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1.
Epibiosis or fouling on living organisms can have direct and indirect detrimental effects, in particular on photosynthetic organisms such as seaweeds. It thus seems reasonable to hypothesize that macroalgae have been selected for the presence or induction of antifouling (AF) defences. The red seaweed Cryptonemia seminervis is usually found in nature with an elevated cover of epibionts. To assess the effect of epibiosis on the susceptibility of this seaweed to herbivory and fouling, the abundance of fouling was evaluated and compared to herbivore consumption (by amphipods and sea urchins) of fouled (bryozoan and sponge) and non-fouled C. seminervis. Attachment of the mussel Perna perna to surfaces treated with extracts from seaweeds with and without epibionts was also assessed. Epibiosis corresponded to ca. 51% of the blade surface of C. seminervis, sometimes covering as much as 90% and up to 51% of the thallus weight, encompassing mainly the bryozoan Membranipora membranacea and an unidentified sponge. Algae colonized by M. membranacea were preferred compared to algae devoid of epibionts, a 'shared doom' effect, either by the amphipod Elasmopus brasiliensis or by the urchin Lytechinus variegatus (p < 0.01). Sponge epibiosis also increased consumption by both herbivores (p < 0.001), suggesting that epibionts may act as lures to herbivores, attracting consumers that otherwise would not feed significantly on the seaweed. Foods containing extracts from fouled C. seminervis were preferred by urchins over the alga devoid of epibionts. However, extracts from fouled alga inhibited mussel attachment when compared to epibiont-free alga. Differences might be a direct detrimental effect of the presence of epibionts. On the other hand, epibiosis may induce the production of AF defences in C. seminervis.  相似文献   

2.
A sample of mud snails Hydrobia ulvae (Prosobranchia) from an intertidal population revealed that the shells of trematode-infected specimens were especially likely to be fouled with epibionts. Experimentally trematode-infected Biomphalaria glabrata (Pulmonata) appeared to be especially prone to develop epigrowth in comparison with uninfected conspecifics as well. These findings suggest an interaction between trematode infections and epibiosis in aquatic gastropods. The two most likely explanations for this are (1) that trematode infections weakens the snails' natural defences against epibionts, or (2) that the defences against epibionts also are effective against invading trematodes, causing snail specimens with a particularly good fouling defence to be less likely to become infected.  相似文献   

3.
Associational resistance and shared doom: effects of epibiosis on herbivory   总被引:7,自引:0,他引:7  
Martin Wahl  Mark E. Hay 《Oecologia》1995,102(3):329-340
The potential for spatial associations between palatable and unpalatable plant species to reduce herbivore pressure on the palatable species has been described as associational resistance, associational refuge or associational defense for numerous terrestrial and marine communities. One of the closest associations between species-epibiosis-has not been thoroughly investigated in this regard. In this study we evaluated how different associations between host seaweeds and epibiotic plants and animals influenced the movement of an omnivorous sea urchin (Arbacia punctulata) to the host and subsequent feeding on the host. A. punctulata showed clear preferences when given pairwise choices between 12 prey species (3 animals, 9 algae). These preferences were consistent and allowed us to rank the six epibiont species and six host species linearly from least to most preferred by A. punculata. Most host-epibiont associations dramatically changed urchin preference, increasing or decreasing urchin grazing on fouled hosts as compared to clean conspecifics. Herbivory on the host increased when the epibiont was more preferred, and decreased when it was less preferred than the unfouled host alga. Taking the host species as a point of reference, we classified epibiosis-caused decrease in herbivory as associational resistance, while epibiont-caused increases in herbivory were defined as shared doom. These epibiont-host-herbivore interactions could select for hosts that facilitate the growth of certain low preference epibionts on their surfaces in situations where the resulting decreases in herbivory would offset the various negative effects of being fouled. In contrast, in situations where herbivores are common, the negative effects of being fouled by palatable epibionts may be much greater than is generally assumed. In our assays, unpalatable hosts fouled by palatable epibionts became much more attractive to urchins and rose several ranks on the urchins' preference hierarchy.  相似文献   

