Latitudinal clines in food distribution correlate with differential migration in the Western Sandpiper

We report that a latitudinal cline in intertidal food distribution is associated with the nonbreeding distribution of the Western Sandpiper (Calidris mauri). This novel result is the first to demonstrate a clear relationship between patterns of differential nonbreeding distribution and food availability for any shorebird species. Within each age class and sex, longer-billed Western Sandpipers winter further south. Moreover, females, the longer-billed sex, tend to winter south of males. Thus, both inter- and intra-sexual clines in bill morphology result in an overall gradient of increasing bill length from north to south. Longer-billed birds are able to extract prey that are buried more deeply in the sediment; therefore, we predicted shifts in the vertical distribution of food resources to coincide with the clines in bill morphology across the nonbreeding range. We tested our prediction by measuring biofilm density and the vertical distribution of macrofaunal invertebrates at six nonbreeding sites. Although no latitudinal trend was observed for biofilm, the vertical distribution of invertebrates was consistent with our prediction and revealed that the greatest relative abundance of surface prey occurred at northern nonbreeding sites and declined with decreasing latitude. We discuss the potential implications of these findings in the context of competing evolutionary hypotheses of differential migration and bill dimorphism in shorebirds.     

Waterbird Communities and Habitat Relationships in Coastal Pastures of Northern California

We examined waterbird assemblages (diversity, composition, and species' densities) in 20 pastures near Humboldt Bay, California, in relation to habitat characteristics (vegetation height, soil penetrability, water depth), abundance of invertebrates (worms and other invertebrates), and presence of livestock. From October 1991 to May 1992 we observed 29 species and 10,776 birds, most (78%) of which foraged. Nonrandom pasture use by birds resulted in a highly clumped spatial distribution (s²:{ovbar>x} ratio = 42.1). Habitat characteristics of pastures correlated with this nonrandom pattern: waterbird diversity and densities of three sandpiper species and one gull species correlated negatively ( r = −0.61 and r = −0.44 to −0.67, respectively) with vegetation height; densities of two plover species correlated negatively ( r = −0.39 and −0.45) with soil penetrability; and waterfowl densities correlated positively with water depth ( r = 0.97). Species composition varied among pastures. Wading birds used pastures with tall vegetation, shorebirds and gulls frequented short-grass pastures, and waterfowl used flooded pastures of waterbirds and their densities increased in association with livestock.In coastal areas where much intertidal habitat has been reclaimed as postureland, pastures offered valuable habitats to nonbreeding waterbirds. We suggest that grazing in coastal pastures can be used to provide a mosaic of vegetation heights, which would yield greater waterbird diversity as well as higher densities of some species.     

Experimental test of the effects of a non-native invasive species on a wintering shorebird

The abundance of nearly one-quarter of the world's shorebird species is declining. At the same time, the number of non-native species in coastal ecosystems is increasing rapidly. In some cases, non-native species may affect negatively the abundance and diversity of shorebird prey species. We conducted an experimental study of the effects of the introduced European green crab (Carcinus maenas) on prey consumption by wintering Dunlin (Calidris alpina) in a central California estuary. We placed green crabs and Dunlin sequentially in field enclosures and measured changes in density of benthic invertebrate prey (e.g. polychaetes and small clams), Dunlin biomass, and gut contents of both Dunlin and crabs and observed foraging behavior of Dunlin. Green crabs significantly affected Dunlin foraging success through both direct and indirect multitrophic linkages. In enclosures with high densities of green crabs, crab foraging reduced the availability of polychaetes, and Dunlin consumed significantly fewer polychaetes compared with Dunlin in enclosures without crabs. High densities of green crabs were also associated with increased availability of small clams. Dunlin consumed significantly more small clams compared with Dunlin in enclosures without crabs. In our literature survey of studies of effects of non-native invasive species on shorebirds, we found three prior experiments that addressed the effect of non-native invasive species on shorebirds. Results of two of these studies showed positive direct effects of non-native invertebrates on shorebirds, 1 showed negative direct effects of a non-native plant on shorebirds through habitat conversion, and none showed indirect effects of non-native invertebrates. We suggest future management of shorebirds explicitly examine how non-native marine species, particularly invertebrates, directly and indirectly affect shorebirds.     

