A high local species richness and biodiversity within high-latitude calcareous aggregates of tube-building polychaetes

In general, biodiversity and species richness follow the latitudinal diversity gradient and decrease from the tropics towards the poles. Exceptions have however been recorded, as for deep coldwater coral reefs at high latitudes, which comprise biodiversity hotspots. Here we assess and characterise the high-latitude (69°N) species richness and diversity of a local shallow-water fauna associated with small calcareous aggregations of a serpulid polychaete. A dense and very species rich fauna was recorded within aggregations of Filograna implexa Berkeley, 1828. Totally 4663 individuals belonging to 99 species (61 solitary, 38 colonial) were recorded in a total aggregation volume of only 4.4 l covering an area less than 0.05 m² of a wreck situated in a tidal stream in North Norway. The number of species within each aggregation was positively related to its size, indicating that the high species diversity may be due to structural heterogeneity, which increase with aggregation size and probably creates new microhabitats and protect against predation. We present a species list including abundance and biomass, pin-point common species and describe a method for sampling such faunas associated with calcareous structures.     

Detecting impacts of non‐native species on associated invertebrate assemblages depends on microhabitat

Invasive plants that displace native floral communities can cause changes to associated invertebrate species assemblages. Using a mini‐review of the literature and our own data we add to the still considerable debate about the most effective methods for testing community‐level impacts by invasive species. In endangered saltmarshes of southeast Australia, the non‐native rush Juncus acutus L. is displacing its native congener J. kraussii Hochst., with concurrent changes to floral and faunal assemblages. In two coastal saltmarshes, we tested the hypothesis that the ability to detect differences in the invertebrate assemblage associated with these congeneric rushes depends on the microhabitat of the plant sampled. We used three sampling methods, each targeting specific microhabitats: sweep netting of the plant stems, vacuum sampling of the plant tussock, and vacuum sampling of the ground directly below the plants. Over 3800 individuals and 92 morphospecies were collected across four main taxa: gastropods, crustaceans, hexapods and arachnids. Detection of differences in invertebrate density, richness and composition associated with native compared with non‐native rushes was dependent on the microhabitat sampled and these differences were spatially variable. For example, at one saltmarsh the stems and tussock of J. acutus had a lower density and richness of total invertebrates and hexapods than those of the native J. kraussii. In contrast, crustaceans on the ground were in greater abundance below J. acutus than J. kraussii. This study demonstrates that on occasions where overall differences in the assemblage are not detected between species, differences may become apparent when targeting different microhabitats of the plant. In addition, separately targeting multiple microhabitats likely leads to a greater probability of detecting impacts of invasion. Comparing the invertebrate assemblage without differentiating between or sampling an array of microhabitats can fail to determine the impact of invasive species. These results highlight that a combination of methods targeting different microhabitats is important for detecting differences within the invertebrate community, even for phylogenetically related species.     

Benthic foraminifera of a Miocene canyon and fan

We present benthic foraminiferal assemblage data from an exhumed Miocene canyon and fan system from the Tabernas Basin (SE Spain). The presence of good indicator taxa and unique assemblages occupying specific environments allows the distinction of slope, canyon and fan environments within the Tabernas Basin by foraminiferal assemblages alone. Five assemblages are defined on the basis of the occurrence of the indicator taxa. Primary control on the distribution of these assemblages is consistent with trends of physical disturbance and consequent defaunation. Barren samples, which are predominantly found in high-energy parts of the proximal canyon, are recognized as representing recently defaunated substrates (i.e. early successional assemblages). High diversity assemblages containing a high abundance of agglutinated taxa are recognised in the undisturbed slope sequences as being the regional equilibrium (“climax”) fauna. Intermediate between these end-members are assemblages with low diversity, dominated by calcareous taxa typically found in the relatively low-energy canyon and fan environments, which are recognized as representing the middle phases of the ecological succession. Two further assemblages, a low diversity assemblage typified by Cassidulina laevigata and Bulimina costata and a very low diversity assemblage dominated (> 10% of all benthic tests) by Globobulimina spp., are restricted to low-energy parts of the canyon and fan, and are absent from the proximal canyon and slope. The composition of these assemblages indicate that nutrient supply/oxygenation is a secondary control on the palaeoecology of the canyon system. A conceptual model for the recolonisation of defaunated substrates in El Buho Canyon is proposed, in which either an oligotrophic climax assemblage or a eutrophic climax assemblage can be achieved at the completion of recolonisation of defaunated substrates, depending on environmental conditions.     