4.
In this work, we studied the effects of predation by birds on the intertidal mole crab Emerita analoga fouled by macroalgae in a sandy beach of central Peru (11° S). The epibiosis affected mostly the larger animals, especially adult females. Epibiosis prevalence for the entire intertidal population was relatively low (1–2%), however, within the size range affected by epibiosis in the intertidal zone (18–23 mm in carapace length), 20–38% of the animals were fouled. Focal observations of birds showed that fouled animals are preferred over those non-fouled of the same size class and hence the same sex, being consumed at a higher rate than their proportion in the intertidal (Chesson’s alpha index of prey selection >0.96), and estimations of mortality rates indicated that more than 35% of the intertidal fouled animals are removed everyday by birds. The effect of epibiosis may be mainly attributed to a higher burrowing time or an increased visual attractive effect of the algae, which make fouled animals more conspicuous to predatory birds, or because of fouling enhances profitability of the animals. The results show that epibiosis has negative effects on E. analoga through increasing predation by birds, which in turn restricts the distribution and abundance of fouled E. analoga in the intertidal zone.  相似文献   

5.
The consequences of invasive species on ecosystem processes and ecological interactions remain poorly understood. Predator–prey interactions are fundamental in shaping species evolution and community structure and can be strongly modified by species introductions. To fully understand the ecological effects of invasive species on trophic linkages it is important to characterize novel interactions between native predators and exotic prey and to identify the impacts of invasive species on the performance of native predators. Although seaweed invasions are a growing global concern, our understanding of invasive algae—herbivore interactions is still very limited. We used a series of feeding experiments between a native herbivore and four invasive algae in the Mediterranean Sea to examine the potential of native sea urchins to consume invasive seaweeds and the impacts of invasive seaweed on herbivore performance. We found that three of the four invasive species examined are avoided by native herbivores, and that feeding behaviour in sea urchins is not driven by plant nutritional quality. On the other hand, Caulerpa racemosa is readily consumed by sea urchins, but may escape enemy control by reducing their performance. Recognizing the negative impacts of C. racemosa on herbivore performance has highlighted an enemy escape mechanism that contributes to explaining how this widespread invasive alga, which is preferred and consumed by herbivores, is not eradicated by grazing in the field. Furthermore, given the ecological and economic importance of sea urchins, negative impacts of invasive seaweeds on their performance could have dramatic effects on ecosystem function and services, and should be accounted for in sea urchin population management strategies.  相似文献   

6.
Coastal zones of the Humboldt Current Upwelling System (HCUS) are composed both of rocky and sandy beaches inhabited by macrozoobenthic communities. These show oscillating changes in the dominance of species; the abundance of the sand crab Emerita analoga is linked to phases of the El Niño Southern Oscillation (ENSO). The biogenic surfaces of these crabs serve as substrate for opportunistic colonizers. This study is the first record of an epibiosis between E. analoga and the rock mussel Semimytilus algosus, detected at a southern Peruvian sandy beach. Mussels fouled a wide size-range of adult E. analoga (7.3%) but they themselves belonged to small-size classes. The largest S. algosus was 17.4 mm in length. Highest permanence of epibionts was found on larger sand crabs (maximum between 24 and 27 mm). Significantly more mussels were found on the ventral surface (39.4%) compared to 10 other surface areas of the sand crab. Possible benefits and disadvantages of the observed epibiosis for both the basibiont and the epibiont are discussed.  相似文献   

7.
Juvenile sporophytes of the giant kelp, Macrocystis pyrifera (L.) C. A. Agardh, were transplanted from local kelp beds to stations located various distances from the outfall from an electrical generating station that was known to cause an increase in the settlement of fouling organisms. Plants near the outfall became heavily fouled by the encrusting bryozoan, Membranipora membranacea (L.), and lost about one-third of their blades during the course of the experiment. Blade loss was significantly correlated with amount of fouling. To test the hypothesis that fouling causes blade loss, we paired fouled and unfouled plants of about the same age, overall length, and number of fronds and placed them at stations in nearby kelp beds and near the outfall. At the stations in the kelp beds, the fouled plants lost blades more rapidly than the unfouled controls. However, at the station near the outfall the “control” plants quickly became fouled so there was little difference in treatments and there was no significant difference in blade loss. Plants fouled by Membranipora suffered greater blade loss than clean plants probably because fouled blades are fragile and break off easily and because fish bite off chunks of blade while foraging on the attached bryozoans.  相似文献   

