Does colonization contribute to spatial patterns of common invertebrates in coralline algal turf?

Abstract The potential of colonization to contribute to the spatial patterns of six common invertebrates in coralline algal turf was investigated on a rocky shore near Sydney, Australia. The species, which included two amphipods (Elasmopus warra, Hyale spp.), a small bivalve (Lasaea australis), a fly larva (Limonia marina), and two microgastropods (Amphithalamus incidata and Eatoniella atropurpurea), had a range of dispersal modes (larval dispersal, crawling, swimming, rafting, and passive transport). Field sampling between May 1997 and November 1999 demonstrated that the amphipods were more abundant in low‐shore areas, the fly larvae and bivalves were more abundant in mid‐shore areas, and the abundances of gastropods did not vary with tidal height. Furthermore, abundances of all species varied among patches separated by tens of metres at one time or another. To test whether rates of colonization could contribute to established patterns of abundance, habitat mimics were deployed for 2‐week periods. The supply of new individuals matched long‐term patterns of abundance at different tidal heights for E. warra and L. marina. Colonization rates also differed among patches separated by tens of metres for three of the six species. Overall, there was little evidence to suggest that common species in coralline turf are limited by colonization on local scales, regardless of their major mode of dispersal. However, the potential for colonization to determine patterns of abundance varied from species to species.     

Role of colonization in spatio-temporal patchiness of microgastropods in coralline turf habitat

The ability of benthic macrofauna to disperse and colonize new habitats throughout their life may contribute substantially to small-scale patchiness in abundances in different habitats. Microgastropods in coralline turf on rocky shores in Australia are very patchy in abundance at different spatio-temporal scales. They therefore represent an ideal assemblage for testing hypotheses about processes of colonization. Patterns and rates of colonization of 10 species of microgastropods were investigated in one intertidal habitat (coralline turf) in Botany Bay, New South Wales, Australia, using artificial substrata, which are considered to be a good mimic of natural coralline turfs. The experiment was designed to test the hypotheses that (1) patterns (abundance of colonizing individuals) and mode (juveniles vs. adults) of colonization depends on the proximity of a patch to a potential source of dispersing colonists (i.e. patch of natural coralline turf), (2) different species show different rates of colonization, and (3) patterns of succession are not repeatable among different patches of natural algae. Seven different plots (natural patches of coralline turf) were randomly chosen in the area of study. Artificial units (called patches) were placed at different distances around each plot (within the plot and 0, 50 or 100 cm away from the edge). Samples were collected 6, 13, and 27 days after the experiment started. Colonization was rapid (i.e. within 6 days) for most species. Pattern (number of individuals) and mode of colonization (adults vs. juveniles) varied among algal plots. Most species responded differently across patches causing no consistent patterns of colonization. Furthermore, patterns of colonization of artificial units were not always synchronous with, or in the same direction as, changes in abundances in the nearest algal plots, which themselves showed no consistent spatio-temporal pattern.     

Littoral macrofauna (secondary) responses to experimental nutrient addition to rocky shore mesocosms and a coastal lagoon

Macrofauna secondary responses to controlled eutrophication within two MARICULT/MAST-III projects, EULIT and COMWEB, are compared. EULIT utilises a nutrient gradient established in eight hard bottom mesocosms, whereas the data from COMWEB originate from a whole-ecosystem study – the case of experimental nutrient addition to Hopavågen lagoon. In both systems, nutrient addition started in May 1998, after initial studies of the background macrofauna communities, thus allowing application of Before-After-Control-Impact-Pairs techniques (BACIP). The main objectives have been to investigate the macrofauna responses to eutrophication in littoral rocky shore ecosystems and to evaluate if similar responses could occur in mesocosm and `field' systems, despite their inherent differences. Apart from a distinct increase in numbers of Littorina littorea L. and some indications of increased abundance of the genus Jaera, no significant changes, caused by 2.5 years of nutrient addition, could be detected in the mesocosm fauna. It is interesting, however, that these two possibly stimulated animal groups have two things in common: (1) both belong to the rather few mobile taxa, which are not flushed out of the system through the mesocosm outlets, (2) both feed on microalgae and green algae in the upper littoral zone, i.e., algal groups that have shown the clearest response to the nutrient addition. In Hopavågen, 1.5 years of nutrient addition only caused modest plant and animal responses in the rocky shore ecosystem, although there was a markedly increased settlement of Mytilus edulis L. at one lagoon site. This increase was probably due to elevated levels of digestible particles (increased phytoplankton production) caused by the nutrient addition. There were also some indications of increased production of filamentous algae at this same lagoon site. All in all, both studies demonstrate only minor responses to increased nutrient levels within the littoral community (both plants and animals). One explanation to this may be that longer time-scales are needed in order for clear-cut changes to occur. Several observations also indicate that wave exposure (significant in both systems) may largely modulate the impact of increased nutrient load on the structure of littoral communities. Another explanation is the high degree of stability among littoral macroalgae-dominated communities with internal biological regulation factors (like grazing) possibly being able to counteract effects.     

