Artificial structures influence fouling on habitat-forming kelps

The addition of artificial structures along urbanised shorelines is a global phenomenon. Such modifications of habitats have important consequences to the abundance of fouling organisms on primary substrata, but the influence on fouling of habitat-formers living on these structures is poorly understood. Fouling of habitat-forming kelps Ecklonia radiata on pier-pilings was compared to that on rocky reefs at three locations in Sydney Harbour. Kelps on pilings supported different assemblages of bryozoans from those on reefs. The abundances of bryozoans on kelps, in particular of the non-indigenous species Membranipora membranacea, were significantly greater on pilings. Differences were consistent in time and space. This indicates that the addition of artificial structures also affects fouling on secondary biogenic substrata, altering biodiversity and potentially facilitating the introduction and dispersal of non-indigenous epibiota. Understanding the processes that cause these patterns is necessary to allow sensible predictions about ecological effects of built structures.     

Artificial structures influence fouling on habitat-forming kelps

The addition of artificial structures along urbanised shorelines is a global phenomenon. Such modifications of habitats have important consequences to the abundance of fouling organisms on primary substrata, but the influence on fouling of habitat-formers living on these structures is poorly understood. Fouling of habitat-forming kelps Ecklonia radiata on pier-pilings was compared to that on rocky reefs at three locations in Sydney Harbour. Kelps on pilings supported different assemblages of bryozoans from those on reefs. The abundances of bryozoans on kelps, inparticular of the non-indigenous species Membranipora membranacea, were significantly greater on pilings. Differences were consistent in time and space. This indicates that the addition of artificial structures also affects fouling on secondary biogenic substrata, altering biodiversity and potentially facilitating the introduction and dispersal of non-indigenous epibiota. Understanding the processes that cause these patterns is necessary to allow sensible predictions about ecological effects of built structures.     

Floating pontoons create novel habitats for subtidal epibiota

Urban structures in the form of pontoons and pilings represent major coastal habitats for marine organisms and understanding the factors causing abundances of organisms to differ between these and natural habitat has been neglected in the study of coastal ecology. It has been proposed that composition of substrata explain differences previously described between subtidal assemblages of epibiota on rocky reef (sandstone) and pontoons (concrete) in Sydney Harbour, Australia. This study tested the hypothesis that differences in the composition of substratum (sandstone vs. concrete) independent of type of habitat (rocky reef vs. pontoon) affects the development of epibiotic assemblages. This was tested by experimentally providing substratum of the two types in both habitats. Epibiotic assemblages were unaffected by the composition of substratum but strongly affected by the type of habitat; demonstrating that pontoons constitute novel habitats for epibiota. This result highlights a need for determining how current ecological understanding of subtidal epibiota, which is heavily based on studies of urban structures (pilings and pontoons), relates to natural reef. Future tests of hypotheses about the nature of these differences will not only contribute to better ecological understanding of epibiota and their use of urban structures as habitats, but also to better predictions of future changes to the ecology of coastal habitats.     

Weak Effects of Epibiota on the Abundances of Fishes Associated with Pier Pilings in Sydney Harbour

Biogenic habitats have profound effects on the distribution and abundances of many organisms. Epibiota are major biogenic components of hard substrata in marine habitats, particularly on artificial structures such as pier pilings, and have the potential to influence organisms associated with these structures. This study tested hypotheses about effects of epibiota on abundances of fishes associated with pilings in Middle Harbour, Sydney. Amount of epibiota had no effect on abundances of most species of fish, suggesting that epibiota are not important resources for these species. Abundances of the hulafish, Trachinops taeniatus, were, however, found to be greater around pilings with large than pilings with small amounts of epibiota at one site, at each of two times. Experimental removals of epibiota from pilings were done to test the hypothesis that epibiota are an important resource to T. taeniatus. Despite great temporal fluctuations in abundances over the duration of the experiment, the response of T. taeniatus to the removal of epibiota from pilings was as predicted, falling to zero following removal of epibiota. Although this suggests that epibiota are an important resource for this species, this pattern was spatially and temporally variable. We suggest that variation in the amount of epibiota has minor effects on the abundances of fishes around pilings.     

