A legacy of invasive sun corals: Distinct mobile invertebrate assemblages at near-reef coral-dominated rubble.

Fast-growing and reproducing sun corals have successfully invaded rocky reefs around the Atlantic Ocean, markedly reducing the diversity of fouling invertebrates and macroalgae, and profoundly changing the composition of reef-associated mobile invertebrates. Here, we address sun-coral rubble depositions and report, for the first time, the effects of sun corals on near-reef soft-bottom invertebrate assemblages. Abundance, richness and diversity were higher at rubble habitats compared to bare sandy grounds, which could be a positive effect of substrate complexity. All those parameters were also higher at rubble patches dominated by sun-coral fragments compared to rubble patches dominated by pebbles or shell fragments, also suggesting possible additive effects of coral-borne chemical attraction (sun-coral specific, as inputs of other coral species were virtually absent). Different epifaunal groups were exclusive of rubble habitats and a subset of those exclusive of sun-coral rubble, explaining the incremental richness across habitats. The relative abundance of the two dominant groups - polychaetes (p) and amphipods (a) - contributed the most to the observed contrasts on community structure, as their proportion (p:a) changed from 10:1 in bare sand to nearly co-dominance in coral rubble. While previous research suggested that spreading sun corals reduce prey supply for fish foraging on reef walls, our results suggest they may increase prey abundance and diversity at the adjacent non-consolidated habitat, possibly reshaping trophic pathways connecting the benthic and the pelagic environment.

The COVID-19 pandemic has altered illegal fishing activities inside and outside a marine protected area.

The global COVID-19 pandemic has presented a unique opportunity to explore the consequences of illegal exploitation on wildlife communities, as it continues to have wide-reaching impacts on multiple sectors, including local and national economies, international trade, and conservation enforcement1. The ongoing reductions in monitoring and enforcement during the pandemic have allowed increased opportunities for illegal, unreported, and unregulated activities, particularly for small-scale fisheries2. Even before the pandemic, policymakers and fisheries managers intent on controlling illegal fishing activities established marine protected areas (MPAs) that restrict or prohibit fishing3. Unfortunately, non-compliance with MPAs is often the rule rather than the exception, and less than 10% of the world's MPAs have managed to effectively reduce infringement4. The COVID-19 pandemic has exacerbated these management challenges: a recent review of MPAs worldwide has revealed a general decline in tourism revenue to operate park services during the pandemic, especially revenue needed for supporting personnel to monitor, patrol, and enforce restrictions2. Here, we compile infraction records of illegal fishing activities by both professional (commercial) and amateur (recreational) boats inside and outside of the Tupinambás Ecological Station and the Alcatrazes Wildlife Refuge (Figure 1A), notable for its high reef-fish biomass and diversity in the Southwestern Atlantic5. We show that illegal exploitation has shifted since the onset of the pandemic, targeting larger, higher-value species that contribute disproportionately to the structure and function of reef-fish communities in the region.

Predator control of marine communities increases with temperature across 115 degrees of latitude.

Early naturalists suggested that predation intensity increases toward the tropics, affecting fundamental ecological and evolutionary processes by latitude, but empirical support is still limited. Several studies have measured consumption rates across latitude at large scales, with variable results. Moreover, how predation affects prey community composition at such geographic scales remains unknown. Using standardized experiments that spanned 115° of latitude, at 36 nearshore sites along both coasts of the Americas, we found that marine predators have both higher consumption rates and consistently stronger impacts on biomass and species composition of marine invertebrate communities in warmer tropical waters, likely owing to fish predators. Our results provide robust support for a temperature-dependent gradient in interaction strength and have potential implications for how marine ecosystems will respond to ocean warming.

Colonization history meets further niche processes: how the identity of founders modulates the way predation structure fouling communities.