The surface ocean productivity response of deeper water benthic foraminifera in the Atlantic Ocean

We test the relationship of deep sea benthic foraminiferal assemblage composition to the surface ocean productivity gradient in the low latitude Atlantic Ocean using 81 surface sediment samples from a water depth range between 2800 and 3500 m. The samples are selected so that the surface ocean productivity gradient, controlling the flux of organic carbon to the seabed, will be the most important environmental variable. The first two principal components of the assemblage data account for 73% of data variance and are clearly linked to the productivity gradient across the Atlantic. These components show that under higher productivity the assemblages contain a higher abundance of Uvigerina peregrina, Melonis barleeanum, Globobulimina spp. and other taxa with probable infaunal microhabitats. Alabaminella weddellensis, a species linked to episodic phytoplankton debris falls, is also important in these assemblages. As productivity decreases there is a regular shift in assemblage composition so that low productivity assemblages are dominated by Globocassidulina subglobosa and several Cassidulina species along with Epistominella exigua. We hypothesize that these taxa are epifaunal to very shallow infaunal since nearly all organic carbon oxidation occurs near the sediment-water interface in low productivity settings. Discriminant function analysis of the foraminiferal assemblages, with groups selected on the basis of surface ocean productivity, shows clear separation among five productivity levels we used. This analysis demonstrates that productivity variations have a strong influence on assemblage composition. Finally, we used two groups of samples from the Rio-Grande Rise representing water depths from 2007 to 2340 m and 2739 to 3454 m to test for effects produced by changing water depth. All these samples are from a low productivity region and represent nearly identical environmental conditions. Although the low productivity nature of all the Rio-Grande Rise samples is obvious, there are assemblage differences between our depth groups. We cannot account for the assemblage differences with changes in organic carbon flux, dissolution effects or other physical/chemical properties of the ocean. Thus there are as yet unidentified factors related to water depth which cause some assemblage variation in the low productivity setting we investigated.     

Short-term responses of phytoplankton to nutrient enrichment and planktivorous fish predation in a temperate South American mesotrophic reservoir

The benthic macroinvertebrate community is an important component of stream diversity, because its members are fundamental connectors among the different trophic levels of running waters. In this study, we assessed alpha and beta diversities of benthic macroinvertebrates in three stream sites and four microhabitats: (i) moss in the air-water interface; (ii) submerged roots of terrestrial plants; (iii) leaf litter deposited in pools; (iv) stones in riffles. We constructed rarefaction curves and compared species richness among microhabitats for each stream site. Additionally, we evaluated which factor, stream site, or microhabitat, was most important in determining variation in assemblage structure, i.e., beta diversity. There was no significant difference among microhabitats in terms of taxa richness evaluated by rarefaction curves. Using partial Constrained Correspondence Analysis (pCCA), we found that microhabitat was most important in determining community composition, accounting for 42.02% of the total variation. Stream sites accounted for 22.27%. In accordance with the pCCA, exploratory multivariate methods (ordination and classification) revealed four distinct groups, corresponding to the four microhabitats, independent of stream sites. Our results indicated that differences among environmental conditions are much more important in the determination of stream assemblage structure than are differences in spatial location. Accordingly, adjacent microhabitats in a single stream site harbor macroinvertebrate assemblages more dissimilar than those found in a single microhabitat at different stream sites. Handling editor: D. Dudgeon     