Seawater-atmosphere O(2) exchange rates in open-top laboratory microcosms: application for continuous estimates of planktonic primary production and respiration

Foraging by predatory fish is thought to be one of the primary ecological processes affecting the abundances of plants and animals in subtidal habitats. The importance of this process was assessed on the subtidal surfaces of urban structures (pontoons and pilings) that represent major coastal habitats for marine organisms. Fish feed with greater intensity on epibiota attached to pilings than pontoons and it was hypothesised that greater predation on pilings explained why the structure of epibiotic assemblages differs between these habitats. I predicted that the structure of epibiotic assemblages would develop differently between pilings and pontoons in the presence of fish (plates open to predation) but not in the absence of fish (plates inside exclusion cages). Results revealed large differences in abundance between pilings and pontoons that were largely independent of the caged and uncaged plates. Predation may be intense (as it appeared on pilings) but unimportant because it does not explain observed abundances of prey (epibiota between pilings and pontoons).     

Indirect effect of nitrogen enrichment modified invertebrate herbivory through altering plant community composition in an alpine meadow

Aims Nitrogen enrichment may affect ‘community invertebrate herbivory’ (hereafter ‘herbivory’) directly by changing plant species’ specific herbivory, or indirectly by altering the composition of natural plant communities. Here, we investigated how community composition altered the community herbivory in natural ecosystems and compared the relative importance of direct and indirect effects of nitrogen addition on community herbivory.     

Characterizing ecosystem-level consequences of biological invasions: the role of ecosystem engineers

Invasions by exotic species represent both threats to ecosystems as well as opportunities to learn more about them. Among the invaders that will have the largest impacts are those that directly modify ecosystems and thus have cascading effects for resident biota. Exotics can affect ecosystems by altering system-level flows, availability, or quality of nutrients, food, and physical resources (e.g. living space, water, heat or light). The invader-mediated control of physical resources, typically achieved through the modification of habitats, has received limited attention in invasion biology. This reflects a general trend in ecology, and only recently has the concept of ecosystem engineering been developed to account for the role of species that shape habitats. Plants and animals in terrestrial and aquatic systems can both create and destroy structure. When introduced into ecosystems, these exotic engineers cause physical state changes with effects that ramify throughout the system. Although the consequences of these modifications are varied and complex, insight gained from general ecological principles offers an opportunity to predict what invaders will do upon their integration into systems. Examples from the literature suggest that introduced ecosystem engineers that increase habitat complexity or heterogeneity tend to cause abundances and/or species richness to rise, while those that decrease complexity tend to have the reverse effect. In assessing such patterns, however, it is critical to also consider spatial scales and the life habits of resident organisms. In addition to providing predictive power, recognition of engineering as a major means by which invasive species affect ecosystems provides a unifying theme for invasion biology and offers a chance to consider more fully the general role of species in ecosystems.     

Effects of two marinas on the composition of fouling assemblages

The composition of fouling assemblages was surveyed inside and near two fully enclosed marinas using settlement plates. The location of a plate, inside or outside the marina, influenced the abundances of four functional groups of fouling organisms (solitary ascidians, arborescent bryozoans, encrusting bryozoans and colonial ascidians). Transplantation of mature assemblages revealed that reductions in the abundance of arborescent bryozoans inside marinas might be explained by increased growth and recruitment of these bryozoans outside the marina. Surveys of settlement revealed decreased recruitment of encrusting bryozoans inside the marinas, a result consistent with patterns of adult abundance. It is proposed that an increased abundance of solitary ascidians inside marinas may be due to decreased competition. A second survey of adult assemblages was performed with multiple ‘Outside’ sites per marina. Effects of location consistent with the first survey were found for arborescent bryozoans, and in one marina area for solitary ascidians and encrusting bryozoans, but not in the other. Although mechanisms can be proposed to explain the effects of the marina (inside or outside) on the abundances of solitary ascidians, arborescent bryozoans and encrusting bryozoans, the second survey revealed that the effects may vary among marinas.     

Mussel beds on different types of structures support different macroinvertebrate assemblages

Abstract Artificial structures, such as seawalls, pilings and pontoons, are common features of urban estuaries. They replace natural structures or add to the amount of hard substratum in an area and provide habitats for many fish and invertebrates. Previous work has concentrated on fish or on the invertebrates that occupy the primary substratum of artificial structures. Mussels often grow on different types of structures (pontoons, pilings, seawalls and natural reefs) and provide a secondary substratum for other organisms to inhabit. Counting and identifying organisms associated with mussel beds is traditionally done to species level, which is very time‐consuming. To save time, organisms in this study were identified to coarse levels of taxonomic resolution (a mix of taxa, such as class, order, family and genus), which showed similar patterns to those when particularly speciose and abundant groups were identified to species. This study tests hypotheses that the distribution and abundance of mobile and sessile organisms that inhabit mussel beds will differ among natural and various types of artificial structures. When the associated assemblages of mussel beds from different types of structures and from different locations were examined, assemblages varied according to the type of structure they inhabited and its location. Assemblages associated with mussels on pontoons differed consistently from those on other types of structures. Patterns in the assemblages were also consistent through time. These data show that the types and amounts of artificial structures added to an environment can affect the types, distribution and abundances of organisms living in biogenic habitats.     

