Intratree vertical variation of fruit density and the nature of contest competition in frugivores

Recently, we demonstrated that the highest densities of fruit pulp are located in the uppermost zones of tree crowns. Since heterogeneous distributions of depletable food is theorized to foster contest competition, we tested three hypotheses involving rank differences among species of arboreal frugivores: (1) In the absence of competitors, species tend to feed in higher strata of tree crowns; (2) interspecific contest competition occurs through monopolization and usurpation of feeding sites in these higher strata; and (3) subordinate species decrease their feeding height and ingestion rate when dominants enter the food patch. To test these hypotheses, we observed chimpanzees (Pan troglodytes), red-tailed monkeys (Cercopithecus ascanius), blue monkeys (Cercopithecus mitis), and gray-cheeked mangabeys (Lophocebus albigena) in Kibale National Park, Uganda. We found that: (1) all four primates fed preferentially in upper tree crowns when alone, (2) dominant species monopolized and aggressively usurped the upper crown when co-feeding with subordinates and the latter retreated below the middle of tree crowns, (3) in the presence of dominant species, subordinate species showed lower standardized feeding height and modified their food intake rates, while dominants were not affected by the subordinate species, (4) subordinates moved down at the arrival of and up at the departure of dominants, and (5) the presence of folivores in the tree did not affect the feeding height of a frugivore, even through folivores were socially dominant. Contrary to expectations, we found that red-tailed monkeys decreased their movements between successive fruits that they ate in the presence of blue monkeys compared to when they were feeding alone, perhaps to avoid disturbing dominants and attracting aggression or because they ingested more semi-ripe and green unripe fruits, i.e., more food of lower quality.     

Habitat complexity reduces the growth of aggressive and dominant brown trout (<Emphasis Type="Italic">Salmo trutta</Emphasis>) relative to subordinates

Animals often prefer areas containing physical structure, and population density often increases with structural complexity, presumably because physical complexity in habitats may offer protection from predators and aggressive competitors. Consequently, increased habitat complexity often results in reduced territory size, lower aggression levels and reduced resource monopolisation by dominants. If behavioural plasticity is limited at early life stages, increased habitat complexity may reduce the relative fitness of aggressive, dominant strategies. Here we tested this hypothesis in an experiment on newly emerged brown trout (Salmo trutta) fry. We show, for the first time, that increased habitat complexity reduces the fitness (i.e. growth rate) of aggressive dominant individuals in relation to subordinates, and that this relation is reversed in simple habitats. Variation in environmental complexity may thus induce fluctuating selective pressures, maintaining behavioural variation in natural populations and allowing subordinate and dominant strategies to coexist.     

Nocturnal feeding migrations of Nerita plicata,N. undata and N. textilis (Prosobranchia: Neritacea) on the rocky shores at Mkomani,Mombasa, Kenya

The vertical zonation of the three common rocky shore neritids at Mkomani, Mombasa, Kenya, Nerita plicata Linnaeus, N. undata Linnaeus, and N. textilis Dillwyn, as a function of feeding migrations and of size, was studied from 28 February to 24 March 1983. These snails perform feeding migrations at night starting at around mid-ebb tide and return to their resting positions with the flood tide. They remain in their resting positions throughout the day until the next nocturnal ebb tide. The direction of migration is sizerelated, with the larger snails of each species moving in the opposite vertical direction to the smaller ones, so that the populations as a whole exhibit no statistically significant net vertical displacement. The larger individuals of two of the species, N. plicata and N. undata, invariably move downwards to their feeding levels, while the smaller individuals move upwards; the larger individuals of N. textilis display a different pattern of migration, moving downwards on and around spring-tide days and upwards on and around neap tide days, while the smaller individuals move in the opposite directions. N. textilis rest above their feeding level around spring tides, and below that level around neap tides. It is demonstrated how these nocturnal migratory feeding rhythms are integrated into the spring-neap and seasonal cycles of the snails' daytime resting positions. The adaptive significance of these migrations is also discussed.     

Habitat choice in the intertidal snail Tegula funebralis

Intraspecific variation in habitat preference was studied in the black turban snail Tegula funebralis (Adams, 1854), at two locations on the northern U.S. Pacific Coast. Studies in 1977 using a mass-marking technique showed that most snails found either above or in permanent tidepools at low tide return to their original habitats within a few days after experimental habitat reversal. This return is not due to homing behavior, but is apparently based on the recognition of ecological characteristics of the two habitats. Experiments in 1978 with individually-marked snails suggest that they prefer specific intertidal levels, and not merely above-pool or in-pool habitats. Theoretical models predict that this behavior could play a major role in the maintenance of genetic polymorphism in a species like T. funebralis, whose intertidal environment is characterized by extreme spatial heterogeneity.     

