Effects of avian grazing on the algal community and small invertebrates in the rocky intertidal zone

In this study of a rocky intertidal habitat in northern Japan, feeding by avian consumers had significant effects on algal assemblages and small herbivorous invertebrates. The effects of the birds on algae were different from those of invertebrate grazers such as urchins and gastropods. The abundance of the dominant algal species decreased during the grazing period, increased again after the grazing period, and indirectly affected algal species richness and evenness. Avian grazing also decreased the density of tube-dwelling amphipods on the dominant alga, but did not change the density of mobile and free-living isopods. These results suggest that avian grazers may act as habitat modifiers rather than exploitative competitors for the small herbivorous crustaceans. Avian herbivores consumed only the upper parts of large algal fronds, apparently reducing the amount of suitable microhabitat for the small herbivorous crustaceans, which are subject to a variety of physical or biological stress. Thus, avian herbivores function as ecosystem engineers, regulating community structure in a manner different to invertebrate herbivores in rocky intertidal habitats.     

The invasive green alga Avrainvillea sp. transforms native epifauna and algal communities on a tropical hard substrate reef

Some introduced species compete directly with native species for resources and their spread can alter communities, while others do not proliferate and remain benign. This study compares community structure and diversity in adjacent areas dominated by the introduced alga Avrainvillea sp. or native algal species on a hard substrate reef. The biomass and species composition of 15 paired plots (30 in total, plot type based on dominance of Avrainvillea sp. or native species) were quantified. Plots dominated by Avrainvillea sp. had a significantly different assemblage of species characterized by lower algal diversity, mostly Dictyota spp. and Laurencia sp., and a higher abundance and diversity of invertebrates, such as small arthropods, polychaetes, and brittlestars. These results suggest that as Avrainvillea sp. becomes more abundant on hard substrate reefs, it will engineer a different community composed of algal epiphytes and an invertebrate assemblage more typically associated with algae in soft sediments.     

Effects of Chiton granosus (Frembly, 1827) and other molluscan grazers on algal succession in wave exposed mid-intertidal rocky shores of central Chile

Molluscan grazers can have important effects on the abundance, colonization rates, and successional pathways of algal assemblages and the entire intertidal community. In general, early successional algae are more readily consumed than corticated algae and kelps, which usually get established later in the community succession. To generalize, however, the effect of different grazers on algal assemblages must be examined on different coasts and under different scenarios. This information could help us understand the mechanisms of ecosystem processes and situations in which general models do not apply. Along the coast of Chile, humans harvest large keyhole limpets, which seem to be the only invertebrate grazers capable of controlling the dominant corticated alga Mazzaella laminarioides, a canopy-forming species that can cover extensive areas of the mid intertidal zone. In this scenario, where large limpets are harvested, the overall effects of the diverse molluscan assemblage of limpets, chitons and snails on algal succession and on corticated algae in particular are not clear. We conducted a 26-month-long experiment to evaluate the effects of molluscan grazers on mid-intertidal algal succession and to isolate the effects of Chiton granosus, the most conspicuous member of the assemblage at these tidal elevations. At sites heavily impacted by humans the molluscan grazer assemblage had strong negative effects on colonization and abundance of green algae such as ulvoids and Blidingia minima. In doing so, the grazer assemblage had a strong negative indirect effect on the establishments of chironomid fly larvae, which were only observed on green algal mats and rarely on bare rock. No significant effects were detected on epilithic microalgae, and effects on sessile invertebrates were highly variable over space and time. C. granosus also had significant negative effects on green algae but did not account for the total grazing pressure exerted by the guild. Limited foraging excursions (ca. 35 cm) from refuges and moderate site (crevice) fidelity in this species may contribute to the patchiness in green algal distribution observed in the field. Nearly 13 months after rock surface were experimentally cleared, M. laminarioides appeared in all experimental plots, but increased over three times faster in enclosures containing C. granosus than in exclosures plots or controls, suggesting that moderate levels of herbivory could actually facilitate the establishment of this alga in the succession and that the green algal cover found in the absence of grazers may delay its establishment.     

