Ecological communities are structured by the interaction of bottom-up forces such as nutrients and temperature, which affect primary production rates, and top-down forces such as herbivory and predation, which alter the abundance and competitive dynamics of lower trophic levels. In marine ecosystems, wave exposure simultaneously influences both bottom-up processes and the ability of herbivores to forage. It is unclear how these forces balance across wave exposure gradients to regulate benthic diversity and community composition. This study examines how herbivory and wave exposure interact to structure benthic communities in the Galapagos. We re-assessed data from a field experiment manipulating urchin and fish herbivore access in a field experiment across four sites and two time periods spanning a wave exposure gradient. Wave-induced water motion was measured in-situ with an acoustic doppler velocimeter. We evaluated the effect of variable access to herbivores across wave exposures on benthic community species richness, multivariate community structure, and co-occurrence networks. Low wave energy conditions yielded overall lower community complexity, where herbivory reduced richness to only grazer-resistant species that were randomly distributed. In contrast, high wave energy supported more complex communities where herbivory increased richness and reduced determinism by limiting competitive exclusion by fast-growing species. Urchins had stronger negative effects on community structure at low-flow sites due to destructive grazing, while fishes enhanced species richness at high-flow sites by limiting species that would otherwise dominate the struggle for space. In this manner wave exposure effectively reversed the impact of top-down control on benthic community diversity, with implications for managing marine ecosystems in a period of intense environmental change.

Antipathes galapagensis is a prevalent habitat-forming black coral in subtidal ecosystems of the Galápagos Marine Reserve (GMR). Despite their ecological importance and status as a CITES-regulated order, little is known about their depth distribution, population structure and ecology in the GMR. Surveys were conducted in 2021 and 2022 at 9 sites in the central Galápagos Archipelago to investigate how black coral densities, occupancy, size, habitat utilization, and epizoan overgrowth varied between 2.0 and 20.0 m depth. The shallowest black corals occurred at 3.4 m depth, one of the shallowest occurrences of an Antipathes spp. in the world. Coral density increased with depth, with a maximum density of 5.2 colonies per m2 observed across the depth range surveyed. Occupancy modeling also yielded curves with increasing probabilities of black coral presence with depth at all sites. Colony height increased with depth at 8 out of 9 sites and was positively correlated with coral density at 6 of 9 sites. Overall, 47

A core challenge in ecology is identifying the factors that determine species distribution and functional diversity of species assemblages. Reef fish are the most diverse group of vertebrates, form taxonomically rich and functionally diverse communities and represent a key source of food for humans. We examine regional distribution patterns of reef fish species richness and functional diversity and investigate how these are determined by historical, biogeographic, energetic, and anthropogenic factors. We compiled data from 3,312 underwater visual censuses performed at 122 locations comprising rocky and coral reefs along the Eastern Tropical Pacific (ETP). We used generalized linear mixed‐effects models (GLMMs) implemented in a Bayesian framework to investigate whether distance from quaternary refugia, distance from mainland, shelf area, primary productivity, sea surface temperature (SST), human population gravity, and conservation status influence reef fish species richness and functional diversity in the ETP. Species richness and functional richness (FRic) peaked towards the center of the ETP and our null model suggests that FRic followed a spatial pattern that would be predicted by species richness. Additionally, functional evenness (FEve) was highest at higher latitudes whereas functional dispersion (FDis) was homogeneous throughout the ETP. Species richness was negatively influenced by shelf area and distance from mainland, but positively influenced by SST and conservation status. FEve was influenced by human population gravity and FDis by shelf area. Reef fish species richness and functional diversity in the ETP exhibited a strong division within the region mainly mediated by SST and human population gravity. Our results also suggest that dominant species within small shelf areas share more common traits than dominant species in large areas. This study uncovers previously unknown regional patterns of reef fish functional diversity and provides new insights into how historical, biogeographic, energetic, and anthropogenic factors influence complementary biodiversity facets.