Foraging in a tidally structured environment by Red Knots (Calidris canutus): ideal, but not free

Besides the "normal" challenge of obtaining adequate intake rates in a patchy and dangerous world, shorebirds foraging in intertidal habitats face additional environmental hurdles. The tide forces them to commute between a roosting site and feeding grounds, twice a day. Moreover, because intertidal food patches are not all available at the same time, shorebirds should follow itineraries along the best patches available at a given time. Finally, shorebirds need additional energy stores in order to survive unpredictable periods of bad weather, during which food patches are covered by extreme tides. In order to model such tide-specific decisions, we applied stochastic dynamic programming in a spatially explicit context. Two assumptions were varied, leading to four models. First, birds had either perfect (ideal) or no (non-ideal) information about the intake rate at each site. Second, traveling between sites was either for free or incurred time and energy costs (non-free). Predictions were generated for three aspects of foraging: area use, foraging routines, and energy stores. In general, non-ideal foragers should feed most intensely and should maintain low energy stores. If traveling for such birds is free, they should feed at a random site; otherwise, they should feed close to their roost. Ideal foragers should concentrate their feeding around low tide (especially when free) and should maintain larger energy stores (especially when non-free). If traveling for such birds is free, they should feed at the site offering the highest intake rate; otherwise, they should trade off travel costs and intake rate. Models were parameterized for Red Knots (Calidris canutus) living in the Dutch Wadden Sea in late summer, an area for which detailed, spatially explicit data on prey densities and tidal heights are available. Observations of radio-marked knots (area use) and unmarked knots (foraging routines, energy stores) showed the closest match with the ideal/non-free model. We conclude that knots make state-dependent decisions by trading off starvation against foraging-associated risks, including predation. Presumably, knots share public information about resource quality that enables them to behave in a more or less ideal manner. We suggest that our modeling approach may be applicable in other systems where resources fluctuate in space and time.     

Effects of salinity on multiplication and transmission of an intertidal trematode parasite

Salinity levels vary spatially in coastal areas, depending on proximity to freshwater sources, and may also be slowly decreasing as a result of anthropogenic climatic changes. The impact of salinity on host–parasite interactions is potentially a key regulator of transmission processes in intertidal areas, where trematodes are extremely common parasites of invertebrates and vertebrates. We investigated experimentally the effects of long-term exposure to decreased salinity levels on output of infective stages (cercariae) and their transmission success in the trematode Philophthalmus sp. This parasite uses the snail Zeacumantus subcarinatus as intermediate host, in which it asexually produces cercariae. After leaving the snail, cercariae encyst externally on hard substrates to await accidental ingestion by shorebirds, which serve as definitive hosts. We found that at reduced salinities (25 or 30 psu), the cercarial output of the parasite was lower, the time taken by cercariae to encyst was longer, fewer cercariae successfully encysted and encysted parasites had lower long-term survival than at normal seawater salinity (35 psu). The strong effect of salinity on the replication and transmission of this parasite suggests that there may be sources and sinks of transmission to birds along coastal areas, depending on local salinity conditions. Also, unless it evolves to adapt to changing conditions, the predicted reduction in salinity as a consequence of climate change may have negative impact on the parasite’s abundance.     