Large gryphaeid oysters as habitats for numerous sclerobionts: a case study from the northern Red Sea

The shell of a living specimen of the Indo-Pacific gryphaeid giant oyster Hyotissa hyotis was colonized by numerous encrusting, boring, nestling and baffling taxa which show characteristic distribution patterns. On the upper valve, sponge-induced bioerosion predominates. On the lower valve intergrowth of chamid bivalves and thick encrusting associations—consisting mostly of squamariacean and corallinacean red algae, acervulinid foraminifera, and scleractinian corals—provides numerous microhabitats for nestling arcid and mytilid bivalves as well as for encrusting bryozoans and serpulids. Such differences between exposed and cryptic surfaces are typical for many marine hard substrata and result from the long-term stable position of the oyster on the seafloor. The cryptic habitats support a species assemblage of crustose algae and foraminifera that, on exposed surfaces, would occur in much deeper water.     

Mass loss and macroinvertebrate colonisation of fish carcasses in riffles and pools of a NW Italian stream

In this study, we analysed the decomposition of trout carcasses in a low-order Apennine stream, with the aim to investigate the mass loss rate in a Mediterranean lotic system, and to examine the influence of microhabitats on the invertebrates colonising fish carcasses. In May 2003, we put 56 dead rainbow trout (Oncorhynchus mykiss) in the stream, placing seven sets (four trout each) in both riffle and pool habitats. At four dates, we removed one trout per set to measure its dry mass and determine the associated macroinvertebrate assemblage. Fifty-eight macroinvertebrate taxa colonised the carcasses, with significant differences between the erosive and depositional microhabitats. Riffle trouts hosted richer and denser colonist communities than pool trouts. Chironomidae, Serratella ignita, Habrophlebia sp., Dugesia sp. and Protonemura sp. were the five most abundant taxa. Decomposition was initially very rapid in both environments and then tapered off over time. The mass loss rate was higher (k= –0.057 day^–1) than that found in other studies. Higher Mediterranean temperatures probably increase the process. Although we found no significant difference between riffles and pools, mass loss was more regular in erosive habitats, underlining the importance of local, small-scale conditions. In small, low-order, heterotrophic streams, fish carcasses represent an important resource and shelter for rich and diversified invertebrate assemblages.     

Distribution patterns of benthic foraminifera across the Ypresian–Lutetian Gorrondatxe section,Northern Spain: Response to sedimentary disturbance

We present a study of benthic foraminiferal assemblages from an Ypresian–Lutetian distal submarine fan system in the lower bathyal Gorrondatxe section (Basque-Cantabrian Basin, northern Spain). The objective of our study is to analyze the benthic foraminiferal distribution patterns and their response to sedimentary disturbance and related factors.Assemblages contain a high percentage of allochthonous taxa, such as asterigerinids and other shallow water taxa, which were transported downslope by turbidity currents.Detailed quantitative analyses, supported by R-mode cluster and Detrended Correspondence Analyses (after removing allochthonous taxa from the foraminiferal counts) allowed us to identify 6 assemblages that are divided into two groups related to the turbidite content in the Gorrondatxe section. Assemblages 1, characteristic of the turbidite-poor intervals with low sedimentary disturbance, include assemblage 1a (with highly diverse common middle–lower bathyal calcareous taxa) assemblage 1b (with common agglutinated taxa, mainly trochamminids), and assemblage 1c (characterized by calcareous taxa that are also common in the turbidite-rich interval).Assemblages 2, characterized by a high dominance, prevail in the turbidite-rich interval, and include assemblage 2a (characterized by the dominance of infaunal bolivinids and epifaunal cibicids), assemblage 2b (typified by moderate to low diversity and dominated by deep-infaunal Globobulimina species), and assemblage 2c (typified by very abundant suspension-feeding astrorhizids). The high abundance of bolivinids and Globobulimina species may be related to an enhanced input of low-quality organic matter transported by turbidity currents to the seafloor, representing different stages of recolonisation after disturbance and different energy regimes. High current activity was probably responsible for the abundance of cibicids, while moderate to low diverse and high dominance assemblages characterize the recolonisation of the substrate after disturbance.We conclude that sedimentary disturbance and other related factors such as current activity, resuspension of sediments at the seafloor, and supply of organic matter (and its quality) played an important role in the distribution of benthic foraminifera in the Gorrondatxe section. The identification of allochthonous taxa emerges as an essential aspect of the study of environments with sedimentary disturbance.     