A core challenge in ecology is identifying the factors that determine species distribution and functional diversity of species assemblages. Reef fish are the most diverse group of vertebrates, form taxonomically rich and functionally diverse communities and represent a key source of food for humans. We examine regional distribution patterns of reef fish species richness and functional diversity and investigate how these are determined by historical, biogeographic, energetic, and anthropogenic factors. We compiled data from 3,312 underwater visual censuses performed at 122 locations comprising rocky and coral reefs along the Eastern Tropical Pacific (ETP). We used generalized linear mixed-effects models (GLMMs) implemented in a Bayesian framework to investigate whether distance from quaternary refugia, distance from mainland, shelf area, primary productivity, sea surface temperature (SST), human population gravity, and conservation status influence reef fish species richness and functional diversity in the ETP. Species richness and functional richness (FRic) peaked towards the center of the ETP and our null model suggests that FRic followed a spatial pattern that would be predicted by species richness. Additionally, functional evenness (FEve) was highest at higher latitudes whereas functional dispersion (FDis) was homogeneous throughout the ETP. Species richness was negatively influenced by shelf area and distance from mainland, but positively influenced by SST and conservation status. FEve was influenced by human population gravity and FDis by shelf area. Reef fish species richness and functional diversity in the ETP exhibited a strong division within the region mainly mediated by SST and human population gravity. Our results also suggest that dominant species within small shelf areas share more common traits than dominant species in large areas. This study uncovers previously unknown regional patterns of reef fish functional diversity and provides new insights into how historical, biogeographic, energetic, and anthropogenic factors influence complementary biodiversity facets.

Tropical mountain ecosystems are threatened by land use pressures, compromising their capacity to provide ecosystem services. Although local patterns and interactions among anthropogenic and biophysical factors shape these socio-ecological systems, the analysis of landscape changes and their driving forces is often qualitative and sector oriented. Using the Driver-Pressure-State-Impact-Response (DPSIR) framework, we characterized land use land cover (LULC) dynamics using Markov chain probabilities by elevation and geographic settings and then integrated them with a variety of publicly available geospatial and temporal data into a Generalized Additive Model (GAM) to evaluate factors driving such landscape dynamics in a sensitive region of the northern Ecuadorian Andes. In previous agricultural land located at lower elevations to the east of the studied territory, we found a significant expansion of floriculture (13 times) and urban areas (25 times), reaching together almost 10% of the territory from 1990 to 2014. Our findings also revealed an unexpected trend of páramo stability (0.75-0.90), but also a 40% reduction of montane forests, with the lowest probability (<0.50) of persistence in the elevation band of 2800-3300 m; agricultural land is replacing this LULC classes at higher elevation. These trends highlight the increasing threat of permanently losing the already vulnerable native mountain biodiversity. GAMs of socio-economic factors, demographic, infrastructure variables, and environmental parameters explained between 21 to 42% of the variation of LULC transitions observed in the study region, where topographic factors was the main drivers of change. The conceptual and methodological approach of our findings demonstrate how dynamic patterns through space and time and their explanatory drivers can assist local authorities and decision makers to improve sustainable resource land management in vulnerable landscapes such as the tropical Andes in northern Ecuador.

Tropical mountain ecosystems are threatened by land use pressures, reducing the capacity of ecosystems to provide a large diversity of benefits to people and to be able to achieve them in the long term. The analysis of land use pressures is often superficial and very general, although they are characterized by numerous interactions and strong differences in their local dynamics. We used a variety of freely available geospatial and temporal data and methods to assess and explain patterns of land use land cover (LULC) change, focusing on native ecosystem dynamics, in a sensitive region of the northern Ecuadorian Andes. Our results demonstrate a dynamic and clear geographical pattern of distinct LULC transitions through time, explained by different combination of socio-economic factors, pressure variables and environmental parameters, from which ecological context variables, such as slope and elevation, were the main drivers of change in this landscape. We found that deforestation of remnant native forest and agricultural expansion still occur in higher elevations located, while land conversion toward anthropic environments were observed in lower elevations to the east of the studied territory. Our findings also reveal an unexpected stability trend of paramo and a successional recovery of previous agricultural land to the west and center of the territory which could be explained by agricultural land abandonment. However, the very low probability of persistence of montane forests in most of the studied landscape, highlights the risk that the remnant montane forests will be permanently lost in a few years, posing a greater threat to the already vulnerable biodiversity and limiting the capacity ecosystem service provisioning. The dynamic patterns through space and time and their explanatory drivers, found in our study, could help improve sustainably resource land management in vulnerable landscapes such as the tropical Andes in northern Ecuador.

