Ecological network analysis of traits reveals variable response capacity to stress.

Response diversity increases the potential 'options' for ecological communities to respond to stress (i.e. response capacity). An indicator of community response diversity is the diversity of different traits associated with their capacity to be resistant to stress, to recover and to regulate ecosystem functions. We conducted a network analysis of traits using benthic macroinvertebrate community data from a large-scale field experiment to explore the loss of response diversity along environmental gradients. We elevated sediment nutrient concentrations (a process that occurs with eutrophication) at 24 sites (in 15 estuaries) with varying environmental conditions (water column turbidity and sediment properties). Macroinvertebrate community response capacity to nutrient stress was dependent on the baseline trait network complexity in the ambient community (i.e. non-enriched sediments). The greater the complexity of the baseline network, the less variable the network response to nutrient stress was; in contrast, more variable responses to nutrient stress occurred with simpler networks. Thus, stressors or environmental variables that shift baseline network complexity also shift the capacity for these ecosystems to respond to additional stressors. Empirical studies that explore the mechanisms responsible for loss of resilience are essential to inform our ability to predict changes in ecological states.

Toward a network perspective in coastal ecosystem management.

Environmental management in coastal ecosystems has been challenged by the complex cumulative effects that occur when many small issues result in large ecological shifts. Current environmental management of these spaces focuses on identifying and limiting problematic stressors via a series of assessment techniques. Whilst there is a strong desire among managers to consider complexity in ecological responses to cumulative effects, current approaches for assessing risk focus on breaking down the issues into multiple cause and effect relationships. However, uncertainty arises when data and information for a place are limited, as is commonly the case, and this creates decision paralysis while more information is generated. Here, we discuss how ecological understanding of network interactions in coastal marine ecosystems can be used as a lens to bring together multiple lines of evidence and create actions. We list and describe four characteristics of marine ecosystem interaction networks including the possibility for; 1) indirect effects, 2) effects that emerge as stressor magnitude increases the number of network components implicated, 3) network interactions that amplify these indirect effects, and 4) feedbacks that reinforce or stabilise against indirect effects. We then link these four characteristics to three case studies of common coastal environmental issues to demonstrate how a general understanding of ecological interaction networks can enhance priorities for stressor management that can be applied even when specific data is limited.

The influence of mussel restoration on coastal carbon cycling.

Increasing responsiveness to anthropogenic climate change and the loss of global shellfish ecosystems has heightened interest in the carbon storage and sequestration potential of bivalve-dominated systems. While coastal ecosystems are dynamic zones of carbon transformation and change, current uncertainties and notable heterogeneity in the benthic environment make it difficult to ascertain the climate change mitigation capacity of ongoing coastal restoration projects aimed at revitalizing benthic bivalve populations. In this study we sought to distinguish between direct and indirect effects of subtidal green-lipped mussels (Perna canaliculus) on carbon cycling, and combined published literature with in-situ experiments from restored beds to create a carbon budget for New Zealand's shellfish restoration efforts. A direct summation of biogenic calcification, community respiration, and sediment processes suggests a moderate carbon efflux (+100.1 to 179.6 g C m-2  year-1 ) occurs as a result of recent restoration efforts, largely reflective of the heterotrophic nature of bivalves. However, an examination of indirect effects of restoration on benthic community metabolism and sediment dynamics suggests that beds achieve greater carbon fixation rates and support enhanced carbon burial compared to nearby sediments devoid of mussels. We discuss limitations to our first-order approximation and postulate how the significance of mussel restoration to carbon-related outcomes likely increases over longer timescales. Coastal restoration is often conducted to support the provisioning of many ecosystem services, and we propose here that shellfish restoration not be used as a single measure to offset carbon dioxide emissions, but rather used in tandem with other initiatives to recover a bundle of valued ecosystem services.

Climate cascades affect coastal Antarctic seafloor ecosystem functioning.

