Shore Shapers: Introducing children and the general public to biogeomorphological processes and geodiversity

Coastal processes and wildlife shape the coast into a variety of eye-catching and enticing landforms that attract people to marvel at, relax and enjoy coastal geomorphology. These landforms also influence biological communities by providing habitat and refuge. There are very few field guides to explain these processes to the general public and children. In contrast, there is a relative wealth of resources and organised activities introducing people to coastal wildlife, especially on rocky shores. These biological resources typically focus on the biology and climatic controls on their distribution, rather than how the biology interacts with its physical habitat. As an outcome of two recent rock coast biogeomorphology projects (detailed at: www.biogeomorph.org/coastal) a multi disciplinary team produced the first known guide to understanding how biogeomorphological processes help create coastal landforms. The ‘Shore Shapers’ guide (shoreshapers.org) is designed to: a. bring biotic geomorphic interactions (how animals, algae and microorganisms protect and shape rock) to life and b. introduce some of the geomorphological and geological controls on biogeomorphic processes and landform development. The guide provides scientific information in an accessible and interactive way – to help sustain children’s interest and extend their learning. We tested a draft version of the guide with children,the general public and volunteers on rocky shore rambles using social science techniques and present the findings, alongside initial results of an evaluation of a newer version of the guide and interactive workshops taking place throughout 2014.

The combined effects of seasonal community succession and adaptive algal physiology on lipid profiles of coastal phytoplankton in the western English channel

Lipids are key constituents of marine phytoplankton, and some fatty acids (key constituents of lipids) are essential dietary components for secondary producers. However, in natural marine ecosystems the interactions of factors affecting seasonal phytoplankton lipid composition are still poorly understood. The aim of this study was to assess the roles of seasonal succession in phytoplankton community composition and nutrient concentrations, on the lipid composition of the phytoplankton community. Fatty acid and polar lipid composition in seston was measured in surface waters at the time series station L4, an inshore station in the Western English Channel, from January to December 2013. Redundancy analyses (RDA) were used to identify factors (abiotic and biotic) that explained the seasonal variability in phytoplankton lipids. RDA demonstrated that nutrients (namely nitrogen) explained the majority of variation in phytoplankton lipid composition, as well as a smaller explanatory contribution from changes in phytoplankton community composition. The physiological adaptations of the phytoplankton community to nutrient deplete conditions during the summer season when the water column was stratified, was further supported by changes in the polar lipid to phytoplankton biomass ratios (also modelled with RDA) and increases in the lipid to chlorophyll a ratios, which are both indicative of nutrient stress. However, the association of key fatty acid markers with phytoplankton groups e.g. 22:6 n-3 and dinoflagellate biomass (predominant in summer), meant there were no clear seasonal differences in the overall degree of fatty acid saturation, as might have been expected from typical nutrient stress on phytoplankton. Based on the use of polyunsaturated fatty acids (PUFA) as markers of ‘food quality’ for grazers, our results suggest that in this environment high food quality is maintained throughout summer, due to seasonal succession towards flagellated phytoplankton species able to maintain PUFA synthesis under surface layer nutrient depletion.

Effects of environment and an introduced invertebrate species on the structure of benthic macroinvertebrate species at the catchment level

