Hidden structural heterogeneity enhances marine hotspots’ biodiversity

Studies in terrestrial and shallow-water ecosystems have unravelled the key role of interspecific interactions in enhancing biodiversity, but important knowledge gaps persist for the deep sea. Cold-water coral reefs are hotspots of biodiversity, but the role of interspecific interactions and “habitat cascades” (i.e. positive effects on focal organisms mediated by biogenic habitat formation) in shaping their biodiversity is unknown. Associations between macrofaunal hosts and epifauna were examined in 47 stations at the Mingulay Reef Complex (northeast Atlantic). In total, 101 (group level) and 340 (species level) unique types of facultative associations formed by 43 hosts and 39 epifaunal species were found. Molluscs and empty polychaete tubes had higher values for the type and number of host-epifaunal associations, the Shannon–Wiener (H) and Margalef (d) indices of the epifauna than the rest of the taxonomic groups (p < 0.05). Hosts’ body size, orientation, surface smoothness, and growth form explained a significant amount of variability (32.96%) in epifauna community composition. Epifaunal species richness (S), H and d were 27.4 (± 2.2%), 56.2 (± 2.8%) and 39.9 (± 2.3%) of the respective values for the total sessile communities living on coral framework. This is intriguing as coral framework is orders of magnitude larger than the size of macrofaunal hosts. It is suggested that bivalves, tunicates and empty polychaete tubes increase habitat heterogeneity and enhance biodiversity through “habitat cascades”, in a similar way that epiphytes do in tropical rainforests. Most macrofaunal habitat suppliers in the studied cold-water coral reef are calcified species and likely susceptible to ocean acidification. This indicates that the impacts of climate change on the total biodiversity, structure and health of cold-water coral reefs may potentially be more severe than previously thought.

Long-term changes in kelp forests in an inner basin of the Salish Sea

Kelp forests form an important biogenic habitat that responds to natural and human drivers. Global concerns exist about threats to kelp forests, yet long-term information is limited and research suggests that trends are geographically distinct. We examined distribution of the bull kelp Nereocystis luetkeana over 145 years in South Puget Sound (SPS), a semi-protected inner basin in a fjord estuary complex in the northeast Pacific Ocean. We synthesized 48 historical and modern Nereocystis surveys and examined presence/absence within 1-km segments along 452 km of shoreline. Compared to the earliest baseline in 1878, Nereocystis extent in 2017 decreased 63%, with individual sub-basins showing up to 96% loss. Losses have persisted for decades, across a range of climate conditions. In recent decades, Nereocystis predominantly occurred along shorelines with intense currents and mixing, where temperature and nutrient concentrations did not reach thresholds for impacts to Nereocystis performance, and high current speeds likely excluded grazers. Losses predominated in areas with elevated temperature, lower nutrient concentrations, and relatively low current velocities. The pattern of long-term losses in SPS contrasts with stability in floating kelp abundance during the last century in an area of the Salish Sea with greater wave exposure and proximity to oceanic conditions. These findings support the hypothesis that kelp beds along wave-sheltered shorelines exhibit greater sensitivity to environmental stressors. Additionally, shorelines with strong currents and deep-water mixing may provide refugia within sheltered systems.

Production of mobile invertebrate communities on shallow reefs from temperate to tropical seas

Primary productivity of marine ecosystems is largely driven by broad gradients in environmental and ecological properties. By contrast, secondary productivity tends to be more variable, influenced by bottom-up (resource-driven) and top-down (predatory) processes, other environmental drivers, and mediation by the physical structure of habitats. Here, we use a continental-scale dataset on small mobile invertebrates (epifauna), common on surfaces in all marine ecosystems, to test influences of potential drivers of temperature-standardized secondary production across a large biogeographic range. We found epifaunal production to be remarkably consistent along a temperate to tropical Australian latitudinal gradient of 28.6°, spanning kelp forests to coral reefs (approx. 3500 km). Using a model selection procedure, epifaunal production was primarily related to biogenic habitat group, which explained up to 45% of total variability. Production was otherwise invariant to predictors capturing primary productivity, the local biomass of fishes (proxy for predation pressure), and environmental, geographical, and human impacts. Highly predictable levels of epifaunal productivity associated with distinct habitat groups across continental scales should allow accurate modelling of the contributions of these ubiquitous invertebrates to coastal food webs, thus improving understanding of likely changes to food web structure with ocean warming and other anthropogenic impacts on marine ecosystems.