Polar seafloor ecosystems are changing rapidly and dramatically, challenging previously held paradigms of extreme dynamical stability. Warming-related declines in polar sea ice are expected to alter fluxes of phytoplankton and under-ice algae to the seafloor. Yet, how changes in food flux cascade through to seafloor communities and functions remains unclear. We leveraged natural spatial and temporal gradients in summertime sea ice extent to better understand the trajectories and implications of climate-related change in McMurdo Sound, Antarctica. McMurdo Sound was expected to be one of the last coastal marine environments on Earth to be affected by planetary warming, but the situation may be changing. Comparing satellite observations of selected coastal sites in McMurdo Sound between 2010-2017 and 2002-2009 revealed more ice-free days per year, and shorter distances to open water during the warmest months each year, in the more recent period. Interdecadal Pacific Oscillation (IPO), Oceanic Niño Index (ONI) and Antarctic Oscillation (AAO) climate indices peaked concurrently between 2014 and 2017 when sea ice breakouts in McMurdo Sound were most spatially and temporally extensive. Increases in sediment chlorophyll a and phaeophytin content (indicating increased deposition of detrital algal food material) were recorded during 2014-2017 at three coastal study sites in McMurdo Sound following the major sea ice breakouts. Soft-sediment seafloor ecosystem metabolism (measured in benthic incubation chambers as dissolved oxygen and inorganic nutrient fluxes) was correlated with sediment algal pigment concentration. Epifaunal invertebrate density, particularly opportunistic sessile suspension feeders, and infaunal community composition also shifted with increased food supply. The ecological characteristics and functions measured at the food-poor sites shifted towards those observed at richer sites at a surprisingly fast pace. These results indicate the sensitivity of the benthos and shed light on Antarctic marine trophic cascades and trajectories of response of iconic high-latitude seafloor habitats to a warming climate.

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.

Cumulative stressors reduce the self-regulating capacity of coastal ecosystems.

Marine ecosystems are prone to tipping points, particularly in coastal zones where dramatic changes are associated with interactions between cumulative stressors (e.g., shellfish harvesting, eutrophication and sediment inputs) and ecosystem functions. A common feature of many degraded estuaries is elevated turbidity that reduces incident light to the seafloor, resulting from multiple factors including changes in sediment loading, sea-level rise and increased water column algal biomass. To determine whether cumulative effects of elevated turbidity may result in marked changes in the interactions between ecosystem components driving nutrient processing, we conducted a large-scale experiment manipulating sediment nitrogen concentrations in 15 estuaries across a national-scale gradient in incident light at the seafloor. We identified a threshold in incident light that was related to distinct changes in the ecosystem interaction networks (EIN) that drive nutrient processing. Above this threshold, network connectivity was high with clear mechanistic links to denitrification and the role of large shellfish in nitrogen processing. The EIN analyses revealed interacting stressors resulting in a decoupling of ecosystem processes in turbid estuaries with a lower capacity to denitrify and process nitrogen. This suggests that, as turbidity increases with sediment load, coastal areas can be more vulnerable to eutrophication. The identified interactions between light, nutrient processing and the abundance of large shellfish emphasizes the importance of actions that seek to manage multiple stressors and conserve or enhance shellfish abundance, rather than actions focusing on limiting a single stressor.

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 changes in species-trait relationships and ecosystem function using a network analysis of traits.

A major challenge in ecology and environmental management is linking changes in community composition to ecosystem functions. We developed the network analysis of traits (NAT) to show changes in community network structure based on the changes in the composition and connectivity between clusters of species that share traits that imply shifts in functional diversity. We tested the application of NAT on a 113 species found on an intertidal sandflat that was subject to experimental nitrogen addition (control [0 g N/m2 ], medium [150 g N/m2 ], and high [600 g N/m2 ]). This allowed us to directly link mechanistic changes in community composition and function with the trait-space network patterns revealed by NAT. We demonstrate that under medium (150 g N/m2 ) N treatment, functional diversity remained consistent, whereas increasing disturbance to high (600 g N/m2 ) N treatment affected the species-trait network structure and caused merging of functional clusters implying a loss of functional trait diversity.