Biotic interactions such as predation and competition can influence aquatic communities at small spatial scales, but they are expected to be overridden by environmental factors at large scales. The continuing threat to freshwater biodiversity of biological invasions indicates that biotic factors do, however, have important structuring roles. In Irish rivers, the native amphipod Gammarus duebeni celticus has become locally extinct, ostensibly through differential predation by the more aggressive and introduced G. pulex. This mechanism explains impacts of G. pulex at within-river spatial scales on native macroinvertebrate community diversity, including declines in ephemeropterans, plecopterans, dipterans and oligochaetes. To determine if these patterns are predictable at larger spatial scales, we assessed patterns in native macroinvertebrate communities across river sites of the Erne catchment in 1998 and 1999, in conjunction with the distribution of G. pulex and G. d. celticus. In both years, G. pulex dominated invaded sites, whereas G. d. celticus occurred at low abundance in uninvaded sites. In both years, invaded sites had lower diversity and fewer pollution sensitive invertebrate species than un-invaded sites. Community ordination in 1998 showed that invaded sites had higher conductivity, smaller substrate particle size and comprised a lower proportion of pollution sensitive taxa including Ephemeroptera and Plecoptera. In contrast, in 1999, conductivity was the only variable explaining site ordination along axis 1, but was unable to separate sites with respect to invasion status. A second explanatory axis separated sites with respect to invasion status, with invaded sites having fewer taxa, including lower abundance of ephemeropterans, dipterans and plecopterans. Laboratory experiments examined the potential role of differential predation between the two Gammarus species in explaining these taxon specific patterns in the field. Survival of the ephemeropterans, Ephemerella ignita and Ecdyonurus venosus and the isopod, Asellus aquaticus, was lower when interacting with G. pulex than with G. d. celticus. This study indicates that G. putex may alter invertebrate community structure at scales beyond those detected within individual rivers. However, effects may be influenced by gradients in physico-chemistry, which may be temporal or depend on catchment characteristics. Invasions by amphipods have increased globally, thus comprehensive assessments of their impacts and of other aquatic invaders, may only be apparent when studies are conducted at a range of spatio-temporal scales.

Coexistence among native and introduced freshwater amphipods (Crustacea); habitat utilization patterns in littoral habitats

Studies of biological invasions predominantly stress threats to biodiversity through the elimination and replacement of native species. However, we must realise that resident communities may often be capable of integrating invaders, leading to patterns of coexistence. Within the past ninety years, three freshwater amphipod species have invaded Northern Ireland the North American Gammarus tigrinus and Crangonyx pseudogracilis, plus the European G. pulex. These species have come into contact with the ubiquitous native species, G. duebeni celticus. This study examined spatiotemporal patterns of stability of single and mixed species assemblages in an invaded lake. Lough Beg and its associated rivers were surveyed in summer 1994 and winter 1995, and a selection of stations re-sampled in summer one and five years later. All possible combinations of the four amphipod species were found. Although species presence/absence was stable between seasons at the scale of the whole lough, it was extremely fluid at the scale of individual sites, 82% of which changed in species composition between seasons. Overall mean amphipod abundance was similar across 5 distinguishable habitat types, but there were differences in species compositions among these habitats. In addition, although co-occurrences of Gammarus species did not differ from random, there was a strong negative association between Gammarus spp. and C. pseudogracilis. This latter pattern was at least in part generated by the better tolerance of C. pseudogracilis to lower water quality. A review of previous studies indicates that the exclusion of C. pseudogracilis by Gammarus species from high water quality areas is likely to involve biotic interaction. Thus, overall, co-existence of the four species, which is clearly dynamic and scale-dependent, appears promoted by spatial and temporal habitat heterogeneity. However, biotic interactions may also play a role in local exclusions. Since the three introduced species have not eliminated the native species, and each successive invasion has not replaced the previous invader, this study demonstrates that freshwater invaders may integrate with native communities leading to coexistence and increased species diversity.

Factors influencing the distribution of native and introduced Gammarus spp. in Irish river systems

Populations of Gammarus duebeni celticus, previously the only amphipod species resident in the rivers of the Lough Neagh catchment, N. Ireland, have been subjected to invasion by G. pulex from the British mainland. Numerous previous studies have investigated the potential behavioural mechanisms, principally differential mutual predation, underlying the replacement of G. d. celticus by G. pulex in Irish waters, and the mutually exclusive distributions of these species in Britain and mainland Europe. However, the relative degree of influence of abiotic versus biotic factors in structuring these amphipod communities remains unresolved. This study used principal component analysis (PCA) to distinguish physico-chemical parameters that have significant roles in determining the current distribution of G. pulex relative to G. d. celticus in L. Neagh rivers. We show that the original domination of rivers by the native G. d, celticus has changed radically, with many sites in several rivers containing either both species or only G. pulex. G. pulex was more abundant than the G. d. celticus in sites with low dissolved oxygen levels. This was reflected in the macroinvertebrate assemblages associated with G. pulex in these sites, which tended to be those tolerant of low biological water quality. The present study thus emphasizes the importance of the habitat template, particularly water quality, for Gammarus spp. interactions. If rivers become increasingly stressed by organic pollution, it is probable the range expansion of G. pulex will continue. Because these two species are not ecological equivalents, the outcomes of G. pulex incursions into G. d. celticus sites may ultimately depend on the prevailing physico-chemical regimes in each site.