Environmental stress impedes predation and herbivory by limiting the ability of animals to search for and consume prey. We tested the contingency of this relationship on consumer traits and specifically hypothesized that herbivore mobility relative to the return time of limiting environmental stress would predict consumer effects. We examined how wave-induced water motion affects marine communities via herbivory by highly mobile (fish) vs. slow-moving (pencil urchin) consumers at two wave-sheltered and two wave-exposed rocky subtidal locations in the Galapagos Islands. The exposed locations experienced 99th percentile flow speeds that were 2-5 times greater than sheltered locations, with mean flow speeds >33 cm/s vs. <16 cm/s, 2-7 times higher standing macroalgal cover and 2-3 times lower cover of crustose coralline algae than the sheltered locations. As predicted by the environmental stress hypothesis (ESH), there was a negative relationship between mean flow speed and urchin abundance and herbivory rates on Ulva spp. algal feeding assays. In contrast, the biomass of surgeonfishes (Acanthuridae) and parrotfishes (Labridae: Scarinae) was positively correlated with mean flow speed. Ulva assays were consumed at equal rates by fish at exposed and sheltered locations, indicating continued herbivory even when flow speeds surpassed maximum reported swimming speeds at a rate of 1-2 times per minute. Modeled variation in fish species richness revealed minimal effects of diversity on herbivory rates at flow speeds <40 cm/s, when all species were capable of foraging, and above 120 cm/s, when no species could forage, while increasing diversity maximized herbivory rates at flow speeds of 40-120 cm/s. Two-month herbivore exclusion experiments during warm and cool seasons revealed that macroalgal biomass was positively correlated with flow speed. Fish limited macroalgal development by 65-91% at one exposed location but not the second and by 70% at the two sheltered locations. In contrast, pencil urchins did not affect algal communities at either exposed location, but reduced macroalgae by 87% relative to controls at both sheltered locations. We propose an extension of the ESH that is contingent upon mobility to explain species-specific changes in feeding rates and consumer effects on benthic communities across environmental gradients.

Cyanobacteria use limiting resources efficiently to take advantage of nutrient pulses, adapt to variable surroundings, and spread; this proliferation is often an indicator of ecosystem stress. We documented the sudden appearance of benthic cyanobacterial mats on a subtidal rocky reef (Roca Cousins, Galapagos Islands) during El Nino in January 2016. At this time, cyanobacteria covered 32.0% of horizontal rock surfaces and 2.6% of rock walls at 6-8 m depth. Monitored photo quadrats and observations indicated that these mats were previously absent from rock walls and horizontal-sloping substrata at this site for 16 years prior to their initial appearance. The cyanobacteria was also observed at 4 other sites in the central Galapagos Islands during 2016-2018. Laboratory experiments testing the effects of temperature (28-31 degrees C) on cyanobacterial growth and survival indicated that survival was higher at 28 than 31 degrees C, suggesting that 31 degrees C may be an upper thermal limit. Over two years in the field, cyanobacterial mats peaked during the warm El Nino (January 2016) and declined during two cold La Nina periods (June 2016, September 2017), ultimately declining to 6.0% cover in January 2018. Regression analysis of the temperature and cyanobacterial percent cover data indicated that temperature explained 56.9% of the variation in cyanobacteria cover in the field over the 2-year period. The cyanobacterial mats may be a consortium of several species as the closest genetic matches confirmed by Sanger sequencing (90-91.5%) were Oscillatoria spongeliae, Merismopedia glauca, and Synechococcus elongatus. Comparison of areas under the cyanobacterial mats to the adjacent uncovered rock substrata suggested that the cyanobacteria had a negative influence on underlying crustose coralline algae (CCA), as the cover of bleached CCA was 1.75 fold higher under the mats while the cover of "healthy" pink-pigmented CCA was 3 fold higher on the uncovered substrata. Short-term field experiments and feeding surveys performed to evaluate predation and to calculate electivity indices indicated that the cyanobacterial mats were avoided by dominant consumers (Eucidaris galapagensis, Pentaceraster cumingi, Nidorellia armata, and Prionurus laticlavius). Taken together, these results imply that the novel appearance of cyanobacterial mats in the Galapagos rocky subtidal zone was facilitated by unusually warm temperatures during the 2014-2017 El Nino (28-29 degrees C) and that the cyanobacteria were regulated by temperature, but not by consumers. Future outbreaks of mat-forming cyanobacteria during El Nino periods may negatively impact the abundance of CCA and have direct and indirect negative effects on other components of marine benthic communities that rely on CCA as either a settlement substratum or food source.

