Nearshore marine biodiversity of Osa Peninsula, Costa Rica: Where the ocean meets the rainforest.

Osa Peninsula in remote southwest Costa Rica harbors 2.5% of global terrestrial biodiversity in only 1,200 km2 and has the largest remaining tract of Pacific lowland wet forest in Mesoamerica. However, little is known about the marine ecosystems of this diverse region. Much of the coastline consists of soft sediment exposed to strong wave action. Three major hard bottom habitat types define this region, including: 1) coral reefs around Isla del Caño Biological Reserve, a no-take marine protected area (MPA) of 52 km2, 2) coastal rocky reefs and islets along the peninsula, including Corcovado National Park, and 3) submerged pinnacles just outside the Isla del Caño MPA. Average coral cover at Isla del Caño was 21%, composed primarily of Porites lobata and Pocillopora elegans. In contrast, coastal rocky reefs were dominated by turf algae (39.8%) and macroalgae (20.7%) with low coral cover (1.1%). Submerged pinnacles were dominated by crustose coralline algae (33.3%) and erect coralline algae (25.7%). Fish assemblage characteristics (species richness, abundance, biomass) were significantly higher at the pinnacles compared to the other habitats and was dominated by schooling species such as Haemulon steindachneri, and the herbivores Kyphosus ocyurus, and Acanthurus xanthopterus. Top predators, primarily Triaenodon obesus, Caranx sexfasciatus, and Lutjanus argentimaculatus, were also most abundant at these pinnacles and accounted for the largest differences in fish trophic structure among habitats. Despite Isla del Caño being fully protected from fishing, biomass was similar to fished areas along the coast and lower than the adjacent submerged pinnacles outside the reserve. Similarly, Corcovado National Park includes 20.3 km2 of no-take MPAs; however, there is limited enforcement, and we noted several instances of fishing within the park. The unique configuration of healthy offshore coral reefs and pinnacles connected to coastal habitats provides corridors for many species including large predators such as sharks and other marine megafauna, which warrants additional protection.

Using stable isotopes analysis to understand ontogenetic trophic variations of the scalloped hammerhead shark at the Galapagos Marine Reserve.

Changes in life-history requirements drive trophic variations, particularly in large marine predators. The life history of many shark species is still poorly known and understanding their dietary ontogeny is a challenging task, especially for highly migratory species. Stable isotope analysis has proven as a useful method for examining the foraging strategies of sharks and other marine predators. We assessed the foraging strategies and ontogenetic changes of scalloped hammerhead sharks, Sphyrna lewini, at Galapagos Marine Reserve (GMR), by analysing δ13C and δ15N signatures in different maturity stages. Our isotopic results suggest ontogenetic shifts in resource use between sub-adult and adult stages, but not between adult and juvenile stages. Carbon isotopic signatures found in the juvenile stage were enriched in contrast to sub-adults (~0.73‰) suggesting a combination of the maternal input and the use of coastal resources around the Galapagos Islands. Adult female sharks also showed enrichment in δ13C (~0.53‰) in comparison to sub-adult stages that suggest feeding in high primary productivity areas, such as the GMR. This study improves the understanding of the trophic ecology and ontogenetic changes of a highly migratory shark that moves across the protected and unprotected waters of the Eastern Tropical Pacific.

Correction: A matter of taste: Spatial and ontogenetic variations on the trophic ecology of the tiger shark at the Galapagos Marine Reserve.

[This corrects the article DOI: 10.1371/journal.pone.0222754.].

Genetic population dynamics of the critically endangered scalloped hammerhead shark (Sphyrna lewini) in the Eastern Tropical Pacific.

