Linking Land and Sea: Different Pathways for Marine Subsidies

Nutrients and energy derived from marine autotrophs subsidize shore ecosystems, increasing productivity and affecting food web dynamics and structure. In this study we examined how the inland reach of such inflow effects depends on vectors carrying the marine inflow inland and on landscape structure. We used stable isotopes of carbon and nitrogen to examine the roles of arthropod vectors in carrying marine-derived nutrients inland in two very different shore ecosystems: shore meadows in Sweden with marine inflows of algae and emerging chironomid midges; and sandy beaches and shore dunes in south-western Australia with marine inflows of algae and seagrass. In a colonization experiment, we found that deposited wrack on the beach is quickly colonized by both grazers and predators. However, in both systems we found a larger inland reach of the marine subsidy than could be accounted for by deposited macrophytes on shores alone, and that dipterans and spiders potentially functioned as vectors for the inflow. Our results indicate that marine inflows are important for near-shore terrestrial ecosystems well above the water’s edge, and that this effect is largely due to arthropod vectors (mainly dipterans and spiders) in both low-productivity sandy beach ecosystems at the Indian Ocean coast of Australia, and more productive shore meadows on the Baltic Sea coast of Sweden. Our findings also suggest that the type of vector transporting marine material inland may be as important as the productivity contrast between ecosystems for explaining the degree of marine influence on the terrestrial system.     

Major food web properties of two sandy beaches with contrasting morphodynamics, and effects on the stability

We determined major structural properties influencing the food webs of two sandy beaches with contrasting morphodynamics in the Atlantic coast of Uruguay: reflective (narrow and steep) and dissipative beaches (wide and flat). Furthermore, we evaluated how these characteristics could influence the stability of the local food webs. To this end, we examined the correlation of several food web properties with different ecosystem types (including freshwater habitats, estuary, marine, and terrestrial environments) using a principal components analysis. Sandy beach food web components included detritus, phytoplankton, zooplankton, benthic invertebrates, fishes, and seabirds. Our results revealed that the dissipative beach presented higher trophic levels, a higher number of trophic species, more links per species, as well as a higher proportion of intermediate trophic species, but lower connectance and proportion of omnivorous species than the reflective beach. The variation in the food web properties was explained by two principal components. Sandy beach food webs contribute mainly to one dimension of the principal components analysis that was determined by the number of trophic species, links per species, the trophic similarity, and the characteristic path length. We suggest that species and link characteristics, such as predominance of scavengers and detritivorous, the relatively high connectance and the short path length are drivers in the food web structure and may play a role in the community dynamic.     

Variation in the abundance of Pseudo-nitzschia and domoic acid with surf zone type

Most harmful algal blooms (HAB) originate away from the shore and, for them to endanger human health, they must be first transported to shore after which they must enter the surf zone where they can be feed upon by filter feeders. The last step in this sequence, entrance into the surf zone, depends on surf zone hydrodynamics. During two 30-day periods, we sampled Pseudo-nitzschia and particulate domoic acid (pDA) in and offshore of a more dissipative surf zone at Sand City, California (2010) and sampled Pseudo-nitzschia in and out of reflective surf zones at a beach and rocky shores at Carmel River State Beach, California (2011). At Sand City, we measured domoic acid in sand crabs, Emerita analoga. In the more dissipative surf zone, concentrations of Pseudo-nitzschia and pDA were an order of magnitude higher in samples from a rip current than in samples collected just seaward of the surf zone and were 1000 times more abundant than in samples from the shoals separating rip currents. Domoic acid was present in all the Emerita samples and varied directly with the concentration of pDA and Pseudo-nitzschia in the rip current. In the more reflective surf zones, Pseudo-nitzschia concentrations were 1–2 orders of magnitude lower than in samples from 125 and 20 m from shore. Surf zone hydrodynamics affects the ingress of Pseudo-nitzschia into surf zones and the exposure of intertidal organisms to HABs on the inner shelf.     