Priority effects on faunal assemblages within artificial seagrass

An enduring challenge for community ecologists is to understand how different assemblages of species are derived from a common pool of potential colonists. Early colonisers can affect the ability of subsequent arrivals to colonise and persist, and thereby influence the development and structure of the resulting assemblage. In two independent experiments, we tested for such historical effects of priority on the assemblages of mobile fauna colonising artificial seagrass initially occupied by epibiota (algae and sessile invertebrates) and/or predatory Palaemonid shrimp. Multivariate analyses detected strong priority effects of both epibiota and Palaemonids on the structure of faunal assemblages as early as 5 days, and up to 45 days after experimental treatments were established, even though initial conditions (i.e. abundances of epibiota and Palaemonids) had become similar among treatments. The abundances of key taxa, identified by SIMPER analyses as those taxa contributing the most to multivariate differences among treatments, were typically enhanced where epibiota were initially present. In contrast, prior colonisation by Palaemonid shrimp produced subtle and variable effects on individual taxa. Nevertheless, these experiments provide evidence for priority effects and demonstrate how variation in the structure of contemporary assemblages can be intimately linked with key historical events that occurred during their development, but may no longer be apparent. Consequently, our ability to interpret variation among contemporary assemblages may be enhanced when contemporary patterns are viewed within an historical context.     

Long-Term Field Study Reveals Subtle Effects of the Invasive Alga Sargassum muticum upon the Epibiota of Zostera marina

Invasive species can alter coastal ecosystems both directly, e.g. through competition for substratum and nutrients, and indirectly. Indirect effects may be mediated by creation of dissimilar or inimical habitats, changes in predator and/or prey assemblages, alterations in associated biota, and perturbations of water movement and thermal regimes. Previous studies have shown that invasive algae can modify native habitat architecture, disrupt intricately linked food webs and alter epibiotic assemblages. In the UK, the seagrass Zostera marina supports a diverse epibiotic assemblage, influencing key factors such as sediment dynamics, depositional regime and trophic linkages. Increasing encroachment of the invasive alga Sargassum muticum into seagrass meadows changes the physical and chemical characteristics of the local environment and creates the potential for changes in the epibionts associated with the seagrass blades, threatening the integrity of the seagrass ecosystem. We investigated the effects of S. muticum invasion upon the epibiota of Z. marina in a drowned river valley in SW England seasonally from spring to autumn over four years in an in-situ manipulative experiment, comparing permanent quadrats with and without artificially introduced S. muticum. Epibiota were weighed, identified to the most detailed operational taxonomic unit (OTU) possible, and unitary organisms were enumerated. Multivariate PERMANOVA+ analysis revealed significant differences in epibiont assemblages between Sargassum treatments. Linear mixed effects models indicated that differences in epibiota assemblage composition were not reflected as significant differences in mean biomass per sample, or number of epibiont OTUs per sample. We conclude that S. muticum invasion into Z. marina meadows may significantly alter the species composition and abundance distribution of epibiotic assemblages found on the blades of the seagrass. Thus S. muticum invasion could have more wide-reaching effects on processes within coastal ecosystems than predicted purely by direct effects.     

delta(15)N as an integrator of the nitrogen cycle

Natural abundances of the rare stable isotope of nitrogen, 15N, are now being used widely in research on N cycling in organisms and ecosystems. 15N natural abundances are used in fundamentally different ways from traditional 15N tracers by integrating N cycle processes via N isotope fractionations and the mixing of various N-containing pools. This approach of using 15N natural abundances still requires certain technical and conceptual advances before it can be used routinely in ecological research. Here, the basis of the natural abundance 15N approach and opportunities for applying it in ecology are reviewed, and recent progress towards overcoming some of the key technical problems and in revealing large-scale patterns in N cycle processes is discussed.     

