Daniel W. Pritchard

CARBON-USE STRATEGIES IN MACROALGAE: DIFFERENTIAL RESPONSES TO LOWERED PH AND IMPLICATIONS FOR OCEAN ACIDIFICATION1

Ocean acidification (OA) is a reduction in oceanic pH due to increased absorption of anthropogenically produced CO2. This change alters the seawater concentrations of inorganic carbon species that are utilized by macroalgae for photosynthesis and calcification: CO2 and HCO3− increase; CO32− decreases. Two common methods of experimentally reducing seawater pH differentially alter other aspects of carbonate chemistry: the addition of CO2 gas mimics changes predicted due to OA, while the addition of HCl results in a comparatively lower [HCO3−]. We measured the short‐term photosynthetic responses of five macroalgal species with various carbon‐use strategies in one of three seawater pH treatments: pH 7.5 lowered by bubbling CO2 gas, pH 7.5 lowered by HCl, and ambient pH 7.9. There was no difference in photosynthetic rates between the CO2, HCl, or pH 7.9 treatments for any of the species examined. However, the ability of macroalgae to raise the pH of the surrounding seawater through carbon uptake was greatest in the pH 7.5 treatments. Modeling of pH change due to carbon assimilation indicated that macroalgal species that could utilize HCO3− increased their use of CO2 in the pH 7.5 treatments compared to pH 7.9 treatments. Species only capable of using CO2 did so exclusively in all treatments. Although CO2 is not likely to be limiting for photosynthesis for the macroalgal species examined, the diffusive uptake of CO2 is less energetically expensive than active HCO3− uptake, and so HCO3−‐using macroalgae may benefit in future seawater with elevated CO2.

Predicting invasive species impacts: a community module functional response approach reveals context dependencies

Summary Predatory functional responses play integral roles in predator–prey dynamics, and their assessment promises greater understanding and prediction of the predatory impacts of invasive species. Other interspecific interactions, however, such as parasitism and higher-order predation, have the potential to modify predator–prey interactions and thus the predictive capability of the comparative functional response approach. We used a four-species community module (higher-order predator; focal native or invasive predators; parasites of focal predators; native prey) to compare the predatory functional responses of native Gammarus duebeni celticus and invasive Gammarus pulex amphipods towards three invertebrate prey species (Asellus aquaticus, Simulium spp., Baetis rhodani), thus, quantifying the context dependencies of parasitism and a higher-order fish predator on these functional responses. Our functional response experiments demonstrated that the invasive amphipod had a higher predatory impact (lower handling time) on two of three prey species, which reflects patterns of impact observed in the field. The community module also revealed that parasitism had context-dependent influences, for one prey species, with the potential to further reduce the predatory impact of the invasive amphipod or increase the predatory impact of the native amphipod in the presence of a higher-order fish predator. Partial consumption of prey was similar for both predators and occurred increasingly in the order A. aquaticus, Simulium spp. and B. rhodani. This was associated with increasing prey densities, but showed no context dependencies with parasitism or higher-order fish predator. This study supports the applicability of comparative functional responses as a tool to predict and assess invasive species impacts incorporating multiple context dependencies.

Predator cue studies reveal strong trait-mediated effects in communities despite variation in experimental designs ☆

Nonconsumptive or trait-mediated effects of predators on their prey often outweigh density-mediated interactions where predators consume prey. For instance, predator presence can alter prey behaviour, physiology, morphology and/or development. Despite a burgeoning literature, our ability to identify general patterns in prey behavioural responses may be influenced by the inconsistent methodologies of predator cue experiments used to assess trait-mediated effects. We therefore conducted a meta-analysis to highlight variables (e.g. water type, predator husbandry, exposure time) that may influence invertebrate prey’s behavioural responses to fish predator cues. This revealed that changes in prey activity and refuge use were remarkably consistent overall, despite wide differences in experimental methodologies. Our meta-analysis shows that invertebrates altered their behaviour to predator cues of both fish that were fed the focal invertebrate and those that were fed other prey types, which suggests that invertebrates were not responding to specific diet information in the fish cues. Invertebrates also altered their behaviour regardless of predator cue addition regimes and fish satiation levels. Cue intensity and exposure time did not have significant effects on invertebrate behaviour. We also highlight that potentially confounding factors, such as parasitism, were rarely recorded in sufficient detail to assess the

