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1.
Although there is a large body of research on food webs in rocky intertidal communities, most of the emphasis has been on the marine benthic components. Effects of avian predation on highly mobile predators such as crabs, remains practically unstudied in rocky shore ecosystems. The crab, Cancer borealis, is an important component of the diet of gulls (Larus marinus, L. argentatus) at the Isles of Shoals, Maine, USA. C. borealis prey include the predatory gastropod Nucella lapillus L., the herbivore Littorina littorea, and mussels Mytilus edulis L. We hypothesized that gulls reduce abundance of C. borealis in the low intertidal and shallow subtidal, thereby allowing C. borealis prey to persist in high numbers. A study of crab tidal migration showed that C. borealis density nearly doubled at high tide compared to low tide; thus, crabs from a large subtidal source population migrate into the intertidal zone during high tides and either emigrate or are removed by gulls during low tides. Results from a small-scale (1 m2) predator caging experiment in the low intertidal zone indicated that enclosed crabs significantly reduced L. littorea abundance when protected from gull predation. In a much larger-scale gull exclusion experiment, densities of C. borealis increased significantly during low and high tides in exclosures relative to the controls. C. borealis density was inversely correlated with changes in the abundance of two mesopredators Carcinus maenas and Nucella lapillus, and with the space-occupier M. edulis. There was a similar negative correlation between abundance of C. borealis and the change in abundance of the herbivore L. littorea, but the trend was not significant. Mortality of tethered L. littorea was associated with C. borealis density across sites. However, preferred algae did not change in response to L. littorea density during the experiment. Thus, we found suggestive, but not conclusive, evidence for a three-level cascade involving gulls, crabs, and L. littorea. Our studies strongly suggest that gulls, as apex predators, generate three-level trophic cascades in rocky intertidal food webs by preventing the highly mobile subtidal predator, C. borealis, from establishing substantial populations in the low-mid intertidal zone thereby indirectly enhancing densities of two key mesopredators (N. lapillus, Carcinus) and blue mussels (M. edulis).  相似文献   

2.
An intensive study of the spatial distribution ofNereis virens (Sars) andNephtys caeca (Fabricius) was conducted during the spring and autumn of 1986 in the lower St. Lawrence estuary. Statistical analysis showed that spatial variations in density, individual body weight and sexual maturity, particularly in the case ofNereis virens, are correlated with the intertidal level, with certain sediment characteristics, and with the thickness of the colonizable sediment layer. The density ofN. virens increases in an offshore-onshore direction, whereas that ofNephtys caeca decreases in the same direction. For both species, mean body weight increases downshore from the upper intertidal level. Other specific relationships exist in relation to sediment characteristics. Sexually matureNereis virens are found only at the lower intertidal level; sexual maturity inNephtys caeca was not studied. There were no changes in spatial distribution patterns between spring and autumn. The spatial distribution ofNereis virens parallels depth contours and may reflect its ability to inhabit environments which become more physically unstable in an offshore-onshore direction. This spatial distribution is consistent with a model whereby larvae are recruited in the upper intertidal zone and juveniles migrate downshore.  相似文献   

3.
Neohaustorius schmitzi Bousfield, the dominant macroscopic invertebrate on sandy beaches in North Carolina, USA, is strictly intertidal, and distributed from mean low tide to high tide, with an average density of 790/m2. Location of greatest densities within the intertidal zone is dependent upon season and sex. An aggregated distribution is shown. N. schmitzi has 2 generations/year: a winter generation which lives about 8 months, and a summer generation which lives about 4 months. The reproductive season extends from February through October, with spring and summer peaks. Each female appears to produce only 1 brood of young. The number of young ranges from 2 to 14, depending upon the length and generation of the female.  相似文献   