8.
Many seaweeds and terrestrial plants induce chemical defences in response to herbivory, but whether they induce chemical defences against competitors (allelopathy) remains poorly understood. We evaluated whether two tropical seaweeds induce allelopathy in response to competition with a reef-building coral. We also assessed the effects of competition on seaweed growth and seaweed chemical defence against herbivores. Following 8 days of competition with the coral Porites cylindrica, the chemically rich seaweed Galaxaura filamentosa induced increased allelochemicals and became nearly twice as damaging to the coral. However, it also experienced significantly reduced growth and increased palatability to herbivores (because of reduced chemical defences). Under the same conditions, the seaweed Sargassum polycystum did not induce allelopathy and did not experience a change in growth or palatability. This is the first demonstration of induced allelopathy in a seaweed, or of competitors reducing seaweed chemical defences against herbivores. Our results suggest that the chemical ecology of coral–seaweed–herbivore interactions can be complex and nuanced, highlighting the need to incorporate greater ecological complexity into the study of chemical defence.  相似文献   

9.
Macroalgae have to cope with multiple natural enemies, such as herbivores and epibionts. As these are harmful for the host, the host is expected to show resistance to them. Evolution of resistance is complicated by the interactions among the enemies and the genetic correlations among resistances to different enemies. Here, we explored genetic variation in resistance to epibiosis and herbivory in the brown alga Fucus vesiculosus, both under conditions where the enemies coexisted and where they were isolated. F. vesiculosus showed substantial genetic variation in the resistance to both epibiosis and grazing. Grazing pressure on the alga was generally lower in the presence than in the absence of epibiota. Furthermore, epibiosis modified the susceptibility of different algal genotypes to grazing. Resistances to epibiosis and grazing were independent when measured separately for both enemies but positively correlated when both these enemies coexisted. Thus, when the enemies coexisted, the fate of genotypes with respect to these enemies was intertwined. Genotypic correlation between phlorotannins, brown-algal phenolic secondary metabolites, and the amount of epibiota was negative, indicating that these compounds contribute to resistance to epibiosis. In addition, phlorotannins correlated also with the resistance to grazing, but this correlation disappeared when grazing occurred in the absence of epibiota. This indicates that the patterns of selection for the type of the resistance as well as for the resistance traits vary with the occurrence patterns of the enemies.  相似文献   

10.
Blue mussels (Mytilus edulis) can alter the strength of byssal attachment and move between and within mussel aggregations on wave‐swept shores, but this movement ability may be limited by epibiont fouling. We quantified the effects of artificial epibiont fouling on the production of byssal threads, attachment strength, and movement in two size classes of blue mussels. In a factorial experiment, large epibiont‐covered mussels produced more functional byssal threads (i.e., those continuous from animal to substrate) after 24 h than large unfouled and small fouled mussels, but not more than small unfouled mussels. Small unfouled mussels formed and released more byssus bundles compared to any other treatment group, which indicates increased movement. Conversely, epibiont fouling resulted in decreased numbers of byssus bundles shed, and therefore reduced movement in small mussels. Epibiont‐covered mussels started producing byssal threads sooner than unfouled mussels, while small mussels began producing byssal threads earlier compared to large mussels. Mean attachment strength from both size classes increased by 9.5% when mussels were artificially fouled, and large mussels had a 34% stronger attachment compared to small mussels. On the other hand, a 2.3% decrease in attachment strength was found with increasing byssus bundles shed. Our results suggest that fouling by artificial epibionts influences byssal thread production and attachment strength in large mussels, whereas epibionts on small mussels impact their ability to move. Mussels are able to respond rapidly to fouling, which carries implications for the dynamics of mussel beds in their intertidal and subtidal habitats, especially in relation to movement of mussels within and among aggregations.  相似文献   