Food supply, grazing activity and growth rate in the limpet Patella vulgata L.: a comparison between exposed and sheltered shores

The limpet Patella vulgata L. is an important microphagous grazer on intertidal rocky shores of north-west Europe, occurring across the wave exposure gradient. Groups of P. vulgata were selected at mid-tide level of two exposed shores and two sheltered, fucoid dominated shores on the Isle of Man, British Isles, and manipulated to equivalent densities and population structure. The level of grazing activity and growth rate were determined over a 1-year period. At the same time, the abundance of epilithic microalgae, measured as the concentration of chlorophyll a, was determined as an estimate of food supply. Microagal abundance showed a seasonal pattern in both exposed and sheltered conditions, with higher levels in winter compared to summer. In both seasons, the microalgal resource was more abundant on the sheltered shore studied. The level of grazing activity in P. vulgata showed a seasonal pattern on the exposed but not the sheltered shores. Averaged over the year, grazing activity on the exposed shores was over double that on sheltered shores. Thus, in sheltered conditions, food supply for limpets was high and grazing activity low; in exposed conditions, food supply was low and grazing activity high. The growth rate of P. vulgata, measured as increase in shell length, showed no significant difference between exposed and sheltered shores. Growth rate was also determined in P. vulgata at natural densities. Although the overall density declined with decreasing exposure to wave action, the density per unit area of grazeable substance was higher in shelter. In these populations, the mean growth rate was over twice as high on exposed compared to sheltered shores.     

Picturing beach erosion and deposition trends using PSInSAR: an example from the non-barred southern west coast of India

Coastal dynamics can rapidly alter beach morphology. In some places, such as the non-barred southern west coast of India, studying changes to beach morphology is a relatively arduous task. Persistent Scatterer Interferometric Synthetic Aperture Radar (PSInSAR), a remote sensing technique that utilizes stacks of radar images for accurate long-term monitoring of ground features, allows for detailed observations of coastal morphodynamics. Twenty-two single look complex (SLC) synthetic aperture radar (SAR) Sentinel-1A images, acquired from 4 March 2016 to 3 June 2017, are used to monitor geomorphological processes such as coastal erosion and deposition. Ground deformation measurements from PSInSAR processing shows the coast stretching?~?70 km between Thaickal and Munambam is highly dynamic, characterized by phases of erosion and deposition. The highest negative displacement of ? 24.9 mm at Thaickal versus the?+?7.6 mm at Chellanam in the north show the co-existing milieus of erosion and deposition. PSInSAR results concur with corresponding Google Earth images. In addition, beach sediment texture and scanning electron microscope grain micro-texture in the beach segment further corroborate temporal phases of erosion and deposition. Two locations are identified as typical erosional sites, while one location typified deposition. Erosion and deposition or rebuilding of beaches, usually correspond respectively with the onset and offset of SW monsoon wave climate in the Arabian Sea. When zones of deposition are mainly located in the proximity of river and/or lake inlets, stretches characterized by erosion are distal to inlets. Linear regression analyses of displacement–time series plots were used to identify general erosion or depositional regimes along beach segments. Results from this study illustrate how PSInSAR is a capable and reliable processing tool for monitoring temporal phases of coastal morpho-dynamics.

     

Spatial subsidies in spider diets vary with shoreline structure: Complementary evidence from molecular diet analysis and stable isotopes

Inflow of matter and organisms may strongly affect the local density and diversity of organisms. This effect is particularly evident on shores where organisms with aquatic larval stages enter the terrestrial food web. The identities of such trophic links are not easily estimated as spiders, a dominant group of shoreline predator, have external digestion. We compared trophic links and the prey diversity of spiders on different shore types along the Baltic Sea: on open shores and on shores with a reed belt bordering the water. A priori, we hypothesized that the physical structure of the shoreline reduces the flow between ecosystem and the subsidies across the sea–land interface. To circumvent the lack of morphologically detectable remains of spider prey, we used a combination of stable isotope and molecular gut content analyses. The two tools used for diet analysis revealed complementary information on spider diets. The stable isotope analysis indicated that spiders on open shores had a marine signal of carbon isotopes, while spiders on reedy shores had a terrestrial signal. The molecular analysis revealed a diverse array of dipteran and lepidopteran prey, where spiders on open and reedy shores shared a similar diet with a comparable proportion of chironomids, the larvae of which live in the marine system. Comparing the methods suggests that differences in isotope composition of the two spider groups occurred because of differences in the chironomid diets: as larvae, chironomids of reedy shores likely fed on terrestrial detritus and acquired a terrestrial isotope signature, while chironomids of open shores utilized an algal diet and acquired a marine isotope signature. Our results illustrate how different methods of diet reconstruction may shed light on complementary aspects of nutrient transfer. Overall, they reveal that reed belts can reduce connectivity between habitats, but also function as a source of food for predators.     

