Implications for the conservation of deep-water corals in the face of multiple stressors: A case study from the New Zealand region.

The waters around New Zealand are a global hotspot of biodiversity for deep-water corals; approximately one sixth of the known deep-water coral species of the world have been recorded in the region. Deep-water corals are vulnerable to climate-related stressors and from the damaging effects of commercial fisheries. Current protection measures do not account for the vulnerability of deep-water corals to future climatic conditions, which are predicted to alter the distribution of suitable habitat for them. Using recently developed habitat suitability models for 12 taxa of deep-water corals fitted to current and future seafloor environmental conditions (under different future climatic conditions: SSP2 - 4.5 and SSP3 - 7.0) we explore possible levels of spatial protection using the decision-support tool Zonation. Specifically, we assess the impact of bottom trawling on predictions of current distributions of deep-water corals, and then assess the effectiveness of possible protection for deep-water corals, while accounting for habitat refugia under future climatic conditions. The cumulative impact of bottom trawling was predicted to impact all taxa, but particularly the reef-forming corals. Core areas of suitable habitat were predicted to decrease under future climatic conditions for many taxa. We found that designing protection using current day predictions alone, having accounted for the impacts of historic fishing impacts, was unlikely to provide adequate conservation for deep water-corals under future climate change. Accounting for future distributions in spatial planning identified areas which may provide climate refugia whilst still providing efficient protection for current distributions. These gains in conservation value may be particularly important given the predicted reduction in suitable habitat for deep-water corals due to bottom fishing and climate change. Finally, the possible impact that protection measures may have on deep-water fisheries was assessed using a measure of current fishing value (kg km-2 fish) and future fishing value (predicted under future climate change scenarios).

Three decades of increasing fish biodiversity across the northeast Atlantic and the Arctic Ocean.

Observed range shifts of numerous species support predictions of climate change models that species will shift their distribution northward into the Arctic and sub-Arctic seas due to ocean warming. However, how this is affecting overall species richness is unclear. Here we analyze 20,670 scientific research trawls from the North Sea to the Arctic Ocean collected from 1994 to 2020, including 193 fish species. We found that demersal fish species richness at the local scale has doubled in some Arctic regions, including the Barents Sea, and increased at a lower rate at adjacent regions in the last three decades, followed by an increase in species richness and turnover at a regional scale. These changes in biodiversity correlated with an increase in sea bottom temperature. Within the study area, Arctic species' probability of occurrence generally declined over time. However, the increase in species from southern latitudes, together with an increase in some Arctic species, ultimately led to an enrichment of the Arctic and sub-Arctic marine fauna due to increasing water temperature consistent with climate change.

Predicting the effects of climate change on deep-water coral distribution around New Zealand-Will there be suitable refuges for protection at the end of the 21st century?

Deep-water corals are protected in the seas around New Zealand by legislation that prohibits intentional damage and removal, and by marine protected areas where bottom trawling is prohibited. However, these measures do not protect them from the impacts of a changing climate and ocean acidification. To enable adequate future protection from these threats we require knowledge of the present distribution of corals and the environmental conditions that determine their preferred habitat, as well as the likely future changes in these conditions, so that we can identify areas for potential refugia. In this study, we built habitat suitability models for 12 taxa of deep-water corals using a comprehensive set of sample data and predicted present and future seafloor environmental conditions from an earth system model specifically tailored for the South Pacific. These models predicted that for most taxa there will be substantial shifts in the location of the most suitable habitat and decreases in the area of such habitat by the end of the 21st century, driven primarily by decreases in seafloor oxygen concentrations, shoaling of aragonite and calcite saturation horizons, and increases in nitrogen concentrations. The current network of protected areas in the region appear to provide little protection for most coral taxa, as there is little overlap with areas of highest habitat suitability, either in the present or the future. We recommend an urgent re-examination of the spatial distribution of protected areas for deep-water corals in the region, utilising spatial planning software that can balance protection requirements against value from fishing and mineral resources, take into account the current status of the coral habitats after decades of bottom trawling, and consider connectivity pathways for colonisation of corals into potential refugia.