Desiccation stress in two intertidal beachrock biofilms

Chlorophyll a fluorescence was used to look at the effect of desiccation on the photophysiology in two beachrock microbial biofilms from the intertidal rock platform of Heron Island, Australia. The photophysiological response to desiccation differed between the beachrock microbial communities. The black biofilm from the upper shoreline, dominated by Calothrix sp., showed a response typical of desiccation-tolerant cyanobacteria, where photosynthesis closed down during air exposure with a rapid and complete recovery upon rehydration. In contrast, the pink biofilm from the mid-intertidal zone, dominated by Blennothrix sp., showed no distinct response to desiccation stress and instead maintained reduced photosynthesis throughout drying and re-wetting cycles. Spatial differences in photosynthetic activity within the black biofilm were evident with a faster recovery rate of photosynthesis in the surface cyanobacteria than in the deeper layers of the biofilm. There was no variation with depth in the pink biofilm. The photophysiological differences in desiccation responses between the beachrock biofilms exemplify the ecological niche specialisation of these complex microbial communities, where the functional differences help to explain their vertical distribution on the intertidal shoreline.     

Relevance of wound-activated compounds produced by diatoms as toxins and infochemicals for benthic invertebrates

Plants evolve the production of wound-activated compounds (WACs) to reduce grazing pressure. In addition, several plant-produced WACs are recognized by various invertebrates, playing the role of infochemicals. Due to co-evolutionary processes, some invertebrates recognize plant infochemicals and use them to identify possible prey, detect the presence of predators or identify algae containing various classes of toxic metabolites. Different metabolites present in the same algae can play the role of toxins, infochemicals or both simultaneously. We investigated the infochemical activity of compounds extracted from three diatoms epiphytes of the seagrass Posidonia oceanica, by conducting choice experiments on invertebrates living in the same community or in close proximity. Furthermore, the specific toxicity of the extracts obtained from the same algae was tested on sea urchin embryos using a standard bioassay procedure, to detect the presence of toxins. The comparison of the two effects demonstrated that invertebrates are subjected to diatom wound-activated toxicants when these algae are not associated with their own habitat, but they are able to recognize volatile infochemicals derived from diatoms associated with their habitats. The specific toxicity of WACs was shown to be inversely correlated to the perceptive ability of invertebrates towards volatile compounds liberated by the same algae. Hence, when the recognition of specific algae by a given invertebrate species evolves, their detrimental effects on the receiving organism may be lost.     

Biofilm cue for larval settlement in Hydroides elegans (Polychaeta): is contact necessary?

Larvae of many sessile marine invertebrates settle in response to surface microbial communities (biofilms), but the effects of soluble compounds from biofilms in affecting larval behavior prior to settlement, attachment, and metamorphosis have been little studied. This question was addressed by videotaping the behavior of competent larvae of the serpulid, Hydroides elegans, above settlement-inducing biofilms. Adult worms were collected in Pearl Harbor, Hawaii, USA in November 2012 and spawned almost immediately. Six-day old larvae were placed in five replicated treatments in small cups: (1) with a natural biofilm; (2) with a natural biofilm on an 8-µm screen, 1 mm above the bottom of a clean cup; (3) with a natural biofilm beneath a clean screen; (4) in a clean cup; and (5) in a clean cup with a clean screen. Using the videotapes, larval swimming speeds and trajectories were quantified within 5 min of the larvae being placed in a treatment. Only larvae that touched a biofilm, i.e., in treatments (1) and (2), slowed their swimming speed and increased the amount of time spent crawling rather than swimming. This shows that under these conditions, any soluble cues emanating from a biofilm do not affect settlement behavior. Furthermore, after 24 h close to 100 % of larva in the two accessible biofilm treatments had metamorphosed and <15 % in treatments that included a biofilm under a clean screen and no biofilm at all, strongly suggesting that soluble cues for settlement were not produced by the biofilms over the longer time period.     