The vertical distribution of Chthamalus montagui and Chthamalus stellatus (Crustacea,Cirripedia) in two areas of the NW Mediterranean Sea

The Mediterranean Sea is characterised by a small tidal range (0.3–1 m). Despite this, intertidal communities are well established and their upper limits often extend above mean high water level. Organisms living in the intertidal region and in the supralittoral zone rely on both tides and wave action to perform their biological functions. Lack of food, desiccation and predation are common stresses in such a harsh environment. The present study deals with the vertical distribution of two species of intertidal barnacles, Chthamalus montagui Southward and Chthamalus stellatus (Poli), which are the main constituents of the barnacle belt along Mediterranean rocky shores. Previous work, carried out in the Atlantic, showed that where the distribution ranges of the two Chthamalus species overlap, C. montagui is more common in the upper barnacle zone while C. stellatus is dominant lower down. The main aims of our study are: (1) to establish if there is a relationship between position and extension of the barnacle belt on the shore and tidal range and/or wave exposure, (2) to test the hypothesis that in the study areas C. montagui is more abundant than C. stellatus high on the shore, and that the pattern is reversed lower down. Barnacle populations were monitored in summer 1998 in the Gulf of Genoa (Ligurian Sea) and in the Gulf of Trieste (North-Adriatic Sea). The two areas differ in tidal range and hydrodynamism, the former presenting quite strong wave action and a tidal range of 30 cm, the latter having limited wave action and 1 m tidal range. Three shores were randomly selected in each gulf and two transects on each shore. Counts of barnacles in 10 ^* 10 cm quadrats were done at different shore heights along each transect. The data was subjected to analysis of variance. Results showed that a more pronounced hydrodynamic regime corresponded to a shift of the barnacle belt towards the higher shore (Gulf of Genoa), while in more sheltered areas (Gulf of Trieste), the barnacle distribution was confined to the intertidal region. The relative spatial distribution of C. montagui and C. stellatus within the barnacle belt varied locally, even between transects on the same shore, and this obscured the distribution pattern along the vertical gradient. Nevertheless, it was still possible to conclude that at mid and high shore in Genoa, C. stellatus was more abundant than C. montagui, while in Trieste the pattern was reversed.     

Predator diversity enhances secondary production and decreases the likelihood of trophic cascades

We manipulated the diversity of top predators in a three trophic level marine food web. The food web included four top benthic marine fish predators (black goby, rock goby, sea scorpion and shore rockling), an intermediate trophic level of small fish, and a lower trophic level of benthic invertebrates. We kept predator density constant and monitored the response of the lower trophic levels. As top predator diversity increased, secondary production increased. We also observed that in the presence of the manipulated fish predators, the density of small gobiid fish (intermediate consumers) was suppressed, releasing certain groups of benthic invertebrates (caprellid amphipods, copepods, nematodes and spirorbid worms) from heavy intermediate predation pressure. We attribute the mechanism responsible for this trophic cascade to a trait-mediated indirect interaction, with the small gobiid fish changing their use of space in response to altered predator diversity. In the absence of top fish predators, a full-blown trophic cascade occurs. Therefore the diversity of predators reduces the likelihood of trophic cascades occurring and hence provides insurance against the loss of an important ecosystem function (i.e. secondary production).     

Effects of microbivore species composition and basal resource enrichment on trophic-level biomasses in an experimental microbial-based soil food web