As marine ecosystems are influenced by global and regional processes, standardized information on community structure has become crucial for assessing broad-scale responses to natural and anthropogenic disturbances. Extensive biogeographic provinces, such as the Brazilian Province in the southwest Atlantic, present numerous theoretical and methodological challenges for understanding community patterns on a macroecological scale. In particular, the Brazilian Province is composed of a complex system of heterogeneous reefs and a few offshore islands, with contrasting histories and geophysical-chemical environments. Despite the large extent of the Brazilian Province (almost 8,000 kilometers), most studies of shallow benthic communities are qualitative surveys and/or have been geographically restricted. We quantified community structure of shallow reef habitats from 0° to 27°S latitude using a standard photographic quadrat technique. Percent cover data indicated that benthic communities of Brazilian reefs were dominated by algal turfs and frondose macroalgae, with low percent cover of reef-building corals. Community composition differed significantly among localities, mostly because of their macroalgal abundance, despite reef type or geographic region, with no evident latitudinal pattern. Benthic diversity was lower in the tropics, contrary to the general latitudinal diversity gradient pattern. Richness peaked at mid-latitudes, between 20°S to 23°S, where it was ~3.5-fold higher than localities with the lowest richness. This study provides the first large-scale description of benthic communities along the southwestern Atlantic, providing a baseline for macroecological comparisons and evaluation of future impacts. Moreover, the new understanding of richness distribution along Brazilian reefs will contribute to conservation planning efforts, such as management strategies and the spatial prioritization for the creation of new marine protected areas.

Climate change increases local climatic variation and unpredictability, which can alter ecological interactions and trigger wildlife disease outbreaks. Here we describe an unprecedented multi-species outbreak of wild fish disease driven by a climate perturbation. The 2015–16 El Niño generated a +2.5 °C sea surface temperature anomaly in the Galapagos Islands lasting six months. This coincided with a novel ulcerative skin disease affecting 18 teleost species from 13 different families. Disease signs included scale loss and hemorrhagic ulcerated patches of skin, fin deterioration, lethargy, and erratic behavior. A bacterial culture isolated from skin lesions of two of the affected fish species was identified by sequencing of the 16S rRNA gene as a Rahnella spp. Disease prevalence rates were linearly correlated with density in three fish species. In January 2016, disease prevalence reached 51.1% in the ring-tailed damselfish Stegastes beebei (n = 570) and 18.7% in the king angelfish Holacanthus passer (n = 318), corresponding to 78% and 86% decreases in their populations relative to a 4.5-year baseline, respectively. We hypothesize that this outbreak was precipitated by the persistent warm temperatures and lack of planktonic productivity that characterize extreme El Niño events, which are predicted to increase in frequency with global warming.