Quantifying temporal variation in heterobranch (Mollusca: Gastropoda) sea slug assemblages: tests of alternate models

Assemblages of heterobranch sea slugs are notoriously variable in space and time which has sometimes led to their exclusion from broader studies of patterns of biodiversity. This variability may also result in a failure to detect underlying spatial and temporal patterns. Assemblage data (species abundances) from three intertidal sites over a period of 13 months, were used to compare the conclusions drawn from three different analytical approaches—a non-specific test for differences among months, and tests of cyclicity, and seriation, using specific model matrices. While no significant difference was detected for the non-specific test (i.e. concluding no significant temporal variation), the data were significantly correlated to both model matrices with the cyclicity model outperforming the seriation model at two of the three sites. The study also highlighted the spatial variability of assemblages over a scale of just a few kilometres. Wider testing using previously published datasets confirmed the utility of these models for exploring specific hypotheses about patterns of temporal change.     

Community structure of an assemblage of tidepool fishes on a tropical eastern Pacific rocky shore, Colombia

The composition, abundance, species richness and diversity of a rocky intertidal fish assemblage in the central zone of the Pacific coast of Colombia were determined. From July to December 2002, 10 tidepools with volumes ranging from 0·087 to 1·350 m³ were surveyed. A total of 893 fishes belonging to 14 species and 12 families were found. The most abundant species were Bathygobius ramosus and Abudefduf concolor , representing 71% of all fishes collected. Bathy gobius ramosus , Malacoctenus zonifer and Gobiesox adustus were residents, Abudefduf concolor , Abudefduf troschelii , Microsphatodon bairdii , Chaenomugil proboscideus , Halichoeres aestuaricola and Echidna nocturna were transients, and Apogon dovii , Sargocentron suborbitalis , Cerdale sp, Rypticus nigripinnis and Ctenochaetus marginatus were considered as accidental visitors. The temporal and spatial variation of assemblage structure was tested. The fish assemblage structure did not differ between months and there was not a clear relationship between tidepool surface area and depth, and community variables. Diversity and species richness were negatively correlated with tidepool height. It is suggested that the tidepool assemblage was stable and resilient in time on short (annual) scales. The constancy in time and space could have been a consequence of eurytopic features of the organisms and the lack of drastic seasonal changes in the tropics.     

Seasonal variation in a low shore Fucus serratus (Fucales,Phaeophyta) population and its epiphytic fauna

Seasonal variation in a Fucus serratus assemblage was studied on the Isle of Man, UK over a 13-month period. Algal cover varied greatly, related to shedding reproductive fronds and winter storms. Peak reproduction was recorded in the autumn. There was little change in density of plants, the population was dominated by a bank of germlings (<20 cm) and an adult mode between 50–70 cm. An indistinct third component of recruiting juveniles was present when gaps in the canopy formed. Seasonal variation in the algae was reflected in the population dynamics of the mobile epiphytes such as Littorina mariae and Lacuna pallidula, which exhibited annual life histories. Sessile species (such as bryozoans) utilized the algae year round, dominating the perennial stipes and lower fronds, which were not shed. Other species utilized the algae in a transient fashion: some feeding on the algae (e.g. Idotea), or other epiphytic animals (e.g. Nucella). Fucus serratus, therefore, provides a variety of spatially and temporally variable niches that explain the diversity of epiphytic animal species that exploit this alga.     

Analysis of the blennioid assemblages associated with different rocky shore habitats in the Ligurian Sea