The non-native turf-forming alga Caulacanthus ustulatus displaces space-occupants but increases diversity

Non-indigenous seaweeds can be found in coastal habitats worldwide yet the ecological effects of only ~6 % of macroalgal introductions have been studied. The turf-forming red alga Caulacanthus ustulatus, a putative introduction from Asia, was discovered in southern California in 1999, yet has received very little attention despite being common in rocky intertidal habitats in the region. The purpose of this study was to evaluate the potential effects of Caulacanthus on native invertebrate and seaweed community composition. Macrofaunal, meiofaunal, and macroalgal community structure and diversity were compared between patches with (non-native) and without Caulacanthus (native) in the upper intertidal zone at 5 locations in southern California. Caulacanthus appears to displace macro invertebrates, such as barnacles, limpets, and periwinkles, while facilitating a more diverse array of meiofauna and macroalgae. This is likely due to the formation of a novel turf habitat in the upper zone where turfs are uncommon in this region naturally; algal turfs can increase habitat complexity, trap sediment, and maintain moisture during low tide which likely benefits meiofauna and seaweeds by providing food, habitat, or refuge from desiccation stress. Subsequent comparisons of invertebrate and seaweed assemblages were conducted in native and non-native patches at one site in the upper intertidal zone as well as in the middle intertidal zone where a native turf zone exists. Despite differences in community composition in the upper intertidal zone, no differences were observed in the middle zone, providing support that the novel turf created by Caulacanthus in the upper zone drives community differences.     

Community composition of the rocky intertidal at Helgoland (German Bight,North Sea)

At the rocky island of Helgoland (North Sea), the distribution and abundances of intertidal communities were assessed and the effects of wave exposure and tidal height on the spatial distribution patterns of the communities were evaluated. Macroalgae and invertebrates were sampled quantitatively along line transects in three intertidal locations, a semi-exposed, an exposed and a sheltered one. The semi-exposed location was characterised by (1) Ulva spp. at the high intertidal (Ulva-community), (2) mussels and periwinkles at the mid intertidal (Mytilus-community) and (3) Corallina officinalis and mainly the large brown alga Fucus serratus at the low intertidal (Fucus-community). The exposed location encompassed the mid and low intertidal; at both zones the Fucus-community occurred. The sheltered location was characterised by (1) barnacles (Balanus-community) and (2) bryozoans, hydrozoans and mainly the large brown alga Ascophyllum nodosum (Ascophyllum-community). At the semi-exposed, but not at the exposed location the communities changed with the intertidal position. A relationship between wave exposure and the occurrence of specific communities was shown for the sheltered location; in contrast, communities of the semi-exposed and the exposed location appear to be little influenced by wave exposure directly. The community concept and the potential causes of distribution patterns of the defined communities are discussed and suggestions for a future monitoring are given. Variations in the communities at different spatial scales speak in favour of a multiple scale sampling design to monitor changes in the intertidal communities at Helgoland. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Marine invertebrates in an algal succession. III. Mechanisms linking habitat complexity with diversity

A diverse community of marine invertebrates is associated with suceessional algal mats in an intertidal boulder field in southern California. Species richness and abundance of invertebrates increased from early to middle suceessional stages, then remained similar into the later stage. These changes are paralleled by those in the physical structure (biomass and surface area) of the algae. Increased complexity of the algal physical structure probably influenced the associated invertebrate community through several mechanisms. (1) It decreased mortality caused by predation from fish and crabs. (2) It reduced the severity of physical stresses, primarily wave shock. (3) It increased the accumulation of those individuals and species transported passively by wave action. (4) In mobile species, selection of algal substrates was largely based on physical aspects of algal structure. While no species tested appeared to select particular algal species. all actively chose algal clumps of high biomass. All four mechanisms result in the accumulation of greater numbers of individuals in algal mats with higher biomass and surface area. Because a larger sample of the available pool of individuals is collected, more species are found in a given area of algal mat when the structure is more complex. The suceessional patterns of change in species richness of the invertebrate community seem to result from this sampling phenomenon. Laboratory experiments and correlations in abundance between pairs of species gave no evidence of any competitive interactions. Any apparent “structure” in this assemblage seems to be the result of the summation of individual responses to changing algal physical structure during succession.     