11.
Toth GB  Karlsson M  Pavia H 《Oecologia》2007,152(2):245-255
Herbivory on marine macroalgae (seaweeds) in temperate areas is often dominated by relatively small gastropods and crustaceans (mesoherbivores). The effects of these herbivores on the performance of adult seaweeds have so far been almost exclusively investigated under artificial laboratory conditions. Furthermore, several recent laboratory studies with mesoherbivores indicate that inducible chemical resistance may be as common in seaweeds as in vascular plants. However, in order to further explore and test the possible ecological significance of induced chemical resistance in temperate seaweeds, data are needed that address this issue in natural populations. We investigated the effect of grazing by littorinid herbivorous snails (Littorina spp.) on the individual net growth of the brown seaweed Ascophyllum nodosum in natural field populations. Furthermore, the capacity for induced resistance in the seaweeds was assessed by removing herbivores and assaying for relaxation of defences. We found that ambient densities of gastropod herbivores significantly reduced net growth by 45% in natural field populations of A. nodosum. Seaweeds previously exposed to grazing in the field were less consumed by gastropod herbivores in feeding bioassays. Furthermore, the concentration of phlorotannins (polyphenolics), which have been shown to deter gastropod herbivores, was higher in the seaweeds that were exposed to gastropod herbivores in the field. This field study corroborates earlier laboratory experiments and demonstrates that it is important to make sure that the lack of experimental field data on marine mesoherbivory does not lead to rash conclusions about the lack of significant effects of these herbivores on seaweed performance. The results strongly suggest that gastropods exert a significant selection pressure on the evolution of defensive traits in the seaweeds, and that brown seaweeds can respond to attacks by natural densities of these herbivores through increased chemical resistance to further grazing.  相似文献   

12.
Twenty-seven species of common seaweeds from the coast of Korea havebeen screened for antifouling activity. The seaweed extracts were tested inlaboratory assays against the marine fouling green alga Enteromorphaprolifera and the blue mussel Mytilus edulis. Tissue growth, sporesettlement, zygote formation and germlings of the E. prolifera wereinhibited by methanol extracts of the seaweed Ishige sinicola (= I. foliacea) and Sargassum horneri. Spore settlement was stronglyinhibited by using extract concentrations as low as 30 g mL-1with I. sinicola and 120 g mL-1 with S. horneri. The repulsive activity of the foot of the mussel was completely inhibited bymethanol extracts of I. sinicola and Scytosiphon lomentaria atconcentrations of 40 g per 10 L drop supplied to eachmussel. These extracts also showed strong antifouling activities onlarval settlement with, respectively, no or only 6% of the spat settlingwhen a test concentration of 0.8 mg mL-1 was used. This work isthe first stage towards the development of novel antifouling agents frommarine macroalgae.  相似文献   

13.
Effects of barnacle epibionts on the periwinkle Littorina littorea (L.)   总被引:1,自引:0,他引:1  
In a sandy bay with mussel beds in the Wadden Sea (Island of Sylt, eastern North Sea), periwinkles Littorina littorea (L.) were often strongly overgrown with the barnacle Balanus crenatus Bruguière in the lower intertidal zone. Consequences of this epibiosis on mobility, reproduction and mortalityof the snail were examined. B. crenatus growing on L. littorea increased snail volume up to 4-fold and weight up to 3.5-fold and crawling speed of fouled L. littorea was significantly slowed down. The epibiotic structure also caused a decrease in reproductive output. In laboratory experiments, egg production of fouled L. littorea was significantly lower than in snails free of barnacles. Presumably, copulation of the periwinkles is hampered by the voluminous and prickly cover of barnacles. Field studies demonstrated an increased mortality of overgrown L. littorea. A decrease in reproductive output and a lower survival of snails with a cover of barnacles suggest that B. crenatus epibionts may have a significant impact on the population of L. littorea. Received: 20 June 1998 / Accepted: 2 October 1998  相似文献   