The scalloped hammerhead shark, Sphyrna lewini, is a Critically Endangered, migratory species known for its tendency to form iconic and visually spectacular large aggregations. Herein, we investigated the population genetic dynamics of the scalloped hammerhead across much of its distribution in the Eastern Tropical Pacific (ETP), ranging from Costa Rica to Ecuador, focusing on young-of-year animals from putative coastal nursery areas and adult females from seasonal aggregations that form in the northern Galápagos Islands. Nuclear microsatellites and partial mitochondrial control region sequences showed little evidence of population structure suggesting that scalloped hammerheads in this ETP region comprise a single genetic stock. Galápagos aggregations of adults were not comprised of related individuals, suggesting that kinship does not play a role in the formation of the repeated, annual gatherings at these remote offshore locations. Despite high levels of fisheries exploitation of this species in the ETP, the adult scalloped hammerheads here showed greater genetic diversity compared with adult conspecifics from other parts of the species' global distribution. A phylogeographic analysis of available, globally sourced, mitochondrial control region sequence data (n = 1818 sequences) revealed that scalloped hammerheads comprise three distinct matrilines corresponding to the three major world ocean basins, highlighting the need for conservation of these evolutionarily unique lineages. This study provides the first view of the genetic properties of a scalloped hammerhead aggregation, and the largest sample size-based investigation of population structure and phylogeography of this species in the ETP to date.

The elasmobranchs of Malpelo Flora and Fauna Sanctuary, Colombia.

Management and conservation actions in marine-protected areas require baselines for monitoring threatened marine fauna such as elasmobranchs. This article provides evidence of the occurrence of 34 species of elasmobranchs (21 sharks and 13 batoids) in the Malpelo Flora and Fauna Sanctuary, Colombia, including five new records of sharks and three of rays. From 1987 to 2021, new records were obtained by underwater visual census using SCUBA, manned submersibles and deep-ocean cameras to depths of up to 2211 m. Of the recorded species, 21 are considered as threatened taxa (64%) by the IUCN, making the Malpelo Flora and Fauna Sanctuary an essential conservation area for this highly threatened group of species.

Marine communities of the newly created Kawésqar National Reserve, Chile: From glaciers to the Pacific Ocean.

The newly created Kawésqar National Park (KNP) and National Reserve (KNR) in southern Chile consists of diverse terrestrial and marine habitats, which includes the southern terminus of the Andes, the Southern Patagonia Ice Fields, sub-Antarctic rainforests, glaciers, fjords, lakes, wetlands, valleys, channels, and islands. The marine environment is influenced by wide ranging hydrological factors such as glacier melt, large terrigenous inputs, high precipitation, strong currents, and open ocean water masses. Owing to the remoteness, rugged terrain, and harsh environmental conditions, little is known about this vast region, particularly the marine realm. To this end, we conducted an integrated ecological assessment using SCUBA and remote cameras down to 600 m to examine this unique and largely unexplored ecosystem. Kelp forests (primarily Macrocystis pyrifera) dominate the nearshore ecosystem and provide habitat for myriad benthic organisms. In the fjords, salinity was low and both turbidity and nutrients from terrigenous sources were high, with benthic communities dominated by active suspension feeders (e.g., Bivalvia, Ascidiacea, and Bryozoa). Areas closer to the Pacific Ocean showed more oceanic conditions with higher salinity and lower turbidity, with benthic communities experiencing more open benthic physical space in which predators (e.g., Malacostraca and Asteroidea) and herbivorous browsers (e.g., Echinoidea and Gastropoda) were more conspicuous components of the community compared to the inner fjords. Hagfish (Myxine sp.) was the most abundant and frequently occurring fish taxa observed on deep-sea cameras (80% of deployments), along with several taxa of sharks (e.g., Squaliformes, Etmopteridae, Somniosidae, Scyliorhinidae), which collectively were also observed on 80% of deep-sea camera deployments. The kelp forests, deep fjords, and other nearshore habitats of the KNR represent a unique ecosystem with minimal human impacts at present. The KNR is part of the ancestral territory of the indigenous Kawésqar people and their traditional knowledge, including the importance of the land-sea connection in structuring the marine communities of this region, is strongly supported by our scientific findings.

Multi-tissue stable isotope analyses reveal temporal changes in the feeding patterns of green turtles in the Galapagos Marine Reserve.