4.
Behavioural patterns of Nereis virens (Sars) were monitored in the laboratory to determine variations in the time budget (i.e., percentage of time spent in various activities) as a function of density and acclimation period (number of days following the introduction of worms into experimental enclosures). Experiments were carried out from 17 June to 13 August 1989, using worms collected from intertidal sand flats of l'Anse à l'Orignal in the lower Saint Lawrence estuary, Canada. Inactivity, locomotion and irrigation were, in that order, the most frequent behaviours for solitary individuals. Correlations between the time spent in the various activities and time of day indicated a general increase in activity during the night (21.00 to 06.00 hrs). Locomotory activity increased at night, inactivity during the daytime. A general sequence of behaviour of N. virens was revealed. Inactivity, locomotion and irrigation were still the most frequent activities for individuals observed in high-density conditions, but their respective importance differed significantly (locomotion, irrigation and inactivity, in that order). There was also less variability in the time spent for each behaviour in the high-density aquarium. Comparisons between behaviour and time of day suggested fewer correlations among the different components of the time budgets under conditions of high density. Finally, the time budget varied but little over time (number of days alloved for acclimation) in the low-density aquarium as opposed to the high-density aquarium. Our study revealed that population density and acclimation are important in determining behavioural patterns of N. virens.  相似文献   

5.
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  相似文献   

6.
Food choice, feeding habitat use and spatial distribution of black-headed gulls ( Larus ridibundus), mew gulls ( L. canus), herring gulls ( L. argentatus) and lesser black-backed gulls ( L. fuscus) were studied in the south-eastern North Sea. At-sea distribution during the breeding period was assessed by transect counts from ships. Clear differences could be established between the four species, with lesser black-backed gulls being widespread and numerous far out at sea and mew and black-headed gulls being restricted to the coastal area; herring gulls showed an intermediate pattern. The diet of these sympatrically breeding gulls was analysed from pellet and faecal samples collected at two colonies. Considering biases in this method, the following components appeared to comprise the principal food for each species: black-headed gulls: bivalves, crustaceans; mew gulls: bivalves, polychaetes; herring gulls: bivalves, crustaceans; and lesser black-backed gulls: fish and crustaceans. Black-headed and mew gulls had much broader dietary niches than herring and lesser black-backed gulls. Based on spatial distribution, food choice and feeding habitat use, three types of foraging strategies are apparent: (1) black-headed and mew gulls forage both intertidally and on land, (2) herring gulls feed most frequently in the intertidal zone and (3) lesser black-backed gulls feed at sea. The foraging behaviour of these four species may well have influenced their current population trends in the study area.  相似文献   

7.
G. Sach 《Marine Biology》1975,31(2):157-160
In 1974, spawning polychaetes, Anaitides mucosa (Oersted, 1843), appeared in great numbers (up to 148 individuals/m2) at the surface of an intertidal flat in the German Waddensea from mid-February until 10th April. A. mucosa forms mucous bags which usually contain more than 10,000 green eggs. The eggs measure 113.4±5.4 μm in diameter, including the fructification membrane (18 measurements). Production if these egg bags was observed 5 times in nature and 4 times in the laboratory. Four to 17 males and 1 female form a ball of interlacing bodies. At the same time a mucous mass is secreted by the female — maybe also by the males — which increases to a diameter of as much as 2 cm within about 8 min. Eggs and sperm are shed into this mass of mucus. The worms appeared at the surface of the flat from early sunset until sunrise from about 3 h before to 1 h after low tide. Judging from observations of worms in the laboratory, it seems possible that the rhythm of sexual activity is controlled endogenously.  相似文献   

8.
Field manipulations in a Maine, USA intertidal mud flat showed that Nereis virens adults were an important factor in regulating the abundance of Corophium volutator, an amphipod which comprised 63% of the total number of individuals in the benthic community. Removal of N. virens adults resulted in an increase in C. volutator abundance, while addition of N. virens adults led to a decrease. There was a significant negative correlation between the abundance of N. virens adults and that of C. volutator. C. volutator was also negatively correlated with the density of all the other infaunal species combined. These results suggest that there is at least a 3-level interactive system (N. virens —C. volutator — other infaunal species) within the infauna in Maine; N. virens is a factor in controlling the abundance of C. volutator, and C. volutator may play a role in regulating the densities of other infaunal species. Models of generalized cropping in soft-bottom systems may be too simple if they ignore complex trophic relationships within the infauna.  相似文献   