Whole-community mutualism: associated invertebrates facilitate a dominant habitat-forming seaweed

Many habitat-forming, or foundation, species harbor diverse assemblages of associated taxa that benefit from the refuges from predators or harsh physical conditions that foundation species provide. Growing numbers of studies show how specific taxa associated with foundation species can benefit their hosts, but the aggregate effects of the entire community of associated species remain poorly understood. Here, we evaluate the role that a diverse assemblage of invertebrates plays in mediating the dominance of a foundation species, the green filamentous seaweed Cladophora columbiana Collins, in rocky intertidal habitats. Cladophora is a fast-growing seaweed with a high nitrogen demand, and we suggest that it persists in nutrient-limited high-intertidal pools because of local-scale nitrogen excretion by the invertebrate taxa living within its filaments. Removal of associated invertebrates resulted in a fourfold increase in the rate of water-column nitrogen depletion by Cladophora, and ammonium concentrations inside Cladophora turfs with invertebrates present were seven times higher than in the adjacent tide-pool water. The ammonium excreted by invertebrate meiofauna far surpassed the nitrogen used by Cladophora, suggesting that all of Cladophora's nitrogen requirements could be met by the invertebrates associated with it. This study links host performance to the total aggregate biomass of mutualists rather than the particular traits of any one species, suggesting the potential for important feedbacks between individual hosts and the communities of associated species that they support.     

Effects of algal grazing and aggressive behaviour of the fishes Pomacentrus lividus and Acanthurus sohal on coral-reef ecology

Aggressive behaviour of the fishes Pomacentrus lividus Bl. Schn. and Acanthurus sohal Forskal from the Red Sea is briefly described, and its effect on intensity of algal grazing by herbivorous fish is demonstrated by settlement experiments. Green filamentous alga settles and grows at shallow depths over large areas of coral reefs, but is cropped by fishes to such an extent that it forms only a thin patchy matting on dead corals. Within pomacentrid territories, the alga forms a thicker matting on loosely cemented coralline rubble. Optimum depth range for growth occurs at less than 20 m. Rich growths of green filamentous alga, such as those which occur within pomacentrid territories or on settlement plates protected by wire netting cages, inhibit settlement of “lithothamnion” and invertebrates. While rasping and grazing fish feeders such as parrot fish and surgeon fish limit the distribution of certain invertebrates such as spirorbids, in shallow water it is also true that, were it not for such active removal of green filamentous alga, “lithothamnion” and many invertebrates would find ewer surfaces suitable for settlement.     

Sibling species or poecilogony in the polychaete<Emphasis Type="Italic"> Scoloplos armiger</Emphasis>?

In marine invertebrates multiple modes of development, or poecilogony, may occur in a single species. However, after close examination, many of such putative cases turned out to be sibling species. A case in point may be the cosmopolitan orbiniid polychaete Scoloplos armiger, which inhabits marine shallow sediments. In addition to the well-known direct, holobenthic development from egg cocoons, pelagic larvae have also been described. Our culture experiments revealed a spatially segregated source of the two developmental modes. All females of an intertidal population produced egg cocoons and no pelagic larvae. All but 2 out of 15 females of an adjacent subtidal population produced pelagic larvae and no egg cocoons. Based on these results we performed a molecular genetic analysis (RAPD-PCR) on three intertidal and four subtidal populations in the North Sea. Selected samples from all sites were analysed also by the AFLP method. We found significantly higher genetic diversity within subtidal than within intertidal populations. This is consistent with a wider dispersal by pelagic larvae and a smaller effective population size when development is holobenthic. Total genetic divergence is not related to distance but to the intertidal/subtidal division. We suggest that S. armiger actually represents two sibling species.     

Selective feeding of littoral harpacticoids on diatom algae: hungry gourmands?