Knowledge of feeding patterns of highly migratory species is critical for understanding their habitat use and informing the management of their populations. The Galapagos Islands are one of the most important nesting and feeding areas for green turtles (Chelonia mydas) across the tropical eastern Pacific, yet little is known about the feeding patterns of this species. The isotopic composition of different tissues has been used to gain insight into the trophic dynamics of mobile aquatic consumers whose trophic behavior is difficult to directly measure. To elucidate the temporal feeding patterns and isotopic niche sizes of Galapagos green turtles, stable isotope analyses were performed on multiple tissues (skin and carapace) collected at the two most important nesting areas in the archipelago: Bachas and Quinta Playa. The δ13 C and δ15 N signatures on the skin and carapace samples from 56 adult females revealed significant differences between tissues (p = .001 and p = .021, respectively) and nesting areas (p = .011 and p = .003, respectively). These differences suggest a shift from oceanic feeding grounds to neritic habitats before nesting. The carapace isotope values indicated an offshore feeding strategy and a greater isotopic niche (SEAc = 1.91‰2 ), whereas the skin isotope values represented an inshore feeding strategy with a narrower niche (SEAc = 1.37‰2 ), likely related to the consumption of specific coastal prey. Our results suggest that Galapagos green turtles feed across different habitats, and this information can be applied to improve the management of this endangered species.

Spatial patterns of continental shelf faunal community structure along the Western Antarctic Peninsula.

Knowledge of continental shelf faunal biodiversity of Antarctica is patchy and as such, the ecology of this unique ecosystem is not fully understood. To this end, we deployed baited cameras at 20 locations along ~ 500 km of the Western Antarctic Peninsula (WAP) at depths from 90 to 797 m. We identified 111 unique taxa, with mud bottom accounting for 90% of the dominant (≥ 50% cover) habitat sampled. Amphipoda comprised 41% of the total maximum number of individuals per camera deployment (MaxN) and occurred on 75% of deployments. Excluding this taxon, the highest MaxN occurred around King George/25 de Mayo Island and was driven primarily by the abundance of krill (Euphausiidae), which accounted for 36% of total average MaxN among deployments around this island. In comparison, krill comprised 22% of total average MaxN at Deception Island and only 10% along the peninsula. Taxa richness, diversity, and evenness all increased with depth and depth explained 18.2% of the variation in community structure among locations, which may be explained by decreasing ice scour with depth. We identified a number of Vulnerable Marine Ecosystem taxa, including habitat-forming species of cold-water corals and sponge fields. Channichthyidae was the most common fish family, occurring on 80% of all deployments. The Antarctic jonasfish (Notolepis coatsorum) was the most frequently encountered fish taxa, occurring on 70% of all deployments and comprising 25% of total MaxN among all deployments. Nototheniidae was the most numerically abundant fish family, accounting for 36% of total MaxN and was present on 70% of the deployments. The WAP is among the fastest warming regions on Earth and mitigating the impacts of warming, along with more direct impacts such as those from fishing, is critical in providing opportunities for species to adapt to environmental change and to preserve this unique ecosystem.

Arthropoda; Crustacea; Decapoda of deep-sea volcanic habitats of the Galapagos Marine Reserve, Tropical Eastern Pacific.

BACKGROUND: The deep-sea biome (> 200 m depth) is the world's last great wilderness, covering more than 65% of the earth's surface. Due to rapid technological advances, deep-sea environments are becoming more accessible to scientific research and ocean exploration around the world and, in recent years, this is also true for the Galapagos Islands. Deep-sea habitats cover the largest proportion of Galapagos Marine Reserve (GMR), yet to date, no comprehensive baseline exists on the biodiversity of the benthic fauna associated with volcanic seafloor formations within this region. Closing this knowledge gap is essential to provide information for decision-making for the management of marine resources within the GMR and assessing any potential changes in biodiversity resulting from climate-driven alterations that deep-sea environments are expected to experience. In 2015, the Charles Darwin Foundation's Seamounts of the GMR Research Project, together with the Galapagos National Park Directorate (GNPD) and Ocean Exploration Trust (OET), conducted a joint expedition on board the EV Nautilus. Using Remotely operated vehicles (ROVs), the aim of the expedition was to characterise the geological formations and biological communities present on seamounts, lava flows and other deep-sea habitats (> 200 m) within the GMR. NEW INFORMATION: We provide the first comprehensive image inventory for the phylum Arthropoda from 260 to 3400 m of depth within the GMR. Past studies on deep-sea macroinvertebrates in the GMR have been limited to voucher samples collected from dredging (restricted to soft bottom environments) or by submersibles (only allowing limited biological sampling). The image inventory, presented here, is based on high-definition video transects conducted by the Hercules ROV on board the EV Nautilus. Images of macroinvertebrate morphospecies were captured, catalogued and identified, thus providing the first known image inventory of in-situ macroinvertebrate species from the deep-sea region of the GMR.We present 32 distinct morphospecies occurrences within the class Malacostraca and order Decapoda. We also report 17 different families, three species that are new records to the GMR, in-situ images of two new species to science recently described and one possible new squat lobster, as well as interesting behavioural observations.