The impact of cumulative stressor effects on uncertainty and ecological risk.

To enable environmental management actions to be more effectively prioritized, cumulative effects between multiple stressors need to be accounted for in risk-assessment frameworks. Ecological risk and uncertainty are generally high when multiple stressors occur. In the face of high uncertainty, transparent communication is essential to inform decision-making. The impact of stressor interactions on risk and uncertainty was assessed using generalized linear models for additive and multiplicative effect of six anthropogenic stressors on the abundance of estuarine macrofauna across New Zealand. Models that accounted for multiplicative stressor interactions demonstrated that non-additive effects dominated, had increased explanatory power (6 to 73 % relative increase between models), and thereby reduced the risk of unexpected ecological responses to stress. Secondly, 3D-plots provide important insights in the direction, magnitude and gradients of change, and aid transparency and communication of complex stressor effects. Notably, small changes in a stressor can cause a disproportionally steep gradient of change for a synergistic effect where the tolerance to stressors are lost, and would invoke precautionary management. 3D-plots were able to clearly identify directional shifts where the nature of the interaction changed from antagonistic to synergistic along increasing stressor gradients. For example, increased nitrogen load and exposure caused a shift from positive to negative effect on the abundance of a deposit-feeding polychaete (Magelona). Assessments relying on model coefficient estimates, which provide one effect term, could not capture the complexities observed in 3D-plots and are at risk of mis-identifying interaction types. Finally, visualising model uncertainty demonstrated that although error terms were higher for multiplicative models, they better captured the uncertainty caused by data availability. Together, the steep gradients of change identified in 3D-plots and the higher uncertainty in model predictions in multiplicative models urges more conservative limits to be set for management that account for risk and uncertainty in multiple stressor effects.

Who is contributing where? Predicting ecosystem service multifunctionality for shellfish species through ecological principles.

A key challenge in environmental management is determining how to manage multiple ecosystem services (ES) simultaneously, to ensure efficient and sustainable use of the environment and its resources. In marine environments, the spatial assessment of ES is lagging as a result of data-scarcity and modelling complexity. Applying mechanistic models to link ecological processes with ecosystem functions and services to assess areas of high ES potential can bridge this gap and accommodate assessments of functional differences between service providers. Here, we applied an ecosystem principles approach to assess ES potential for food provision, water quality regulation, nitrogen removal, and sediment stabilisation, provided by two estuarine bivalves (Austrovenus stutchburyi and Paphies australis) that differ in habitat association (broad and narrow distributions), to gain insight into the utility of these models for local-scale management. Maps of individual ES displayed differing patterns related to habitat associations of the species providing them, with variation in the quantities of services being delivered and locations of importance. Areas of importance for the provision of multiple services (number of services provided and their combined intensity per species) were assessed using hotspot analyses, which suggested that areas of high shellfish density at the harbour entrances were important for ES multifunctionality. A targeted management approach that includes environmental context, rather than a focus solely on the protection of high-density shellfish areas, is required to sustain the provision of individual ES.

Sampling frequency, duration and the Southern Oscillation influence the ability of long-term studies to detect sudden change.