Community assembly relies on deterministic niche-based processes (e.g., biotic interactions), and stochastic sources of unpredictable variation (e.g., colonization history), that combined will influence late-stage community structure. When community founders present distinct functional traits and a colonization-competition trade-off is not operating, initial colonization can result in late-stage assemblages of variable diversity and composed by different species sets, depending if early colonizers facilitate or inhibit subsequent colonization and survival. By experimentally manipulating the functional identity of founders and predators access during the development of fouling communities, we tested how founder traits constrain colonization history, species interactions and thereby regulate community diversity. We used as founders functionally different fouling organisms (colonial and solitary ascidians, and arborescent and flat-encrusting bryozoans) to build experimental communities that were exposed or protected against predation using a caging approach. Ascidians and bryozoans are pioneer colonizers in benthic communities and also good competitors, but the soft-body of ascidians makes them more susceptible to predators than mineralized bryozoans. When ascidians were founders, their dominance (but not richness) was reduced by predation, resulting in no effects of predators on overall diversity. Conversely, when bryozoans were founders, both space limitation and predator effects resulted in species-poor communities, with reduced number and cover of ascidian species and high overall dominance at the end of the experiment. We, thus, highlight that current species interactions and colonization contingencies related to founder identity should not be viewed as isolated drivers of community organization, but rather as strongly interacting processes underlying species distribution patterns and diversity.

Climate drives the geography of marine consumption by changing predator communities.

The global distribution of primary production and consumption by humans (fisheries) is well-documented, but we have no map linking the central ecological process of consumption within food webs to temperature and other ecological drivers. Using standardized assays that span 105° of latitude on four continents, we show that rates of bait consumption by generalist predators in shallow marine ecosystems are tightly linked to both temperature and the composition of consumer assemblages. Unexpectedly, rates of consumption peaked at midlatitudes (25 to 35°) in both Northern and Southern Hemispheres across both seagrass and unvegetated sediment habitats. This pattern contrasts with terrestrial systems, where biotic interactions reportedly weaken away from the equator, but it parallels an emerging pattern of a subtropical peak in marine biodiversity. The higher consumption at midlatitudes was closely related to the type of consumers present, which explained rates of consumption better than consumer density, biomass, species diversity, or habitat. Indeed, the apparent effect of temperature on consumption was mostly driven by temperature-associated turnover in consumer community composition. Our findings reinforce the key influence of climate warming on altered species composition and highlight its implications for the functioning of Earth's ecosystems.

Evidence of surplus carrying capacity for benthic invertebrates with the poleward range extension of the tropical seagrass Halophila decipiens in SE Brazil.

Seagrasses may enhance the abundance and diversity of benthic invertebrates through trophic facilitation. We investigated this potential ecological function for two seagrasses in SE Brazil: Halodule emarginata, a native species, and Halophila decipiens, a tropical seagrass recently established in the region. At Halophila sites, the organic matter (or carbon) in sediments decreased steadily from seagrass patches to isolated bare grounds, indicating surplus primary production. This was not observed at Halodule sites. At one of the two Halophila sites, localized trophic enrichment was also consistently linked to increased invertebrate abundance within patches, chiefly through increased carrying capacity of small mesoherbivores. Rather than spillover, edge effects were observed at bordering bare habitats, where polychaete predators were abundant. The transition from seagrass edges to isolated bare habitats was marked by an increase of the density of sipunculid worms. The current spread of Halophila may thus change the spatial distribution of benthic ecological functions.

Decadal losses of canopy-forming algae along the warm temperate coastline of Brazil.

The loss of canopy-forming seaweeds from urbanized coasts has intensified in response to warming seas and non-climatic pressures such as population growth and declining water quality. Surprisingly, there has been little information on the extent of historical losses in the South-western Atlantic, which limits our ability to place this large marine ecosystem in a global context. Here, we use meta-analysis to examine long-term (1969-2017) changes to the cover and biomass of Sargassum spp. and structurally simple algal turfs along more than 1,000 kilometres of Brazil's warm temperate coastline. Analysis revealed major declines in canopy cover that were independent of season (i.e., displaying similar trends for both summer and winter) but varied with coastal environmental setting, whereby sheltered bays experienced greater losses than coastal locations. On average, covers of Sargassum spp. declined by 2.6% per year, to show overall losses of 52% since records began (ranging from 20% to 89%). This contrasted with increases in the cover of filamentous turfs (24% over the last 27 years) which are known to proliferate along human-impacted coasts. To test the relative influence of climatic versus non-climatic factors as drivers of this apparent canopy-to-turf shift, we examined how well regional warming trends (decadal changes to sea surface temperature) and local proxies of coastal urbanization (population density, thermal pollution, turbidity and nutrient inputs) were able to predict the changes in seaweed communities. Our results revealed that the most pronounced canopy losses over the past 50 years were at sites exhibiting the greatest degree of coastal warming, the highest population growth and those located in semi-enclosed sheltered bays. These findings contribute knowledge on the drivers of canopy loss in the South-western Atlantic and join with global efforts to understand and mitigate declines of marine keystone species.