In diverse tropical webs, trophic cascades are presumed to be rare, as species interactions may dampen top-down control and reduce their prevalence. To test this hypothesis, we used an open experimental design in the Galápagos rocky subtidal that enabled a diverse guild of fish species, in the presence of each other and top predators (sea lions and sharks), to attack two species of sea urchins grazing on benthic algae. Time-lapse photography of experiments on natural and experimental substrates revealed strong species identity effects: only two predator species-blunthead triggerfish (Pseudobalistes naufragium) and finescale triggerfish (Balistes polylepis)-drove a diurnal trophic cascade extending to algae, and they preferred large pencil urchins (Eucidaris galapagensis) over green urchins (Lytechinus semituberculatus). Triggerfish predation effects were strong, causing a 24-fold reduction of pencil urchin densities during the initial 21 hours of a trophic cascade experiment. A trophic cascade was demonstrated for pencil urchins, but not for green urchins, by significantly higher percent cover of urchin-grazed algae in cages that excluded predatory fish than in predator access (fence) treatments. Pencil urchins were more abundant at night when triggerfish were absent, suggesting that this species persists by exploiting a nocturnal predation refuge. Time-series of pencil urchin survivorship further demonstrated per capita interference effects of hogfish and top predators. These interference effects respectively weakened and extended the trophic cascade to a fourth trophic level through behavioral modifications of the triggerfish-urchin interaction. We conclude that interference behaviors capable of modifying interaction strength warrant greater attention as mechanisms for altering top-down control, particularly in speciose food webs.

Replicate flow measurements made with the Acoustic Doppler current Profiler (ADP).

Background The Galapagos Whale Shark Project is a multi-institutional effort initiated in 2011 with the aim to characterize the presence, population structure and movement patterns of whale sharks within and around the Galapagos Marine Reserve. This specific study aimed to understand their occurrence, residency and habitat use around Darwin Island, located at the northern tip of the Galapagos Archipelago, where large individuals had been reported to be seasonally abundant. Approach This study followed a diversified methodology approach that included: analysis of a local underwater visual census database of pelagic species (2007–2013) to establish seasonality in their occurrence; specific whale shark surveys (2011–2013) to collect information about shark's size (laser photogrammetry), sex, behavior and signs of potential pregnancy; photo-ID records (2011–2013) obtained during these surveys to determine residency and abundance; and the deployment of acoustic tags for continuous tracking around Darwin Island to assess habitat use at the study site. Results Whale shark presence at Darwin Island follows a seasonal pattern. During the cool season (July–December), a strongly female-biased whale shark population, composed mostly (91.8%) by large individuals (11.35 m ± 0.12 m (TL ± SE)), pass through the study site. The great majority of these individuals show clear distended bellies, which could be a sign of a potential pregnancy. Population dynamics models for these apparently pregnant sharks estimated the presence of 3.76 ± 0.90 (mean±SE) sharks in the study area per day with an individual residence time of 2.09 ± 0.51 (mean±SE) days. Assuming constancy in these rates for the entire cool season, we can estimate a net abundance of 695 ± 166 (SE; 95%CI 442–1110) apparently pregnant whale sharks per season. Movement patterns analysis of four apparently pregnant individuals revealed an intense use of Darwin's Arch, where no feeding or specific behavior has been recorded or could be inferred from their dive profiles, together with periodic excursions around the island's vicinity. Sharks showed a preference for intermediate depths (20–30 m) with occasional dives mostly to mid-water, remaining the majority of their time at water temperatures between 24–25°C. Conclusions The lack of evidence of specific behavior observed at Darwin Island, together with the short residence time and strong intra-seasonal abundance and high turnover rate, indicate that this location is not an aggregation site but an important stopover in a migration. In the case of adult R. typus individuals observed, this migration might involve reproductive purposes, as all but one were apparently gravid.

Relationship between maximum bottom depth within 15 km of the study sites and the frequency of EUC water masses at the study sites.

Ecosystem-based management (EBM) is an emerging tool that considers humans as an integral part of the ecosystem (Arkema et al. 2006). EBM is different from other marine management tools (i.e., marine protected areas (MPAs), fishing regulations, quotas) because they typically deal with only one sector, resource, or impact. Primarily due to this, these strategies are not suitable because they fail to acknowledge the complex dynamics that affect social-ecological interactions. Instead, EBM attempts to embrace the complexity that drives the interactions between humans, their multiple impacts, and their environment (McLeod et al. 2005; Tallis et al. 2010). EBM assesses how multiple sectors and cumulative impacts interact to affect the capacity of marine systems to deliver benefits to humans (Arkema et al. 2006; Ruckelshaus et al. 2008). 1 The main goal of EBM is to build resilient social-ecological systems that can secure the long-term provision of ecosystem services and goods to humans (McLeod et al. 2005).