14.
This work reports and illustrates secondary colonisation of Hexaplex (Trunculariopsis) trunculus shells by polychaetes in the Ria Formosa lagoon (Algarve coast—southern Portugal). Information is presented in terms of qualitative and quantitative data on epibiotic polychaetes, fouling frequency, degree of epibiosis, preferential areas for colonisation and incidence of shell damages. A total of 2,880 polychaetes belonging to 10 families were identified. Besides Spirorbidae (not quantified), the most abundant families were Serpulidae (67.5%), Sabellidae (10.2%), Sabellariidae (9.6%) and Cirratulidae (8.5%). From a total of 2,264 T. trunculus individuals analysed for epibiosis, 28.4% were non-fouled, 66.8% were intermediately fouled and 4.8% were heavily fouled. The fouling frequency, degree of epibiosis and incidence of damages in the shell spire increased markedly with T. trunculus shell length. Some epibiotic polychaetes were located preferentially on/or nearby particular features of T. trunculus shells, which probably facilitate settlement and tube construction, in addition to provide some protection against tube damage. The main consequences of epibiosis (benefits and disadvantages) for the gastropod basibiont (T. trunculus) are discussed. The present study apparently indicates that besides burrowing into soft bottom substrates, sporadic inter-tidal exposure and mutual predation on epibiosis, T. trunculus lacks other typical antifouling defence adaptations. Handling editor: K. Martens  相似文献   

15.
Marine invertebrates settle, attach, and/or metamorphose in response to signals from several sources, including seaweeds. In response to the aquaculture challenge of producing constant numbers of juveniles from cultured species, natural inducers have been screened for their ability to improve those processes. However, few chemical inducers of attachment of invertebrates have been identified, and even less of these were secondary metabolites. The goal of this work was to isolate the natural products responsible for induction activity using bioassay-guided fractionation of the organic extract of the brown seaweed Stypopodium zonale and the attachment of juveniles of the common brown mussel, Perna perna, as a model. The meroditerpene epitaondiol, identified by comparison of spectral data with the literature, promoted as much as 4.7 times more mussel attachment compared to controls at the natural concentration found in this alga (0.041% of the crude extract or 0.012% of algal dry weight). This is the first report showing that a seaweed produces terpenoid compounds as cues for invertebrate attachment, and future studies evaluating this action on settlement of mussels in the field are expected to improve aquaculture technology by increasing mussel spat production.  相似文献   

16.
This study aimed to improve understanding of the strategies developed by the Mediterranean seaweed Taonia atomaria to chemically control bacterial epibiosis. An experimental protocol was optimized to specifically extract algal surface-associated metabolites by a technique involving dipping in organic solvents whilst the integrity of algal cell membranes was assessed by fluorescent microscopy. This methodology was validated using mass spectrometry-based profiles of algal extracts and analysis of their principal components, which led to the selection of methanol as the extraction solvent with a maximum exposure time of 15 s. Six compounds (AF) were identified in the resulting surface extracts. Two of these surface-associated compounds (B and C) showed selective anti-adhesion properties against reference bacterial strains isolated from artificial surfaces while remaining inactive against epibiotic bacteria of T. atomaria. Such specificity was not observed for commercial antifouling biocides and other molecules identified in the surface or whole-cell extracts of T. atomaria.  相似文献   

17.
Human activities, including overfishing and species introductions, have had a dramatic impact on benthic communities in the Gulf of Maine within the past two decades. Prior to the 1970s, the climax community in the shallow subtidal was composed of Laminaria spp. kelp beds with an understory of arborescent red algae. In the 1980s, a population explosion of the green sea urchin, Strongylocentrotus droebachiensis, created an alternate community state, urchin barrens. Recently, a new community has been observed in former urchin barrens and kelp beds. This assemblage is principally composed of the introduced species: Codium fragile subsp. tomentosoides (green alga), Membranipora membranacea (bryozoan), Diplosoma listerianum (tunicate), Bonnemaisonia hamifera (red alga) and the opportunistic species Mytilus edulis (mussel) and Desmarestia aculeata (brown alga). In addition to changes in relative abundance, many of these species have greatly expanded their distribution and habitat selection. A model detailing mechanisms for the transition of the traditional kelp bed and urchin barren communities to others is presented and implications for this new community are discussed. This revised version was published online in July 2006 with corrections to the Cover Date.  相似文献   