Role of behaviour in biological invasions and species distributions; lessons from interactions between the invasive Gammarus pulex and the native G-duebeni (Crustacea : Amphipoda)

In attempting to understand the distributions of both introduced species and the native species on which they impact, there is a growing trend to integrate studies of behaviour with more traditional life history/ecological approaches. The question of what mechanisms drive the displacement of the freshwater amphipod Gammarus duebeni by the often introduced G pulex is presented as a case study Patterns of displacement are well documented throughout Europe, but the speed and direction of displacement between these species can be varied. From early studies proposing interspecific competition as causal in these patterns, I review research progress to date. I show there has been no evidence for interspecific competition operating, other than the field patterns themselves, a somewhat tautological argument. Rather, the increased recognition of behavioural attributes with respect to the cannibalistic and predatory nature of these species gave rise to a series of studies unravelling the processes driving field patterns. Both species engage in 'intraguild predation' (IGP), with moulting females particularly vulnerable to predation by congeneric males. G pulex is more able both to engage in and avoid this interaction with G duebeni. However, several factors mediate the strength and asymmetry of this IGP, some biotic (e.g. parasitism) and others abiotic (e.g. water chemistry). Further, a number of alternative hypotheses that may account for the displacement (hybridization; parasite transmission) have been tested and rejected. While interspecific competition has been modelled mathematically and found to be a weak interaction relative to IGP, mechanisms of competition between these Gammarus species remain largely untested empirically. Since IGP may be finely balanced in some circumstances, I conclude that the challenge to detect interspecific competition remains and we require assessment of its role, if any, in the interaction between these species. Appreciation of behavioural attributes and their mediation should allow us to more fully understand, and perhaps predict, species introductions and resultant distributions.

The replacement of a native freshwater amphipod by an invader: roles for environmental degradation and intraguild predation

We assessed the extent to which an invader, Gammarus pulex (Crustacea: Amphipoda), has replaced a native, Gammarus duebeni celticus, over a 13-year period in a European river system and some of the abiotic and biotic factors that could account for this. Between 1988 and 2001, 56% of mixed-species sites had become invader-only sites, whereas no mixed sites had become native only again. The native dominated areas of higher dissolved oxygen and water quality, with the reciprocal true for the invader. Field transplant experiments revealed that native survivorship was lower in areas where it had been replaced than in areas where the invader does not yet occur. In invader-only areas, native survivorship was lower than that of the invader when kept separately and lowest when both species were kept together. We also observed predation of the native by the invader. Laboratory oxygen manipulation experiments revealed that at 30% saturation, the native's survivorship was two thirds that of the invader. We conclude that decreasing water quality favours replacement of the native by the invader.

Differential drift and parasitism in invading and native Gammarus spp. (Crustacea : Amphipoda)

Invading and native species often interact directly, such as by predation, producing patterns of exclusion and coexistence. Less direct factors, such as interactions with the broader abiotic and biotic environment, may also contribute to such patterns, but these have received less recognition. In Northern Ireland, the North American Gammarus tigrinus has invaded freshwaters populated with the native Gammarus duebeni celticus, with intraguild predation between the two implicated in their relative success. However, these species also engage in day and night

Differential physico-chemical tolerances and intraguild predation among native and invasive amphipods (Crustacea); a field study