Long-term changes in kelp forests in an inner basin of the Salish Sea

Understanding the historical extent of biogenic habitats can provide insight into the nature of human impacts and inform restoration and conservation actions. Kelp forests form an important biogenic habitat that responds to natural and human drivers. Global concerns exist about threats to kelp forests, yet long term information is limited and research suggests that trends are geographically distinct. We examined distribution of the bull kelp Nereocystis luetkeana over 145 years in South Puget Sound (SPS), a semi-protected inner basin in a fjord estuary complex in the northeast Pacific Ocean. We synthesized 48 historical and modern Nereocystis surveys and examined presence/absence within 1-km shoreline segments along 452 km of shoreline. Over the last 145 years, Nereocystis has been documented in 26% of the shoreline segments. Its extent decreased 62% basin-wide between the 1870s and 2017, with extreme losses in the two out of three sub-basins (96% in Central and 83% in West). In recent years, almost all Nereocystis occurred in the East sub-basin. In the majority of segments where Nereocystis disappeared, the most recent observation was 4 decades ago, or earlier. Multiple natural and human factors that are known to impact kelp could have contributed to observed patterns, but limited data exist at the spatial and temporal scale of this study. In some areas, recent environmental conditions approached thresholds associated with decreased kelp performance. Longstanding Nereocystis losses occurred exclusively in areas with relatively low current velocities. Remaining Nereocystis predominantly occurred in areas where circulation is stronger. Exceptions to this pattern demonstrate that additional factors outside the scope of this study contributed to trajectories of Nereocystis persistence or loss.

Living on the Edge: Variation in the Abundance and Demography of a Kelp Forest Epibiont

The ecology of an epibiont may depend not only on the dynamics of its biogenic habitat but also on microclimate variation generated within aggregations of its host, a process called physical ecosystem engineering. This study explored variation in the abundance and demography of Membranipora, a suspension-feeding bryozoan, within forests of giant kelp (Macrocystis pyrifera) off the coast of Santa Barbara, California, USA. First, we assessed differences in Membranipora abundance between the edge and interior of kelp forests. The occurrence of Membranipora on kelp blades and its percent cover on occupied blades were higher along forest edges than interiors. Second, we conducted observational studies and field experiments to understand spatial variation in substrate longevity, colony mortality, larval recruitment, and colony growth rates. A higher density of recruits and colonies occurred along forest edges than interiors, suggesting kelp acts like a sieve, whereby larvae settle to edge blades first. Moreover, growth rates along the edge were up to 45% higher than forest interiors. Reduced current speeds, combined with feeding by exterior colonies, may have lowered the uptake of suspended food particles by interior colonies. These results suggest that variation in Membranipora abundance is due in part to differences in colony growth between forest edges and interiors, and not solely the result of recruitment limitation. Our results highlight the importance of ecosystem engineers in influencing the ecological dynamics of epiphytic flora and fauna in marine systems.

A human impact metric for coastal ecosystems with application to seagrass beds in Atlantic Canada

Coastal biogenic habitats are particularly vulnerable to cumulative human impacts from both terrestrial and marine realms. Yet the broad spatial scale used in current global or regional approaches of quantifying multiple anthropogenic stressors are not relevant to the local or bay-wide scales affecting most coastal biogenic habitats. To fill this gap, we developed a standardized human impact metric to quantify the magnitude of anthropogenic impacts to coastal ecosystems more broadly, and biogenic habitats in particular. We applied this metric to 180 seagrass beds (Zostera marina), an important biogenic habitat prioritized for marine protection, across Atlantic Canada. Our impact metric includes five bay-scale and four local-scale terrestrial and marine-based impacts. Results show that seagrass beds and coastal bays in Atlantic Canada exist across a wide gradient of human impacts. Considerable differences in the range and intensity of impacts within and between regions provide insight into where coastal bays and seagrass ecosystems are expected to be most and least affected by individual or cumulative human threats. We discuss implications for management and conservation planning, and the general application of our impact metric to other coastal regions and habitats in Canada and beyond.

