Habitat configurations shape the trophic and energetic dynamics of reef fishes in a tropical-temperate transition zone: implications under a warming future.

Understanding the extent to which species' traits mediate patterns of community assembly is key to predict the effect of natural and anthropogenic disturbances on ecosystem functioning. Here, we apply a trait-based community assembly framework to understand how four different habitat configurations (kelp forests, Sargassum spp. beds, hard corals, and turfs) shape the trophic and energetic dynamics of reef fish assemblages in a tropical-temperate transition zone. Specifically, we tested (i) the degree of trait divergence and convergence in each habitat, (ii) which traits explained variation in species' abundances, and (iii) differences in standing biomass (kg ha-1), secondary productivity (kg ha-1 day-1) and turnover (% day-1). Fish assemblages in coral and kelp habitats displayed greater evidence of trait convergence, while turf and Sargassum spp. habitats displayed a higher degree of trait divergence, a pattern that was mostly driven by traits related to resource use and thermal affinity. This filtering effect had an imprint on the trophic and energetic dynamics of reef fishes, with turf habitats supporting higher fish biomass and productivity. However, these gains were strongly dependent on trophic guild, with herbivores/detritivores disproportionately contributing to among-habitat differences. Despite these perceived overall gains, turnover was decoupled for fishes that act as conduit of energy to higher trophic levels (i.e. microinvertivores), with coral habitats displaying higher rates of fish biomass replenishment than turf despite their lower productivity. This has important implications for biodiversity conservation and fisheries management, questioning the long-term sustainability of ecological processes and fisheries yields in increasingly altered marine habitats.

Persistent thermally driven shift in the functional trait structure of herbivorous fishes: Evidence of top-down control on the rebound potential of temperate seaweed forests?

Extreme climatic events can reshape the functional structure of ecological communities, potentially altering ecological interactions and ecosystem functioning. While these shifts have been widely documented, evidence of their persistence and potential flow-on effects on ecosystem structure following relaxation of extreme events remains limited. Here, we investigate changes in the functional trait structure - encompassing dimensions of resource use, thermal affinity, and body size - of herbivorous fishes in a temperate reef system that experienced an extreme marine heatwave (MHW) and subsequent return to cool conditions. We quantify how changes in the trait structure modified the nature and intensity of herbivory-related functions (macroalgae, turf, and sediment removal), and explored the potential flow-on effects on the recovery dynamics of macroalgal foundation species. The trait structure of the herbivorous fish assemblage shifted as a result of the MHW, from dominance of cool-water browsing species to increased evenness in the distribution of abundance among temperate and tropical guilds supporting novel herbivory roles (i.e. scraping, cropping, and sediment sucking). Despite the abundance of tropical herbivorous fishes and intensity of herbivory-related functions declined following a period of cooling after the MHW, the underlying trait structure displayed limited recovery. Concomitantly, algal assemblages displayed a lack of recovery of the formerly dominant foundational species, the kelp Ecklonia radiata, transitioning to an alternative state dominated by turf and Sargassum spp. Our study demonstrates a legacy effect of an extreme MHW and exemplified the value of monitoring phenotypic (trait mediated) changes in the nature of core ecosystem processes to predict and adapt to the future configurations of changing reef ecosystems.

Effects of human footprint and biophysical factors on the body-size structure of fished marine species.

Marine fisheries in coastal ecosystems in many areas of the world have historically removed large-bodied individuals, potentially impairing ecosystem functioning and the long-term sustainability of fish populations. Reporting on size-based indicators that link to food-web structure can contribute to ecosystem-based management, but the application of these indicators over large (cross-ecosystem) geographical scales has been limited to either fisheries-dependent catch data or diver-based methods restricted to shallow waters (<20 m) that can misrepresent the abundance of large-bodied fished species. We obtained data on the body-size structure of 82 recreationally or commercially targeted marine demersal teleosts from 2904 deployments of baited remote underwater stereo-video (stereo-BRUV). Sampling was at up to 50 m depth and covered approximately 10,000 km of the continental shelf of Australia. Seascape relief, water depth, and human gravity (i.e., a proxy of human impacts) were the strongest predictors of the probability of occurrence of large fishes and the abundance of fishes above the minimum legal size of capture. No-take marine reserves had a positive effect on the abundance of fishes above legal size, although the effect varied across species groups. In contrast, sublegal fishes were best predicted by gradients in sea surface temperature (mean and variance). In areas of low human impact, large fishes were about three times more likely to be encountered and fishes of legal size were approximately five times more abundant. For conspicuous species groups with contrasting habitat, environmental, and biogeographic affinities, abundance of legal-size fishes typically declined as human impact increased. Our large-scale quantitative analyses highlight the combined importance of seascape complexity, regions with low human footprint, and no-take marine reserves in protecting large-bodied fishes across a broad range of species and ecosystem configurations.