We used field surveys and transplant experiments to elucidate the relative roles of physico-chemical regime and intraguild predation in determining the generally mutually exclusive distributions of native and invader freshwater amphipod species. Field surveys showed that the native Gammarus duebeni celticus dominates the shoreline of Lough Neagh, N. Ireland, with some co-occurrence with the N. American invader G. tigrinus. However, the latter species dominates the deeper areas of the mid-Lough. Transplant experiments showed no difference in survival of the native and invader in single species 'bioassay tubes' placed along the shoreline. However, there was significantly higher survival of the invader compared with the native in single species tubes placed in the mid-Lough. In mixed species tubes on the shoreline, the native killed and ate the invader, with no reciprocal interaction, leading to significant reductions of the invader. However, the invader had significantly higher survival than the native in mixed species tubes in the mid-Lough, with no evidence. of predation between the two species. These results indicate that, whereas differential intraguild predation may determine domination of the shoreline by the native, differential physico-chemical tolerances may be major determinants of the domination of the mid-Lough by the invader. This study emphasises the need to consider the habitat template in conjunction with biotic interactions before attempting to draw conclusions about mechanisms determining relative distribution patterns of native and invasive species.

Differential physico-chemical tolerances of amphipod species revealed by field transplantations

Physico-chemical regimes of river systems are major determinants of the distributions and relative abundances of macroinvertebrate taxa. Other factors, however, such as biotic interactions, may co-vary with changes in physico-chemistry and concomitant changes in community composition. Thus, direct cause and effect relationships may not always be established from field surveys. Equally, however, laboratory studies may suffer from lack of realism in extrapolation to the field. Here, we use balanced field transplantation experiments to elucidate the role of physico-chemical regime in determining the generally mutually exclusive distributions of two amphipod taxa, Gammarus (two species) and Crangonyx pseudogracilis. Within two river systems in Ireland, the former species dominate stretches of well oxygenated, high-quality water, whereas the latter dominates stretches of poorly oxygenated, low-quality water. G. pulex and G. duebeni celticus did not survive in bioassay tubes in areas dominated by C. pseudogracilis, which itself survived in tubes in such areas. However, both C. pseudogracilis and Gammarus spp. survived equally well in tubes in areas dominated by Gammarus spp. Physicochemical regime thus limits the movement of Gammarus spp. into C. pseudogracilis areas, but some other factor excludes C. pseudogracilis from Gammarus spp. areas. Since previous laboratory experiments showed high predation rates of Gammarus spp. on C. pseudogracilis, we propose that predation by the former causes exclusion of the latter. Hence, presumed effects of physico-chemical regime on macroinvertebrate presence/abundance may often require experimental field testing and appreciation of alternative explanations.

Limits of life in hostile environments: no barriers to biosphere function?

Environments that are hostile to life are characterized by reduced microbial activity which results in poor soil- and plant-health, low biomass and biodiversity, and feeble ecosystem development. Whereas the functional biosphere may primarily be constrained by water activity (a w) the mechanism(s) by which this occurs have not been fully elucidated. Remarkably we found that, for diverse species of xerophilic fungi at a w values of = 0.72, water activity per se did not limit cellular function. We provide evidence that chaotropic activity determined their biotic window, and obtained mycelial growth at water activities as low as 0.647 (below that recorded for any microbial species) by addition of compounds that reduced the net chaotropicity. Unexpectedly we found that some fungi grew optimally under chaotropic conditions, providing evidence for a previously uncharacterized class of extremophilic microbes. Further studies to elucidate the way in which solute activities interact to determine the limits of life may lead to enhanced biotechnological processes, and increased productivity of agricultural and natural ecosystems in arid and semiarid regions.

4 ◦C and beyond: what did this mean for biodiversity in the past?

How do the predicted climatic changes (IPCC, 2007) for the next century compare in magnitude and rate to those that Earth has previously encountered? Are there comparable intervals of rapid rates of temperature change, sea-level rise and levels of atmospheric CO2 that can be used as analogues to assess possible biotic responses to future change? Or are we stepping into the great unknown? This perspective article focuses on intervals in time in the fossil record when atmospheric CO2 concentrations increased up to 1200 ppmv, temperatures in mid- to high-latitudes increased by greater than 4 ?C within 60 years, and sea levels rose by up to 3 m higher than present. For these intervals in time, case studies of past biotic responses are presented to demonstrate the scale and impact of the magnitude and rate of such climate changes on biodiversity. We argue that although the underlying mechanisms responsible for these past changes in climate were very different (i.e. natural processes rather than anthropogenic), the rates and magnitude of climate change are similar to those predicted for the future and therefore potentially relevant to understanding future biotic response. What emerges from these past records is evidence for rapid community turnover, migrations, development of novel ecosystems and thresholds from one stable ecosystem state to another, but there is very little evidence for broad-scale extinctions due to a warming world. Based on this evidence from the fossil record, we make four recommendations for future climate-change integrated conservation strategies.