We studied benthic harpacticoid grazing on diatom algae from two sites on the White Sea intertidal sandflat. Diatoms from sediments and from harpacticoid gut contents were sampled in situ, identified and counted, and grazing rates were calculated by two ways: (1) using potential daily ration estimations and (2) from the gut content and gut-residence time data. Paraleptastacus kliei did not contain any diatoms in the guts and presumably fed on other objects (bacteria or flagellates). Two other dominating species studied, Heterolaophonte minuta and Huntemannia jadensis, contained an average of 604 and 222 diatom cells per specimen. Diet composition differed significantly from the natural algal community. Two diatoms of intermediate cell size (Nitzschia palea var. debilis and Navicula sp.) contributed 92–97% of gut content for H. minuta and 58–81% for Hn. jadensis, whereas these diatoms amounted to only about 10% of biomass in native community. Mean consumption rates were estimated as 50–200 μg of wet biomass/day/cm², so the harpacticoids grazed only between 3 and 11% of the total microalgae biomass per day. The grazing impact on the two preferred diatom populations, however, was much more intensive, 10–30% per day for Navicula sp. and 55–228% for N. palea. Therefore, native harpacticoid populations demonstrate highly selective feeding and could be strongly limited by their food in spite of seemingly plentiful total abundance of microphytobenthos. This disproportionally high grazing pressure upon some species apparently could affect the structure of microalgae communities resulting in low relative abundance of mid-sized forms. We hypothesize that a very dynamic spatio-temporal distribution of epibenthic harpacticoids (short-living micropatches) may be the possible adaptation to such local food limitation.     

Feeding preferences of periwinkles among four species ofFucus

A widely accepted view of intertidal community organizatiton in the NW Atlantic proposes that fucoid vegetation is maintained by the actions of predators which remove species competitively superior toFucus species. Herbivory is an important component of these predatory interactions, but has been studied largely with reference to the interaction betweenF. vesiculosus andLittorina littorea. There are many species of fucoids and herbivorous invertebrates on the shores of the NW Atlantic and this paper reports field and laboratory experiments performed in 1987 (in or near Halifax, Canada) on the effects of grazing by three species ofLittorina on adults and juveniles of four species ofFucus. In laboratory experiments, portions ofFucus species were presented singly (no choice) or together (multiple choice) to single species ofLittorina. AdultF. distichus was grazed to only a small extent.F. evanescens andF. vesiculosus adult tissues were heavily grazed in most laboratory experiments.F. spiralis adults were heavily grazed in no choice experiments, but were grazed only slightly in multiple choice tests. When adult fucoid tissues were exposed to a mixed suite of grazers in the field, onlyF. vesiculosus was grazed heavily. Tests done on adult tissues showed clearly thatFucus species are highly variable in their vunerability to grazers. Moreover, we found variability among periwinkle species in grazing rates on adultFucus. We also found a great disparity in the feeding rates of different periwinkle species on juveniles ofFucus. Therefore, it is not possible to extrapolate from a single periwinkle species/single fucoid species interaction when attempting to identify vegetation structuring processes.     

First monospecific bloom of the harmful raphidophyte Chattonella antiqua (Hada) Ono in Alexandria waters related to water quality and copepod grazing

The massive red tide bloom of Chattonella antiqua that occurred in Alexandria waters during late August/early September 2006 was monospecific, of very high density, and of wide spatial distribution, and was accompanied by mass fish and invertebrates mortalities. During the bloom, surface water temperature ranged between 26.5 and 28.5 °C and salinity between 23 and 27 psu. The bloom started under very high nitrate and ammonia levels; where the uptake of ammonia seemed to be faster than that of nitrate, and the bloom seemed to avoid the copepod grazing. Mass fish and invertebrates mortality was observed. Few cells of Heterosigma species have been reported for the first time in the Alexandria waters. Several environmental constraints (including physical factors), nutrient loading, copepod grazing and comparison with other data for C. antiqua blooms in Alexandria waters are also discussed.     

Carrion-feeding on the sediment surface at nocturnal low tides by the polychaete<Emphasis Type="Italic"> Phyllodoce mucosa</Emphasis>

Harsh physical conditions in the intertidal zone are the cause of an ample amount of dead macroinvertebrates, which constitute a food source for carrion-feeders. In the European Wadden Sea, this trophic guild includes decapod crustaceans and fish when the tide is in, while during nocturnal low tides the polychaete Phyllodoce mucosa is attracted in large numbers by dead mollusks, crabs or worms on the sediment surface. Within 10 s worms emerged to the surface, crawled as far as 15 m on mucus trails towards the carcass, sucked in tissue up to one-third of their own weight, and then quickly retreated to below the surface. Abundance of P. mucosa was highest in the lower intertidal zone and winter. The seaward high abundance pattern, however, did not continue into the shallow subtidal. In summer, few were attracted during daytime or when the tide was in. However, up to 447 worms aggregated at a single crushed mussel within 20 min at dusk during low-tide exposure. This study suggests that during winter carrion-feeding is an important trophic niche on cold-temperate, intertidal mud flats occupied by a phyllodocid polychaete that is segregated in feeding time from most other scavengers and benefits from cold-sensitive benthic invertebrates.Communicated by O. Kinne, Oldendorf/Luhe     