LONG‐TERM CHANGES IN THE STRUCTURE OF INTERTIDAL ASSEMBLAGES AFTER INVASION BY SARGASSUM MUTICUM (PHAEOPHYTA)1

The macroalgal assemblages at the low intertidal zone were studied at three localities on the north coast of Spain between 1977 and 2002. Two of these localities were invaded at the end of the 1980s by the brown seaweed Sargassum muticum (Yendo) Fensholt (Phaeophyta, Sargassaceae), whereas the third locality remained free of the invader. In 2002, distinct algal assemblages were noticed in invaded and noninvaded localities. No major changes were detected in the noninvaded locality. Apart from the obvious presence of S. muticum, the changes observed in the invaded localities included a significant reduction in abundance of the previous dominant species (the red alga Gelidium spinosum (S. G. Gmelin) P. C. Silva) as well as an increased number of species and diversity, increased primary productivity, and variations in the seasonal abundance patterns of some species. We speculate that the arrival of S. muticum had a negative effect on the dominant native G. spinosum, probably related to competition for light. This resulted in indirect positive effects on other species of the assemblage (such as Bifurcaria bifurcata R. Ross). Other small epiphytic opportunistic species might also have been benefited from the presence of S. muticum, because the invader has a rich associated epiphytic assemblage.     

Individual and combined effects of fish predation and bed disturbance on stream benthic communities: a streamside channel experiment

1. The composition and spatiotemporal dynamics of biological communities are influenced by biotic processes, such as predation and competition, but also by physical disturbances, such as floods in running waters. However, the interplay of disturbance with predation is still poorly understood, especially in frequently disturbed streams. Further, different predator species can affect prey communities in different ways depending on their feeding mode and efficiency. 2. We investigated the individual and combined effects of flood‐induced bed disturbance and fish predation on the benthos for 4 weeks in 18 streamside channels fed by a flood‐prone New Zealand river. Bed movements caused by floods were simulated by tumbling the substratum in half the channels. Six channels each were stocked with introduced brown trout (Salmo trutta) or native upland bully (Gobiomorphus breviceps) or had fish excluded. We studied algal biomass and both invertebrate density and daytime activity on surface stones on several dates after the disturbance, invertebrate community composition in the substrata of the entire channels on day 28 and leaf decomposition rates over the 28‐day period. 3. Disturbance affected algal biomass and density, richness and activity of surface stone invertebrates, and overall density and richness of channel invertebrates. Presence or absence of fish, by contrast, did not influence overall invertebrate standing stocks when subsurface substrata were included but did affect invertebrate densities on surface stones in 45% of all analysed cases and invertebrate activity on surface stones in all cases. Leaf decomposition rates were not influenced at all by the experimental manipulations. 4. Native upland bullies featured more often than exotic brown trout in causing invertebrate density changes and equally often in causing changes to grazer behaviour. Overall, our results imply that fish predation can have strong effects on the benthic invertebrate community in frequently disturbed streams, especially via behavioural changes.     

Cultivation of the brown alga Sargassum horneri: sexual reproduction and seedling production in tank culture under reduced solar irradiance in ambient temperature

As a large conspicuous intertidal brown alga, individuals of Sargassum horneri can reach a length of more than 7 m with a fresh weight of 3 kg along the coasts of the Eastern China Sea. The biomass of this alga as a vital component in coastal water ecology has been well documented. In recent years, a steady disappearance of the algal biomass along the once densely populated coastal areas of the Eastern China Sea has drawn attention in China. Efforts have been made to reconstruct the subtidal algal flora or even to grow the alga by use of long-lines. As part of the efforts to establish an efficient technique for producing seedlings of S. horneri, in this investigation a series of culture experiments were carried out in indoor raceway and rectangular tanks under reduced solar irradiance at ambient temperature in 2007–2008. The investigation demonstrated that: (1) sexual reproduction of S. horneri could be accelerated in elevated temperature and light climates, at least 3 months earlier than in the wild; (2) eggs of S. horneri had the potential to be fertilized up to 48 h, much longer than that of known related species; (3) suspension and fixed culture methods were both effective in growing the seedlings to the long-line cultivation stage; and (4) the life cycle of S. horneri in culture could be shortened to 4.5 months, thus establishing this alga as an appropriate model for investigating sexual reproduction in dieocious species of this genus.     

