Nuanced impacts of the invasive aquatic plant Crassula helmsii on Northwest European freshwater macroinvertebrate assemblages.

Invasive alien species are considered one of the greatest threats to global biodiversity, and are particularly problematic in aquatic systems. Given the foundational role of macrophytes in most freshwaters, alien aquatic plant invasions may drive strong bottom-up impacts on recipient biota. Crassula helmsii (New Zealand pygmyweed) is an Australasian macrophyte, now widespread in northwest Europe. Crassula helmsii rapidly invades small lentic waterbodies, where it is generally considered a serious threat to native biodiversity. The precise ecological impacts of this invasion remain poorly understood, however, particularly with respect to macroinvertebrates, which comprise the bulk of freshwater faunal biodiversity. We conducted a field study of ponds, ditches and small lakes across the core of C. helmsii's invasive range (United Kingdom, Belgium and the Netherlands), finding that invaded sites had higher macroinvertebrate taxon richness than uninvaded sites, and that many infrequent and rare macroinvertebrates co-occurred with C. helmsii. Alien macroinvertebrates were more abundant in C. helmsii sites, however, particularly the North American amphipod Crangonyx pseudogracilis. At the order level, water beetle (Coleoptera) richness and abundance were higher in C. helmsii sites, whereas true fly (Diptera) abundance was higher in uninvaded sites. Taxonomic and functional assemblage composition were both impacted by invasion, largely in relation to taxa and traits associated with detritivory, suggesting that the impacts of C. helmsii on macroinvertebrates are partly mediated by the availability and palatability of its detritus. The nuanced effects of C. helmsii on macroinvertebrates found here should encourage further quantitative research on the impacts of this invasive plant, and perhaps prompt a more balanced re-evaluation of its effects on native aquatic macrofauna.

The influence of light and temperature on detritus degradation rates for kelp species with contrasting thermal affinities.

Kelp detritus fuels coastal food webs and may play an important role as a source of organic matter for natural carbon sequestration. Here, we conducted ex situ and in situ manipulations to evaluate the role of temperature and light availability in the breakdown of detrital material. We examined degradation rates of two North Atlantic species with contrasting thermal affinities: the 'warm water' kelp Laminaria ochroleuca and the 'cool water' Laminaria hyperborea. Detrital fragments were exposed to different temperatures in controlled conditions and across an in situ gradient of depth, corresponding to light availability. Overall, degradation rates (i.e. changes in Fv/Fm and biomass) were faster under lower light conditions and at higher temperatures, although responses were highly variable between plants and fragments. Crucially, as L. ochroleuca degraded faster than L. hyperborea under some conditions, a climate-driven substitution of the 'cool' for the 'warm' kelp, which has been observed at some locations, will likely increase detritus turnover rates and alter detrital pathways in certain environments. More importantly, ocean warming combined with decreased coastal water quality will likely accelerate kelp detritus decomposition, with potential implications for coastal food webs and carbon cycles.

Tropical CO2 seeps reveal the impact of ocean acidification on coral reef invertebrate recruitment.

Rising atmospheric CO2 concentrations are causing ocean acidification by reducing seawater pH and carbonate saturation levels. Laboratory studies have demonstrated that many larval and juvenile marine invertebrates are vulnerable to these changes in surface ocean chemistry, but challenges remain in predicting effects at community and ecosystem levels. We investigated the effect of ocean acidification on invertebrate recruitment at two coral reef CO2 seeps in Papua New Guinea. Invertebrate communities differed significantly between 'reference' (median pH7.97, 8.00), 'high CO2' (median pH7.77, 7.79), and 'extreme CO2' (median pH7.32, 7.68) conditions at each reef. There were also significant reductions in calcifying taxa, copepods and amphipods as CO2 levels increased. The observed shifts in recruitment were comparable to those previously described in the Mediterranean, revealing an ecological mechanism by which shallow coastal systems are affected by near-future levels of ocean acidification.

The effects of warming on the ecophysiology of two co-existing kelp species with contrasting distributions.