Predator traits determine food-web architecture across ecosystems.

Predator-prey interactions in natural ecosystems generate complex food webs that have a simple universal body-size architecture where predators are systematically larger than their prey. Food-web theory shows that the highest predator-prey body-mass ratios found in natural food webs may be especially important because they create weak interactions with slow dynamics that stabilize communities against perturbations and maintain ecosystem functioning. Identifying these vital interactions in real communities typically requires arduous identification of interactions in complex food webs. Here, we overcome this obstacle by developing predator-trait models to predict average body-mass ratios based on a database comprising 290 food webs from freshwater, marine and terrestrial ecosystems across all continents. We analysed how species traits constrain body-size architecture by changing the slope of the predator-prey body-mass scaling. Across ecosystems, we found high body-mass ratios for predator groups with specific trait combinations including (1) small vertebrates and (2) large swimming or flying predators. Including the metabolic and movement types of predators increased the accuracy of predicting which species are engaged in high body-mass ratio interactions. We demonstrate that species traits explain striking patterns in the body-size architecture of natural food webs that underpin the stability and functioning of ecosystems, paving the way for community-level management of the most complex natural ecosystems.

What's in a tide pool? Just as much food web network complexity as in large open ecosystems.

Understanding the fundamental laws that govern complex food web networks over large ecosystems presents high costs and oftentimes unsurmountable logistical challenges. This way, it is crucial to find smaller systems that can be used as proxy food webs. Intertidal rock pool environments harbour particularly high biodiversity over small areas. This study aimed to analyse their food web networks to investigate their potential as proxies of larger ecosystems for food web networks research. Highly resolved food webs were compiled for 116 intertidal rock pools from cold, temperate, subtropical and tropical regions, to ensure a wide representation of environmental variability. The network properties of these food webs were compared to that of estuaries, lakes and rivers, as well as marine and terrestrial ecosystems (46 previously published complex food webs). The intertidal rock pool food webs analysed presented properties that were in the same range as the previously published food webs. The niche model predictive success was remarkably high (73-88%) and similar to that previously found for much larger marine and terrestrial food webs. By using a large-scale sampling effort covering 116 intertidal rock pools in several biogeographic regions, this study showed, for the first time, that intertidal rock pools encompass food webs that share fundamental organizational characteristics with food webs from markedly different, larger, open and abiotically stable ecosystems. As small, self-contained habitats, intertidal rock pools are particularly tractable systems and therefore a large number of food webs can be examined with relatively low sampling effort. This study shows, for the first time that they can be useful models for the understanding of universal processes that regulate the complex network organization of food webs, which are harder or impossible to investigate in larger, open ecosystems, due to high costs and logistical difficulties.

Environmental health assessment of warming coastal ecosystems in the tropics - Application of integrative physiological indices.

According to climate science, ocean warming is one of the current and future greatest threats to coastal ecosystems. Projection scenarios for the end of this century show that tropical intertidal ecosystems are particularly at risk. In this study we optimized and tested a holistic method for bio-monitoring present and projected thermal pressure in such ecosystems, in order to assess organism vulnerability to ocean warming. Several species representative of different animal groups (fish, crustaceans and gastropods) were collected from the field and subjected to an experimental trial for 28 days, testing two temperatures: control (present seawater summer temperature) and elevated temperature (+3 °C, projected seawater temperature anomaly for 2100). Muscle samples were collected weekly to quantify several biomarkers of: i) macromolecular damage (protein unfolding and denaturation, and lipid peroxidation), ii) reactive oxygen species (ROS) scavengers (antioxidant enzymes), and iii) body condition (energy reserves and body mass). These biomarkers were combined in integrated biomarker response (IBR) indices, either in three separate stress response categories (as previously defined) or in a unique combined analysis of overall physiological performance. Both approaches suggest that temperature affected IBRs, with increasing temperatures significantly impairing the overall health of individuals. Biomarkers of lower levels of biological organization indicated deleterious effects of temperature, whereas biomarkers of higher levels suggested maintenance of performance after chronic exposure. Overall indices combining the estimates of biomarkers across levels of biological organization are essential to predict the vulnerability of species, or populations, to climate warming. Such indices may assist managers and stakeholders in the establishment of monitoring programs and environmental policies toward the conservation of fragile coastal systems.