Mangrove fish-communities in tropical Queensland,Australia: Spatial and temporal patterns in densities,biomass and community structure

Regular daylight sampling over 13 mo (February 1985–February 1986) in and adjacent to intertidal forested areas, in small creeks and over accreting mudbanks in the mainstream of a small mangrove-lined estuary in tropical northeastern Queensland, Australia, yielded 112 481 fish from 128 species and 43 families. Species of the families Engraulidae, Ambassidae, Leiognathidae, Clupeidae and Atherinidae were numerically dominant in the community. The same species, with the addition ofLates calcarifer (Latidae).Acanthopagrus berda (Sparidae) andLutjanus agentimaculatus (Lutjanidae) dominated total community biomass. During high-tide periods, intertidal forested areas were important habitats for juvenile and adult fish, with grand mean (±1 SE) density and biomass of 3.5±2.4 fish m^–3 and 10.9±4.5 g m^–3, respectively. There was evidence of lower densities and less fish species using intertidal forests in the dry season (August, October), but high variances in catches masked any significant seasonality in mean fish biomass in this habitat. On ebb tides, most fish species (major families; Ambassidae, Leiognathidae, Atherinidae, Melanotaeniidae) moved to small shallow creeks, where mean (±1 SE) low-tide density and biomass were 31.3±12.4 fish m^–2 and 29.0±12.1 g m^–2, respectively. Large variances in catch data masked any seasonality in densities and biomasses, but the mean number of species captured per netting in small creeks was lowest in the dry season (July, August). Species of Engraulidae and Clupeidae, which dominated high-tide catches in the forested areas during the wet season, appeared to move into the mainstream of the estuary on ebbing tides and were captured over accreting banks at low tide. Accreting banks supported a mean (±1 SE) density and biomass of 0.4±0.1 fish m^–2 and 1.7±0.3 g m^–2, respectively, at low tide. There were marked seasonal shifts in fish community composition in the estuary, and catches in succeeding wet seasons were highly dissimilar. Comparison of fish species composition in this and three other mangrove estuaries in the region revealed significant geographic and temporal (seasonal) variation in fish-community structure. Modifications and removal of wetlands proposed for north Queensland may have a devastating effect on the valuable inshore fisheries of this region, because mangrove forests and creeks support high densities of fish, many of which are linked directly, or indirectly (via food chains) to existing commercial fisheries.Contribution No. 493 from the Australian Institute of Marine Science     

Fish assemblages in seagrass beds are influenced by the proximity of mangrove forests

Mangrove forests and seagrass beds frequently occur as adjacent habitats in the temperate waters of southeastern Australia. At low tide when fish cannot occupy mangroves they might utilise adjacent habitats, including seagrass. We first sampled small fish from seagrass beds close to and far from mangroves in the Pittwater estuary, NSW, Australia. Seagrass beds close to mangroves had a greater density of fish species than beds far from mangroves (close: mean 16.0 species net⁻¹, SE 1.0; far: 13.2, 1.3; P < 0.05). In particular, juvenile fish were in greater densities near to than far from mangroves (close: 5.3, 0.4; far: 3.1, 0.4; P < 0.05). We then sampled the mangrove forests during the high tide and seagrass beds during the low tide, in beds along a continuum of distances from mangroves. Multivariate analysis showed that fish assemblages differed with distance from mangroves, and the differences were attributed to the composition of the fish assemblage (i.e. presence/absence of fish species), not the abundances of individual species. In particular, fish that utilise mangrove forests at high tide were found in greater species densities and species richness in seagrass nearer to mangroves. A negative relationship was found between the density of mangrove-utilising fish species and the distance of the bed from mangroves (R ² = 0.37, P < 0.05). This confirms the important connectivity between mangroves and seagrass for fish in temperate Australian waters.     