优势种去除对崇明东滩盐沼湿地生态系统的影响

盐沼生态系统环境梯度明显,物种组成较简单,是研究生物多样性与生态系统功能关系的理想对象。本研究以崇明东滩盐沼湿地为研究区域,研究优势种去除对植物群落结构以及底栖动物群落的影响。结果表明:(1)去除处理仅对植物群落分株密度有极显著效应(P0.01)。去除组和对照组物种组成差异随时间增加而减小,处理效应逐渐减弱。(2)去除组底栖动物密度均低于对照组,但差异不显著。(3)盐沼植物群落特征与底栖动物群落有密切关系,植物密度、冠层高度与底栖动物密度相关性极显著。去除优势种后,植物群落分株密度升高,群落内剩余物种占比有所上升,次优势种对群落的补偿效应具有较大贡献;而底栖动物群落密度下降,其生物量和多样性指数的变化趋势与密度并不一致。上述结果表明生物多样性变化影响了盐沼湿地生态系统植物群落和底栖动物群落结构,进而可能影响物质循环和能量流动过程。     

Seasonal changes in an intertidal annual algal assemblage in northern Japan: The role of pre-emption and grazing on algal replacement

The pattern and process of seasonal changes in an intertidal annual algal assemblage were examined at Hiura, northern Japan. Short-term field experiments (<2months duration) were set up to quantify the effects of both grazing and pre-emption on species replacement in the assemblage in three different seasons. An 8-month field experiment was set up to quantify long-term effects, including the indirect effects of both grazing and competitive dominance on the community structure. Results suggested that seasonal change in the algal assemblage resulted from the interaction of abiotic environmental change, competition and grazing. The relative contribution of these factors varied within a short period, presumably as a result of seasonal changes in physical environmental stress, free space availability and grazing pressure. From February to March, when grazer density was low and there was much free space available for algae, the dominant species shifted from foliose green alga Monostroma angicava to filamentous red alga Bangia atropurpurea, because B.atropurpurea grew faster than M.angicava. This species replacement was not influenced strongly by biological interaction but by temporal changes in abiotic environmental conditions. From April to mid May, when there was less free space available for algae in the natural community, the dominant B.atropurpurea decreased with increasing foliose red alga Porphyra yezoensis, because only P.yezoensis was able to invade an area pre-empted by algae. Grazing did not affect this species replacement. After mid May, the two dominant species, P.yezoensis and B.atropurpurea, decreased. Their decline was mainly caused by desiccation stress and was partially affected by grazing.     

Comparative physiological studies on the growth of cryptic species of Bostrychia intricata (Rhodomelaceae,Rhodophyta) in various salinity and temperature conditions

Inter‐ and intra‐specific physiological variations of intertidal macroalgae have been well investigated. However, studies on physiological responses of cryptic algal species have been poorly documented. Bostrychia intricata is a widespread marine red alga in the Southern Hemisphere, and has many cryptic species. We investigated the effect of different salinities and temperatures on the specific growth rate of three cryptic species (N2, N4 and N5) of B. intricata from New Zealand. Our data indicated that all cryptic species grew at the full range of salinities and temperatures tested, but exhibited a significant difference in their specific growth rates. Cryptic species N4 had a higher growth rate than the other two cryptic species under all experimental conditions, whereas cryptic species N2 occasionally showed a higher growth rate than cryptic species N5 at high salinities and lower temperatures. The distinct physiological properties of these cryptic species may explain their distribution pattern (a wider distribution of cryptic species N4 than N2 and N5) in New Zealand. The physiological divergence between the cryptic species could be related to their levels of evolutionary divergence, with more similar physiology between cryptic species, which share a more recent common ancestor (N2 and N5). Our findings underline that morphologically indistinguishable cryptic algal species are different in many other aspects and are truly independent entities.     