The differences among blennioid assemblages (families Blenniidae and Tripterygiidae) on different habitats were assessed at two localities of the Ligurian Sea, namely Arenzano and Riva Trigoso. The assemblage composition and species relative density were evaluated visually on four different habitats of diverse wave exposure and substratum orientation (macro‐habitat characteristics): two vertical intertidal and subtidal habitats (exposed and sheltered rockwalls) and two horizontal subtidal habitats (semi‐exposed flat rock and boulders and pebbles). Each habitat was also characterized in relation to micro‐habitat features, such as substratum complexity, heterogeneity and amount of algae cover. Patterns of differences among habitats in assemblage variables and fish density, and the influences of macro‐ and micro‐habitat features on these patterns were studied at small (within localities) and large (across localities) spatial scales. Higher values of species richness (S), diversity and evenness (J) were generally associated with vertical habitats, as a result of a positive correlation with substratum orientation. The presence of an intertidal zone in the rockwall habitats may partially explain the observed differences in assemblage variables between vertical and horizontal habitats. The strength of relationships between S, and J and the other investigated habitat variables (exposure, complexity, heterogeneity and algae cover) varied greatly depending on spatial scale. All these relationships were positive, except for complexity. Significant variation in the assemblage total density among habitats was recorded only at Arenzano, where a larger number of fishes were counted on rockwalls rather than on the horizontal habitats. The positive effect of orientation on fish total density was strictly dependent on spatial scale. Fish total density showed a negative correlation with complexity and a positive correlation with heterogeneity, both relationships being unaffected by spatial scale. The unexpected relationship with complexity was probably due to the fact that, in the most complex habitat (i.e. boulder and pebbles), the potential positive effect of high complexity on fish density might be overcome by the negative influence of other environmental features, such as horizontal orientation and low wave exposure. Complexity and heterogeneity thus seemed good predictors of fish total density, but their role needs to be carefully interpreted. The most marked differences in species composition and relative density were found between rockwalls and the other habitats, mostly due to an unbalanced distribution of some stenoecious species. Variations in species relative density were related to different combinations of both macro‐ and micro‐habitat features, and these relationships usually changed depending on spatial scale.     

Epiphytes on the rocky intertidal red alga RhodomelaLarix (Turner) C. Agardh: Negative effects on the host and food for herbivores?

The diversity of epiflora and fauna associated with a dominant turf-forming alga was examined in intertidal communities on the central Oregon coast. Epiphytes associated with the red alga, Rhodomelalarix (Turner) C. Agardh, were examined by surveying intertidal areas for the presence of epiphytes, and by following changes in epiphyte cover in marked quadrats of R. larix. The alga is host for at least 17 species of sessile plants and animals. To determine the role of some of the larger epiphytes in the community, Rhodomela plants were marked and monitored and herbivore feeding was examined. Data suggest that epiphytes decrease the growth rate of their host, increase the probability of axis breakage and decrease reproductive output. Epiphytes provide food for littorine snails and gammarid amphipods that live in the beds of the plant. Amphipods were found to decrease epiphyte cover on R. larix in laboratory tanks, suggesting that these herbivores may have beneficial effects on the host plant.     

Seasonality of macroalgal assemblages in a wave-eroded intertidal granite shore at Chengshan Cape, on the northwest coast of the Yellow Sea

An ecological survey was carried out at bimonthly intervals for 2 years from 2000 to 2002 in the intertidal granite shore at Chengshan Cape, on the northwest coast of the Yellow Sea. Significant differences in macroalgal composition were evident in different seasonal assemblages. The highest species richness was found from April to August, and the lowest in October. In all the seasonal assemblages surveyed, Rhodophyta showed the highest species richness, Phaeophyta ranked second and Chlorophyta last. Macroalgal biomass peaked in August (246 gm^-2). From December to February cold-water species, such asEnteromorpha intestinalis, Porphyra spp., Gelidium amansii, Halymennia sinensis, Laminaria japonica and Polysiphonia urceolata, dominated the assemblage.In June warm-temperate and subtropical species became dominant. Species diversity peaked in April and then declined throughout the year until December. The seasonal assemblages surveyed could be grouped into an Enteromorpha-Corallina-Ulva dominated winter-spring association, a Sargassum-Ulva-Corallina dominated summer association and an Ulva-Gymnogongrus-Sargassum dominated autumn association.Regression analysis showed that changes in seawater temperature were the main influence on macroalgal biomass. Assemblage structure is influenced by substrate heterogeneity resulting from wave action at Chengshan Cape.     

The effect of sieve mesh size on the abundance and composition of macrophyte-associated macrofaunal assemblages

Sampling efficiency of several sieve mesh sizes (2, 1, 0.5, 0.2 and 0.1 mm) was tested in invertebrate assemblages associated to Sargassum stenophyllum. Samples of this brown algae were collected in southeastern Brazil to determine whether different sampling devices resulted in differences on the abundance and composition of the macrofauna. The 1 and 2 mm mesh sizes sampled less organisms, resulting in smaller abundances, densities, number of species and diversity, when compared to the other three mesh sizes, that achieved similar results. The most efficient sieve was the 0.5 mm mesh, sampling 85.8% and 94.5% of gammaridean and caprellidean amphipods, respectively, and 93.1% of gastropods. However, polychaetes and isopods were best sampled with the 0.2 mm sieve. Our results suggest that caution should be exercised when deciding on which sieve to use, as mesh sizes commonly used in benthic studies (1 and 0.5 mm) may result in very different estimates of diversity and abundance, as well as community structure patterns. This revised version was published online in July 2006 with corrections to the Cover Date.     