The northeast Atlantic has warmed significantly since the early 1980s, leading to shifts in species distributions and changes in the structure and functioning of communities and ecosystems. This study investigated the effects of increased temperature on two co-existing habitat-forming kelps: Laminaria digitata, a northern boreal species, and Laminaria ochroleuca, a southern Lusitanian species, to shed light on mechanisms underpinning responses of trailing and leading edge populations to warming. Kelp sporophytes collected from southwest United Kingdom were maintained under 3 treatments: ambient temperature (12 °C), +3 °C (15 °C) and +6 °C (18 °C) for 16 days. At higher temperatures, L. digitata showed a decline in growth rates and Fv/Fm, an increase in chemical defence production and a decrease in palatability. In contrast, L. ochroleuca demonstrated superior growth and photosynthesis at temperatures higher than current ambient levels, and was more heavily grazed. Whilst the observed decreased palatability of L. digitata held at higher temperatures could reduce top-down pressure on marginal populations, field observations of grazer densities suggest that this may be unimportant within the study system. Overall, our study suggests that shifts in trailing edge populations will be primarily driven by ecophysiological responses to high temperatures experienced during current and predicted thermal maxima, and although compensatory mechanisms may reduce top-down pressure on marginal populations, this is unlikely to be important within the current biogeographical context. Better understanding of the mechanisms underpinning climate-driven range shifts is important for habitat-forming species like kelps, which provide organic matter, create biogenic structure and alter environmental conditions for associated communities.

Ocean acidification affects fish spawning but not paternity at CO2 seeps.

Fish exhibit impaired sensory function and altered behaviour at levels of ocean acidification expected to occur owing to anthropogenic carbon dioxide emissions during this century. We provide the first evidence of the effects of ocean acidification on reproductive behaviour of fish in the wild. Satellite and sneaker male ocellated wrasse (Symphodus ocellatus) compete to fertilize eggs guarded by dominant nesting males. Key mating behaviours such as dominant male courtship and nest defence did not differ between sites with ambient versus elevated CO2 concentrations. Dominant males did, however, experience significantly lower rates of pair spawning at elevated CO2 levels. Despite the higher risk of sperm competition found at elevated CO2, we also found a trend of lower satellite and sneaker male paternity at elevated CO2 Given the importance of fish for food security and ecosystem stability, this study highlights the need for targeted research into the effects of rising CO2 levels on patterns of reproduction in wild fish.

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.

The future of the northeast Atlantic benthic flora in a high CO2 world.

Seaweed and seagrass communities in the northeast Atlantic have been profoundly impacted by humans, and the rate of change is accelerating rapidly due to runaway CO2 emissions and mounting pressures on coastlines associated with human population growth and increased consumption of finite resources. Here, we predict how rapid warming and acidification are likely to affect benthic flora and coastal ecosystems of the northeast Atlantic in this century, based on global evidence from the literature as interpreted by the collective knowledge of the authorship. We predict that warming will kill off kelp forests in the south and that ocean acidification will remove maerl habitat in the north. Seagrasses will proliferate, and associated epiphytes switch from calcified algae to diatoms and filamentous species. Invasive species will thrive in niches liberated by loss of native species and spread via exponential development of artificial marine structures. Combined impacts of seawater warming, ocean acidification, and increased storminess may replace structurally diverse seaweed canopies, with associated calcified and noncalcified flora, with simple habitats dominated by noncalcified, turf-forming seaweeds.

Using species-area relationships to inform baseline conservation targets for the deep North East Atlantic.

Demands on the resources of the deep-sea have increased in recent years. Consequently, the need to create and implement a comprehensive network of Marine Protected Areas (MPAs) to help manage and protect these resources has become a global political priority. Efforts are currently underway to implement MPA networks in the deep North East Atlantic. To ensure these networks are effective, it is essential that baseline information be available to inform the conservation planning process. Using empirical data, we calculated conservation targets for sessile benthic invertebrates in the deep North East Atlantic for consideration during the planning process. We assessed Species-Area Relationships across two depth bands (200-1100 m and 1100-1800 m) and nine substrata. Conservation targets were predicted for each substratum within each depth band using z-values obtained from fitting a power model to the Species-Area Relationships of observed and estimated species richness (Chao1). Results suggest an MPA network incorporating 10% of the North East Atlantic's deep-sea area would protect approximately 58% and 49% of sessile benthic species for the depth bands 200-1100 m and 1100-1800 m, respectively. Species richness was shown to vary with substratum type indicating that, along with depth, substratum information needs to be incorporated into the conservation planning process to ensure the most effective MPA network is implemented in the deep North East Atlantic.