Why less complexity produces better forecasts: An independent data evaluation of kelp habitat models

Understanding how species are distributed in the environment is increasingly important for natural resource management, particularly for keystone and habitat forming species, and those of conservation concern. Habitat suitability models are fundamental to developing this understanding; however their use in management continues to be limited due to often-vague model objectives and inadequate evaluation methods. Along the Northeast Pacific coast, canopy kelps (Macrocystis pyrifera and Nereocystis luetkeana) provide biogenic habitat and considerable primary production to nearshore ecosystems. We investigated the distribution of these species by examining a series of increasingly complex habitat suitability models ranging from process-based models based on species' ecology to complex Generalised Additive Models applied to purpose-collected survey data. Seeking limits on model complexity, we explored the relationship between model complexity and forecast skill, measured using both cross-validation and independent data evaluation. Our analysis confirmed the importance of predictors used in models of coastal kelp distributions developed elsewhere (i.e., depth, bottom type, bottom slope, and exposure); it also identified additional important factors including salinity, and interactions between exposure and salinity, and slope and tidal energy. Comparative results showed that cross-validation can lead to over-fitting, while independent data evaluation clearly identified the appropriate model complexity for generating habitat forecasts. Our results also illustrate that, depending on the evaluation data, predictions from simpler models can out-perform those from more complex models. Collectively, the insights from evaluating multiple models with multiple data sets contribute to the holistic assessment of model forecast skill. The continued development of methods and metrics for evaluating model forecasts with independent data, and the explicit consideration of model objectives and assumptions, promise to increase the utility of model forecasts to decision makers.

Caprellid amphipods (Caprella spp.) are vulnerable to both physiological and habitat-mediated effects of ocean acidification

Ocean acidification (OA) is one of the most significant threats to marine life, and is predicted to drive important changes in marine communities. Although OA impacts will be the sum of direct effects mediated by alterations of physiological rates and indirect effects mediated by shifts in species interactions and biogenic habitat provision, direct and indirect effects are rarely considered together for any given species. Here, we assess the potential direct and indirect effects of OA on a ubiquitous group of crustaceans: caprellid amphipods (Caprella laeviuscula and Caprella mutica). Direct physiological effects were assessed by measuring caprellid heart rate in response to acidification in the laboratory. Indirect effects were explored by quantifying caprellid habitat dependence on the hydroid Obelia dichotoma, which has been shown to be less abundant under experimental acidification. We found that OA resulted in elevated caprellid heart rates, suggestive of increased metabolic demand. We also found a strong, positive association between caprellid population size and the availability of OA-vulnerable O. dichotoma, suggesting that future losses of biogenic habitat may be an important indirect effect of OA on caprellids. For species such as caprellid amphipods, which have strong associations with biogenic habitat, a consideration of only direct or indirect effects could potentially misestimate the full impact of ocean acidification.

Individual-based simulation of the spatial and temporal dynamics of macroinvertebrate functional groups provides insights into benthic community assembly mechanisms

The complexity and scales of the processes that shape communities of marine benthic macroinvertebrates has limited our understanding of their assembly mechanisms and the potential to make projections of their spatial and temporal dynamics. Individual-based models can shed light on community assembly mechanisms, by allowing observed spatiotemporal patterns to emerge from first principles about the modeled organisms. Previous work in the Rance estuary (Brittany, France) revealed the principal functional components of its benthic macroinvertebrate communities and derived a set of functional relationships between them. These elements were combined here for the development of a dynamic and spatially explicit model that operates at two spatial scales. At the fine scale, modeling each individual’s life cycle allowed the representation of recruitment, inter- and intra-group competition, biogenic habitat modification and predation mortality. Larval dispersal and environmental filtering due to the tidal characteristics of the Rance estuary were represented at the coarse scale. The two scales were dynamically linked and the model was parameterized on the basis of theoretical expectations and expert knowledge. The model was able to reproduce some patterns of α- and β-diversity that were observed in the Rance estuary in 1995. Model analysis demonstrated the role of local and regional processes, particularly early post-settlement mortality and spatially restricted dispersal, in shaping marine benthos. It also indicated biogenic habitat modification as a promising area for future research. The combination of this mechanism with different substrate types, along with the representation of physical disturbances and more trophic categories, could increase the model’s realism. The precise parameterization and validation of the model is expected to extend its scope from the exploration of community assembly mechanisms to the formulation of predictions about the responses of community structure and functioning to environmental change.