Late Miocene biota from the Abad Member of the Carboneras-Nijar Basin (Spain,Andalusia): A bathyal fossil assemblage pre-dating the Messinian salinity crisis

Marly sediments of the early Messinian Abad Member of the Turre Formation from the northeastern sector of the Carboneras-Nijar Basin (southern Spain) have yielded a rich fossil assemblage, of which 60 taxa are documented herein. Besides nannoflora and microfauna, this assemblage includes the first autochthonous macrofauna described from the Abad Member. Based on the calcareous nannofossil assemblage, in particular the occurrence of the zonal index taxon Amaurolithus primus, the sediments are assigned to the Mediterranean calcareous nannofossil zone CNM17, corresponding to the latest Tortonian to earliest Messinian interval. This matches the age range generally reported for the Abad Member. Palaeoecological evidence from calcareous nannofossils (20 autochthonous taxa), planktic and benthic foraminifera (12 taxa), Porifera (3 taxa), Octocorallia (Keratoisis), Serpulidae (4 taxa), Bivalvia (5 taxa), Gastropoda (2 taxa), Brachiopoda (7 taxa), Cirripedia (Faxelepas) and Vertebrata (5 taxa) indicates an upper bathyal environment with an influx of neritic elements for the Abad Member near Carboneras. Additionally, several faunal components may represent allochthonous/parautochthonous elements from adjacent habitats, which were transported into the deep marine setting by turbiditic mass flows. Although similarities exist, the fossil assemblage from the marls is compositionally significantly different from the biota previously documented from a nearby exposed olistostrome, the ‘red breccia’. Similar fossil assemblages from the Mediterranean have so far mainly been reported from the Pliocene-Pleistocene of southern Italy and Greece. The Carboneras fauna thus adds to our knowledge of the development of these habitats and their biota prior to the Messinian salinity crisis. Beyond the novel palaeoenvironmental data, the range of the dyscoliid brachiopod Ceramisia meneghiniana, previously known only from the Pliocene of Italy, is extended to the Miocene of Spain. The cirripede crustacean Pycnolepas paronai De Alessandri, 1895 is transferred to the hitherto monospecific genus Faxelepas Gale, 2015, whereby the range of the latter (previously Maastrichtian to Danian) is extended to the late Miocene.     

Horizontal gradient in fish assemblage structures in and around a seagrass habitat: some implications for seagrass habitat conservation

Horizontal gradients in fish assemblage structures in and around a seagrass habitat were evaluated by visual observations. The assemblage structures clearly differed between open microhabitats lacking seagrass, i.e., nearby sand and the area adjoining the outer margin of the seagrass bed (referred to as “outer gap”), and microhabitats with seagrass within the overall seagrass bed, although not differing among the latter microhabitats, including both edge and core portions. Such open microhabitats were found to be not always inferior, but nearly equal to or even sometimes greatly superior in fish species’ diversity and/or abundance to the microhabitats with seagrass. In particular, the outer gap was always ranked first in total species’ number and had outstanding abundance in spring. Similar open microhabitats adjoining seagrass walls facing the sand patch within the seagrass bed (referred to as “inner gap”) in spring were also characterized by higher fish species and individual numbers. The 11 most abundant fishes showed four discrete distribution patterns (three recognized herein and one implied by precedent studies), such contributing to a horizontal gradient in the fish assemblage structure. While no fishes showed a preference for the edge or core of the seagrass bed, group-forming juveniles of several species favored gap microhabitats, suggesting that, in addition to the traditionally recognized edge and core microhabitats, the concept of gap microhabitats should be included in seagrass conservation ecology.     