9.
Seasonal variations in the microphytobenthic diatom community were investigated in an intertidal sand flat of a tropical marine environment influenced by monsoons. Cores of sediments were collected along the beach gradient: low tide, mid tide and high tide zone up to a depth of 15 cm.. Diatom abundance was lowest during the monsoons and highest during the post-monsoons and the early pre-monsoon season throughout the intertidal transect. Diatom diversity was highest at the mid tide, followed by the high and low tide zones. Diatoms were viable up to a depth of 15 cm throughout the intertidal transect. The diatom community included the pennates, the permanent residents of this area, centric genera, which lead an attached mode of life and also some planktonic genera, brought in from ambient waters. Among the pennates, Navicula and Amphora were the dominant genera whereas in the case of centrics, Thalassiosira dominated the community throughout the intertidal transect down to a 15 cm depth. . Grain size fractions, which served as predictors of some diatom genera changed with tidal zones. The effect of winds on the resuspension of the pennate diatoms was evident only at the low tide zone down to a depth of 5 cm . Chlorophyll a concentration proved to be a good predictor of both pennate and centric diatom abundance at the low tide zone down to a depth of 10 cm and at the mid tide zone down to a depth of 5 cm.. However, even though chlorophyll a concentrations failed to reveal any positive correlation with the diatom abundance at both the deeper sediment layers and the high tide zone, the fact that viable cells were present at these areas reveal that the diatoms adopt survival strategies, contributing significantly to the carbon budgets of such unstable habitats.  相似文献   

10.
The effects of infaunal mode of life on sediment properties, microbial reaction rates, as well as abundance and composition of bacterial communities were studied in sediment surrounding burrows (mucus lining, oxidised wall, ambient anoxic and surface sediment) of two closely related, but behaviourally different, nereidid polychaete worms: the facultative suspension-feeder Nereis (Hediste) diversicolor and the obligate deposit-feeder Nereis (Neanthes) virens. Burrow sediment of the two species was collected from two adjacent (50 m distance) shallow sandy locations (Kertinge Nor, Denmark). The burrow lining and wall of both polychaete species were enriched in organic matter originating from mucous secretions by the inhabitants and phytoplankton trapped through irrigation. This was more evident for N. diversicolor that shows a significantly higher irrigation rate than N. virens. Both the organic matter mineralisation rates (based on anaerobic incubations) and bacterial abundance were higher along the burrow linings and walls. However, accumulation of porewater TCO2 and dissolved organic carbon in sediments adjacent to burrows increased most rapidly in the presence of N. diversicolor, suggesting higher heterotrophic activity associated with this species. Surprisingly, bacterial abundance was lower around burrows of N. diversicolor than those from N. virens indicating that burrow environments from the first species harbour a more active bacterial community. Molecular fingerprints of the 16S rRNA gene from bacterial communities showed that the composition of the burrow linings and walls resembled the ambient anoxic sediment rather than the oxic sediment surface. On the other hand, the bacterial fingerprints of the sediment surrounding the burrows of the two polychaete species were markedly different suggesting either a site-specific difference in sediment parameters or a significant species-specific impact of the burrow inhabitants.  相似文献   

11.
D. Dean 《Marine Biology》1978,45(2):165-173
There have been many previous reports of the sandworm Nereis virens Sars swimming in the water column. This behavior usually has been attributed to reproductive processes. Sandworms were found swimming in surface waters at night on ebb tides during many nights of January, February and March in a Maine (USA) estuary. None of the specimens examined contained gametes or possessed other characteristic spawning or pre-spawning modifications. Several age classes were found, with worms measuring 9 to 38 cm in length, weighing 0.5 to 19.8 g, and having 82 to 187 segments. The greatest numbers of worms were observed during near-average tides on evenings in which low tides occurred a few hours after sunset but prior to moonrise. Up to 83 worms per minute were observed swimming seaward through a 20 m transect, while none were observed swimming landward at any stage of the tide. It is concluded that sandworms swimming during winter nights is unrelated to reproduction and that it is an inherent behavior pattern.Contribution No. 102 of the Ira C. Darling Center, Walpole, Maine 04573, USA.  相似文献   