18.
In the Northwest Atlantic, overgrowth of the competitively dominant, native kelps by an invasive bryozoan Membranipora membranacea increases frond erosion, which has facilitated the establishment and spread of the invasive macroalga Codium fragile ssp fragile. To document the spread of both introduced species along the Atlantic coast of Nova Scotia from initial introduction points (the ‘epicentre’) southwest of Halifax, we conducted video-surveys of shallow rocky habitats along the southwestern shore of Nova Scotia (100 km linear distance, encompassing the range of M. membranacea) in 2000, and then along the entire Atlantic coast in 2007 (650 km). Membranipora membranacea was observed continuously throughout the surveyed ranges in 2000 and 2007, wherever kelps were present, suggesting natural dispersal via planktonic larvae. Codium fragile was observed along 95 km of the surveyed range in 2000 and along 445 km in 2007, with a relatively patchy distribution beyond the epicentre, suggesting a combination of natural and anthropogenic dispersal mechanisms. Rockweed-dominated (Fucus spp.) or mixed algal assemblages common outside the epicentre may alter the interaction between M. membranacea and C. fragile, since seaweeds other than kelp are not subject to defoliation by the bryozoan. Percent cover of kelp at the epicentre generally increased from 2000 to 2007, while that of C. fragile generally decreased. Codium fragile was the dominant canopy alga at 54% of sites in 2000 and at only 15% of sites in 2007. These findings indicate that, at near decadal timescales, C. fragile does not prevent re-colonization by native kelps.  相似文献   

19.
Natural within‐thallus concentrations of elatol produced by Laurencia obtusa (Huds.) J. V. Lamour. inhibit herbivory and prevent fouling. However, elatol occurs in larger amounts within the thallus compared with the quantities from the surface of this alga. We evaluated whether the surface elatol concentrations inhibit both herbivory and fouling and whether the content of corps en cerise can be transferred to the external cell walls. Surface elatol concentrations did not inhibit herbivory by sea urchins, settlement of barnacle larvae, or mussel attachment. Evidence of a connection between the corps en cerise, where elatol is probably stored, and the cell wall of L. obtusa was based on channel‐like membranous connections that transport vesicles from the corps to the cell wall region. Therefore, L. obtusa presents a specific process of chemical transport between the cell storage structures and the plant surface. We hypothesized that if high amounts of elatol are capable of inhibiting herbivory and fouling, if the tested organisms are ecologically relevant, and if elatol really occurs on the surface of L. obtusa and this seaweed can transport this compound to its surface, the low natural concentration of defensive chemicals on the surface of L. obtusa is probably not absolute but may be variable according to environmental conditions. We also hypothesized that herbivory and fouling would not exert the same selective force for the production of defensive chemicals on L. obtusa’s surface since the low concentrations of elatol were inefficient to inhibit either processes or distinguish selective pressures.  相似文献   

20.
Epibiosis, the colonization of biogenic surfaces by epibiotic organisms such as bacteria, filamentous algae, and sessile invertebrates, poses a major threat to the fitness and survival of macroorganisms which could potentially be fouled. Fouling of artificial submerged structures can also cause severe economic problems, making the need for refined bioassays to determine the efficacy of potential antifouling compounds even more important. The aim of this study was to use the distinct swimming behaviour of zoospores of the fouling brown alga Hincksia irregularis to develop a new laboratory antifouling bioassay to test the effect of marine natural products. Spores were exposed to different concentrations of aqueous and organic extracts from body walls of sympatric echinoderms (Asteroidea: Luidia clathrata, Astropecten articulatus; Ophiuroidea: Astrocyclus caecilia). Computer-assisted motion analysis was used to distinguish between the straight and fast swimming movements of undisturbed spores (controls) and the helical and erratic swimming patterns of chemically irritated spores, using the quantitative parameters rate of direction change (RCD) and swimming speed (SPD). The ratio RCD/SPD of spore swimming paths at extract treatments compared to controls can be used to quantify the detrimental effect of echinoderm extracts. Echinoderm extracts had significant effects on spore swimming behaviour at concentrations three orders of magnitude lower than that present naturally in the echinoderm body walls (mg extract/dry weight echinoderm body wall). Comparative studies on spore settlement and germination under similar treatment conditions show that changes in spore swimming behaviour reflect decreased fitness and survivourship of algal spores. It is suggested that this bioassay can be used to screen potential antifouling extracts and compounds at very low concentrations, making this assay particularly suitable for detection of concentration dependent effects and for bioassay-guided fractionation of extracts to identify active compounds.  相似文献   

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