Temporal variation of amphipod assemblages associated with Sargassum filipendula (Phaeophyta) and its epiphytes in a subtropical shore

The phytal assemblages change in response to variation in biological and environmental conditions. In the present study, we evaluated the temporal variation of amphipod assemblages associated with a Sargassum filipendula bed in a subtropical shore, in relation to variation of the host alga, its epiphytes and local environmental conditions. Samples of S. filipendula with associated amphipods, water temperature, water movement and suspended solids were obtained monthly from June 2000 to May 2001. We recorded 24 species of amphipods associated with S. filipendula. Species richness varied throughout the year, with maximum values in October 2000 and minimum in April 2001. Total amphipod density gradually increased during the sampling period, with the highest value in March 2001. Amphipod diversity and evenness were both positively influenced by epiphyte load and negatively by temperature, with higher values during summer months. Total density and tube-builder density were also positively influenced by temperature, whereas nestler density was influenced by epiphyte load. Individual amphipod species showed significant density fluctuations over the year. The canonical correspondence analysis performed explained 88.2% of the variation, with a strong correlation of water movement, temperature and suspended solids with the first axis, and a strong effect of epiphyte load on both the first and the second axes. The temporal structural variation of the studied algal bed strongly influenced amphipod diversity and assemblage composition, possibly through direct and indirect effects.     

Compound‐specific isotope analysis of benthic foraminifer amino acids suggests microhabitat variability in rocky‐shore environments

The abundance and biomass of benthic foraminifera are high in intertidal rocky‐shore habitats. However, the availability of food to support their high biomass has been poorly studied in these habitats compared to those at seafloor covered by sediments. Previous field and laboratory observations have suggested that there is diversity in the food preferences and modes of life among rocky‐shore benthic foraminifera. In this study, we used the stable nitrogen isotopic composition of amino acids to estimate the trophic position, trophic niche, and feeding strategy of individual foraminifera species. We also characterized the configuration and structure of the endobiotic microalgae in foraminifera using transmission electron microscopy, and we identified the origin of endobionts based on nucleotide sequences. Our results demonstrated a large variation in the trophic positions of different foraminifera from the same habitat, a reflection of endobiotic features and the different modes of life and food preferences of the foraminifera. Foraminifera did not rely solely on exogenous food sources. Some species effectively used organic matter derived from endobionts in the cell cytoplasm. The high biomass and species density of benthic foraminifera found in intertidal rocky‐shore habitats are thus probably maintained by the use of multiple nitrogen resources and by microhabitat segregation among species as a consequence.     

The role of small mammals and birds in transport of matter through the shore zone of lakes

An attempt was made to estimate the role of small mammals and birds in the cycling of nutrients in the littoral zone of lakes. We estimated the approximate amounts of food, organic matter and elements ingested and egested during the breeding season by waterfowl. Small mammal densities were estimated, and their movements studied between different patches of a mosaic landscape, including the lake's shore region. Studies on passerine birds were aimed at determining the territories of some species in alder forest. Special attention was paid to the range of penetration of the reed belts by forest species.     

Typology of macrofaunal assemblages: a tool for the management of running waters in The Netherlands

A survey was carried out at 156 sites, situated in streams in the province of Overijssel (The Netherlands), to describe the macroinvertebrate assemblages and their environments. Fifty-six environmental variables were measured once at each sampling site. The main aim was to describe a typology of stream for this region.Different multivariate analysis techniques (clustering and ordination) were used in combination with ecological information on individual taxa to derive and describe site groups in terms of taxonomic composition and mean environmental conditions. The resulting site groups were termed cenotypes.Eleven cenotypes were distinguished. Differences between cenotypes were attributed to (combinations of) environmental key factors, namely, dimensions (width and depth), morphological (profile shape) and hydraulic stream characteristics, duration of drought and load of organic material.The typology offers a basis to be used for regional stream management and nature conservation. Efforts to improve stream ecosystems should be directed at the physical and hydraulic conditions.Typological studies are needed (especially in semi-natural landscapes) if we are to better understand, manage and conserve freshwater biota.     