Effects of Polyester Microfibers on Microphytobenthos and Sediment-Dwelling Infauna.

Microfibers often dominate sediment microplastic samples, but little is known about their ecological effects on benthic organisms and functions. Polyethylene terephthalate) (PET) microfibers were added to 36 sediment chambers at six concentrations (0-0.5 g kg-1 sediment) to assess the effects on microphytobenthos (MPB), a key deposit-feeding bivalve, Macomona liliana, and sediment nutrient pools. MPB photosynthesis was promoted in 18 chambers through a 12 h light/dark cycle. Another 18 chambers were maintained under dark conditions to inhibit photosynthesis. After 35 days of MPB growth and stabilization, four M. liliana were added to each chamber for a further 40 days. MPB biomass and composition were examined alongside M. liliana biochemical and behavioral properties and porewater dissolved inorganic nutrient concentrations. Increasing microfibers resulted in lower MPB biomass, fewer diatom-associated fatty acids (FAs), and an increase in cyanobacteria. The changes in MPB coincided with up to 75% lower energy reserves and reduced burrowing activity in M. liliana. In the light, nitrate + nitrate (NOx) was significantly elevated and related to M. liliana and MPB biochemical properties. Ammoniu (NH4+) concentrations increased but were variable in both the light and the dark. Our results suggest that increasing microfiber concentrations influence the interactions between M. liliana and MPB and affect biogeochemical processing in coastal marine sediments.

Co-occurrence patterns and the large-scale spatial structure of benthic communities in seagrass meadows and bare sand.

BACKGROUND: Species distribution models are commonly used tools to describe diversity patterns and support conservation measures. There is a wide range of approaches to developing SDMs, each highlighting different characteristics of both the data and the ecology of the species or assemblages represented by the data. Yet, signals of species co-occurrences in community data are usually ignored, due to the assumption that such structuring roles of species co-occurrences are limited to small spatial scales and require experimental studies to be detected. Here, our aim is to explore associations among marine sandy-bottom sediment inhabitants and test for the structuring effect of seagrass on co-occurrences among these species across a New Zealand intertidal sandflat, using a joint species distribution model (JSDM). RESULTS: We ran a JSDM on a total of 27 macrobenthic species co-occurring in 300,000 m2 of sandflat. These species represented all major taxonomic groups, i.e. polychaetes, bivalves and crustaceans, collected in 400 sampling locations. A number of significant co-occurrences due to shared habitat preferences were present in vegetated areas, where negative and positive correlations were approximately equally common. A few species, among them the gastropods Cominella glandiformis and Notoacmea scapha, co-occurred randomly with other seagrass benthic inhabitants. Residual correlations were less apparent and mostly positive. In bare sand flats shared habitat preferences resulted in many significant co-occurrences of benthic species. Moreover, many negative and positive residual patterns between benthic species remained after accounting for habitat preferences. Some species occurring in both habitats showed similarities in their correlations, such as the polychaete Aglaophamus macroura, which shared habitat preferences with many other benthic species in both habitats, yet no residual correlations remained in either habitat. CONCLUSIONS: Firstly, analyses based on a latent variable approach to joint distributions stressed the structuring role of species co-occurrences beyond experimental scales. Secondly, results showed context dependent interactions, highlighted by species having more interconnected networks in New Zealand bare sediment sandflats than in seagrass meadows. These findings stress the critical importance of natural history to modelling, as well as incorporating ecological reality in SDMs.

Benthic responses to an Antarctic regime shift: food particle size and recruitment biology.