Previous theoretical and empirical evidence suggests that species composition within trophic levels may profoundly affect the response of trophic-level biomasses to enhanced basal resources. To test whether species composition of microbivorous nematodes has such an effect in microbial-based soil food webs, I created three microcosm food webs, consisting of bacteria, fungi, bacterial-feeding nematodes (Acrobeloides tricornus, Caenorhabditis elegans), fungal-feeding nematodes (Aphelenchus avenae, Aphelenchoides sp.) and a predatory nematode (Prionchulus punctatus). The food webs differed in species composition at the second trophic level: food web A included A. tricornus and Aph. avenae, food web B included C. elegans and Aphelenchoides sp., and food web AB included all four species. I increased basal resources by adding glucose to half of the replicates of each food web, and sampled microcosms destructively four times during a 22-week experiment to estimate the biomass of organisms at each trophic level. Microbivore species composition significantly affected bacterivore and fungivore biomass but not bacterial, fungal or predator biomass. Greatest bacterivore and fungivore biomass was found in food web A, intermediate biomass in food web AB, and smallest biomass in food web B. Basal resource addition increased the biomass of microbes and microbivores but did not affect predator biomass. Importantly, microbivore species composition did not significantly modify the effect of additional resources on trophic-level biomasses. The presence of a competitor reduced the biomass of A. tricornus and Aph. avenae, in that the biomass of these species was less in food web AB than in food web A, whereas the biomass of C. elegans and Aphelenchoides sp. was not affected by their potential competitors. The biomass of Aph. avenae increased with additional resources in the absence of the competitor only, while the biomass of A. tricornus and Aphelenchoides sp. increased also in the presence of their competitors. The results imply that microbivore species composition may determine the second-level biomass in simple microbe-nematode food webs, but may not significantly affect biomass at other levels or modify the response of trophic-level biomasses to enhanced basal resources. The study also shows that even if the role of predation in a food web is diminished, the positive response of organisms to increased resource availability may still be hindered by competition. Received: 22 June 1998 / Accepted: 28 August 1998     

Physical and biological conditions on a steep intertidal gradient at Rottnest Island,Western Australia

Seven species of grazing molluscs, two littorinids, one nerite, three limpets and one chiton, lived on the vertical intertidal rock wall at the landward edge of the coastal limestone platforms at Rottnest Island, Western Australia. On the average, the vertical ranges of these species overlapped broadly, although a consistent zonation pattern was conspicuous from the platform surface upwards for 2 m. On these vertical intertidal shores, physical and biological conditions were predicted quantitatively from easily made measurements of vertical height on the shore; the percentage of time any shore level was immersed in seawater, the percent weight loss of plaster clods, the standing crop of algae, and the growth rate of the limpet, Notoacmea onychitis, all decreased linearly or semi-logarithmically with increasing height on the shore. The standing crop of animals was greatest, largely due to the presence of the chiton, between 40 and 70 cm from the platform surface and decreased rapidly down the shore and more gradually towards higher shore levels. We interpret this information and the positive correlation between algal production rate, and egestion rate of the animal community at various levels of the shore as evidence supporting the idea that food may be in short supply on these shores.     

Effects of defoliation intensity on soil food-web properties in an experimental grassland community

We established a greenhouse experiment based on replicated mini‐ecosystems to evaluate the effects of defoliation intensity on soil food‐web properties in grasslands. Plant communities, composed of white clover (Trifolium repens), perennial ryegrass (Lolium perenne) and plantain (Plantago lanceolata) with well‐established root and shoot systems, were subjected to five defoliation intensity treatments: no trimming (defoliation intensity 0, or DI 0), and trimming of all plant material to 35 cm (DI 1), 25 cm (DI 2), 15 cm (DI 3) and 10 cm (DI 4) above soil surface every second week for 14 weeks. Intensification of defoliation reduced shoot production and standing shoot and root mass of plant communities but increased their root to shoot ratio. Soil microbial activity and biomass decreased with intensification of defoliation. Concentrations of NO₃–N in soil steadily increased with intensifying defoliation, whereas NH₄–N concentrations did not vary between treatments. Numbers of microbi‐detritivorous enchytraeids, bacterial‐feeding rotifers and bacterial‐feeding nematodes steadily increased with intensifying defoliation, while the abundance of fungal‐feeding nematodes was significantly enhanced only in DI 3 and DI 4 relative to DI 0. The abundance of herbivorous nematodes per unit soil mass was lower in DI 3 and DI 4 than in DI 0, DI 1 and DI 2, but when calculated per unit root mass, their abundance tended to increase with defoliation intensity. The abundance of omnivorous and predatory nematodes appeared to be highest in the most intensely defoliated systems. The ratio of abundance of fungal‐feeding nematodes to that of bacterial‐feeding nematodes was not significantly affected by defoliation intensity. The results infer that defoliation intensity may significantly alter the structure of soil food webs in grasslands, and that defoliation per se is able to induce patterns observed in grazing studies in the field. The results did not support hypotheses that defoliation per se would cause a shift between the bacterial‐based and fungal‐based energy channels in the decomposer food web, or that herbivore and detritivore densities in soil would be highest under intermediate defoliation. Furthermore, our data for microbes and microbial feeders implies that the effects of defoliation intensity on soil food‐web structure may depend on the duration of defoliation and are therefore likely to be dynamic rather than constant in nature.     