12.
The effects of two fixed photoperiodic regimes on the rate of somatic growth and segment proliferation in the semelparous polychaete Nereis (Neanthes) virens Sars has been investigated. The two photoperiods (16 h light:8 h dark and 8 h light:16 h dark, hereafter LD 16:8 and LD 8:16) were close to the extremes experienced naturally by N. (N.) virens in the course of a year. The experiments were carried out during the ambient winter and all worms (including four-setiger larvae raised out of season) showed elevated feeding, growth and enhanced segment-proliferation rates when reared under LD 16:8, compared to worms reared under LD 8:16. The rate of replacement of lost segments after caudal ablation was also higher under LD 16:8. The number of segments present when the segment proliferation rate had fallen to zero was higher in individuals grown from birth under LD 16:8 compared to those grown under LD 8:16. Feeding activity showed a degree of spontaneous recovery under LD 8:16 from January onwards, but remained lower than under LD 16:8. This difference continued into the following spring, and was not attributable to differences in sexual maturity. We conclude that the rate of feeding in N. (N.) virens and other indicators of somatic growth rate such as rates of caudal regeneration and segment proliferation are directly influenced by the photoperiod, but that the seasonal cycle of growth also includes a circa-annual component modifying the response to static photoperiods according to the time of year. Received: 20 October 1998 / Accepted: 4 February 1999  相似文献   

13.
The cytochrome P-450 dependent mixed-function oxygenase (MFO) system ofNereis virens, found in the endoplasmic reticulum of the intestinal epithelium, has similar components and cofactor requirements when compared to other invertebrates. Worms exposed to food containing benzo(a)pyrene or a mixture of polychlorinated biphenyls (Aroclor 1254) had significantly higher MFO activity and cytochrome P-450 content than unexposed ones.N. virens from a site in Porland, Maine, USA, which was heavily contaminated with oil, and nearby clean reference sites were studied. The weight, MFO activity and presence or absence of gametes were determined. The MFO activity in worms from the oil-contaminated site was approximately six times that of worms from the reference sites; worms from the reference areas were six times larger than oil-contaminated worms. Worms from the oil-contaminated area lacked or had undeveloped gametes in spring, summer and fall. In contrast, many of the worms collected from the reference sites at the same times had fully developed gametes.  相似文献   

14.
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  相似文献   

15.
R. T. Kneib 《Marine Biology》1987,96(2):215-223
Postlarval and juvenile grass shrimp (Palaemonetes pugio Holthuis) ≦15 mm total length (TL) were abundant at low tide in shallow aquatic microhabitats (i.e. puddles and films of residual tidal water) in the intertidal zone of a salt marsh on Sapelo Island, Georgia, USA from 1982 to 1984. The highest concentrations of young P. pugio occurred at 190 to 200 cm above mean low water, ∼10 to 20 cm below mean high water. The intertidal distributions of young grass shrimp expanded and contracted with changes in tidal amplitude. Postlarval grass shrimp (6 to 8 mm TL) continuously recruited into the intertidal marsh population from May until October, but densities varied in a regular pattern with peaks in abundance occurring at ∼2-week intervals, corresponding to spring tide periods in the lunartidal cycle. Although present nearly year-round in the intertidal marsh, juveniles (9 to 15 mm TL) were most abundant from August to October. Apparent growth rates of individuals up to 15 mm TL averaged 0.268±0.026 (mean±95% C.I.) mm d-1 from May to October and 0.070±0.032 mm d-1 in November and December. Unlike larger aquatic organisms, which can forage in the emergent marsh only when it is flooded by the tide, juvenile grass shrimp have constant access to intertidal resources. Although potentially important predators in this system, the role of young P. pugio in the trophic organization of salt marsh benthic invertebrate assemblages has yet to be examined. Contribution No. 576 of the University of Georgia Marine Institute  相似文献   