Persistence and space preemption explain species-specific founder effects on the organization of marine sessile communities.

Community assembly may not follow predictable successional stages, with a large fraction of the species pool constituted by potential pioneering species and successful founders defined through lottery. In such systems, priority effects may be relevant in the determination of trajectories of developing communities and hence diversity and assemblage structure at later advanced states. In order to assess how different founder species may trigger variable community trajectories and structures, we conducted an experimental study using subtidal sessile assemblages as model. We manipulated the identity of functionally different founders and initial colony size (a proxy of the time lag before the arrival of later species), and followed trajectories. We did not observe any effects of colony size on response variables, suggesting that priority effects take place even when the time lag between the establishment of pioneering species and late colonizers is very short. Late community structure at experimental panels that started either with the colonial ascidian Botrylloides nigrum, or the arborescent bryozoan Bugula neritina, was similar to control panels allowed natural assembling. In spite of high potential for fast space domination, and hence negative priority effects, B. nigrum suffered high mortality and did not persist throughout succession. Bugula neritina provided complex physical microhabitats through conspecific clustering that have enhanced larval settlement of late species arrivals, but no apparent facilitation was observed. Differently, panels founded by the encrusting bryozoan Schizoporella errata led to different and less diverse communities compared to naturally assembled panels, evidencing strong negative priority effects through higher persistence and space preemption. Schizoporella errata founder colonies inhibited further conspecific settlement, which may greatly relax intraspecific competition, allowing resource allocation to colony growth and space domination, thus reducing the chances for the establishment of other species.

Ecological traps in shallow coastal waters-Potential effect of heat-waves in tropical and temperate organisms.

Mortality of fish has been reported in tide pools during warm days. That means that tide pools are potential ecological traps for coastal organisms, which happen when environmental changes cause maladaptive habitat selection. Heat-waves are predicted to increase in intensity, duration and frequency, making it relevant to investigate the role of tide pools as traps for coastal organisms. However, heat waves can also lead to acclimatization. If organisms undergo acclimatization prior to being trapped in tide pools, their survival chances may increase. Common tide pool species (46 species in total) were collected at a tropical and a temperate area and their upper thermal limits estimated. They were maintained for 10 days at their mean summer sea surface temperature +3°C, mimicking a heat-wave. Their upper thermal limits were estimated again, after this acclimation period, to calculate each species' acclimation response. The upper thermal limits of the organisms were compared to the temperatures attained by tide pool waters to investigate if 1) tide pools could be considered ecological traps and 2) if the increase in upper thermal limits elicited by the acclimation period could make the organisms less vulnerable to this threat. Tropical tide pools were found to be ecological traps for an important number of common coastal species, given that they can attain temperatures higher than the upper thermal limits of most of those species. Tide pools are not ecological traps in temperate zones. Tropical species have higher thermal limits than temperate species, but lower acclimation response, that does not allow them to survive the maximum habitat temperature of tropical tide pools. This way, tropical coastal organisms seem to be, not only more vulnerable to climate warming per se, but also to an increase in the ecological trap effect of tide pools.

Current conditions and colonization history asymmetrically shape the organization of shallow sessile communities after simulated state shifts.

Historical processes affecting biological organization are rarely considered when predicting the effects of disturbance on community structure. In order to assess the relative importance of historical and post-disturbance conditions as determinants of community structure, we undertook reciprocal transplants, at different successional stages, of sessile communities developing at recreational piers that were previously observed to show contrasting fish predation pressure and settlement rate in the São Sebastião Channel, Brazil. Regardless the direction of state shift, after 15 weeks communities converged to the destination site structure, substantially drifting away from the path observed at origin, therefore revealing high susceptibility to environmental change. Although converging, transplanted communities never matched the destination standard in both transplant directions, suggesting that history still mattered, as providing some legacy that lasted, at least, for 15 weeks. The taxonomic groups resisting community drift were hard-bodied invertebrates, which could eventually provide some resilience to these communities through ecosystem engineering.