Ecologists have long acknowledged the importance of context dependency related to position along spatial gradients. It is also acknowledged that broad-scale climate patterns can directly and indirectly alter population dynamics. What is not often addressed is whether climate patterns such as the Southern Oscillation interact with population-level temporal patterns and affect the ability of time-series data, such as long-term state of the environment monitoring programmes, to detect change. Monitoring design criteria generally focus on number of data points, sampling frequency and duration, often derived from previous information on species seasonal and multi-year temporal patterns. Our study questioned whether the timing of any changes relative to Southern Oscillation, interacting with species populations dynamics, would also be important. We imposed a series of simulated reductions on macrofaunal abundance data collected regularly over 29 years from two sites, using species selected for observed differences in temporal dynamics. We hypothesized that (1) high within-year sampling frequency would increase detection ability for species with repeatable seasonality cycles and (2) timing of the reduction in abundance relative to the Southern Oscillation was only likely to affect detection ability for long-lived species with multi-year cyclic patterns in abundance. However, regardless of species population dynamics, we found both within-year sampling frequency and the timing of the imposed reduction relative to the Southern Oscillation Index affected detection ability. The latter result, while apparently demonstrating a confounding influence on monitoring, offers the opportunity to improve our ability to detect and interpret analyses of monitoring data, and thus our ability to make recommendations to managers.

Increasing climate-driven taxonomic homogenization but functional differentiation among river macroinvertebrate assemblages.

Global change is increasing biotic homogenization globally, which modifies the functioning of ecosystems. While tendencies towards taxonomic homogenization in biological communities have been extensively studied, functional homogenization remains an understudied facet of biodiversity. Here, we tested four hypotheses related to long-term changes (1991-2016) in the taxonomic and functional arrangement of freshwater macroinvertebrate assemblages across space and possible drivers of these changes. Using data collected annually at 64 river sites in mainland New Zealand, we related temporal changes in taxonomic and functional spatial ß-diversity, and the contribution of individual sites to ß-diversity, to a set of global, regional, catchment and reach-scale environmental descriptors. We observed long-term, mostly climate-induced, temporal trends towards taxonomic homogenization but functional differentiation among macroinvertebrate assemblages. These changes were mainly driven by replacements of species and functional traits among assemblages, rather than nested species loss. In addition, there was no difference between the mean rate of change in the taxonomic and functional facets of ß-diversity. Climatic processes governed overall population and community changes in these freshwater ecosystems, but were amplified by multiple anthropogenic, topographic and biotic drivers of environmental change, acting widely across the landscape. The functional diversification of communities could potentially provide communities with greater stability, resistance and resilience capacity to environmental change, despite ongoing taxonomic homogenization. Therefore, our study highlights a need to further understand temporal trajectories in both taxonomic and functional components of species communities, which could enable a clearer picture of how biodiversity and ecosystems will respond to future global changes.

Mapping the estuarine ecosystem service of pollutant removal using empirically validated boosted regression tree models.

Humans rely on the natural environment and benefit from the goods and services provided by natural ecosystems. Quantification and mapping of ecosystem services (ES) is required to better protect valued ES benefits under pressure from anthropogenic activities. The removal of excess nitrogen, a recognized catchment-derived pollutant, by biota in estuarine soft sediments is an important ES that potentially ameliorates the development of eutrophication symptoms. Here, we quantified estuarine benthic sediment characteristics and denitrification enzyme activity (DEA), a proxy of inorganic N removal, at 109 sites in four estuaries to develop a general ("global") model for predicting DEA. Our initial global model for linking DEA and environmental characteristics had good explanatory power, with sediment mud content having the strongest influence on DEA (60%), followed by sediment organic matter content (≈35%) and sediment chlorophyll a content (≈5%). Predicted and empirically evaluated DEA values in a fifth estuary (Whitford, n = 90 validation sites) were positively correlated (r = 0.77), and the fit and certainty of the model (based on two types of uncertainty measures) increased further after the validation sites were incorporated into it. The model tended to underpredict DEA at the upper end of its range (at the muddier, more organically enriched sites), and the relative roles of the three environmental predictors differed in Whitford relative to the four previously sampled estuaries (reducing the explained deviance relative to the initial global model). Our detailed quantification of DEA and methodological description for producing empirically validated maps, complete with uncertainty information, represents an important first step in the construction of nutrient pollution removal ES maps for use in coastal marine spatial management. This technique can likely be adapted to map other ecosystem functions and ES proxies worldwide.

Investigating changes in estuarine ecosystem functioning under future scenarios.