Polar ecosystems are bellwether indicators of climate change and offer insights into ecological resilience. In this study, we describe contrasting responses to an apparent regime shift of two very different benthic communities in McMurdo Sound, Antarctica. We compared species-specific patterns of benthic invertebrate abundance and size between the west (low productivity) and east (higher productivity) sides of McMurdo Sound across multiple decades (1960s-2010) to depths of 60 m. We present possible factors associated with the observed changes. A massive and unprecedented shift in sponge recruitment and growth on artificial substrata observed between the 1980s and 2010 contrasts with lack of dramatic sponge settlement and growth on natural substrata, emphasizing poorly understood sponge recruitment biology. We present observations of changes in populations of sponges, bryozoans, bivalves, and deposit-feeding invertebrates in the natural communities on both sides of the sound. Scientific data for Antarctic benthic ecosystems are scant, but we gather multiple lines of evidence to examine possible processes in regional-scale oceanography during the eight years in which the sea ice did not clear out of the southern portion of McMurdo Sound. We suggest that large icebergs blocked currents and advected plankton, allowed thicker multi-year ice, and reduced light to the benthos. This, in addition to a possible increase in iron released from rapidly melting glaciers, fundamentally shifted the quantity and quality of primary production in McMurdo Sound. A hypothesized shift from large to small food particles is consistent with increased recruitment and growth of sponges on artificial substrata, filter-feeding polychaetes, and some bryozoans, as well as reduced populations of bivalves and crinoids that favor large particles, and echinoderms Sterechinus neumayeri and Odontaster validus that predominantly feed on benthic diatoms and large phytoplankton mats that drape the seafloor after spring blooms. This response of different guilds of filter feeders to a hypothesized shift from large to small phytoplankton points to the enormous need for and potential value of holistic monitoring programs, particularly in pristine ecosystems, that could yield both fundamental ecological insights and knowledge that can be applied to critical conservation concerns as climate change continues.

Monitoring for tipping points in the marine environment.

Increasingly studies are reporting sudden and dramatic changes in the structure and function of communities or ecosystems. The prevalence of these reports demonstrates the importance for management of being able to detect whether these have happened and, preferably, whether they are likely to occur. Ecological theory provides the rationale for why such changes occur and a variety of statistical indicators of approach that have generic properties have been developed. However, whether the theory has successfully translated into monitoring programmes is unknown. We searched the literature for guidelines that would drive design of monitoring programmes able to detect past and approaching tipping points and analysed marine monitoring programmes in New Zealand. We found very few guidelines in the ecological, environmental or monitoring literature, although both simulation and marine empirical studies suggest that within-year sampling increases the likelihood of detecting approaching tipping points. The combination of the need to monitor both small and medium scale temporal dynamics of multiple variables to detect tipping points meant that few marine monitoring programmes in New Zealand were fit for that purpose. Interestingly, we found many marine examples of studies detecting past and approaching TP with fewer data than was common in the theoretical literature. We, therefore, suggest that utilizing ecological knowledge is of paramount importance in designing and analyzing time-series monitoring for tipping points and increasing the certainty for short-term or infrequent datasets of whether a tipping point has occurred. As monitoring plays an important role in management of tipping points by providing supporting information for other locations about when and why a tipping point may occur, we believe that monitoring for tipping points should be promoted.

Changes in the location of biodiversity-ecosystem function hot spots across the seafloor landscape with increasing sediment nutrient loading.

Declining biodiversity and loss of ecosystem function threatens the ability of habitats to contribute ecosystem services. However, the form of the relationship between biodiversity and ecosystem function (BEF) and how relationships change with environmental change is poorly understood. This limits our ability to predict the consequences of biodiversity loss on ecosystem function, particularly in real-world marine ecosystems that are species rich, and where multiple ecosystem functions are represented by multiple indicators. We investigated spatial variation in BEF relationships across a 300 000 m2 intertidal sandflat by nesting experimental manipulations of sediment pore water nitrogen concentration into sites with contrasting macrobenthic community composition. Our results highlight the significance of many different elements of biodiversity associated with environmental characteristics, community structure, functional diversity, ecological traits or particular species (ecosystem engineers) to important functions of coastal marine sediments (benthic oxygen consumption, ammonium pore water concentrations and flux across the sediment-water interface). Using the BEF relationships developed from our experiment, we demonstrate patchiness across a landscape in functional performance and the potential for changes in the location of functional hot and cold spots with increasing nutrient loading that have important implications for mapping and predicating change in functionality and the concomitant delivery of ecosystem services.