Extreme heatwave drives topography-dependent patterns of mortality in a bed-forming intertidal barnacle,with implications for associated community structure

Heatwave frequency and intensity will increase as climate change progresses. Intertidal sessile invertebrates, which often form thermally benign microhabitats for associated species, are vulnerable to thermal stress because they have minimal ability to behaviourally thermoregulate. Understanding what factors influence the mortality of biogenic species and how heatwaves might impact their ability to provide habitat is critical. Here, we characterize the community associated with the thatched barnacle, Semibalanus cariosus (Pallass, 1788), in British Columbia (BC), Canada. Then, we investigate what site-level and plot-level environmental factors explained variations in barnacle mortality resulting from an unprecedented regional heatwave in BC, Canada. Furthermore, we used a manipulative shading experiment deployed prior to the heatwave to examine the effect of thermal stress on barnacle survival and recruitment and the barnacle-associated community. We identified 50 taxa inhabiting S. cariosus beds, with variations in community composition between sites. Site-scale variables and algal canopy cover did not predict S. cariosus mortality, but patch-scale variation in substratum orientation did, with more direct solar irradiance corresponding with higher barnacle mortality. The shading experiment demonstrated that S. cariosus survival, barnacle recruitment, and invertebrate community diversity were higher under shades where substratum temperatures were lower. Associated community composition also differed between shaded and non-shaded plots, suggesting S. cariosus was not able to fully buffer acute thermal stress for its associated community. While habitat provisioning by intertidal foundation species is an important source of biodiversity, these species alone may not be enough to prevent substantial community shifts following extreme heatwaves. As heatwaves become more frequent and severe, they may further reduce diversity via the loss of biogenic habitat, and spatial variation in these impacts may be substantial.     

The influence of fish predation on the abundance cycles of an algal turf invertebrate fauna

Summary A fauna of small invertebrates in a perennial algal turf habitat showed marked seasonal fluctuations in abundance. These were characterised by a summer (December) peak in the abundance of gammarid amphipods and an autumn (April) peak in polychaete densities. Invertebrate abundance was lowest during the period February–March which coincided with an influx of very high densities of juveniles (0⁺ year class) of the sparid fish Chrysophrys auratus. Both juvenile C. auratus and also the mullid Upeneichthys porosus achieved their highest densities over coralline turf areas and fed on the associated invertebrates. Gammarid amphipods constituted the main food items of each species. All these observations suggested a key role of fish predators in determining the observed seasonal patterns of invertebrate abundance.This hypothesis was investigated by the use of replicated fish exclusion shields and cages which excluded both species from the algal turf and also permitted an assessment of Upeneichthys porosus feeding in the absence of Chrysophrys auratus. The experiment ran from November until June and covered the period of invertebrate and fish abundance changes. Little evidence of a fish predation effect on either the densities of or the timing of abundance peaks of the invertebrate fauna was detected. Sediment analysis of the cage and control experimental sites revealed no evidence of a cage effect on the microhabitat. It is suggested that seasonal changes in the abundance of invertebrates occurs independently of the high densities of predatory fish recorded in this habitat.     

Microfauna associated with algal mats in melt ponds of the Ross Ice Shelf

Summary The microfauna of benthic algal mats in eight ponds of diverse chemistry on the Ross Ice Shelf was examined. Tardigrades, nematodes and rotifers were successfully extracted from algal material by a modified Baermann funnel technique, which was robust enough to use under field conditions. Seven species of rotifer (of which 6 were bdelloids) and one species of nematode and tardigrade were found, all of which are cosmopolitan to Antarctic lentic ecosystems. Philodina sp. was the only invertebrate to occur in all ponds sampled. Rotifers were the most abundant invertebrates with average densities in all ponds of 4.26×10⁵ individuals/m², whereas nematodes were least abundant with densities of 4.96×10⁴ individuals/m². Total invertebrate densities and densities of individual taxa were significantly different in each of the 8 ponds, and of those measured environmental parameters were most strongly associated with conductivity. Cores taken from four ponds 2 cm into underlying substratum revealed that most animals live within the algal mats, with underlying sediment supporting a minimal population.     