The contribution of seagrass beds (<Emphasis Type="Italic">Zostera noltii</Emphasis>) to the function of tidal flats as a juvenile habitat for dominant,mobile epibenthos in the Wadden Sea

Since the substantial loss of subtidal eelgrass (Zostera marina L.) in the 1930s, seagrass beds in the Wadden Sea are limited to the intertidal zone and dominated by Z. noltii Hornem. This study deals with the effect of vegetated tidal flats on quantities of mobile epifauna and proves empirically the function of seagrass canopies as a refuge for marine animals remaining in the intertidal zone at ebb tide. Drop-trap samples were taken in the Sylt-Rømø Bight, a shallow tidal basin in the northern Wadden Sea, on vegetated and unvegetated tidal flats during July and August 2002, and during the entire growth period of Z. noltii from May to September in 2003. The species composition in Z. noltii and bare sand flats showed minor differences since only two isopod species (Idotea baltica and I. chelipes) occurred on Z. noltii flats exclusively. Juvenile shore crabs (Carcinus maenas L.), brown shrimps (Crangon crangon L.) and common gobies (Pomatoschistus microps Krøyer) were also found abundantly on bare sand flats. However, the results showed significantly higher abundances and production of these dominant species on vegetated tidal flats. Additionally, the analyses of faunal size classes indicated higher percentages of small individuals in the seagrass bed during the entire sampling period. Despite drastic diurnal fluctuations of dissolved oxygen at low tide, faunal density in the residual water layer remaining in seagrass canopies at ebb tide was found to be consistently higher than that found in artificially created tide-pool units. Although species composition of mobile epifauna did not basically differ between vegetated and unvegetated tidal flats, Z. noltii beds are considered to contribute quantitatively to the function of tidal flats, as an extended juvenile habitat for some of the most important species of the Wadden Sea food web.Communicated by O. Kinne, Oldendorf/Luhe     

Amphipods and isopods in the rocky intertidal: dispersal and movements during high tide

Animals in the intertidal, both mobile and sessile, generally exhibit some zonation pattern, in which each species shows a preference for, or is confined to, some height levels. The study of zonation patterns is, however, almost exclusively based on surveys made during low tide, when many animals are relatively inactive. We studied zonation patterns of amphipods and isopods on rocky shores in southwestern Iceland, both by traditional sampling at low tide as well as by sampling during high tide. The distributional patterns seen at high tide differed significantly from that at low tide. One amphipod, Anonyx sarsi, was common around baits at all levels at high tide but absent from the intertidal at low tide. Several other species were either relatively more common or tended to be recorded higher, or in one instance, lower on the shore when the tide was in than at low tide. There was also evidence of some species changing habitats within the intertidal with the tidal cycle. Many species, however, moved little away from their respective zones occupied at low tide, and for some species, including some capable of rapid swimming, very limited mobility was indicated. We conclude that low-tide surveys of the intertidal give an incomplete picture of the community structure, and even key species may be missed in such surveys.Communicated by L. Hagermann, Helsingør     

Rhythmic vertical migration of the gastropod Cerithidea decollata in a Kenyan mangrove forest

In Mida Creek, Kenya (3°20′S, 40°5′E), at high water, the snail Cerithidea decollata dwells on the trunks of mangrove trees (Avicennia marina), while during low water it migrates to the ground, foraging at various distances from the trunk, where it aggregates again well before the incoming tide. Snails from the upper shore level are 150–200 m distant from those living at the lower shore level and they cluster at lower heights on trunks. In any case, sufficient height is usually attained to avoid being submersed. An experiment was designed (February and October 2005), exchanging individuals from different shore levels subject to different tide regimes, in order to test whether snails rely on internal information or on external, direct cues, to adapt their behaviour to local conditions. Results show that C. decollata mostly rely on internal information, presumably based on an internal clock. When individuals from upper and lower shore levels were exchanged, their internal clocks continued to govern when to ascend the home trunk and how high to climb for five to six successive tides, after which the behaviour was reset to the new local conditions.     