The life history of the whale shark (Rhincodon typus), including its reproductive ecology, still remains largely unknown. Here, we present results from the first whale shark population study around Darwin Island, Galapagos Marine Reserve. Following a diversified approach we characterized seasonal occurrence, population structure and size, and described habitat use of whale sharks based on fine scale movements around the island. Whale shark presence at Darwin Island was negatively correlated with Sea Surface Temperature (SST), with highest abundance corresponding to a cool season between July and December over six years of monitoring. From 2011 to 2013 we photo-identified 82 whale sharks ranging from 4 to 13.1 m Total Length (TL). Size distribution was bimodal, with a great majority (91.5%) of adult female individuals averaging 11.35 m±0.12 m (TL±SE), all but one showing signs of a potential pregnancy. Population dynamics models for apparently pregnant sharks estimated the presence of 3.76±0.90 (mean ± SE) sharks in the study area per day with an individual residence time of 2.09±0.51 (mean ± SE) days. Movement patterns analysis of four apparently pregnant individuals tracked with acoustic tags at Darwin Island revealed an intense use of Darwin's Arch, where no feeding or specific behavior has been recorded, together with periodic excursions around the island's vicinity. Sharks showed a preference for intermediate depths (20-30 m) with occasional dives mostly to mid-water, remaining the majority of their time at water temperatures between 24-25°C. All of our results point to Darwin Island as an important stopover in a migration, possibly with reproductive purposes, rather than an aggregation site. Current studies carried out in this area to investigate regional scale movement patterns may provide essential information about possible pupping grounds for this enigmatic species.

Stakeholders increasingly expect ecosystem assessments as part of advice on fisheries management. Quantitative models to support fisheries decision-making may be either strategic (big picture', direction-setting and contextual) or tactical (focused on management actions on short timescales), with some strategic models informing the development of tactical models. We describe and review Models of Intermediate Complexity for Ecosystem assessments' (MICE) that have a tactical focus, including use as ecosystem assessment tools. MICE are context- and question-driven and limit complexity by restricting the focus to those components of the ecosystem needed to address the main effects of the management question under consideration. Stakeholder participation and dialogue is an integral part of this process. MICE estimate parameters through fitting to data, use statistical diagnostic tools to evaluate model performance and account for a broad range of uncertainties. These models therefore address many of the impediments to greater use of ecosystem models in strategic and particularly tactical decision-making for marine resource management and conservation. MICE are capable of producing outputs that could be used for tactical decision-making, but our summary of existing models suggests this has not occurred in any meaningful way to date. We use a model of the pelagic ecosystem in the Coral Sea and a linked catchment and ocean model of the Gulf of Carpentaria, Australia, to illustrate how MICE can be constructed. We summarize the major advantages of the approach, indicate opportunities for the development of further applications and identify the major challenges to broad adoption of the approach.

Climate change presents significant challenges to modelling and managing aquatic resources. Equilibrium assumptions common in many modelling approaches need to be replaced by formulations that allow for changing baselines and integration of ongoing changes and adaptations by species, ecosystems and humans. As ecosystems change, so will the ways humans use, monitor and manage them. Consequently, adaptive management loops and supporting tools deserve more prominence in the management toolbox. Models are critical tools for providing an early understanding of the challenges to be faced by integrating observations and examining possible solutions. We review modelling tools currently available to incorporate the effect of climate change on marine and freshwater ecosystems, and the implications for management of natural resources. System non-linearity can confound interpretations and hence adaptive management responses are needed that are robust to unexpected outcomes. An improvement in the ability to model the effects of climate change from a social and economic perspective is necessary. The outputs from 'end-to-end' and socio-ecological models can potentially inform planning, in both Australia and the Pacific region, about how best to build resilience to climate change. In this context, the importance of well directed data-collection programs is also emphasised. Lessons from this region, which is advanced with regard to modelling approaches, can guide increased use of models to test options for managing aquatic resources worldwide.