Long-term environmental monitoring for assessment of change: measurement inconsistencies over time and potential solutions.

The importance of long-term environmental monitoring and research for detecting and understanding changes in ecosystems and human impacts on natural systems is widely acknowledged. Over the last decades, a number of critical components for successful long-term monitoring have been identified. One basic component is quality assurance/quality control protocols to ensure consistency and comparability of data. In Norway, the authorities require environmental monitoring of the impacts of the offshore petroleum industry on the Norwegian continental shelf, and in 1996, a large-scale regional environmental monitoring program was established. As a case study, we used a sub-set of data from this monitoring to explore concepts regarding best practices for long-term environmental monitoring. Specifically, we examined data from physical and chemical sediment samples and benthic macroinvertebrate assemblages from 11 stations from six sampling occasions during the period 1996-2011. Despite the established quality assessment and quality control protocols for this monitoring program, we identified several data challenges, such as missing values and outliers, discrepancies in variable and station names, changes in procedures without calibration, and different taxonomic resolution. Furthermore, we show that the use of different laboratories over time makes it difficult to draw conclusions with regard to some of the observed changes. We offer recommendations to facilitate comparison of data over time. We also present a new procedure to handle different taxonomic resolution, so valuable historical data is not discarded. These topics have a broader relevance and application than for our case study.

Multiple stressors, nonlinear effects and the implications of climate change impacts on marine coastal ecosystems.

Global climate change will undoubtedly be a pressure on coastal marine ecosystems, affecting not only species distributions and physiology but also ecosystem functioning. In the coastal zone, the environmental variables that may drive ecological responses to climate change include temperature, wave energy, upwelling events and freshwater inputs, and all act and interact at a variety of spatial and temporal scales. To date, we have a poor understanding of how climate-related environmental changes may affect coastal marine ecosystems or which environmental variables are likely to produce priority effects. Here we use time series data (17 years) of coastal benthic macrofauna to investigate responses to a range of climate-influenced variables including sea-surface temperature, southern oscillation indices (SOI, Z4), wind-wave exposure, freshwater inputs and rainfall. We investigate responses from the abundances of individual species to abundances of functional traits and test whether species that are near the edge of their tolerance to another stressor (in this case sedimentation) may exhibit stronger responses. The responses we observed were all nonlinear and some exhibited thresholds. While temperature was most frequently an important predictor, wave exposure and ENSO-related variables were also frequently important and most ecological variables responded to interactions between environmental variables. There were also indications that species sensitive to another stressor responded more strongly to weaker climate-related environmental change at the stressed site than the unstressed site. The observed interactions between climate variables, effects on key species or functional traits, and synergistic effects of additional anthropogenic stressors have important implications for understanding and predicting the ecological consequences of climate change to coastal ecosystems.

The role of time and species identities in spatial patterns of species richness and conservation.

Many conservation actions are justified on the basis of managing biodiversity. Biodiversity, in terms of species richness, is largely the product of rare species. This is problematic because the intensity of sampling needed to characterize communities and patterns of rarity or to justify the use of surrogates has biased sampling in favor of space over time. However, environmental fluctuations interacting with community dynamics lead to temporal variations in where and when species occur, potentially affecting conservation planning by generating uncertainty about results of species distribution modeling (including range determinations), selection of surrogates for biodiversity, and the proportion of biodiversity composed of rare species. To have confidence in the evidence base for conservation actions, one must consider whether temporal replication is necessary to produce broad inferences. Using approximately 20 years of macrofaunal data from tidal flats in 2 harbors, we explored variation in the identity of rare, common, restricted range, and widespread species over time and space. Over time, rare taxa were more likely to increase in abundance or occurrence than to remain rare or disappear and to exhibit temporal patterns in their occurrence. Space-time congruency in ranges (i.e., spatially widespread taxa were also temporally widespread) was observed only where samples were collected across an environmental gradient. Fifteen percent of the taxa in both harbors changed over time from having spatially restricted ranges to having widespread ranges. Our findings suggest that rare species can provide stability against environmental change, because the majority of species were not random transients, but that selection of biodiversity surrogates requires temporal validation. Rarity needs to be considered both spatially and temporally, as species that occur randomly over time are likely to play a different role in ecosystem functioning than those exhibiting temporal structure (e.g., seasonality). Moreover, temporal structure offers the opportunity to place management and conservation activities within windows of maximum opportunity.