Las pesquerías marinas de los ecosistemas costeros en muchas áreas del mundo históricamente han removido a individuos de gran tamaño, potencialmente perjudicando el funcionamiento ambiental y la sostenibilidad a largo plazo de las poblaciones de peces. Los reportes sobre los indicadores basados en el tamaño que se vinculan con la estructura de la red alimenticia pueden contribuir al manejo basado en el ecosistema, aunque la aplicación de estos indicadores a grandes (inter-ecosistemas) escalas geográficas ha estado limitada a datos de captura dependientes de las pesquerías o métodos basados en el buceo restringidos a aguas someras (<20 m), lo cual puede representar erróneamente la abundancia de peces de gran tamaño capturados para la pesca. Obtuvimos los datos de la estructura del tamaño corporal de 82 teleósteos marinos demersales focalizados por razones recreativas o comerciales tomados de 2,904 despliegues de video estéreo subacuático remoto con cebo (stereo-BRUV, en inglés). El muestreo se realizó hasta los 50 metros de profundidad y abarcó aproximadamente 10,000 km del talud continental de Australia. El relieve marino, la profundidad del agua y la gravedad humana (es decir, un indicador de los impactos humanos) fueron los pronosticadores más sólidos de la probabilidad de incidencia de los peces de gran tamaño y de la abundancia de peces por encima del tamaño legal mínimo de captura. Las reservas marinas de protección total tienen un efecto positivo sobre la abundancia de los peces que están por encima del tamaño legal, aunque el efecto varió según el grupo de especies. Como contraste, los peces de tamaño sublegal fueron pronosticados de mejor manera usando gradientes de la temperatura de la superficie marina (media y varianza). En las áreas con un impacto humano reducido, los peces de gran tamaño corporal tenían hasta tres veces mayor probabilidad de aparecer y los peces de tamaño legal eran aproximadamente cinco veces más abundantes. Para los grupos de especies conspicuas con afinidades contrastantes de hábitat, ambiente y biogeografía, la abundancia de peces de tamaño legal normalmente declinó conforme aumentó el impacto humano. Nuestros análisis cuantitativos a gran escala resaltan la importancia conjunta que tienen la complejidad marina, las regiones con una huella humana reducida y las reservas marinas de protección total para la protección de los peces de gran tamaño corporal en una extensa gama de especies y configuraciones ecosistémicas. Efectos de la Huella Humana y los Factores Biofísicos sobre la Estructura del Tamaño Corporal de Especies Marinas Capturadas para la Pesca.

Does the benthic biota or fish assemblage within a large targeted fisheries closure differ to surrounding areas after 12 years of protection in tropical northwestern Australia?

A large (~2450 km2) offshore (~75 km) targeted fisheries closure (TFC) area was implemented on the North West Shelf of Australia (NWS) in 1998 as part of a suite of management controls to address overfishing concerns, and in the process to potentially mitigate any impacts of trawling to benthic habitats. Twelve years later, the benthic biota and fish assemblages in the TFC were assessed using stereo-video and compared with adjacent areas that have been consistently fished with a range of commercial fishing methods. The remote nature of the region has meant that these areas would be inaccessible to recreational fishers. After 12 years of protection there were significant differences between the TFC and comparable fished areas in both the composition and the height of biogenic structures, however the magnitude of these differences were subtle, except for branching soft corals, which were significantly taller in the TFC area. Despite the relatively young age of the TFC, significant differences in the fish abundance and biomass compositions were driven by the slower growing, longer lived and inherently less productive fishery target species. The abundance of Lutjanus sebae (red emperor) and Epinephelus multinotatus (Rankin cod), and the associated biomass of L. sebae and Pristipomoides multidens (goldband snapper) were all greater within the TFC. However, neither the abundance or biomass of the relatively shorter lived and more productive fishery species (e.g. the bluespotted emperor Lethrinus punctulatus and the brownstripe snapper Lutjanus vitta) were greater within the TFC. Growth rates of benthic biota across the NWS are unknown, however the limited detectable differences in benthic biota between the TFC and fished areas, suggests that either recovery of the benthic biota is slow and may not yet be at a threshold for detection and/or alternatively that current fishing activities are not causing adverse impacts to biogenic structures. These large, offshore targeted fishery closures provide a useful reference point to examine the natural variability, growth and recovery of benthic biota and fish assemblages after the cessation of fishing.