Nahrungsbeziehungen der Turbellarien in Küstensalzwiesen

Turbellarians represent one of the most characteristic and often also most numerous elements of the salt-marsh biocoenosis. The analysis of their food relations promised, therefore, insights into the food web in salt marshes. Two questions are of particular importance: (1) Is autochthonous or allochthonous food more important? (2) Are there noticeable relations to the aerial phase of the biotope? Gut-content surveys, feeding-experiments and information from literature indicate, that the main food of salt-marsh turbellarians consists of oligochetes, nematodes and diatoms. There exist certain relations between the taxonomic categories of turbellarians and their food sources, and, furthermore, between the type of pharynx and the kind of food. Oligochetes are 4 and nematodes up to 14 times more abundant than turbellarians. Nematodes are also likely to be the most important food competitors for turbellarians. Enemies of salt-marsh turbellarians are the polychaeteNereis diversicolor and sporozoan parasites. The main food of salt-marsh turbellarians consists of organisms existing in sufficient numbers in the biotope; a dependency on allochthonous food is therefore unlikely. Allochthonous food (e. g. copepods from adjacent mud flats) may, however, at times enlarge the food resources available. Little evidence for the use of air-living organisms as food, and no evidence for air-living enemies, was found. The aquatic food chain in salt marshes ends at the level of flatworm-eating polychaetes or, where these are absent, already at that of the turbellarians themselves. Although the biomass (standing crop) of primary producers reaches its maximum in the most protected part of the beach (salt marshes), maximum population density of turbellarians is found in parts of maximum exposure. This fact is considered to be due to the great amount of allochthonous food available at the upper edge of the surf zone.     

Bottom‐up control in the benthos of ocean‐exposed sandy beaches?

Bottom‐up control is a fundamental structuring force in food webs. Food webs of ocean‐exposed sandy beaches are predicted to be bottom‐up controlled systems, underpinned by imported organic matter rather than in situ primary production. This ecological model of resource‐based regulation of biological assemblages is juxtaposed against a prevailing paradigm built around a dominance of physical drivers in sandy beach ecosystems. Surprisingly, given the apparently ubiquitous energetic subsidies of beach food webs, the central premise of bottom‐up control has not been tested. Here we experimentally manipulated in situ nutrient levels on a sandy beach to test food web responses at the levels of primary producers (benthic microalgae) and their grazers (meiofauna). The meiofauna community as a whole appeared most strongly influenced by the local physical environment, particularly changes in sediment grain size – this supports the traditional ‘environmental control paradigm’. We also detected a significant, positive response of two consumer groups of the meiofauna (nematodes, ostracods) to nutrient enrichment that supports a model of biological, bottom‐up control. Although the predicted response of elevated producer biomass following nutrient enrichment was not detected, intense grazing pressure on new, stimulated production may have masked positive responses by the primary producers. Multichannel regulation of food webs is likely for many exposed sandy beaches, albeit an often lower importance of in situ bottom‐up forces compared with stronger environmental control.     

Mortalities caused by off-road vehicles (ORVs) to a key member of sandy beach assemblages,the surf clam <Emphasis Type="Italic">Donax deltoides</Emphasis>

Sandy beaches are prime recreational areas, but human use of beaches is not without ecological consequences. Driving of off-road vehicles on beaches for recreational pursuits is perhaps the physically most severe form of direct anthropogenic disturbance on sandy shores. Potential management and conservation interventions lack, however, data on how sensitive beach species are to vehicle impacts. We therefore experimentally quantified the link between beach traffic and lethal damages caused by vehicles to sandy shore invertebrates, using surf clams (Donax deltoides) as the biological response variable. Although clams had some tolerance against vehicles at low traffic volumes (5 vehicle passes), more than half of them were killed at higher traffic volumes (75 passes) in situations where cars traversed soft sand and turned across the beach face. Overall, both traffic volume and driver behaviour (i.e. straight vs. turning vehicle tracks) determined the incidence of direct crushing of clams under vehicles. Our data demonstrate that recreational use of ORVs is a source of mortality for beach invertebrates, but equally caution against extrapolating impact data from hard-shelled clams to potentially more sensitive soft-bodied species. Robust management interventions that seek to mitigate ecological damage from beach traffic will therefore require information on the functional relationship between the form, intensity and frequency of human disturbance and the biological responses for entire faunal assemblages on sandy shores. Handling editor: T. P. Crowe     