Diel eclosion rhythm of a sublittoral population of the marine insect Pontomyia pacifica

Within the scope of studies on adaptation of insects to intertidal and sublittoral environments, correlations between the reproduction period of the short-lived chironomid Pontomyia pacifica Tokunaga and tidal conditions were examined at the only known Japanese location of this species in 1980. The larval habitat is situated, except for deeper tide pools in the lower midlittoral zone, mainly within the inner sublittoral zone, whose upper area is covered by Hypnea choroides, the dominating algae during the summer. The aerial adults emerge on the surface of the open sea and swarm there independent of the tidal situation above the submersed sublittoral habitat. Eclosion always started during dusk and all reproductive activities were ended within 2.5 h after sunset. The adaptation of P. pacifica (sublittoral habitat, diel eclosion after sunset with mass concentration of the adults on the water surface, no semilunar or lunar timing of the reproduction period and sinking egg masses) corresponds with those of the convergent marine chironomid Clunio balticus from Europe. On the basis of a few laboratory observations with P. pacifica and detailed experiments with C. balticus, it is supposed that the diel eclosion of P. pacifica is also controlled by an endogenous, circadian timing mechanism and the 24-h light-dark cycle as an environmental time cue.Communicated by O. Kinne, Oldendorf/Luhe     

Adaptive droving behavior observed in the fiddler crab Uca vocans vocans

Droving is conspicuous in Uca vocans vocans in summer. The crabs burrowing on the upper habitat have a higher tendency to wander compared to the ones burrowing on the lower habitat. Most of the wandering crabs captured on the low tide levels are relatively large and male. Larger crabs and males prefer to burrow on the upper zones of U. vocans habitat, but the smaller ones and females prefer the lower habitat. The upper level of the U. vocans habitat has relatively low N-content compared to the water's edge. Therefore droving is advantageous in crabs that have burrows on the upper level. Female and smaller resident crabs have faster feeding motions than male and larger ones, and can satisfy their feeding demands more rapidly. Therefore, for males and larger crabs it is advantageous to move away from the burrow area and forage in areas of higher food content near the low tide level where the number of feeding motions increases.     

Interactions between two introduced species: Zostera japonica (dwarf eelgrass) facilitates itself and reduces condition of Ruditapes philippinarum (Manila clam) on intertidal flats

Dwarf eelgrass (duckgrass; Zostera japonica) and Manila clams (Ruditapes philippinarum) are two introduced species that co-occur on intertidal flats of the northeast Pacific. Through factorial manipulation of clam (0, 62.5, 125 clams m⁻²) and eelgrass density (present, removed by hand, harrowed), we examined intra- and interspecific effects on performance, as well as modification of the physical environment. The presence of eelgrass reduced water flow by up to 40% and was also observed to retain water at low tide, which may ameliorate desiccation and explain why eelgrass grew faster in the presence of conspecifics (positive feedback). Although shell growth of small (20–50 mm) clams was not consistently affected by either treatment in this 2-month experiment, clam condition improved when eelgrass was removed. Reciprocally, clams at aquaculture densities had no effect on eelgrass growth, clam growth and condition, or porewater nutrients. Overall, only Z. japonica demonstrated strong population-level interactions. Interspecific results support an emerging paradigm that invasive marine ecosystem engineers often negatively affect infauna. Positive feedbacks for Z. japonica may characterize its intraspecific effects particularly at the stressful intertidal elevation of this study (+1 m above mean lower low water).     

Changes in morphology and physiology of an East Mediterranean sponge in different habitats

The sponge Tetilla sp. (Tetractinomorpha: Tetillidae) is a common species in the eastern Mediterranean. This sponge inhabits four different habitat types differing in wave impact and irradiance levels. Two of these habitats (a shallow cave and deep water) are characterized by relatively calm water, whereas the other two (shallow exposed site and tide pools) are in turbulent water with high energy flow. The present study examined the influence of physical (depth, illumination and water motion) and biotic factors on morphology, skeletal plasticity and reproductive traits among the four spatially separated populations. Sponges from tidal pools had significantly larger body volume than sponges from deep water and from shallow caves (ANOVA: tidal-deep P<0.0001; tidal-shallow caves P<0.05). Sponges from exposed habitats were significantly larger than deep-water sponges (ANOVA: P=0.01). In addition, individuals from tide pools and from the exposed habitat had a significantly higher proportion of structural silica than sponges from the calmer deep water and from the cave sites. Oxea spicules in sponges from the calm habitats were significantly shorter than in those from the tidal pools and the exposed habitats. The percentage of spicules out of a sponges dry weight in individuals transplanted from deep (calm) to shallow (turbulent) water significantly increased by 21.9±12.9%. The new spicule percentage did not differ significantly from that of sponges originally from shallow water. Oocyte diameter differed significantly between habitats. The maximal size of mature eggs was found in deep-water sponges in June (97±5 m). In the shallow habitats, a smaller maximal oocyte diameter was found in the cave, in May (56.5±3 m). Furthermore, oocyte density in shallow-water sponges was highest in May and decreased in June (with 88.2±9 and 19.3±9 oocytes mm^–2, respectively). At the same time (June), oocyte density of deep-water sponges had just reached its maximum (155±33.7 oocytes mm^–2). The difference in oocyte size and density between deep- and shallow-water individuals indicates an earlier gamete release in the shallow sponge population. The results suggest that plasticity in skeletal design of this sponge indicates a trade off between spicule production and investment in reproduction.Communicated by O. Kinne, Oldendorf/Luhe     