Environmental constraints upon locomotion and predator-prey interactions in aquatic organisms: an introduction

Environmental constraints in aquatic habitats have become topics of concern to both the scientific community and the public at large. In particular, coastal and freshwater habitats are subject to dramatic variability in various environmental factors, as a result of both natural and anthropogenic processes. The protection and sustainable management of all aquatic habitats requires greater understanding of how environmental constraints influence aquatic organisms. Locomotion and predator-prey interactions are intimately linked and fundamental to the survival of mobile aquatic organisms. This paper summarizes the main points from the review and research articles which comprise the theme issue 'Environmental constraints upon locomotion and predator-prey interactions in aquatic organisms'. The articles explore how natural and anthropogenic factors can constrain these two fundamental activities in a diverse range of organisms from phytoplankton to marine mammals. Some major environmental constraints derive from the intrinsic properties of the fluid and are mechanical in nature, such as viscosity and flow regime. Other constraints derive from direct effects of factors, such as temperature, oxygen content of the water or turbidity, upon the mechanisms underlying the performance of locomotion and predator-prey interactions. The effect of these factors on performance at the tissue and organ level is reflected in constraints upon performance of the whole organism. All these constraints can influence behaviour. Ultimately, they can have an impact on ecological performance. One issue that requires particular attention is how factors such as temperature and oxygen can exert different constraints on the physiology and behaviour of different taxa and the ecological implications of this. Given the multiplicity of constraints, the complexity of their interactions, and the variety of biological levels at which they can act, there is a clear need for integration between the fields of physiology, biomechanics, behaviour, ecology, biological modelling and evolution in both laboratory and field studies. For studies on animals in their natural environment, further technological advances are required to allow investigation of how the prevailing physico-chemical conditions influence basic physiological processes and behaviour.     

Ecosystem engineers in the pelagic realm: alteration of habitat by species ranging from microbes to jellyfish

Ecosystem engineers are species that alter the physical environment in ways that create new habitat or change the suitability of existing habitats for themselves or other organisms. In marine systems, much of the focus has been on species such as corals, oysters, and macrophytes that add physical structure to the environment, but organisms ranging from microbes to jellyfish and finfish that reside in the water column of oceans, estuaries, and coastal seas alter the chemical and physical environment both within the water column and on the benthos. By causing hypoxia, changing light regimes, and influencing physical mixing, these organisms may have as strong an effect as species that fall more clearly within the classical category of ecosystem engineer. In addition, planktonic species, such as jellyfish, may indirectly alter the physical environment through predator-mediated landscape structure. By creating spatial patterns of habitats that vary in their rates of mortality due to predation, planktonic predators may control spatial patterns and abundances of species that are the direct creators or modifiers of physical habitat.     

Opportunistic predation by small fishes on epibiota of jetty pilings in urban waterways

Epibiota were sampled on nine small jetties in the tidal, urban canals of south-east Queensland, Australia, to determine if the small fishes that are associated with these jetties prey on the epibiota on the pilings of the jetties and whether these fishes depend on the epibiota as a source of food. Epibiota was dominated by barnacles, filamentous and foliose algae and ranged in thickness from 4 to 11 mm. The two species of fishes that associated most closely with jetty pilings, Pandaka lidwilli (Gobiidae) and Monodactylus argenteus (Monodactylidae), were sampled twice during the day and twice during the night for analysis of stomach contents. During the day, the diet of P. lidwilli was dominated by amphipods ( c. 70%, by mass of organic content), with copepods, bivalves and bryozoans each contributing <10%. At night, amphipods contributed less ( c. 45%) and copepods more ( c. 35%). The diet of M. argenteus was dominated by filamentous algae (55%) and amphipods (20%) during the day and filamentous algae (70%) and barnacle cirri (23%) at night. Epibiota, therefore, made a substantial contribution to the diet of the fishes but were not the sole source of food for either species. As jetties were the only structures that supported epibiota in the area, fishes probably sourced their epibiota from the pilings of the jetties. Whether fishes depended on the epibiota was, therefore, tested using a manipulative before-after-control-impact (BACI) study. Three jetties were assigned randomly to each of three treatments: (1) epibiota removed from pilings, (2) epibiota cut and damaged (a procedural control) and (3) epibiota left undisturbed. Abundances of P. lidwilli and M. argenteus around jetty pilings remained similar across all treatments from before to after the removal of epibiota. These results indicate that although fishes consumed epibiota on the jetties, they did not depend on the epibiota of the jetties for food.     