16.
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).  相似文献   

17.
When feeding on blue mussels (Mytilus edulis), oystercatchers (Haematopus ostralegus) either stab into the mollusc’s gaping valves or hammer through its dorsal or ventral shell. Whilst the selectivity of hammering and stabbing oystercatchers for specific prey morphologies has been well studied, the way in which the effects of environment on M. edulis morphology can in turn affect feeding methods of H. ostralegus is very poorly understood. Based on morphological analyses on randomly selected shells from three intertidal zones, this study failed to detect differences in morphology or distribution of dorsally and ventrally hammered shells but confirms the finding of previous authors that hammering oystercatchers select thinner mussels than stabbing birds. Additionally, we show that this difference in optimal prey morphology can lead to spatial patterns in oystercatcher feeding behaviour. Whilst at the low intertidal and higher mid intertidal zones, characterised by comparatively thick shells, most empty shells had apparently been stabbed, hammering was the dominant feeding behaviour at the lower mid intertidal zone, where shells were thinner. Preference of hammering birds for smaller mussels was not ubiquitous. Sagittal shell shape was predominantly influenced by allometric growth effects and had only minor effect on prey selection. All oystercatchers preferred less inflated mussels, with the degree of shell inflation gradually increasing with higher intertidal elevation. Our results illustrate the importance of small-scale patterns in prey ecophenotypes in determining the distribution and feeding dynamics of wading birds.  相似文献   

18.
In Deception Bay, northern Australia, during 1979–1981, a study was made of the distribution of Scylla serrata (Forskal) in an area having a broad intertidal zone. Juveniles (20 to 99 mm carapace width) were resident in the mangrove zone, remaining there during low tide. The majority of subadult crabs (100 to 149 mm) migrated into the intertidal zone to feed at high tide and retreated to subtidal waters at low tide. Adults (150 mm and larger) were caught mainly subtidally and only small numbers were captured in the intertidal at high tide. Few crabs were captured in the coolest months (May to August). Adults were captured on the flats mainly in the warmest months (January to April), but subadults could be captured over the entire summer (September to March). Juveniles were found in the upper intertidal throughout the year.  相似文献   

19.
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  相似文献   

20.
The gastro-intestinal contents of the sea gull Laurus brunicephalus Jerdon were found to possess antibacterial activity. This activity could be traced to heavy accumulations of the marine blue-green alga Trichodesmium erythraeum in the gut. During 1969, a bloom of T. erythraeum began in Porto Novo waters about the middle of February, and attained a peak during the second week of March. It has been reported previously by the author that antibacterial properties are exhibited by T. erythraeum maintained in laboratory cultures, as well as in water samples collected from a red tide area. It was also reported (Ramamurthy, 1970) that the gut contents in 2 pelagic fishes, Hilsa kanagurta and Rastrelliger kanagurta, collected during the same red tide bloom period, possessed antibacterial properties. During this period large numbers of these fishes were consumed by sea gulls L. brunicephalus. In view of this finding, experimental procedures were adopted to determine whether extracts of T. erythraeum occurring in the gut of the sea gulls might exhibit antibacterial activity. It was found that T. erythraeum collected from the gut of L. brunicephalus could inhibit both gram positive and gram negative bacteria. Gastro-intestinal extracts from L. brunicephalus collected during the non-bloom period of T. erythraeum showed heavy microbial growth of bacteria and fungi. Evidently, antibacterial or sterile conditions prevail in the gut of these tropical sea gulls in a manner similar to that observed in Polar penguins by Sieburth (1959, 1961).  相似文献   

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