Stable isotopes, mesocosms and gut content analysis demonstrate trophic differences in two invasive decapod crustacea

1. Improving our understanding of dietary differences among omnivorous, benthic crustacea can help to define the scope of their trophic influence in benthic food webs. In this study, we examined the trophic ecology of two non‐native decapod crustaceans, the Chinese mitten crab (Eriocheir sinensis) (CMC) and the red swamp crayfish (Procambarus clarkii) (RSC), in the San Francisco Bay ecosystem to describe their food web impacts and explore whether these species are functionally equivalent in their impacts on aquatic benthic communities. 2. We used multiple methods to maximise resolution of the diet of these species, including N and C stable isotope analysis of field data, controlled feeding experiments to estimate isotopic fractionation, mesocosm experiments, and gut content analysis (GCA). 3. In experimental enclosures, both CMC and RSC caused significant declines in terrestrially derived plant detritus (P < 0.01) and algae (P < 0.02) relative to controls, and declines in densities of the caddisfly Gumaga nigricula by >50% relative to controls. 4. Plant material dominated gut contents of both species, but several sediment‐dwelling invertebrate taxa were also found. GCA and mesocosm results indicate that CMC feed predominantly on surface‐dwelling invertebrates, suggesting that trophic impacts of this species could include a shift in invertebrate community composition towards sediment‐dwelling taxa. 5. Stable isotope analysis supported a stronger relationship between CMC and both algae and algal‐associated invertebrates than with allochthonous plant materials, while RSC was more closely aligned with terrestrially derived detritus. 6. The trophic ecology and life histories of these two invasive species translate into important differences in potential impacts on aquatic food webs. Our results suggest that the CMC differs from the RSC in exerting new pressures on autochthonous food sources and shallow‐dwelling invertebrates. The crab's wide‐ranging foraging techniques, use of intertidal habitat, and migration out of freshwater at sexual maturity increases the distribution of the impacts of this important invasive species.     

A long-term comparison of the benthic algal flora of Clare Island, County Mayo, western Ireland

The marine benthic algal flora of Clare Island, off County Mayo, western shore of Ireland, was investigated; collections of intertidal and subtidal marine algae were made at 16 sites along the eastern and southern shores in the years 1990, 1993 and 2000–2002. The data and observations obtained were compared with the results of a similar survey conducted by Arthur Disbrowe Cotton in 1910–1911. Considering the results of the original survey and the new survey together, the marine algal flora of the island currently totals 293 species; 224 species were recorded by Cotton in the original survey, whereas 223 species were identified in the present study. Most species are common to the original and the new list and the main differences are easily explainable; the new survey used SCUBA diving, which allowed the collection of several subtidal species not collected in 1910, and Cotton reported several microscopic green and brown algae, usually difficult to recognise in the field, which were not rediscovered. The most remarkable differences consist in the current presence of some large intertidal brown algae (Bifurcaria bifurcata, Cystoseira foeniculacea and Cystoseira nodicaulis) that were not reported in the survey of 1910. Two algae, Codium fragile subsp. tomentosoides and Asparagopsis armata, were introduced in Europe after the original survey. At present, the benthic algal assemblages of Clare Island still have basically the same structure and distribution as in 1910 and, if compared with other coastal areas of Europe, the intertidal marine environment of Clare Island appears remarkably well conserved.     

黄河三角洲芦苇湿地底栖无脊椎动物与环境因子的关系研究——以石油开采区与淡水补给区为例

分别于2018年5月和8月对黄河三角洲芦苇湿地19(淡水补给区11处,石油开采区8处)处采样点的底栖无脊椎动物和水体理化指标进行调查采样,运用统计方法分析两个区域物种组成、优势种、多样性、群落结构以及与环境因子的关系。结果表明:两季共采集到底栖无脊椎动物54种,主要以水生昆虫、腹足纲和软甲纲为主,淡水补给区和石油开采区各类群组成差异明显。独立样本T检验表明淡水补给区和石油开采区水体理化指标间差异显著(P0.05)。双因子方差分析显示,昆虫纲和腹足纲密度在两区域差异显著(P0.05),软甲纲和腹足纲密度在季节上差异明显(P0.05)。底栖无脊椎动物优势种共10种,淡水补给区指示物种8种,而石油开采区未发现有指示物种。聚类和非参数多维排序(nM DS)显示,底栖无脊椎动物群落结构相似性较低; RDA结果表明:淡水补给区底栖无脊椎动物群落结构主要受Cond,TDS,Sal,pH,Eh,HCO_3~-,SO_4~(2-)等环境因子的影响。石油开采区底栖无脊椎动物群落结构影响较大的环境因子为HCO_3~-、NH_4-N。     