Non-indigenous invasive bivalves as ecosystem engineers

Several non-indigenous bivalve species have been colonising aquatic ecosystems worldwide, in some cases with great ecological and economic impacts. In this paper, we focus on the ecosystem engineering attributes of non-indigenous invasive bivalves (i.e., the capacities of these organisms to directly or indirectly affect the availability of resources to other species by physically modifying the environment). By reviewing the ecology of several invasive bivalves we identify a variety of mechanisms via which they modify, maintain and/or create habitats. Given the usually high densities and broad spatial distributions of such bivalves, their engineering activities can significantly alter ecosystem structure and functioning (e.g., changes in sediment chemistry, grain size, and organic matter content via bioturbation, increased light penetration into the water column due to filter feeding, changes in near bed flows and shear stress due to the presence of shells, provision of colonisable substrate and refuges by shells). In addition, changes in ecosystem structure and functioning due to engineering by invasive bivalves often have very large economic impacts. Given the worldwide spread of non-indigenous bivalves and the varied ways in which they physically modify habitats, their engineering effects should receive more serious consideration in restoration and management initiatives.     

Trophic interception: how a boundary-foraging organism influences cross-ecosystem fluxes

The rate at which subsidies move between habitats is a function not only of those factors that facilitate such transfers, but also of factors that limit or restrict the movement of subsidies. The interruption or redirection of subsidies by organisms foraging at the boundary between habitats, or trophic interception, has the potential to substantially restructure the food webs of recipient habitats. In this study we describe how a waterstrider, Aquarius remigis , limits the transfer of a subsidy across the land-water boundary. Prey interception varied with the type of stream habitat; on average, waterstriders in isolated pools intercepted 71.8% of experimental prey inputs of all sizes compared with 21.5% in connected pools and 0.8% in riffles. Across all stream habitat types, waterstriders consumed 43.2% of the experimental inputs of the smallest prey used, the prey size class most similar to natural inputs in our study area. We estimate that foraging waterstriders may intercept 0.3–1.2 kg of terrestrial prey subsidy for every 100 m of stream channel during three summer months, resources that could otherwise support 13–58 young-of-the-year salmonids. In controlled trials, waterstriders significantly altered the amount of terrestrial prey directly consumed by fish, while fish also altered waterstrider interception of those prey. Interestingly, when waterstriders and fish were present together, more terrestrial prey were lost to the streambed than when either or both of these consumers were absent, making this resource available to benthic detritivores, and facilitating the direct incorporation of terrestrial nutrients into stream detrital webs. Overall, we demonstrate that organisms that forage at habitat boundaries can control the quantity and quality of subsidies arriving in recipient habitats, potentially altering food web structures within those habitats.     

Direct and indirect effects of ocean acidification and warming on a marine plant–herbivore interaction

The impacts of climatic change on organisms depend on the interaction of multiple stressors and how these may affect the interactions among species. Consumer–prey relationships may be altered by changes to the abundance of either species, or by changes to the per capita interaction strength among species. To examine the effects of multiple stressors on a species interaction, we test the direct, interactive effects of ocean warming and lowered pH on an abundant marine herbivore (the amphipod Peramphithoe parmerong), and whether this herbivore is affected indirectly by these stressors altering the palatability of its algal food (Sargassum linearifolium). Both increased temperature and lowered pH independently reduced amphipod survival and growth, with the impacts of temperature outweighing those associated with reduced pH. Amphipods were further affected indirectly by changes to the palatability of their food source. The temperature and pH conditions in which algae were grown interacted to affect algal palatability, with acidified conditions only affecting feeding rates when algae were also grown at elevated temperatures. Feeding rates were largely unaffected by the conditions faced by the herbivore while feeding. These results indicate that, in addition to the direct effects on herbivore abundance, climatic stressors will affect the strength of plant–herbivore interactions by changes to the susceptibility of plant tissues to herbivory.     

Reproductive success in varying light environments: direct and indirect effects of light on plants and pollinators

Plant populations often exist in spatially heterogeneous environments. Light level can directly affect plant reproductive success through resource availability or by altering pollinator behavior. It can also indirectly influence reproductive success by determining floral display size which may in turn influence pollinator attraction. We evaluated direct and indirect effects of light availability and measured phenotypic selection on phenological traits that may enhance pollen receipt in the insect-pollinated herb Campanulastrum americanum. In a natural population, plants in the sun had larger displays and received 7 times more visits than plants in the shade. Using experimental arrays to separate the direct effects of irradiance on insects from their response to display size, we found more visits to plants in the sun than in the shade, but no association between number of visits each flower received and display size. Plants in the sun were not pollen limited but pollen-augmented shade flowers produced 50% more seeds than open-pollinated flowers. Phenological traits, which may influence pollen receipt, were not under direct selection in the sun. However, earlier initiation and a longer duration of flowering were favored in the shade, which may enhance visitation in this pollen-limited habitat.