Lateral or along-shore distribution of the polychaetous annelids of an intertidal,sandy habitat

The lateral or along-shore distribution of the polychaetous annelids of a subtropical, sandy, intertidal habitat in Florida (USA) was studied on three transects (covering 2.5 km of shoreline), sampled quarterly for 2 years. Although the average species number and density of individuals were not significantly different at specific tidal levels, the species composition was highly variable over time. Composition of density dominants was identical along-shore, and down-shore density distribution of selected species was also identical. Analysis by entire transects rather than by individual stations decreased along-shore heterogeneity by 10 to 15%.     

Habitat use and group size of pied cormorants (<Emphasis Type="Italic">Phalacrocorax varius</Emphasis>) in a seagrass ecosystem: possible effects of food abundance and predation risk

Both food abundance and predation risk may influence habitat use decisions. However, studies of habitat use by birds in marine environments have focused only on food abundance. I investigated the possible influences of food abundance and predation risk from tiger sharks (Galeocerdo cuvier) on habitat use by pied cormorants (Phalacrocorax varius) over two spatial scales and on cormorant group size. Cormorants were usually solitary, but group size was highest in shallow habitats during months when shark density was low. Regardless of season, cormorant density within shallow habitats was higher over seagrass than sand, and cormorants were distributed between these two microhabitats proportional to prey density. Therefore, cormorants appear to respond to prey abundance at a relatively narrow spatial scale (i.e., tens of meters). At the habitat-patch scale (~1 km), the density of cormorants and their prey (teleosts) was higher in shallow habitats than in deep ones, but the density of cormorants was influenced by an interaction between water temperature (i.e., season) and habitat. There was decreased use of shallow habitats as water temperature, and the density of tiger sharks, increased. When shark density was low, cormorants were distributed across habitats roughly in proportion to the abundance of fish, suggesting that cormorants respond to food abundance at the scale of habitat patches. However, as shark abundance increased, the relative density of cormorants dropped in the dangerous shallow habitats such that there was a greater density of cormorants relative to their food in deep habitats when sharks were abundant. This suggests that pied cormorants trade-off food and risk by accepting lower energetic returns to forage in safer habitats. This study provides the first evidence that marine habitat selection by birds may be influenced by such a trade-off, and provides further evidence that tiger sharks are important in determining habitat use of their prey and mediating indirect interactions within Shark Bay.Communicated by P. W. Sammarco, Chauvin     

Rhythms in larval release of the sublittoral crab Neopanope sayi and the supralittoral crab Sesarma cinereum (Decapoda: Brachyura)

Ovigerous females of the subtidal xanthid crab Neopanope sayi (Smith) and the high intertidal grapsid crab Sesarma cinereum (Bosc) were collected during the summers of 1986 and 1987 in the Beaufort, North Carolina (USA), area and brought into the laboratory, where rhythms in larval release were monitored. When crabs with late-stage embryos were put under a 14 h light:10 h dark cycle in an otherwise constant-environment room, an apparent tidal rhythm in release of larvae was observed for both species, with N. sayi releasing near the time of day and night high tides, and S. cinereum releasing around the time of night high tides. The time of sunset relative to high tide was a complicating factor, since larval release for both species was often concentrated around sunset when evening high tides fell several hours before sunset. When a group of N. sayi and S. cinereum were brought into the laboratory and placed under constant lowlevel light for 5 d, the release rhythm of the population persisted, thus implying that the rhythm is endogenous. Larval release near the time of high tide and often at night is common among brachyurans living in tidal areas, regardless of specific adult habitat, suggesting a common functional advantage. Possibilities include transport of larvae from areas where predation and the likelihood of stranding and exposure to low-salinity waters are high, as well as a reduced probability of predation on adult females. Results of the present study suggest that the importance of release after darkness may increase with increasing tidal height of the adult.     