Coexistence in the intertidal: interactions between the non‐indigenous New Zealand mud snail Potamopyrgus antipodarum and the native estuarine isopod Gnorimosphaeroma insulare

Nonindigenous species (NIS) that achieve high densities in their recipient communities are expected to have negative effects on native competitors. However, recent reviews have shown that competitors, unlike predators and parasites, have rarely been documented to cause extinctions. There is a need for better understanding of competitive interactions between nonindigenous species and native competitors across systems where NIS reach high densities. In this paper we examine competitive interactions between the exotic, invasive New Zealand mud snail Potamopyrgus antipodarum and native estuarine invertebrates. P. antipodarum is often described as a freshwater snail but can occur in brackish water as well; it has been established in the Columbia River Estuary for over ten years. We addressed competition in this estuarine system using three approaches: field surveys of the benthic invertebrate community, stable isotope analysis of overlap in resource use by common invertebrates, and a laboratory competition experiment that tested the strength of competition between P. antipodarum and the isopod, Gnorimosphaeroma insulare, which overlap in resource use. We found no evidence of negative competitive impacts of P. antipodarum on native benthic invertebrates in this estuarine system. Densities of P. antipodarum and common native epibenthic invertebrates are positively correlated in the intertidal, likely due to shared habitat preferences. The competition experiment showed that the effect of interspecific competition from P. antipodarum on the foraging and survival of G. insulare was significantly less than the effect of intraspecific competition between isopods. The presence of the isopod G. insulare reduced foraging in P. antipodarum, but this change did not result in reduced survival and growth of snails. Hence, interspecific competition is weak despite high densities achieved by the invading species. Finally, we discuss several possible explanations for the observation that P. antipodarum does not have an obvious negative competitive impact on native benthic invertebrates.     

Disturbance and organization of macroalgal assemblages in the Northwest Atlantic

Large seaweeds are often structurally dominant in subtidal and intertidal rocky shore benthic communities of the N.W. Atlantic. The mechanisms by which these algal assemblages are maintained are surprisingly different in the two habitats. In the subtidal community, kelps are dominant space competitors in the absence of strong grazing interactions. In contrast, the large perennial seaweeds of intertidal zones (fucoids and Chondrus crispus) are competitively inferior to both sessile filter feeders and ephemeral, pioneer algal species. Intertidal seaweed beds are maintained by carnivory of whelks, which reduces filter feeder populations, and by herbivorous periwinkles which reduce ephemeral algal populations. Through most of the intertidal zone, disturbance, both biological and physical, dictates which species shall compete and equilibrium conditions obtain subsequently.The roles of subtidal consumers are quite different. Sea urchins are the major algal herbivores and these voracious animals maintain an equilibrium state in which large tracts of subtidal coralline pavement are kept free of kelp forests. Urchins do not seem to play a successional facilitative role for kelps in the way that periwinkles do for fucoids in the intertidal. Control of herbivore populations is thus a key to the maintenance of subtidal foliose algal beds. It is clear that parasitic amoebas can decimate sea urchin populations so that kelp forest dominance is assured. However, the importance of carnivory in limiting urchins in the subtidal community is unclear in the absence of appropriate manipulation experiments. It is possible that carnivorous decapods and fin fish control sea urchin populations and hence foliose algal abundance, but this must remain speculative. The seaweed-dominated state of the subtidal system is an alternative equilibrium condition to the urchin/coralline alga configuration. The structure of the kelp beds is relatively uniform in responding to frequent small-scale, infrequent large-scale, or no, disturbance.