Tri-trophic consequences of UV-B exposure: plants, herbivores and parasitoids.

In this paper we demonstrate a UV-B-mediated link between host plants, herbivores and their parasitoids, using a model system consisting of a host plant Brassica oleracea, a herbivore Plutella xylostella and its parasitoid Cotesia plutellae. Ultraviolet-B radiation (UV-B) is a potent elicitor of a variety of changes in the chemistry, morphology and physiology of plants and animals. Recent studies have demonstrated that common signals, such as jasmonic acid (JA), play important roles in the mechanisms by which plants respond to UV-B and to damage by herbivores. Plant responses elicited by UV-B radiation can affect the choices of ovipositing female insects and the fitness of their offspring. This leads to the prediction that, in plants, the changes induced as a consequence of UV damage will be similar to those elicited in response to insect damage, including knock-on effects upon the next trophic level, predators. In our trials female P. xylostella oviposited preferentially on host plants grown in depleted UV-B conditions, while their larvae preferred to feed on tissues from UV-depleted regimes over those from UV-supplemented ones. Larval feeding patterns on UV-supplemented tissues met the predictions of models which propose that induced defences in plants should disperse herbivory; feeding scars were significantly smaller and more numerous--though not significantly so--than those on host plant leaves grown in UV-depleted conditions. Most importantly, female parasitoids also showed a clear pattern of preference when given the choice between host plants and attendant larvae from the different UV regimes; however, in the case of the female parasitoids, the choice was in favour of potential hosts foraging on UV-supplemented tissues. This study demonstrates the potential for UV-B to elicit a variety of interactions between trophic levels, most likely mediated through effects upon host plant chemistry.

Do developmental mode and dispersal shape abundance-occupancy relationships in marine macroinvertebrates?

1. Dispersal is a crucial process in maintaining population structures in many organisms, and is hypothesized as a process underlying the interspecific relationship between abundance and distribution. Here we examined whether there was a link between the dispersal and developmental modes of marine macroinvertebrates and the slopes and elevations of interspecific abundance-occupancy relationships. We predicted that if within-site retention of larvae ranks in the order brooders > lecithotrophs > planktotrophs, for any given level of mean abundance, occupancy should increase in the order brooders < lecithotrophs < planktotrophs. We also predicted that propensity to form metapopulations should be greater for planktonic dispersers (i.e. lecithotrophs and planktotrophs combined) than for non-planktonic (i.e. brooders), resulting in steeper abundance-occupancy relationships for the former. 2. Predictions were tested using a data set for 362 subtidal marine macroinvertebrates occurring across 446 1-km(2) grid squares around the British Isles; analyses were performed on the data set as a whole and for separate phyla. 3. The total data set had a Z-transformed effect size of 0.79, within the confidence intervals described by Blackburn et al. (2006; Journal of Animal Ecology, 75, 1426-1439), and was consistently present with relatively homogeneous effect size in separate analyses of polychaetes, crustaceans, molluscs and echinoderms. 4. In all cases, planktonic dispersing organisms showed an abundance-occupancy relationship with greater elevation than that for non-planktonic organisms; in polychaetes the elevation of slopes was in the rank order planktotrophs > lecithotrophs > brooders. No differences between the slopes of the abundance-occupancy relationship were apparent for different dispersal modes either within, or across phyla. 5. We conclude that dispersal capacity may play an important part in determining the elevation of the abundance-occupancy relationship, the corollary of low dispersal in the marine realm being greater local retention of larvae and greater local population abundance at low extents of geographical distribution.

Herbivore-induced infochemicals influence foraging behaviour in two intertidal predators.