Mechanisms of habitat segregation between an invasive and an indigenous mussel: settlement,post-settlement mortality and recruitment

The mussel Mytilus galloprovincialis is highly invasive worldwide, but displays varying degrees of local and regional coexistence with indigenous mussels through spatial habitat segregation. We investigated the roles of settlement, post-settlement mortality, juvenile growth and recruitment in partial habitat segregation between the invasive M. galloprovincialis and the indigenous mussel Perna perna on the south coast of South Africa. We used two study locations, Plettenberg Bay and Tsitsikamma, 70 km apart, with two sites (separated by 300–400 m) per location, each divided into three vertical zones. There were no significant effects in Tsitsikamma, where daily settlement and monthly recruitment were significantly lower than in Plettenberg Bay. In Plettenberg Bay, settlement (primary and secondary) and recruitment of both species decreased upshore. Post-settlement mortality was measured over two consecutive 6-day periods during a spring tide and a neap tide. For both species mortality was low on the low-shore. High-shore mortality was consistently low for M. galloprovincialis, but increased dramatically for P. perna during spring tide. No data were obtained for growth of P. perna, but juvenile M. galloprovincialis grew more slowly farther upshore. P. perna recruited mainly in spring and summer, with a peak in summer far greater than for M. galloprovincialis. Recruitment of M. galloprovincialis was more protracted, continuing through autumn and winter. Thus local coexistence is due to a combination of pre- and post-recruitment factors differing in importance for each species. P. perna is excluded from the high-shore by recruitment failure (low settlement, high mortality). High survival and slow growth in juveniles may allow large densities of M. galloprovincialis to accumulate there, despite low settlement rates. With no differences between species in settlement or mortality on the low-shore, exclusion of M. galloprovincialis from that zone is likely to be by post-recruitment processes, possibly strengthened by periodic heavy recruitments of P. perna. At larger scales, larval retention and protracted recruitment contribute to the success of M. galloprovincialis at Plettenberg Bay, while recruitment limitation may explain why M. galloprovincialis is less successful at other sites.     

Persistent locomotor rhythmicity in the prawns Penaeus indicus and P. monodon

The locomotory activity of adult Penaeus indicus Milne Edwards and P. monodon (Fabricius) collected from the Vellar estuary, South India, in 1983 was monitored continuously for a period of 40 d in the laboratory. Both species exhibited persistent circatidal and circadian rhythmicity, the former displayed peak activity during predicated times of high water in their original habitat. The latter exhibited peak activity during periods corresponding to in situ night-time, with almost no activity during the day, except at the time of high tide. The tidal and daily rhythms were evident for only 3 to 4 d and 5 to 6 d, respectively, under laboratory conditions. The amplitude of the rhythms displayed variations in each semilunar period, reflecting the synchrony and dissynchrony of the tidal and daily rhythms. Activity was correlated with phases of the moon, peaking at times of new and full moon, and showing a scattered pattern with smaller peaks during the first and third quarters.     

Impact of Land-Based Sources On Marine Environment: Phytoplankton Blooms Within an Annual Cycle

The discharge water into the western region of Alexandria (Egypt), creates suitable conditions for the acceleration of the phytoplankton growth; daily replenishment of nutrients and development of density stratified water column. Water transparency and ammonia concentrations seem to be affecting the variations of the numerical standing crop. Eight phytoplankton bloom pulses of different causative species were observed. Asterionella glacialis represents a newly recorded red tide species in the neritic waters of Alexandria.     

Estimate of removal rate ofNereis virens (Polychaeta: Nereidae) from an intertidal mudflat by gulls (Larus spp.)

The feeding behavior of herring gulls (Larus argentatus), ringed-billed gulls (L. delawarensis) and great blackbacked gulls (L. marinus) on an intertidal mudflat in Maine, USA, was investigated. Remains of fish, mussels, crabs, insects, and the polychaeteNereis virens were recovered from gull feces. Forty-three percent of the fecal samples containedN. virens jaws, setae, or both. A comparison of jaws from fecal material and from worms collected from the natural community demonstrates that gulls preferentially preyed upon larger worms. Feeding was largely confined to 3 h around low tide, and birds fed mostly in the low intertidal and below mean low water where the largest worms were located. Individual birds remained on the flat for a mean of 28.0 min and consumed a mean of 19.2 worms per visit. It was calculated that gulls remove a mean of 808 largeN. virens from the flat per tide from June to October, representing an estimated 0.04% of the standing crop of largeN. virens.