Large scale patterns and trophic structure of southern African rocky shores: the roles of geographic variation and wave exposure

In this study we revise the biogeographic delimitation, and large-scale patterns of community structure of the intertidal rocky shores of southern Africa. We use binary (presence/absence) and per-species biomass data collected at fifteen localities and thirty-seven different rocky sites, encompassing the shores of southern Namibia, South Africa and southern Mozambique. Multivariate analyses revealed that the shores of southern Africa (south of 25°) can be divided into three main biogeographic provinces: the west coast or Namaqua province, the south coast or Agulhas province and the east coast or Natal province. The biomass structure of the intertidal rocky shores communities of southern Africa varied at a large scale, corresponding to biogeographic differences, while local-scale variation accorded with the intensity of local wave action. The average biomass of west coast communities was on average significantly greater than that of the south and east provinces. At a local scale, the community biomass on exposed shores was an order of magnitude greater than on sheltered shores, within all biogeographic provinces. Semi-exposed shores exhibited intermediate average biomass. The trophic structure of these communities varied significantly with wave action: autotrophs, filter-feeders and invertebrate predators were more prevalent on wave exposed than sheltered shores, whereas grazers were more abundant on sheltered and semi-exposed shores. Exposed shores were consistently dominated by far fewer species than semi-exposed and sheltered shores, independently of biogeographic differences. Within all biogeographic provinces semi-exposed and sheltered shores were more diverse than exposed shores. West coast intertidal communities therefore had high levels of biomass, but were consistently species-poor. Several working hypotheses that could explain these large and small-scale patterns are presented.     

Orientation of the sandhopperTalitrus saltator (Amphipoda,Talitridae) living on dynamic sandy shores

Summary The littoral sandhopperTalitrus saltator Montagu maintains its position along the beach mainly by means of an innate solar orientation. Different populations show different directional tendencies depending on the direction of the sealand axis. Experiments were designed which aimed to disentangle the innate and acquired components of the escape orientation. Therefore, the solar orientation of experienced adults living on shores subject to fluctuations of the shoreline and of their laboratory-born (inexperienced) offspring were compared. The innate sun compass orientation shown by the young is modified by learning during life under natural conditions, depending on the variability of the shore. Results showed: (1) an improvement in the solar orientation in individuals living on relatively constant shores; (2) a loss of precision in individuals living on highly dynamic shores; (3) a change to a new ecologically efficient escape direction in individuals living on a relatively stable lagoon shore.Abbreviation TED theoretical escape direction     

Contribution of nematodes to the structure and function of the soil food web

As carbon and energy flow through the soil food web they are depleted by the metabolic and production functions of organisms. To be sustained, a "long" food web, with a large biomass at higher trophic levels, must receive a high rate of rhizodeposition or detrital subsidy, or be top-populated by organisms of slow growth and long life cycle. Disturbed soil food webs tend to be bottom heavy and recalcitrant to restoration due to the slow growth of upper predator populations, physical and chemical constraints of the soil matrix, biological imbalances, and the relatively low mobility and invasion potential of soil organisms. The functional roles of nematodes, determined by their metabolic and behavioral activities, may be categorized as ecosystem services, disservices or effect-neutral. Among the disservices attributable to nematodes are overgrazing, which diminishes services of prey organisms, and plant-damaging herbivory, which reduces carbon fixation and availability to other organisms in the food web. Unfortunately, management to ameliorate potential disservices of certain nematodes results in unintended but long-lasting diminution of the services of others. Beneficial roles of nematodes may be enhanced by environmental stewardship that fosters greater biodiversity and, consequently, complementarity and continuity of their services.     