Morphology and microhabitat preferences of benthic foraminifera from the northwest Atlantic Ocean

The distribution of Rose Bengal stained calcareous benthic foraminifera was determined in six ☐ cores raised from water depths between 200 and 3000 m on the Nova Scotian continental margin and Gulf of Maine. The taxa can be separated into four microhabitats within the surficial sediments. Epifaunal taxa are generally found in the top cm, intermediate infaunal taxa are found from about 1 to 4 cm and deep infaunal taxa are found at > 4 cm sediment depth in at least one ☐ core. A fourth group, shallow infaunal taxa, is found in the top 2 cm and is inferred to be infaunal based on wall porosity characteristics and test shapes similar to infaunal taxa. The epifaunal, shallow infaunal and intermediate infaunal taxa maintain their positions within the sediments from core to core, whereas the deep infaunal taxa are found at progressively shallower sediment depths in cores within increasing organic carbon contents from shallower water depths.Each microhabitat category has distinct morphological characteristics. Epifaunal taxa have plano-covex or biconvex cross sections, trochospiral coiling and large pores absent or found on only one side. Shallow infaunal taxa have uniserial, triserial, or planispiral coiling, with surface ornamentation present on a number of taxa. The intermediate infaunal taxa have rounded peripheries, pores over the entire test and planispiral coiling, with the exception ofCibicidoides bradyi which has trochospiral coiling. The deep infaunal taxa have, in general, planispiral or triserial coiling with cylindrical or ovate shaped tests.     

Fine‐scale patterns of species and phylogenetic turnover in a global biodiversity hotspot: Implications for climate change vulnerability

Question: What is the relative importance of environmental and spatial factors for species compositional and phylogenetic turnover? Location: High‐rainfall zone of the Southwest Australian Floristic Region (SWAFR). Methods: Correlates of species compositional turnover were assessed using quadrat‐based floristic data, and establishing relationships with environmental and spatial factors using canonical correspondence analyses and Mantel tests. Between‐quadrat phylogenetic distance measures were computed and examined for correlations with environmental and spatial attributes. Processes structuring pa2t2terns of beta diversity were also evaluated within four broad floristic assemblages defined a priori. Results: Floristic diversity was strongly related to environmental attributes. A low significance of spatial variables on assemblage patterns suggested no evident effect of dispersal limitations. Species compositional turnover was especially high within the swamp and outcrop assemblage. Phylogenetic turnover was closely coupled to species compositional turnover, implying the occurrence of many locally endemic and phylogenetically relict taxa. Beta diversity patterns within assemblages were also significantly correlated with the local environment, and relevant correlates differed between floristic assemblage types. Conclusion: Phylogenetic diversity in the SWAFR high‐rainfall zone is clustered within edaphic microhabitats in a generally subdued landscape. A clustered rather than dispersed distribution of phylogenetic diversity increases the probability of significant plant diversity loss during periods of climate change. Climate change susceptibility of the region's flora is accordingly estimated to be high. We highlight the conservation significance of swamp and outcrops that are characterized by distinct hydrological properties and may provide refugial habitat for plant diversity during periods of moderate climate change.     

Phylogenetic relationships of Serpulidae (Annelida: Polychaeta) based on 18S rDNA sequence data, and implications for opercular evolution

Phylogenetic relationships of (19) serpulid taxa (including Spirorbinae) were reconstructed based on 18S rRNA gene sequence data. Maximum likelihood, Bayesian inference, and maximum parsimony methods were used in phylogenetic reconstruction. Regardless of the method used, monophyly of Serpulidae is confirmed and four monophyletic, well-supported major clades are recovered: the Spirorbinae and three groups hitherto referred to as the Protula-, Serpula-, and Pomatoceros-group. Contrary to the taxonomic literature and the hypothesis of opercular evolution, the Protula-clade contains non-operculate (Protula, Salmacina) and operculate taxa both with pinnulate and non-pinnulate peduncle (Filograna vs. Vermiliopsis), and most likely is the sister group to Spirorbinae. Operculate Serpulinae and poorly or non-operculate Filograninae are paraphyletic. It is likely that lack of opercula in some serpulid genera is not a plesiomorphic character state, but reflects a special adaptation.     