Herbivore-induced defences appear ubiquitous across most biomes and habitats. Yet the direct correlation between induced changes in host plant chemistry and the population dynamics of the herbivore remain untested in many systems. In plant-herbivore interactions in the terrestrial environment, indirect or tritrophic interactions appear a successful way in which changes in the host plant chemistry induced by prior herbivory can impact on herbivore populations via increased success of natural enemies. This set of interactions remains untested in the marine system. Here, we present work from experiments using orthogonal contrasts of plants with different prior treatments (control, mechanical damage or herbivory) and the presence or absence of herbivores on the foraging behaviour of a crab, Carcinus maenas, and a fish, Lipophrys pholis. These experiments were carried out using a novel flow-through flume, i.e. as a choice chamber supplied by turbulent water from independent cue sources. Our results show that in the Ascophyllum nodosum (plant)-Littorina obtusata (herbivore) system infochemicals from induced plants can directly influence predator foraging behaviour. L. pholis was attracted to the presence of a feeding L. obtusata, but was also more attracted to odours from herbivore-induced tissue than odours from mechanically damaged or naïve A. nodosum. C. maenas was more attracted to odours from herbivore-induced tissue compared to naïve tissue, regardless of the presence of L. obtusata. This is the first demonstration of such behavioural consequences of herbivore-induced changes in plants for marine systems.

Soil microbial and faunal community responses to bt maize and insecticide in two soils.

The effects of maize (Zea mays L.), genetically modified to express the Cry1Ab protein (Bt), and an insecticide on soil microbial and faunal communities were assessed in a glasshouse experiment. Soil for the experiment was taken from field sites where the same maize cultivars were grown to allow comparison between results under glasshouse conditions with those from field trials. Plants were grown in contrasting sandy loam and clay loam soils, half were sprayed with a pyrethroid insecticide (deltamethrin) and soil samples taken at the five-leaf stage, flowering, and maturity. The main effect on all measured parameters was that of soil type and there were no effects of Bt trait or insecticide on plant growth. The Bt trait resulted in more soil nematodes and protozoa (amoebae), whereas insecticide application increased plant Bt concentration and altered nematode community structure. The only significant effects on soil microbial community structure, microarthropods, and larvae of a nontarget root-feeding Dipteran, were due to soil type and plant growth stage. The results indicate that, although there were statistically significant effects of the Bt trait on soil populations, they were small. The relative magnitude of the effect could best be judged by comparison with the insecticide treatment, which was representative of current best practice. The Bt trait had no greater effect than the insecticide treatment. Results from this glasshouse experiment were in broad agreement with conclusions from field experiments using the same plant material grown in the same soils.

Induced resistance in intertidal macroalgae modifies feeding behaviour of herbivorous snails.

Herbivory in terrestrial and marine systems can induce changes in plant chemistry affecting the foraging behaviour of herbivores. A model based on terrestrial plant-herbivore interactions predicts herbivory-induced changes in leaf chemistry to be manifested in (1) increased herbivore mobility, (2) increased feeding dispersal and (3) reduced tissue consumption by herbivores. This study is the first to demonstrate that herbivory-induced changes in the tissue chemistry of the brown seaweed Ascophyllum nodosum elicit the same response in the feeding behaviour of the gastropod Littorina obtusata as predicted for herbivorous insects, providing good evidence for the model's validity across different ecosystems. The potential benefit of increased feeding dispersal to terrestrial plants as suggested by the model is the prevention of concentrated damage to apical tissues thereby preserving the plant's ability to compete for light; A. nodosum does not conform to these predictions. Increased dispersal of feeding damage on A. nodosum away from primary frond tissues would reduce the likelihood of frond breakage implying a fitness benefit of induced resistance.

Root damage and water stress: treatments affecting the exploitation of the buds of common ash Fraxinus excelsior L., by larvae of the ash bud moth Prays fraxinella Bjerk. (Lep., Yponomeutidae).

Damage to the buds of the common ash (Fraxinus excelsior L.) by larvae of the ash bud moth Prays fraxinella Bjerk. was investigated in trees growing in the presence, or absence, of an adjacent ditch. The presence of a ditch was correlated with increased damage levels due to bud moth. Saplings were used in an experiment to compare the effects of artificial drought and root damage with those of ditches. Both root damage and drought elicited analogous responses to those observed in trees growing near a ditch, being associated with elevated levels of bud moth exploitation and reduced growth of shoots. There exists a documented link between ditches, root disturbance, and the dieback of hedgerow ash trees in Great Britain; the response of P. fraxinella to environmental disturbances associated with dieback is hypothesised to be evidence of the possible role of moth damage as a causal factor in dieback.