Camouflage through colour change: mechanisms, adaptive value and ecological significance.

Animals from a wide range of taxonomic groups are capable of colour change, of which camouflage is one of the main functions. A considerable amount of past work on this subject has investigated species capable of extremely rapid colour change (in seconds). However, relatively slow colour change (over hours, days, weeks and months), as well as changes arising via developmental plasticity are probably more common than rapid changes, yet less studied. We discuss three key areas of colour change and camouflage. First, we review the mechanisms underpinning colour change and developmental plasticity for camouflage, including cellular processes, visual feedback, hormonal control and dietary factors. Second, we discuss the adaptive value of colour change for camouflage, including the use of different camouflage types. Third, we discuss the evolutionary-ecological implications of colour change for concealment, including what it can tell us about intraspecific colour diversity, morph-specific strategies, and matching to different environments and microhabitats. Throughout, we discuss key unresolved questions and present directions for future work, and highlight how colour change facilitates camouflage among habitats and arises when animals are faced with environmental changes occurring over a range of spatial and temporal scales.This article is part of the themed issue 'Animal coloration: production, perception, function and application'.

Interspecies comparison of the mechanical properties and biochemical composition of byssal threads.

Several bivalve species produce byssus threads to provide attachment to substrates, with mechanical properties highly variable among species. Here, we examined the distal section of byssal threads produced by a range of bivalve species (Mytilus edulis, Mytilus trossulus, Mytilus galloprovincialis, Mytilus californianus, Pinna nobilis, Perna perna, Xenostrobus securis, Brachidontes solisianus and Isognomon bicolor) collected from different nearshore environments. Morphological and mechanical properties were measured, and biochemical analyses were performed. Multivariate redundancy analyses on mechanical properties revealed that byssal threads of M. californianus, M. galloprovincialis and P. nobilis have very distinct mechanical behaviours compared with the remaining species. Extensibility, strength and force were the main variables separating these species groups, which were highest for M. californianus and lowest for P. nobilis Furthermore, the analysis of the amino acid composition revealed that I. bicolor and P. nobilis threads are significantly different from the other species, suggesting a different underlying structural strategy. Determination of metal contents showed that the individual concentration of inorganic elements varies, but that the dominant elements are conserved between species. Altogether, this bivalve species comparison suggests some molecular bases for the biomechanical characteristics of byssal fibres that may reflect phylogenetic limitations.

Shape, colour plasticity, and habitat use indicate morph-specific camouflage strategies in a marine shrimp.

BACKGROUND: Colour and shape polymorphisms are important features of many species and may allow individuals to exploit a wider array of habitats, including through behavioural differences among morphs. In addition, differences among individuals in behaviour and morphology may reflect different strategies, for example utilising different approaches to camouflage. Hippolyte obliquimanus is a small shrimp species inhabiting different shallow-water vegetated habitats. Populations comprise two main morphs: homogeneous shrimp of variable colour (H) and transparent individuals with coloured stripes (ST). These morphs follow different distribution patterns between their main algal habitats; the brown weed Sargassum furcatum and the pink-red weed Galaxaura marginata. In this study, we first investigated morph-specific colour change and habitat selection, as mechanisms underlying camouflage and spatial distribution patterns in nature. Then, we examined habitat fidelity, mobility, and morphological traits, further indicating patterns of habitat use. RESULTS: H shrimp are capable of changing colour in just a few days towards their algal background, achieving better concealment in the more marginal, and less preferred, red weed habitat. Furthermore, laboratory trials showed that habitat fidelity is higher for H shrimp, whereas swimming activity is higher for the ST morph, aligned to morphological evidence indicating these two morphs comprise a more benthic (H) and a more pelagic (ST) life-style, respectively. CONCLUSIONS: Results suggest that H shrimp utilise a camouflage strategy specialised to a limited number of backgrounds at any one time, whereas ST individuals comprise a phenotype with more generalist camouflage (transparency) linked to a more generalist background utilisation. The coexistence within a population of distinct morphotypes with apparently alternative strategies of habitat use and camouflage may reflect differential responses to substantial seasonal changes in macroalgal cover. Our findings also demonstrate how colour change, behaviour, morphology, and background use all interact in achieving camouflage.