Feeding ecology of a carnivorous bladderwort (Utricularia uliginosa,Lentibulariaceae)

The terrestrial carnivorous bladderwort, Utricularia uliginosa Vahl. (Lentibulariaceae) was studied to determine the species assemblage present in traps of these plants in situ across four sites over 15 months. The immediate soil environment was also sampled to determine the fauna present, and to compare the fauna present in traps with the fauna in the environment. The soil fauna consisted of 10 taxon types, which occupied either pelagic, epibenthic or interstitial microhabitats. All were found in traps of U. uliginosa, with the main prey being interstitial taxa followed by epibenthic and occasionally pelagic taxa. Numbers of individuals of the two most abundant soil taxa (nematodes, Elaphoidella) varied independently across the four sites over the 15 months of the study, as did numbers of Elaphoidella in the traps of U. uliginosa. Numbers of nematodes in the traps of U. uliginosa showed significant differences among sites, but not differences among times. Comparison of the trap fauna with the soil fauna revealed differences in relative abundance between soil samples and trap samples for two of the three taxa examined. There was an under‐representation of nematodes in the traps relative to numbers in surrounding soil. There was an over‐representation of the copepod Elaphoidella in the traps of U. uliginosa relative to numbers in soil at some of the times of sampling. Acarina were equally abundant in soil and trap samples. The patterns observed for Elaphoidella and nematodes may be due to selectivity in trapping by U. uliginosa, and/or differences in digestibility of the prey. Elaphoidella individuals were found to be attracted to U. uliginosa in a behavioural experiment. This may contribute to the over‐representation of Elaphoidella in the traps of U. uliginosa in the field.     

Large‐scale ecosystem engineering by flamingos and fiddler crabs on West‐African intertidal flats promote joint food availability

Although the ecosystem engineering concept is well established in ecology, cases of joint engineering by multiple species at large scales remain rare. Here, we combine observational studies and exclosure experiments to investigate how co‐occurring greater flamingos Phoenicopterus roseus and fiddler crabs Uca tangeri promote their own and each other's food availability by creating a spatially complex mosaic of depressions (bowls, gullies) and hummocks (plateaus, mounds) in the intertidal zone. This results in a mosaic of microhabitats with different tidal inundation regimes. These microhabitats are spatially organized with labyrinth‐like patterns in the high intertidal zone and spotted patterns in the lower intertidal, both of which likely arise from biophysical interactions between these organisms and hydrodynamic forces. We show that the resulting spatial complexity is vital for biofilm production. The depression microhabitats were wetter and richer in organic matter and biofilms compared with hummocks. Excluding flamingos and crabs resulted in an increase in biofilm biomass over the shorter term (six months), but a decrease over the longer term (after one year). Moreover, our results strongly suggest that these biogeomorphological microhabitats in the mosaics were maintained by the feeding activities of flamingos and to a lesser extent crabs. During a period of flamingo exclusion, all the spotted patterns filled up with sediment, while the exclusion of crabs led to gradual sediment accumulation in the labyrinth‐like patterns. Collectively, these findings provide empirical evidence for large‐scale joint promotion of food availability by multiple taxa in a marine ecosystem.     

