperature shifts (−0.2–18.1 °C). Approximately half of intertidal and subtidal detections were between 9–13 °C and 1–3 °C, respectively. Stomach contents and deep diving suggested feeding in both inter-/subtidal zones. We suggest that char tolerate cold water at depth to capture prey in the subtidal zone, then seek warmer water to enhance feeding/digestion physiology.     

Reintroduction of eelgrass (Zostera marina) in the Dutch Wadden Sea; review of research and suggestions for management measures

Eelgrass (Zostera marina L.) in the Dutch Wadden Sea historically covered an area varying from 65–150 km² in the eulittoral as well as the sublittoral zones. At present, this area comprises less than 1 km² eulittoral eelgrass stands, with an associated decrease in habitat diversity. The causes for this decline are presumably connected with the ‘wasting disease’ and the closure of the former Zuiderzee in the early 1930s resulting in increased tidal range and increased currents. After a slight recovery of the eelgrass populations on the intertidal flats a definite decline started in the early 1970s, possibly connected to increased turbidity. The present water quality and turbidity do not negatively influence eelgrass growth up to a depth of at least 0.6m below Mean Sea Level. Based on mesocosm experiments and field experiments it is concluded that re-establishment of eelgrass should be possible in sheltered bays and on unexposed tidal flats. The most suitable depths for a reintroduction are those between 0 and 20–40 cm below mean sea level.     

Reintroduction of eelgrass ( Zostera marina ) in the Dutch Wadden Sea; Review of research and suggestions for management measures

Eelgras (Zostera marina L.) in the Dutch Wadden Sea historically covered an area varying from 65–150 km² in the eulittoral as well as the sublittoral zones. At present, this area comprises less than 1 km² eulittoral eelgrass stands, with an associated decrease in habitat diversity. The causes for this decline are presumably connected with the ‘wasting disease’ and the closure of the former Zuiderzee in the early 1930s resulting in increased tidal range and increased currents. After a slight recovery of the eelgrass populations on the intertidal flats a definite decline started in the early 1970s, possibly connected to increased turbidity. The present water quality and turbidity do not negatively influence eelgrass growth up to a depth of at least 0.6m below Mean Sea Level. Based on mesocosm experiments and field experiments it is concluded that re-establishment of eelgrass should be possible in sheltered bays and on unexposed tidal falts. The most suitable depths for a reintroduction are those between 0 and 20–40 cm below mean sea level.     

Rhythmicity of larval release in three species of intertidal brachyuran crabs (Crustacea: Brachyura) from Inhaca Island (Mozambique)

 A study of rhythmicity of larval release in three species of intertidal brachyuran crabs, based on laboratory and field experiments, was undertaken at Inhaca Island, southern Mozambique, using Leptodius exaratus and Macrophthalmus grandidieri from December 1994 to January 1995, and Arcotheres palaensis from April to July 1995. L. exaratus and M. grandidieri showed a semi-lunar cycle in larval release. The release of larvae for L. exaratus, a species having conspicuous larvae, occurred in the first half of the night, after the post-crepuscular high tide, which suggests maximisation of protection of larvae from visual predation. The larval release activity matched the late spring and early neap tides. Results from the field were similar to those from the laboratory. M. grandidieri, having inconspicuous larvae, did not show a pattern related to the light–dark cycle and hatched during spring tides (around full and new moons) to maximise larval dispersion. A. palaensis, living inside the host mussel which inhabits the lowest section of the intertidal zone, did not show a relation with moon phase, tidal or light–dark cycles. Received: 16 February 1999 / Accepted: 8 December 1999     

Impact of Arenicola marina (Polychaeta) on the microbial assemblages and meiobenthos in a marine intertidal flat

The benthic microbial food web can be responsible for a large proportion of benthic carbon cycling yet there are few data on the trophic interactions between this food web and macrobenthos. A large-scale field experiment was conducted to investigate effects of eliminating the polychaete Arenicola marina on benthic microbes (prokaryotes, heterotrophic and autotrophic protists) and metazoan meiofauna in a marine intertidal flat of the North Sea, Germany. Over a period of 2 years, quantity and composition of micro- and meiobenthos from unmanipulated sites were compared to those from sites deplete of lugworms. These grazer treatments were cross-classified with different sediment characteristics (low- and mid-intertidal areas). Lugworm removal resulted in an initial increase in abundance of prokaryotes and nanoflagellates, which became less pronounced in the second year. Ciliates were not affected quantitatively, but in the absence of lugworms, diversity and the proportion of carnivorous forms increased. Meiobenthos (nematodes, ostracods and copepods) were affected only moderately. The observed changes are probably due to a combination of release from grazing/predation pressure, changes in the species composition of higher trophic levels (namely large polychaetes) and altered environmental conditions (such as depth of the oxygenated layer and sediment grain size). Spatial differences between sites of different tidal exposure/grain size appeared to be as large as temporal differences during the 2 years following the manipulation of the system. We conclude that in intertidal sediments, indirect effects due to habitat transformation are as important as direct biological interactions (grazing pressure and competition) for the dynamics of the benthic microbial food web.