Characterization of deep-sea benthic invertebrate megafauna of the Galapagos Islands.

The deep sea represents the largest and least explored biome on the planet. Despite the iconic status of the Galapagos Islands and being considered one of the most pristine locations on earth, the deep-sea benthic ecosystems of the archipelago are virtually unexplored in comparison to their shallow-water counterparts. In 2015, we embarked on a multi-disciplinary scientific expedition to conduct the first systematic characterization of deep-sea benthic invertebrate communities of the Galapagos, across a range of habitats. We explored seven sites to depths of over 3,300 m using a two-part Remotely Operated Vehicle (ROV) system aboard the E/V Nautilus, and collected 90 biological specimens that were preserved and sent to experts around the world for analysis. Of those, 30 taxa were determined to be undescribed and new to science, including members of five new genera (2 sponges and 3 cnidarians). We also systematically analysed image frame grabs from over 85 h of ROV footage to investigate patterns of species diversity and document the presence of a range of underwater communities between depths of 290 and 3,373 m, including cold-water coral communities, extensive glass sponge and octocoral gardens, and soft-sediment faunal communities. This characterization of Galapagos deep-sea benthic invertebrate megafauna across a range of ecosystems represents a first step to study future changes that may result from anthropogenic impacts to the planet's climate and oceans, and informed the creation of fully protected deep-water areas in the Galapagos Marine Reserve that may help preserve these unique communities in our changing planet.

Sharks, rays, and chimaeras of the Revillagigedo National Park: An update of new and confirmed records.

The Revillagigedo Archipelago National Park is the largest no-take Marine Protected Area in North America and it is essential for the conservation of globally endangered marine megafauna, including cetaceans, sea birds and cartilaginous fishes. The present paper reports the occurrence of 28 chondrichthyan species, including 19 sharks, seven batoids and two chimaeras in the Revillagigedo Archipelago, comprising eight new records and a brief revision of confirmed species to date.

Kelp forests at the end of the earth: 45 years later.

The kelp forests of southern South America are some of the least disturbed on the planet. The remoteness of this region has, until recently, spared it from many of the direct anthropogenic stressors that have negatively affected these ecosystems elsewhere. Re-surveys of 11 locations at the easternmost extent of Tierra del Fuego originally conducted in 1973 showed no significant differences in the densities of adult and juvenile Macrocystis pyrifera kelp or kelp holdfast diameter between the two survey periods. Additionally, sea urchin assemblage structure at the same sites were not significantly different between the two time periods, with the dominant species Loxechinus albus accounting for 66.3% of total sea urchin abundance in 2018 and 61.1% in 1973. Time series of Landsat imagery of the region from 1998 to 2018 showed no long-term trends in kelp canopy over the past 20 years. However, ~ 4-year oscillations in canopy fraction were observed and were strongly and negatively correlated with the NOAA Multivariate ENSO index and sea surface temperature. More extensive surveying in 2018 showed significant differences in benthic community structure between exposed and sheltered locations. Fish species endemic to the Magellanic Province accounted for 73% of all nearshore species observed and from 98-100% of the numerical abundance enumerated at sites. Fish assemblage structure varied significantly among locations and wave exposures. The recent creation of the Yaganes Marine Park is an important step in protecting this unique and biologically rich region; however, the nearshore waters of the region are currently not included in this protection. There is a general lack of information on changes in kelp forests over long time periods, making a global assessment difficult. A complete picture of how these ecosystems are responding to human pressures must also include remote locations and locations with little to no impact.

A matter of taste: Spatial and ontogenetic variations on the trophic ecology of the tiger shark at the Galapagos Marine Reserve.