Biostratigraphic and paleoclimatic significance of a new Pliocene foraminiferal fauna from the central Arctic Ocean

A Pliocene benthic foraminiferal fauna containing a previously unknown species association was found in the basal section of a piston core collected from the crest of Northwind Ridge (NWR) in the central Arctic Ocean. The fauna is dominated by Epistominella exigua, Cassidulina reniforme, Eponides tumidulus, Cibicides scaldisiensis, Lagena spp., Cassidulina teretis, Eponides weddellensis, Bolivina arctica, and Patellina corrugata. The presence of Cibicides scaldisiensis in the assemblage and the occurrence of Cibicides grossus higher in the core are indicative of an early Pliocene age. The morphologically distinctive species Cibicidoides sp. 795 of McNeil (in press) which occurs in the NWR core sample was previously known only from Oligocene through Miocene deposits in the Beaufort-Mackenzie Basin of Arctic Canada. Ehrenbergina sp. A and Cibicidoides aff. C. sp. 795, also present in the core, are new and endemic to the Arctic late Miocene and early Pliocene. These species, and possibly others, are survivors of the late Miocene (Messinian) sea-level crisis, which caused a significant faunal turnover in the Arctic Ocean. The predominantly calcareous assemblage indicates deposition above the calcium carbonate compensation depth in an upper bathyal environment. Paleogeographic affinities for the bulk of the assemblage indicate probable connections between the Arctic and the North Atlantic Oceans, but the endemic species identify environmental differences or partial isolation of the western Arctic Ocean. The species association suggests a cold but milder paleoclimate than that which existed during Pleistocene glacial intervals.     

The effect of turbidity state and microhabitat on macroinvertebrate assemblages: a pilot study of six shallow lakes

Shallow lakes can occur in two alternative stable states, a clear-water state and a turbid state. This is associated with separate assemblages of fish, zooplankton and plants. Little is known about whether macroinvertebrate assemblages differ across both stable states. This study investigated this in a connected set of three turbid and three clear-water shallow lakes. To overcome confounding effects of differences in spatial structure of macrophytes in turbid and clear-water lakes, we sampled three microhabitats that occurred in both alternative stable states: open water, sago pondweed (Potamogeton pectinatus) and reed (Phragmites australis). Univariate analyses indicated no differences in the number of organisms, taxon richness or diversity between turbid and clear-water lakes. Multivariate analysis, however, showed significant differences in the macroinvertebrate community structure of both stable states. Nine taxa explained a significant amount of the variation between both lake types, of which seven preferred the clear-water lakes. The number of organisms and the taxon richness were higher in reed than in the other microhabitats, but diversity and evenness did not differ among the microhabitats. Multivariate analyses could separate all three microhabitats. Eight taxa, mainly detritus feeders and collector–gatherers, explained most of the variation in the data and preferred the reed microhabitat. The effects of stable state (6.8% explained variance) and microhabitat (13.1% explained variance) on the macroinvertebrate assemblages were largely independent from each other (1.5% shared variance). Although macroinvertebrates are not implemented in the initial theory of stable states, our results show clearly different assemblages across both stable states.     

Modifications of the specific diversity and feeding guilds in an intertidal sediment colonized by an eelgrass meadow (Zostera marina) (Brittany,France)

A sandy beach located in a bay of north Brittany (France) is partly colonized by an eelgrass meadow. The overall macrobenthic fauna, i.e. epifauna, endofauna and fish, living in this seagrass and in non-vegetated sands was compared in terms of species, abundance and biomass in the different feeding guilds. Seagrass makes this habitat more complex by increasing both its surface by 5.45–fold and volume by 4.79–fold. Eighteen species among the 23 living in the non-vegetated sands were observed in the seagrass assemblage. Their abundance and biomass were increased by 7.33–fold and 3.54–fold, respectively. The occurrence of Zostera was associated with the presence of 124 new species and 34 new feeding guilds. In the Zostera meadow, a spatial segregation exists among the taxa from the different microhabitats explored for food-quest. The balanced distribution of biomass within the various microhabitats may suggest that an optimization in the exploitation of feeding resources goes hand in hand with a higher complexity of the trophic web. It looks as if the architecturally increasing complexity of the habitat resulting from the development of a seagrass meadow on sediment both stimulates the abundance and biomass of the indigenous macrobenthic assemblage, increases the specific and functional biodiversity and favours the specialization of feeding guilds.