Nematode Indicators of Organic Enrichment

The organisms of the soil food web, dependent on resources from plants or on amendment from other sources, respond characteristically to enrichment of their environment by organic matter. Primary consumers of the incoming substrate, including bacteria, fungi, plant-feeding nematodes, annelids, and some microarthropods, are entry-level indicators of enrichment. However, the quantification of abundance and biomass of this diverse group, as an indicator of resource status, requires a plethora of extraction and assessment techniques. Soluble organic compounds are absorbed by bacteria and fungi, while fungi also degrade more recalcitrant sources. These organisms are potential indicators of the nature of incoming substrate, but current methods of biomass determination do not reliably indicate their community composition. Guilds of nematodes that feed on bacteria (e.g., Rhabditidae, Panagrolaimidae) and fungi (e.g., Aphelenchidae, Aphelenchoididae) are responsive to changes in abundance of their food. Through direct herbivory, plant-feeding nematodes (e.g., many species of Tylenchina) also contribute to food web resources. Thus, analysis of the nematode community of a single sample provides indication of carbon flow through an important herbivore channel and through channels mediated by bacteria and fungi. Some nematode guilds are more responsive than others to resource enrichment. Generally, those bacterivores with short lifecycles and high reproductive potential (e.g., Rhabditidae) most closely mirror the bloom of bacteria or respond most rapidly to active plant growth. The feeding habits of some groups remain unclear. For example, nematodes of the Tylenchidae may constitute 30% or more of the individuals in a soil sample; further study is necessary to determine which resource channels they portray and the appropriate level of taxonomic resolution for this group. A graphic representation of the relative biomass of bacterivorous, fungivorous, and herbivorous nematodes provides a useful tool for assessing the importance of the bacterial, fungal, and plant resource channels in an extant food web.     

Distribution and abundance of co-occurring chthamalid barnacles Chthamalus montagui and Chthamalus stellatus (Crustacea,Cirripedia) on the southwest coast of Portugal

Two co-occurring chthamalid barnacle species occur on the rocky shores of continental Portugal: Chthamalus stellatus (Poli) and Chthamalus montaguiSouthward. In the present study, patterns of distribution and abundance (density and percentage cover), both vertically (tidal level) and horizontally (wave action), were studied for each species on headlands (more wave action) and not on headlands (less wave action) and at two spatial scales (shore and site) and three tidal levels (low, mid and upper). Chthamalus montagui was significantly more abundant than C. stellatus in both degrees of wave action and at all tidal levels. Despite this great difference in abundance and considering each species individually, C. montagui was more abundant at mid tide-level, while C. stellatus was more abundant at the low tide-level, and C. montagui was less abundant on headlands, while C. stellatus was more abundant on headlands. Small-scale variability (between sites, tens of meters) of abundance of each species was detected, while differences on a larger scale (between shores, tens of kilometres) were not significant.     

Seasonal patterns of food availability: influences on the reproductive output and body condition of the herbivorous crab Grapsus albolineatus

 Rocky shores in Hong Kong experience marked seasonal differences in climate resulting in seasonal changes in macroalgal assemblages. The tropical rocky shore crab, Grapsus albolineatus, feeds selectively on filamentous algae through the year but the abundance of these algae and foliose algae is greatly reduced during the summer when encrusting algae dominate the shores and the crab’s diet. This switch in diet may have implications for the reproductive output of this crab. Standing crop of algae varied greatly through the year, peaking in March and April, when the nutritional quality of the algae was also highest. On the shore, available algal protein and energy were both lowest in July. The crab selected an algal diet rich in nutrients than that available to it on the shore for all months of the year except September to December for protein, and July and August for energy. The input of animal matter considerably increased the protein content of the diet, but made little difference to the energy content. Growth and body condition were greatest during March and June, coinciding with the peak in algal biomass. Storage of nutrients in the hepatopancreas of G. albolineatus commenced in November, coinciding with the increase in biomass and quality of algae on the shores, and then, peaking in May and June, these nutrients were utilised by the reproductive organs during the reproductive season which ran from April to November. The middle of the reproductive season coincided with the period when the standing crop of algae on the shore was at its lowest levels and poorest quality. Cycles of growth, reproduction and storage in G. albolineatus appear to be directly influenced by seasonal patterns of algal food availability. Nutrient storage in the hepatopancreas, during periods when food is nutritionally rich and is most abundant, does not guarantee reproductive success of the crab, but appears to be a prerequisite. Selection of an optimal diet (energy- and protein-rich) in a seasonal environment by ingesting abundant nutrient-rich algal species and the opportunistic consumption of animal matter strongly influences growth and reproductive output of G. albolineatus. Received: 18 March 1996 / Accepted: 17 July 1996