Shore exposure affects mussel population structure and mediates the effect of epibiotic algae on mussel survival in SW Ireland

On rocky shores, the relative importance of abiotic and biotic processes that regulate community structure are thought to vary with levels of shore exposure. This can lead to characteristic features found on sheltered and exposed shores. This study identified differences in the population structure of mussels on exposed and sheltered rocky shores on Atlantic coasts of south-west Ireland. Direct interactions between epibiotic algae and their host mussels were also examined to test if potential effects varied with shore exposure. Mussel beds on sheltered shores were less dense and comprised larger mussels with greater rates of individual survival and growth than those on exposed shores. The results of a field experiment showed that algal epibionts had a negative effect on mussel survival on sheltered shores but not on exposed shores. Surprisingly, the presence of algal epibionts had no effect on mussel growth on either shore type. These findings contrast with those of previous studies. The effects of shore exposure and algal epibionts on Mussels may be species-specific and may interact with other factors across different regions. This study shows that predictions of effects of exposure on mussel populations and their epibionts should only be based on specific experimental evidence and cannot be generalised across regions. (C) 2009 Elsevier Ltd. All rights reserved.

The extent of grazing release from epiphytism for Sargassum muticum (Phaeophyceae) within the invaded range

The overall biotic pressure on a newly introduced species may be less than that experienced within its native range, facilitating invasion. The brown alga Sargassum muticum (Yendo) Fensholt is a conspicuous and successful invasive species originally from Japan and China. We compared S. muticum and native macroalgae with respect to the biotic pressures of mesoherbivore grazing and ectocarpoid fouling. In Strangford Lough, Northern Ireland, S. muticum thalli were as heavily overgrown with seasonal blooms of epiphytic algae as native macroalgal species were. The herbivorous amphipod Dexamine spinosa was much more abundant on S. muticum than on any native macroalga. When cultured with this amphipod, S. muticum lost more tissue than three native macroalgae, Saccharina latissima (Linnaeus) Lane et al., Halidrys siliquosa (Linnaeus) Lyngbye and Fucus serratus Linnaeus. Sargassum muticum cultured with both ectocarpoid fouling and amphipods showed a severe impact, consistent with our previous findings of large declines in the density of S. muticum observed in the field during the peak of fouling. Despite being a recent introduction into the macroalgal community in Strangford Lough, S. muticum appears to be under biotic pressure at least equal to that on native species, suggesting that release from grazing and epiphytism does not contribute to the invasiveness of this species in Strangford Lough.

Hydrophobic substances induce water stress in microbial cells

Ubiquitous noxious hydrophobic substances, such as hydrocarbons, pesticides and diverse industrial chemicals, stress biological systems and thereby affect their ability to mediate biosphere functions like element and energy cycling vital to biosphere health. Such chemically diverse compounds may have distinct toxic activities for cellular systems; they may also share a common mechanism of stress induction mediated by their hydrophobicity. We hypothesized that the stressful effects of, and cellular adaptations to, hydrophobic stressors operate at the level of water : macromolecule interactions. Here, we present evidence that: (i) hydrocarbons reduce structural interactions within and between cellular macromolecules, (ii) organic compatible solutes-metabolites that protect against osmotic and chaotrope-induced stresses-ameliorate this effect, (iii) toxic hydrophobic substances induce a potent form of water stress in macromolecular and cellular systems, and (iv) the stress mechanism of, and cellular responses to, hydrophobic substances are remarkably similar to those associated with chaotrope-induced water stress. These findings suggest that it may be possible to devise new interventions for microbial processes in both natural environments and industrial reactors to expand microbial tolerance of hydrophobic substances, and hence the biotic windows for such processes.