Increased connectivity and depth improve the effectiveness of marine reserves.

Marine reserves are a key tool for the conservation of marine biodiversity, yet only ~2.5% of the world's oceans are protected. The integration of marine reserves into connected networks representing all habitats has been encouraged by international agreements, yet the benefits of this design has not been tested empirically. Australia has one of the largest systems of marine reserves, providing a rare opportunity to assess how connectivity influences conservation success. An Australia-wide dataset was collected using baited remote underwater video systems deployed across a depth range from 0 to 100 m to assess the effectiveness of marine reserves for protecting teleosts subject to commercial and recreational fishing. A meta-analytical comparison of 73 fished species within 91 marine reserves found that, on average, marine reserves had 28% greater abundance and 53% greater biomass of fished species compared to adjacent areas open to fishing. However, benefits of protection were not observed across all reserves (heterogeneity), so full subsets generalized additive modelling was used to consider factors that influence marine reserve effectiveness, including distance-based and ecological metrics of connectivity among reserves. Our results suggest that increased connectivity and depth improve the aforementioned marine reserve benefits and that these factors should be considered to optimize such benefits over time. We provide important guidance on factors to consider when implementing marine reserves for the purpose of increasing the abundance and size of fished species, given the expected increase in coverage globally. We show that marine reserves that are highly protected (no-take) and designed to optimize connectivity, size and depth range can provide an effective conservation strategy for fished species in temperate and tropical waters within an overarching marine biodiversity conservation framework.

Overwintering tropical herbivores accelerate detritus production on temperate reefs.

The tropicalization of temperate marine ecosystems can lead to increased herbivory rates, reducing the standing stock of seaweeds and potentially causing increases in detritus production. However, long-term studies analysing these processes associated with the persistence of tropical herbivores in temperate reefs are lacking. We assessed the seasonal variation in abundances, macrophyte consumption, feeding modes and defecation rates of the range-extending tropical rabbitfish Siganus fuscescens and the temperate silver drummer Kyphosus sydneyanus and herring cale Olisthops cyanomelas on tropicalized reefs of Western Australia. Rabbitfish overwintered in temperate reefs, consumed more kelp and other macrophytes in all feeding modes, and defecated more during both summer and winter than the temperate herbivores. Herbivory and defecation increased with rabbitfish abundance, but this was dependent on temperature, with higher rates attained by big schools during summer and lower rates in winter. Still, rabbitfish surpassed temperate herbivores, leading to a fivefold acceleration in the transformation of macrophyte standing stock to detritus, a function usually attributed to sea urchins in kelp forests. Our results suggest that further warming and tropicalization will not only increase primary consumption and affect the habitat structure of temperate reefs but also increase detritus production, with the potential to modify energy pathways.

Network of small no-take marine reserves reveals greater abundance and body size of fisheries target species.

No-take marine reserves (NTRs), i.e. areas with total fishing restrictions, have been established worldwide aiming to promote biodiversity and ecosystem conservation. Brazil has 3.3% of its exclusive economic zone protected by 73 different NTRs, however, most of them currently lack scientific knowledge and understanding of their ecological role, particularly regarding rocky reefs in subtropical regions. In this context, this study aimed to contrast a network of NTRs with comparable fished sites across a coastal biogeographic gradient to investigate the effect of fishing and habitat variability on the abundance and body size of rocky reef fish. We used Baited Remote Underwater stereo-Video (stereo-BRUVs) and Diver Operated stereo-Video (stereo-DOVs) systems to simultaneously sample reef fish and habitat. Model selection and results identified habitat and biogeographic variables, such as distance from shore, as important predictor variables, explaining several aspects of the fish assemblage. The effect of protection was important in determining the abundance and body size of targeted species, in particular for epinephelids and carangids. Conversely, species richness was correlated with habitat complexity but not with protection status. This is the first study using these survey methods in the Southwestern Atlantic, demonstrating how a network of NTRs can provide benchmarks for biodiversity conservation and fisheries management.

A simple function for full-subsets multiple regression in ecology with R.