Experimenting with ecosystem interaction networks in search of threshold potentials in real-world marine ecosystems.

Thresholds profoundly affect our understanding and management of ecosystem dynamics, but we have yet to develop practical techniques to assess the risk that thresholds will be crossed. Combining ecological knowledge of critical system interdependencies with a large-scale experiment, we tested for breaks in the ecosystem interaction network to identify threshold potential in real-world ecosystem dynamics. Our experiment with the bivalves Macomona liliana and Austrovenus stutchburyi on marine sandflats in New Zealand demonstrated that reductions in incident sunlight changed the interaction network between sediment biogeochemical fluxes, productivity, and macrofauna. By demonstrating loss of positive feedbacks and changes in the architecture of the network, we provide mechanistic evidence that stressors lead to break points in dynamics, which theory predicts predispose a system to a critical transition.

Polyester microfiber impacts on coastal sediment organic matter consumption.

As plastic pollution continues to accumulate at the seafloor, concerns around benthic ecosystem functionality heightens. This research demonstrates the systematic effects of polyester microfibers on seafloor organic matter consumption rates, an important benthic ecosystem function connected to multiple reactions and processes. We used a field-based assay to measure the loss of organic matter, both with and without polyester microfiber contamination. We identified sediment organic matter content, mud content, and mean grain size as the main drivers of organic matter consumption, however, polyester microfiber contamination decoupled ecosystem relationships and altered observed organic matter cycling dynamics. Organic matter consumption rates varied across horizontal and vertical spaces, highlighting that consumption and associated plastic effects are dependent on environmental heterogeneity at both small (within sites) and larger (between sites) scales. Our results emphasize the important role habitat heterogeneity plays in seafloor organic matter consumption and the associated effects of plastic pollution on ecosystem function.

Biogenic habitat transitions influence facilitation in a marine soft-sediment ecosystem.

Habitats are often defined by the presence of key species and biogenic features. However, the ecological consequences of interactions among distinct habitat-forming species in transition zones where their habitats overlap remain poorly understood. We investigated transition zone interactions by conducting experiments at three locations in Mahurangi Harbour, New Zealand, where the abundance of two habitat-forming marine species naturally varied. The two key species differed in form and function: One was a sessile suspension-feeding bivalve that protruded from the sediment (Atrina zelandica; Pinnidae); the other was a mobile infaunal urchin that bioturbated sediment (Echinocardium cordatum; Spatangoida). The experimental treatments established at each site reflected the natural densities of the species across sites (Atrina only, Echinocardium only, Atrina and Echinocardium together, and plots with neither species present). We identified the individual and combined effects of the two key species on sediment characteristics and co-occurring macrofauna. After five months, we documented significant treatment effects, including the highest abundance of co-occurring macrofauna in the Atrina-only treatments. However, the facilitation of macrofauna by Atrina (relative to removal treatments) was entirely negated in the presence of Echinocardium at densities >10 individuals/m2. The transitional areas in Mahurangi Harbour composed of co-occurring Atrina and Echinocardium are currently widespread and are probably more common now than monospecific patches of either individual species, due to the thinning of dense Atrina patches into sparser mixed zones during the last 10-15 years. Thus, although some ecologists avoid ecotones and habitat edges when designing experiments, suspecting that it will skew the extrapolation of results, this study increased our understanding of benthic community dynamics across larger proportions of the seascape and provided insights into temporal changes in community structure associated with patch dynamics. Particularly in situations where non-abrupt habitat transitions are commonplace, documentation of community dynamics in individual biogenic habitats and in mixed transition zones is required in order to scale-up and generalize results.