Relating bird host distribution and spatial heterogeneity in trematode infections in an intertidal snail—from small to large scale

Shorebird abundance and spatial distribution of larval trematodes in the New Zealand mudsnail, Zeacumantus subcarinatus, were investigated in soft-sediment intertidal bays within Otago Harbour, South Island, New Zealand. In a small-scale study, recruitment of trematodes to caged sentinel snails and the prevalence of infection in free-living snails were examined across a grid of fifteen 50×25 m plots arranged in a representative area of an intertidal bay, in relation to within-plot shorebird abundance (definitive hosts) and tidal height. In a large-scale study, natural spatial variation of larval trematodes in Z. subcarinatus was examined across 12 bays in relation to local abundance of shorebirds. Our results revealed that trematode prevalence in snails was positively correlated with bird abundance across bays (R ²=0.503, P=0.006). In contrast, despite a difference in bird abundance between tidal heights, there was no evidence that trematode prevalence reflected the spatial distribution of birds in the small-scale study, suggesting that factors related to differences in submersion time may override the differential input of trematode eggs from birds.     

Influence of seasonal food abundance and quality on the feeding habits of an opportunistic feeder,the intertidal crab <Emphasis Type="Italic">Pachygrapsus marmoratus</Emphasis>

The abundance of trophic sources on the intertidal zone is discontinuous and their supply can vary both in a predictable or unpredictable way. The Mediterranean semi-terrestrial crab Pachygrapsus marmoratus, is known, as adult, to entirely rely on the intertidal trophic sources, and, consequently, it faces the fluctuations of nutritional sources and quality. To clarify the relationships between the feeding habits of an Italian population of P. marmoratus and the temporal variation of its food sources, we carried out a 2-year sampling protocol. Data on seasonal variation in composition of intertidal food item assemblages, on the average content in N and C of the commonest algae, on seasonal changes in crabs feeding habits were collected and compared using a suite of multivariate and univariate techniques. Results showed that P. marmoratus takes advantage of the recruitment phase of the most common invertebrates, affecting and controlling the abundance even of those species whose adults are out of its reach. It can act both as a herbivore, and as a carnivore that shifts between bivalves, more abundant in spring/summer, and the periwinkle Melaraphe neritoides, during winter. In conclusion, we provide evidences on the food choice of this common rocky shore species, which is very plastic and capable of relying on many trophic sources, possibly influencing the abundance and/or the population structure of a number of intertidal populations.     

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

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

Substrate-dependent bacterivory by intertidal benthic copepods

The trophic importance of bacteria to harpacticoid copepods in intertidal areas remains poorly understood, and so do the mechanisms of bacterial feeding. It is, for instance, unclear whether harpacticoids directly target bacterial cells or merely co-ingest them with substrates to which bacterial cells may be attached. Here, we investigate bacterial uptake and substrate requirement for four mud intertidal species (Microarthridion littorale, Platychelipus littoralis, Delavalia palustris and Nannopus palustris) by means of ¹³C-labeled bacteria and biomarker fatty acids (FA). Bacterial uptake strongly depended on grazing on a primary food source but bacterial ingestion rates were low, and no clear indication of copepods directly targeting bacteria was found. Delavalia was the only species that accumulated bacteria-derived FA and gained in polyunsaturated FA (PUFA) probably through bioconversion of bacteria-derived FA. In general, however, our results suggest that bacteria represent a relatively minor and low-quality food for intertidal harpacticoid copepods.