Full-subsets information theoretic approaches are becoming an increasingly popular tool for exploring predictive power and variable importance where a wide range of candidate predictors are being considered. Here, we describe a simple function in the statistical programming language R that can be used to construct, fit, and compare a complete model set of possible ecological or environmental predictors, given a response variable of interest and a starting generalized additive (mixed) model fit. Main advantages include not requiring a complete model to be fit as the starting point for candidate model set construction (meaning that a greater number of predictors can potentially be explored than might be available through functions such as dredge); model sets that include interactions between factors and continuous nonlinear predictors; and automatic removal of models with correlated predictors (based on a user defined criterion for exclusion). The function takes continuous predictors, which are fitted using smoothers via either gam, gamm (mgcv) or gamm4, as well as factor variables which are included on their own or as two-level interaction terms within the gam smooth (via use of the "by" argument), or with themselves. The function allows any model to be constructed and used as a null model, and takes a range of arguments that allow control over the model set being constructed, including specifying cyclic and linear continuous predictors, specification of the smoothing algorithm used, and the maximum complexity allowed for smooth terms. The use of the function is demonstrated via case studies that highlight how appropriate model sets can be easily constructed and the broader utility of the approach for exploratory ecology.

Drivers of reef shark abundance and biomass in the Solomon Islands.

Remote island nations face a number of challenges in addressing concerns about shark population status, including access to rigorously collected data and resources to manage fisheries. At present, very little data are available on shark populations in the Solomon Islands and scientific surveys to document shark and ray diversity and distribution have not been completed. We aimed to provide a baseline of the relative abundance and diversity of reef sharks and rays and assess the major drivers of reef shark abundance/biomass in the Western Province of the Solomon Islands using stereo baited remote underwater video. On average reef sharks were more abundant than in surrounding countries such as Fiji and Indonesia, yet below that of remote islands without historical fishing pressure, suggesting populations are relatively healthy but not pristine. We also assessed the influence of location, habitat type/complexity, depth and prey biomass on reef shark abundance and biomass. Location was the most important factor driving reef shark abundance and biomass with two times the abundance and a 43% greater biomass of reef sharks in the more remote locations, suggesting fishing may be impacting sharks in some areas. Our results give a much needed baseline and suggest that reef shark populations are still relatively unexploited, providing an opportunity for improved management of sharks and rays in the Solomon Islands.

Addressing Criticisms of Large-Scale Marine Protected Areas.

Designated large-scale marine protected areas (LSMPAs, 100,000 or more square kilometers) constitute over two-thirds of the approximately 6.6% of the ocean and approximately 14.5% of the exclusive economic zones within marine protected areas. Although LSMPAs have received support among scientists and conservation bodies for wilderness protection, regional ecological connectivity, and improving resilience to climate change, there are also concerns. We identified 10 common criticisms of LSMPAs along three themes: (1) placement, governance, and management; (2) political expediency; and (3) social-ecological value and cost. Through critical evaluation of scientific evidence, we discuss the value, achievements, challenges, and potential of LSMPAs in these arenas. We conclude that although some criticisms are valid and need addressing, none pertain exclusively to LSMPAs, and many involve challenges ubiquitous in management. We argue that LSMPAs are an important component of a diversified management portfolio that tempers potential losses, hedges against uncertainty, and enhances the probability of achieving sustainably managed oceans.

Tropicalization strengthens consumer pressure on habitat-forming seaweeds.

Ocean warming is driving species poleward, causing a 'tropicalization' of temperate ecosystems around the world. Increasing abundances of tropical herbivores on temperate reefs could accelerate declines in habitat-forming seaweeds with devastating consequences for these important marine ecosystems. Here we document an expansion of rabbitfish (Siganus fuscescens), a tropical herbivore, on temperate reefs in Western Australia following a marine heatwave and demonstrate their impact on local kelp forests (Ecklonia radiata). Before the heatwave there were no rabbitfish and low rates of kelp herbivory but after the heatwave rabbitfish were common at most reefs and consumption of kelp was high. Herbivory increased 30-fold and kelp abundance decreased by 70% at reefs where rabbitfish had established. In contrast, where rabbitfish were absent, kelp abundance and herbivory did not change. Video-analysis confirmed that rabbitfish were the main consumers of kelp, followed by silver drummers (Kyphosus sydneyanus), a temperate herbivore. These results represent a likely indirect effect of the heatwave beyond its acute impacts, and they provide evidence that range-shifting tropical herbivores can contribute to declines in habitat-forming seaweeds within a few years of their establishment.

Fish wariness is a more sensitive indicator to changes in fishing pressure than abundance, length or biomass.