Distinct community dynamics at two artificial habitats in a recreational marina.

Man-made facilities along coastlines modify water circulation and sedimentation dynamics which can affect the structure of marine benthic and pelagic communities. To test how environmental heterogeneity associated with a recreational marina affects the structure of the fouling community and the benthic-pelagic link, we conducted an experiment in which predation effects on recruitment and community structure were assessed in two artificial habitats: inside the marina, an area of calm waters and often disturbed by boating activity, and the breakwater, a more hydrodynamic area. Using visual censuses and video footages we also described the predation pressure and the identity of predators on the two areas. Inside the marina, the recruitment of ascidians and serpulids, but not of bryozoans, was restricted in some occasions, possibly due to reduced water circulation. Predation, mainly by the silver porgy fish Diplodus argenteus, reduced the survivor of didemnid ascidians on both areas, but predation intensity was 40 times higher in the breakwater than inside the marina. While the two artificial habitats did not necessarily support distinct communities, low recruitment coupled to weak predation inside the marina, a less dynamic environment, likely imply lower resilience and more susceptibility to disturbance.

Effect of warming rate on the critical thermal maxima of crabs, shrimp and fish.

The threat of global warming has prompted numerous recent studies on the thermal tolerance of marine species. A widely used method to determine the upper thermal limit has been the Critical Thermal Maximum (CTMax), a dynamic method, meaning that temperature is increased gradually until a critical point is reached. This method presents several advantages over static methods, however, there is one main issue that hinders interpretation and comparison of CTMax results: the rate at which the temperature is increased. This rate varies widely among published protocols. The aim of the present work was to determine the effect of warming rate on CTMax values, using different animal groups. The influence of the thermal niche occupied by each species (intertidal vs subtidal) and habitat (intertidal vs subtidal) was also investigated. CTMax were estimated at three different rates: 1°Cmin(-1), 1°C30min(-1) and 1°Ch(-1), in two species of crab, Eurypanopeus abbreviatus and Menippe nodifrons, shrimp Palaemon northropi and Hippolyte obliquimanus and fish Bathygobius soporator and Parablennius marmoreus. While there were significant differences in the effect of warming rates for some species, for other species warming rate produced no significant differences (H. obliquimanus and B. soporator). While in some species slower warming rates lead to lower CTMax values (P. northropi and P. marmoreus) in other species the opposite occurred (E. abbreviatus and M. nodifrons). Biological group has a significant effect with crabs' CTMax increasing at slower warming rates, which did not happen for shrimp and fish. Subtidal species presented lower CTMax, at all warming rates tested. This study highlights the importance of estimating CTMax values at realistic rates that species encounter in their environment and thus have an ecological value.

Environmentally driven shift between alternative female morphotypes in the mottled shore crab.

Precocious maturity is an important life history trait and might be advantageous if the juvenile habitat is risky. Larvae of the mottled shore crab Pachygrapsus transversus settle to the benthic habitat at a very large size, undergo a brief juvenile development and mature within a few months at a size about a fourth of the asymptotic maximum size for this species. This strategy may rely on the capacity of this species to molt to a juvenile-like morphotype (mI) in which reproduction is suppressed. In the laboratory, winter temperature triggered the puberty molt for a large proportion of juveniles, and still allowed high growth rates if combined with long photoperiod. This would result in a large number of juvenile crabs to join the adult reproductive stock in spring, at the beginning of the breeding season. Adult morphs (mII) grow faster under winter conditions, and therefore might be able to direct resources to reproduction during summer. Yet, females held in captivity without any interaction with conspecifics failed to maintain their reproductive status and often reversed to mI stages. In contrast, when a potential mate was presented, all crabs held their mII status, regardless of whether interaction involved visual, visual + chemical, or visual + chemical + tactile cues. Males discriminate female morphs, and physical interactions, including the inspection of mate receptivity and copulation, took longer when they were interacting with mII females. More than a trade-off between growth and reproduction, sustaining a breeding condition in P. transversus females is apparently a bet for successful mating in the presence of a suitable male conspecific.