frair: an R package for fitting and comparing consumer functional responses

Summary Consumer-resource interactions (i.e. the functional response) underpin decades of ecological advancements. However, selecting, fitting and comparing functional response models using appropriate methods remains a non-trivial endeavour. The R package frair provides tools for selecting and differentiating various forms of consumer functional response models, a consistent interface for fitting and visualising response curves, and a selection of statistically robust methods for comparing fitted parameters. Using real data from crustacean predator-prey systems, we demonstrate the utility of frair, highlighting best practice and common analytical mistakes.

Light Limitation within Southern New Zealand Kelp Forest Communities

Light is the fundamental driver of primary productivity in the marine environment. Reduced light availability has the potential to alter the distribution, community composition, and productivity of key benthic primary producers, potentially reducing habitat and energy provision to coastal food webs. We compared the underwater light environment of macroalgal dominated shallow subtidal rocky reef habitats on a coastline modified by human activities with a coastline of forested catchments. Key metrics describing the availability of photosynthetically active radiation (PAR) were determined over 295 days and were related to macroalgal depth distribution, community composition, and standing biomass patterns, which were recorded seasonally. Light attenuation was more than twice as high in shallow subtidal zones along the modified coast. Macroalgal biomass was 2–5 times greater within forested sites, and even in shallow water (2m) a significant difference in biomass was observed. Long-term light dose provided the best explanation for differences in observed biomass between modified and forested coasts, with light availability over the study period differing by 60 and 90 mol photons m−2 at 2 and 10 metres, respectively. Higher biomass on the forested coast was driven by the presence of larger individuals rather than species diversity or density. This study suggests that commonly used metrics such as species diversity and density are not as sensitive as direct measures of biomass when detecting the effects of light limitation within macroalgal communities.

Strangford Lough and the SeaGen Tidal Turbine

The background to and outcomes of the Environmental Monitoring Programme (EMP) required by statutory regulators for the deployment of the SeaGen tidal turbine in Strangford Lough, Northern Ireland, an area with many conservation designations, are described. The EMP, which was set within the context of an adaptive management approach, considered possible effects of the device on local populations of seals and harbour porpoises, representative seabirds and benthic communities. The studies on seals were carried out on both local and regional scales. The ecological studies were complemented by detailed field and hydrodynamic modelling investigations together with a programme of mitigation measures designed to reduce collisions between seals and turbine rotors. In general only minor statistically significant changes in abundance, distribution and animal behaviour patterns were recorded, principally associated with small distributional shifts close to the turbine structure and with the likelihood that these changes were ecologically of little significance. The seal–rotor collision mitigation studies provided a base for the establishment of acceptable collision risk strategies. The EMP highlighted observational, methodological and statistical challenges in assessing the environmental consequences of marine energy devices. A brief review of related studies in Strangford Lough is included.

Survival in low light: photosynthesis and growth of a red alga in relation to measured in situ irradiance