Sharks are top predators across ocean food webs and have a major ecological role in marine ecosystems. Investigating the trophic ecology of this group of species is thus essential to understand ecosystem functioning and inform specific management actions aimed at shark conservation. The Galapagos Islands represent one of the last ocean wildernesses, where populations of sharks and other top marine predators come close to a pristine status. Here we provide the first study on the trophic ecology of the tiger shark (Galeocerdo cuvier) within the Galapagos Marine Reserve (GMR), using a combination of stable isotope analysis, satellite tracking, and passive acoustic telemetry to investigate ontogenetic and spatial variations at two regions. The mean estimated δ13C and δ15N at Isabela island (western region) were -13.9 ± 0.5‰ and 13.7 ± 0.7‰; and for Santa Cruz island (central region) were -13.8 ± 0.3‰ and 13.4 ± 0.7‰, respectively. Green sea turtles (Chelonia mydas) were the main prey item for large tiger sharks (>280 cm TL), while smaller sharks mainly fed on squid and pelagic fish. Tiger sharks exhibited a high degree of philopatry around green sea-turtle nesting areas, with the majority of sharks detected around green sea-turtle nesting areas for at least 10 months after their capture date, and some individuals were even present during the entire three-year study period. Although we did not report statistically significant differences between the two regions, isotopic and electronic tagging data suggest that tiger sharks in the Galapagos could be segregated into specific populations separated by geographical scales of <100 km. The high productivity of the archipelago, along with the protection from industrial fishing granted by the GMR, result in abundant and predictable sources of prey. This high food abundance, combined with the presence of suitable habitats throughout the tiger shark life cycle, might result in a reduction of migratory behaviours when compared to movement patterns of tiger sharks in other ocean basins. Additional studies using genetic tools could provide further evidence on the presence of separate management units, as it has been recently revealed for other shark species inhabiting the GMR.

Marine biodiversity from zero to a thousand meters at Clipperton Atoll (Île de La Passion), Tropical Eastern Pacific.

Clipperton Atoll (Île de La Passion) is the only atoll in the Tropical Eastern Pacific (TEP) ecoregion and, owing to its isolation, possesses several endemic species and is likely an important stepping stone between Oceania, the remainder of the TEP, including other oceanic islands and the west coast of Central America. We describe the biodiversity at this remote atoll from shallow water to depths greater than one thousand meters using a mixture of technologies (SCUBA, stereo baited remote underwater video stations, manned submersible, and deep-sea drop cameras). Seventy-four unique taxa of invertebrates were identified during our expedition. The majority (70%) of these taxa were confined to the top 400 m and consisted mostly of sessile organisms. Decapod crustaceans and black corals (Antipatharia) had the broadest depth ranges, 100-1,497 m and 58-967 m, respectively. Decapods were correlated with the deepest depths, while hard corals were correlated with the shallow depths. There were 96 different fish taxa from 41 families and 15 orders, of which 70% were restricted to depths <200 m. While there was a decreasing trend in richness for both fish and invertebrate taxa with depth, these declines were not linear across the depth gradient. Instead, peaks in richness at ∼200 m and ∼750 m coincided with high turnover due to the appearance of new taxa and disappearance of other taxa within the community and is likely associated with the strong oxygen minimum zone that occurs within the region. The overall depth effect was stronger for fishes compared with invertebrates, which may reflect ecological preferences or differences in taxonomic resolution among groups. The creation of a no-take marine reserve 12 nautical miles around the atoll in 2016 will help conserve this unique and relatively intact ecosystem, which possesses high predator abundance.

Ecosystem effects of fishing & El Niño at the Galápagos Marine Reserve.