Identifying the most sensitive indicators to changes in fishing pressure is important for accurately detecting impacts. Biomass is thought to be more sensitive than abundance and length, while the wariness of fishes is emerging as a new metric. Periodically harvested closures (PHCs) that involve the opening and closing of an area to fishing are the most common form of fisheries management in the western Pacific. The opening of PHCs to fishing provides a unique opportunity to compare the sensitivity of metrics, such as abundance, length, biomass and wariness, to changes in fishing pressure. Diver-operated stereo video (stereo-DOV) provides data on fish behavior (using a proxy for wariness, minimum approach distance) simultaneous to abundance and length estimates. We assessed the impact of PHC protection and harvesting on the abundance, length, biomass, and wariness of target species using stereo-DOVs. This allowed a comparison of the sensitivity of these metrics to changes in fishing pressure across four PHCs in Fiji, where spearfishing and fish drives are common. Before PHCs were opened to fishing they consistently decreased the wariness of targeted species but were less likely to increase abundance, length, or biomass. Pulse harvesting of PHCs resulted in a rapid increase in the wariness of fishes but inconsistent impacts across the other metrics. Our results suggest that fish wariness is the most sensitive indicator of fishing pressure, followed by biomass, length, and abundance. The collection of behavioral data simultaneously with abundance, length, and biomass estimates using stereo-DOVs offers a cost-effective indicator of protection or rapid increases in fishing pressure. Stereo-DOVs can rapidly provide large amounts of behavioral data from monitoring programs historically focused on estimating abundance and length of fishes, which is not feasible with visual methods.

Baseline seabed habitat and biotope mapping for a proposed marine reserve.

Seabed mapping can quantify the extent of benthic habitats that comprise marine ecosystems, and assess the impact of fisheries on an ecosystem. In this study, the distribution of seabed habitats in a proposed no-take Marine Reserve along the northeast coast of Great Barrier Island, New Zealand, was mapped using underwater video combined with bathymetry and substratum data. As a result of the boundary extending to the 12 nautical mile Territorial Limit, it would have been the largest coastal Marine Reserve in the country. Recreational and commercial fisheries occur in the region and would be expected to affect species' abundance. The seabed of the study area and adjacent coastal waters has been trawled up to five times per year. Benthic communities were grouped by multivariate cluster analysis into four biotope classes; namely (1) shallow water macroalgae Ecklonia sp. and Ulva sp. on rocky substrata (Eck.Ulv); and deeper (2) diverse epifauna of sponges and bryozoans on rocky substrata (Por.Bry), (3) brittle star Amphiura sp. and sea anemone Edwardsia sp. on muddy sand (Amph.Edw), and (4) hydroids on mud (Hyd). In biotopes Por.Bry, Amph.Edw and Hyd, there where boulders and rocks were present, and diverse sponge, bryozoan and coral communities. Fifty species were recorded in the deep water survey including significant numbers of the shallow-water hexactinellid glass sponges Symplectella rowi Dendy, 1924 and Rossella ijimai Dendy, 1924, the giant pipe demosponge Isodictya cavicornuta Dendy, 1924, black corals, and locally endemic gorgonians. The habitats identified in the waters to the northeast of Great Barrier Island are likely to be representative of similar depth ranges in northeast New Zealand. This study provides a baseline of the benthic habitats so that should the area become a Marine Reserve, any habitat change might be related to protection from fishing activities and impacts, such as recovery of epifauna following cessation of trawling. The habitat map may also be used to stratify future sampling that would aim to collect and identify epifauna and infauna for identification, and thus better describe the biodiversity of the area.

The tropicalization of temperate marine ecosystems: climate-mediated changes in herbivory and community phase shifts.

Climate-driven changes in biotic interactions can profoundly alter ecological communities, particularly when they impact foundation species. In marine systems, changes in herbivory and the consequent loss of dominant habitat forming species can result in dramatic community phase shifts, such as from coral to macroalgal dominance when tropical fish herbivory decreases, and from algal forests to 'barrens' when temperate urchin grazing increases. Here, we propose a novel phase-shift away from macroalgal dominance caused by tropical herbivores extending their range into temperate regions. We argue that this phase shift is facilitated by poleward-flowing boundary currents that are creating ocean warming hotspots around the globe, enabling the range expansion of tropical species and increasing their grazing rates in temperate areas. Overgrazing of temperate macroalgae by tropical herbivorous fishes has already occurred in Japan and the Mediterranean. Emerging evidence suggests similar phenomena are occurring in other temperate regions, with increasing occurrence of tropical fishes on temperate reefs.