Reduced light availability for benthic primary producers as a result of anthropogenic activities may be an important driver of change in coastal seas. However, our knowledge of the minimum light requirements for benthic macroalgae limits our understanding of how these changes may affect primary productivity and the functioning of coastal ecosystems. This knowledge gap is particularly acute in deeper water, where the impacts of increased light attenuation will be most severe. We examined the minimum light requirements of Anotrichium crinitum, which dominates near the maximum depth limit for macroalgae throughout New Zealand and Southern Australia, and is a functional analog of rhodophyte macroalgae in temperate low‐light (deep‐water) habitats throughout the world. These data show that A. crinitum is a shade‐adapted seaweed with modest light requirements for the initiation of net photosynthesis (1.49–2.25 μmol photons · m−2 · s−1) and growth (0.12–0.19 mol photons · m−2 · d−1). A. crinitum maintains high photosynthetic efficiency and pigment content and a low C:N ratio throughout the year and can maintain biomass under sub‐compensation (critical) light levels for at least 5 d. Nevertheless, in situ photon flux is less than the minimum light requirement for A. crinitum on at least 103 d per annum and is rarely sufficient to saturate growth. These findings reinforce the importance of understanding the physiological response of macroalgae at the extremes of environmental gradients and highlight the need to establish minimum thresholds that modification of the subtidal light environment should not cross.

Restricted use of nitrate and a strong preference for ammonium reflects the nitrogen ecophysiology of a light‐limited red alga

Ammonium and nitrate are important sources of inorganic nitrogen for coastal primary producers. Nitrate has higher energy requirement for uptake and assimilation, compared with ammonium, suggesting that it might be a more efficient nitrogen source for slow‐growing, light‐limited macroalgae. To address this hypothesis, we examined the nitrogen ecophysiology of Anotrichium crinitum, a rhodophyte macroalgae common in low‐light habitats in New Zealand. We measured seasonal changes in seawater nitrate and ammonium concentrations and the concentration of nitrate and ammonium stored internally by A. crinitum. We determined the maximal uptake rates of nitrate and ammonium seasonally and grew A. crinitum in the laboratory with these nitrogen sources under two ecologically relevant saturating light levels. Our results show that field‐harvested A. crinitum has a high affinity for ammonium and although it will grow when supplied exclusively with nitrate, internal nitrate pools are low and it is unable to take up nitrate without several days of acclimation to saturating light. Our data predict that A. crinitum would be able to sustain growth with ammonium as the sole source of nitrogen, a strategy that would help it survive under low‐light conditions that prevail in the field.

Coupled hydrodynamic and wastewater plume models of Belfast Lough, Northern Ireland: A predictive tool for future ecological studies

Wastewater outfalls provide a natural laboratory for the study of nutrient dynamics in coastal seas, however if properly designed and operated their impact can be difficult to detect. A model was developed and applied to investigate the effect of variation in hydrodynamic conditions on the transport and dilution of a treated wastewater plume in Belfast Lough, Northern Ireland. To validate these predictions we measured the physiochemical properties of the waters surrounding the outfall with a specific focus on inherent plume tracers likely to be relevant to the study of macroalgae (salinity, nitrogen and phosphorus). The model performed well and our data show high dilution of the plume, even under neap-tide conditions. This provides a spatially and temporally defined predictive framework for future studies investigating the compliance of Northern Irelands coastal waters with European Water Framework Directive objectives and for feasibility studies investigating macroalgal aquaculture near wastewater outfalls.

Ecotones as indicators: boundary properties in wetland-woodland transition zones

Ecotones, representing the transition zones between species or communities, have been suggested as focal points for detecting early shifts in vegetation composition due to anthropogenic impact. Here we examined if changes in ecotone location or properties can be used as reliable indicators of hydrological change in temperate wetland communities. We examined 38 woodland-wetland-woodland transitions, distributed across four sites with different anthropogenic disturbance histories and hydrological traits. We tested whether: 1) the ecotones are associated with environmental gradients, and 2) the location or properties of these ecotones change with disturbance history. Well-defined ecotones were associated with steep gradients in soil depth and soil moisture. Most ecotones showed a change in vegetation from hydrophytic to dryland species. There was also some evidence that in highly disturbed sites the link between ecotones and environmental gradients was less apparent. By sampling across boundaries we can better understand the factors controlling the distribution of species. This allows us to make better predictions about the impacts of anthropogenic change in wetland communities. By investigating the transitions between different vegetation communities we were able to highlight key indicators that could be used to monitor these ecologically sensitive and diverse wetland communities.