Grassland invertebrate assemblages in managed landscapes: Effect of host plant and microhabitat architecture

Abstract Grasslands are often considered as two‐dimensional habitats rather than complex, multilayered habitats. However, native grasslands are complex habitats, with multiple layers of annual and perennial grasses, sedges, shrubs and mosses. Vegetation complexity, including plant type, quality and three‐dimensional structure is important for providing a variety of food and habitat resources for insects. Grazing by domestic livestock can affect these processes through the loss or fragmentation of habitats, as well as altering the vertical and horizontal vegetation structure. This study aimed to investigate the role of host plants and microhabitat architecture for determining foliage invertebrate assemblages. Different plant species supported distinct invertebrate assemblages and less complex host plants supported fewer invertebrate individuals and species. Manipulations of plant architecture changed the species composition of invertebrates, with most species found in more complex vegetation. This study illustrates the importance of host diversity and pasture complexity for invertebrate communities. Management practices that encourage a heterogeneous environment with diverse and structurally complex pastures should also sustain a more diverse and functional invertebrate assemblage.     

Effects of an alien invertebrate species and wave action on prey selection by African black oystercatchers (Haematopus moquini)

Abstract Shorebirds foraging in the intertidal have been shown to exert a significant effect on assemblage level processes; this is particularly true of the oystercatcher–limpet–algae system. The African black oystercatcher (Haematopus moquini) is endemic to the southern African coastline, where it plays a significant role in ecosystem processes as a rocky‐shore predator, especially of mussels and limpets. This understanding was based on studies of a rocky shore environment that has since been considerably modified following invasion of an alien mussel (Mytilus galloprovincialis). This invasion has not only changed the relative proportions of different food types on the shore, but has also greatly increased overall food biomass. We tested the previous model that food selection by oystercatchers reflected prey abundance and that intake by male and female oystercatchers differed owing to bill morphology. We predicted that this difference would persist despite the changed nature of the food base. We also predicted that wave action would modify prey selection as a result of both its influence on prey behaviour and its impact on searching and handling times of the birds. Overall, both sexes consumed more limpets than expected by encounter rate alone, but contrary to prediction, the relative proportions of different prey types taken post invasion did not differ between the sexes. Dietary convergence is interpreted as a result of greatly increased food biomass on the shore, which is also reflected in increased oystercatcher densities since the invasion. Also contrary to prediction there was no evidence that waves acted as indirect modifiers of the interaction between oystercatchers and their prey. The results of this study indicate that models of trophic cascades will need to be altered in the event of a significant change in a trophic level, which then effects behavioural changes in the key predator.     

Seasonal changes in an intertidal annual algal assemblage in northern Japan: The role of pre-emption and grazing on algal replacement

The pattern and process of seasonal changes in an intertidal annual algal assemblage were examined at Hiura, northern Japan. Short-term field experiments (<2months duration) were set up to quantify the effects of both grazing and pre-emption on species replacement in the assemblage in three different seasons. An 8-month field experiment was set up to quantify long-term effects, including the indirect effects of both grazing and competitive dominance on the community structure. Results suggested that seasonal change in the algal assemblage resulted from the interaction of abiotic environmental change, competition and grazing. The relative contribution of these factors varied within a short period, presumably as a result of seasonal changes in physical environmental stress, free space availability and grazing pressure. From February to March, when grazer density was low and there was much free space available for algae, the dominant species shifted from foliose green alga Monostroma angicava to filamentous red alga Bangia atropurpurea, because B.atropurpurea grew faster than M.angicava. This species replacement was not influenced strongly by biological interaction but by temporal changes in abiotic environmental conditions. From April to mid May, when there was less free space available for algae in the natural community, the dominant B.atropurpurea decreased with increasing foliose red alga Porphyra yezoensis, because only P.yezoensis was able to invade an area pre-empted by algae. Grazing did not affect this species replacement. After mid May, the two dominant species, P.yezoensis and B.atropurpurea, decreased. Their decline was mainly caused by desiccation stress and was partially affected by grazing.     

Reduced survey intensity and its consequences for marine reserve selection

There has been much interest in the potential of short-cuts in biodiversity surveys (e.g. physical surrogates, indicator groups, and lower taxonomic resolution) in systematic processes to select networks of representative marine reserves. This study tested the consequences for reserve selection of reducing survey intensity in intertidal rocky shores in southeast Australia. Using a reference data set of species' distributions based on surveys of two replicate sites in each of 15 locations, a reduction in survey intensity was simulated by randomly eliminating the data from one of the replicate sites in each location. A complementarity-based reserve selection algorithm was used to determine the number of locations required to represent all species once in a reserve network and the irreplaceability value of locations. A reduction in survey intensity led to increases in: the size of reserve networks (of between 8 and 17%); the irreplaceability value of locations; and the number of irreplaceable locations. These changes were caused by a reduction in the observed range sizes of species in the data sets simulating a reduced survey intensity.