The Galápagos Archipelago is home to a diverse range of marine bioregions due to the confluence of several cold and warm water currents, resulting in some of the most productive tropical marine ecosystems in the world. These ecosystems are strongly influenced by El Niño events which can reduce primary production by an order of magnitude, dramatically reducing energy available throughout the food web. Fisheries are an important component of the local economy, although artisanal and illegal overfishing have dramatically reduced the productivity of invertebrate and finfish resources in recent decades, resulting in reductions in catches for local fishers. The regionally-endemic sailfin grouper (Myctereoperca olfax), locally known as bacalao, was once the most important fished species in the Galápagos, but is now listed as vulnerable by the IUCN due to its limited range and dramatic declines in catch over time. It is unknown how reduction of this predatory species has affected ecosystem structure and function. In the absence of stock assessments, we used an estimate of unfished bacalao biomass from fishers' ecological knowledge along with unfished biomass estimates of other heavily exploited stocks-lobster (Panulirus penicillatus and P. gracilis) and sea cucumber (Isostichopus fuscus)-to create historical, unfished versions of existing modern day ecosystem models. We used the unfished and modern versions of the ecosystem models to test the ecosystem effects of bacalao exploitation at the Bolivar Channel, located in the cold, west upwelling bioregion of the archipelago during both El Niño and non El Niño years, and at Floreana Island, in the warmer, central bioregion. Fishers' ecological knowledge indicates that at present, the biomass of bacalao is at least seven times lower than when unfished. This reduced bacalao biomass is linked with a greatly reduced ecosystem role compared to when unfished, and ecosystem role is further reduced in El Niño years. Allowing bacalao populations to rebuild to at least half of unfished biomass would partially restore their role within these ecosystems, while also resulting in greater fisheries catches. Comparing ecosystem impacts caused by fishing and El Niño, fishing has had a greater negative impact on bacalao ecosystem role than regular El Niño events.

Correction: Mangroves in the Galapagos islands: Distribution and dynamics.

[This corrects the article DOI: 10.1371/journal.pone.0209313.].

Mangroves in the Galapagos islands: Distribution and dynamics.

Mangrove forests provide valuable coastal protection from erosion, habitat for terrestrial and marine species, nursery grounds for commercial fisheries and are economically important for tourism. Galapagos' mangroves usually grow directly on solid lava and fragmented rocky shores, thereby stabilizing the sediment and facilitating colonisation by other plants and many animals. However, until very recently, only inaccurate data described mangrove coverage and its distribution. We mapped mangroves using freely available Google Earth Very High Resolution images based on on-screen classification and compared this method to three semi-automatic classification algorithms. We also analysed mangrove change for the period 2004-2014. We obtained an area of 3657.1 ha of fringing mangrove that covers 35% of the coastline. Eighty percent of mangrove cover is found in Isabela island, and 90% in the western and central south-eastern bioregions. The overall accuracy of mangrove classification was 99.1% with a Kappa coefficient of 0.97 when validated with field data. On-screen digitization was significantly more accurate than other tested methods. From the semi-automated methods, Maximum Likelihood Classification with prior land-sea segmentation yielded the best results. During the 2004-2014 period, mangrove coverage increased 24% mainly by expansion of existing mangroves patches as opposed to generation of new patches. We estimate that mangrove cover and growth are inversely proportional to the geological age of the islands. However, many other factors like nutrients, currents or wave exposure protection might explain this pattern. The precise localization of mangrove cover across the Galapagos islands now enables documenting whether it is changing over time.

El Niño drives a widespread ulcerative skin disease outbreak in Galapagos marine fishes.

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.

Deep-sea hydrothermal vents as natural egg-case incubators at the Galapagos Rift.

The discovery of deep-sea hydrothermal vents in 1977 challenged our views of ecosystem functioning and yet, the research conducted at these extreme and logistically challenging environments still continues to reveal unique biological processes. Here, we report for the first time, a unique behavior where the deep-sea skate, Bathyraja spinosissima, appears to be actively using the elevated temperature of a hydrothermal vent environment to naturally "incubate" developing egg-cases. We hypothesize that this behavior is directly targeted to accelerate embryo development time given that deep-sea skates have some of the longest egg incubation times reported for the animal kingdom. Similar egg incubating behavior, where eggs are incubated in volcanically heated nesting grounds, have been recorded in Cretaceous sauropod dinosaurs and the rare avian megapode. To our knowledge, this is the first time incubating behavior using a volcanic source is recorded for the marine environment.

Correction: Largest global shark biomass found in the northern Galápagos Islands of Darwin and Wolf.

[This corrects the article DOI: 10.7717/peerj.1911.].