Estuaries are subject to disturbance by land-based sediment and nutrient inputs, resulting in changes to the ecosystems and the functions and services that they support. Spatial mapping tools that identify how functional hotspots in the estuary may shift in location and intensity under different disturbance scenarios highlight to managers the trajectory of change and the value of active management and restoration, but to date these tools are only available in the most intensively researched ecosystems. Using empirical data derived from long-term monitoring and multi-habitat field experiments we developed future scenarios representing different impacts of environmental degradation on estuarine ecosystem functions that are important for supporting ecosystem services. We used the spatial prioritization software Zonation in a novel fashion to assess effects of different disturbance scenarios on critical soft-sediment ecosystem processes (nutrient fluxes and sediment erodibility measures) that are influenced by macrofaunal communities and local environment conditions. We compared estimates of current conditions with three scenarios linked to changes in land-use and resulting downstream impacts on estuarine ecosystems to determine how disturbance influences the distribution of high value areas for ecosystem function. Scenarios investigated the implications of habitat degradation associated with sediment deposition and declines in large sediment-dwelling animal abundance whose behavior has important influences on ecosystem function. Our analyses demonstrate decreases in the majority of ecosystem processes under scenarios associated with disturbances. These results suggest that it is important to restore biodiversity and ecosystem function and that the application of Zonation in this context offers a simple, rapid and cost-effective way of identifying priority actions and locations for restoration, and how these shift due to multiple impacts.

Linking Traits across Ecological Scales Determines Functional Resilience.

Under globally accelerating rates of ecosystem degradation, maintaining ecosystem function is a priority to avoid loss of valuable ecosystem services. Two factors are important: changes to the disturbance regime (stresses imposed) and resilience of biodiversity and ecosystem functions (the ecosystem's capacity to respond to change). Various attributes at different scales of ecological organisation confer resilience (from individual species to communities at landscape scales), but it is critical to understand how these attributes interact to inform how ecosystem function changes with disturbances that vary in intensity, spatial extent, and frequency. Individual species attributes influence their resistance, while attributes at the landscape-scale influence recovery of communities and function. Understanding resilience to disturbances requires defining the characteristics of a resilient community at multiple scales.

Environmental filtering of native and non-native stream macrophyte assemblages by habitat disturbances in an agricultural landscape.

Understanding how inter-specific variation in functional traits affects native and non-native species responses to stream disturbances, is necessary to inform management strategies, providing tools for biomonitoring, conservation and restoration. This study used a functional trait approach to characterise the responses of macrophyte assemblages to reach-scale disturbances (measured by lack of riparian shading, altered hydromorphology and eutrophication), from 97 wadeable stream sites in an agriculturally impacted region of New Zealand. To determine whether macrophyte assemblages differed due to disturbances, we examined multidimensional assemblage functional structure in relation to eleven functional traits and further related two functional diversity indices (entropy and originality) to disturbances. Macrophyte assemblages showed distinct patterns in response to disturbances, with riparian shading and hydromorphological conditions being the strongest variables shaping macrophyte functional structure. In the multidimensional space, most of the non-native species were associated with disturbed conditions. These species had traits allowing faster colonisation rates (higher number of reproductive organs and larger root-rhizome system) and superior competitive abilities for resources (tall and dense canopy, heterophylly and greater preferences for light and nitrogen). In addition, lack of riparian shading increased the abundance of functionally distinct species (i.e. entropy), and eutrophication resulted in the growth of functionally unique species (i.e. originality). We demonstrated that stream reach-scale habitat disturbances were associated to a dominance of more productive species, equating to a greater abundance of non-native species. This, can result in a displacement of native species, habitat alterations, and changes to higher trophic level assemblages. Our results suggests that reach-scale management efforts such as the conservation and restoration of riparian vegetation that provides substantial shading and hydromorphologically diverse in-stream habitat, would have beneficial direct and indirect effects on ecosystem functioning, and contribute to the mitigation of land-use impacts.