Patterns and drivers of benthic macroinvertebrate assemblages in the kelp forests of southern Patagonia.

The kelp forests of southern Patagonia have a large diversity of habitats, with remote islands, archipelagos, peninsulas, gulfs, channels, and fjords, which are comprised of a mixture of species with temperate and sub-Antarctic distributions, creating a unique ecosystem that is among the least impacted on Earth. We investigated the distribution, diversity, and abundance of marine macroinvertebrate assemblages from the kelp forests of southern Patagonia over a large spatial scale and examined the environmental drivers contributing to the observed patterns in assemblage composition. We analyzed data from 120 quantitative underwater transects (25 x 2 m) conducted within kelp forests in the southern Patagonian fjords in the Kawésqar National Reserve (KNR), the remote Cape Horn (CH) and Diego Ramírez (DR) archipelagos of southern Chile, and the Mitre Peninsula (MP) and Isla de los Estados (IE) in the southern tip of Argentina. We observed rich assemblages of macroinvertebrates among these kelp forests, with a total of 185 unique taxa from 10 phyla and 23 classes/infraorders across the five regions. The number of taxa per transect was highest at IE, followed by MP, CH, and KNR, with the lowest number recorded at DR. The trophic structure of the macroinvertebrate assemblages was explained mostly by wave exposure (28% of the variation), followed by salinity (12%) and the KNR region (11%). KNR was most distinct from the other regions with a greater abundance of deposit feeders, likely driven by low salinity along with high turbidity and nutrients from terrigenous sources and glacial melt. Our study provides the first broad-scale description of the benthic assemblages associated with kelp forests in this vast and little-studied region and helps to establish baselines for an area that is currently lightly influenced by local anthropogenic factors and less impacted by climate change compared with other kelp forests globally.

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.

Assemblage structure and spatial diversity patterns of kelp forest-associated fishes in Southern Patagonia.

Knowledge of the ecology of the fish fauna associated with kelp (primarily Macrocystis pyrifera) forests in Southern Patagonia is scarce, especially in how abiotic and biotic variables influence their structure, diversity, and distribution. This information is important for the management and conservation of this unique ecosystem, which has minimal anthropogenic impacts at present. We analyzed data from 122 quantitative underwater transects conducted within kelp forests at 61 stations from Chile's southern Patagonian fjords to the Cape Horn and Diego Ramirez archipelagos and the southern tip of Argentina, including the Mitre Peninsula and Isla de los Estados. In total, 25 fish species belonging to 13 families were observed. Multivariate analysis indicated that there are significant differences in fish assemblage structure among locations and wave exposures, which was driven primarily by Patagonotothen sima and Paranotothenia magellanica, which occurred on exposed and semi-exposed stations. P. cornucola was mainly distributed across sheltered stations of the Kawésqar National Park. Temperature, salinity, depth, and kelp density influenced fish assemblage structure, with the highest diversity in areas with the lowest temperature and greater depth at Isla de los Estados. In contrast, species richness, diversity, abundance, and biomass were all lower in areas with high density of the understory kelp Lessonia spp., which might be driven by the absence of P. tessellata, P. squamiceps and P. cornucola, the most important species in terms of occurrence, abundance, and biomass. Our study provides the first broad-scale description of the fish assemblages associated with kelp forests along the southern cone of South America based on non-invasive visual transects, improving our knowledge of the distribution of fish assemblages across several environmental conditions in this vast and little-studied area.

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.

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.

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

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

Marine biodiversity at the end of the world: Cape Horn and Diego Ramírez islands.

The vast and complex coast of the Magellan Region of extreme southern Chile possesses a diversity of habitats including fjords, deep channels, and extensive kelp forests, with a unique mix of temperate and sub-Antarctic species. The Cape Horn and Diego Ramírez archipelagos are the most southerly locations in the Americas, with the southernmost kelp forests, and some of the least explored places on earth. The giant kelp Macrocystis pyrifera plays a key role in structuring the ecological communities of the entire region, with the large brown seaweed Lessonia spp. forming dense understories. Kelp densities were highest around Cape Horn, followed by Diego Ramírez, and lowest within the fjord region of Francisco Coloane Marine Park (mean canopy densities of 2.51 kg m-2, 2.29 kg m-2, and 2.14 kg m-2, respectively). There were clear differences in marine communities among these sub-regions, with the lowest diversity in the fjords. We observed 18 species of nearshore fishes, with average species richness nearly 50% higher at Diego Ramírez compared with Cape Horn and Francisco Coloane. The number of individual fishes was nearly 10 times higher at Diego Ramírez and 4 times higher at Cape Horn compared with the fjords. Dropcam surveys of mesophotic depths (53-105 m) identified 30 taxa from 25 families, 15 classes, and 7 phyla. While much of these deeper habitats consisted of soft sediment and cobble, in rocky habitats, echinoderms, mollusks, bryozoans, and sponges were common. The southern hagfish (Myxine australis) was the most frequently encountered of the deep-sea fishes (50% of deployments), and while the Fueguian sprat (Sprattus fuegensis) was the most abundant fish species, its distribution was patchy. The Cape Horn and Diego Ramírez archipelagos represent some of the last intact sub-Antarctic ecosystems remaining and a recently declared large protected area will help ensure the health of this unique region.

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

Overfishing has dramatically depleted sharks and other large predatory fishes worldwide except for a few remote and/or well-protected areas. The islands of Darwin and Wolf in the far north of the Galapagos Marine Reserve (GMR) are known for their large shark abundance, making them a global scuba diving and conservation hotspot. Here we report quantitative estimates of fish abundance at Darwin and Wolf over two consecutive years using stereo-video surveys, which reveal the largest reef fish biomass ever reported (17.5 t [Formula: see text] on average), consisting largely of sharks. Despite this, the abundance of reef fishes around the GMR, such as groupers, has been severely reduced because of unsustainable fishing practices. Although Darwin and Wolf are within the GMR, they were not fully protected from fishing until March 2016. Given the ecological value and the economic importance of Darwin and Wolf for the dive tourism industry, the current protection should ensure the long-term conservation of this hotspot of unique global value.

Marine Biodiversity in Juan Fernández and Desventuradas Islands, Chile: Global Endemism Hotspots.

The Juan Fernández and Desventuradas islands are among the few oceanic islands belonging to Chile. They possess a unique mix of tropical, subtropical, and temperate marine species, and although close to continental South America, elements of the biota have greater affinities with the central and south Pacific owing to the Humboldt Current, which creates a strong biogeographic barrier between these islands and the continent. The Juan Fernández Archipelago has ~700 people, with the major industry being the fishery for the endemic lobster, Jasus frontalis. The Desventuradas Islands are uninhabited except for a small Chilean military garrison on San Félix Island. We compared the marine biodiversity of these islands across multiple taxonomic groups. At San Ambrosio Island (SA), in Desventuradas, the laminarian kelp (Eisenia cokeri), which is limited to Desventuradas in Chile, accounted for >50% of the benthic cover at wave exposed areas, while more sheltered sites were dominated by sea urchin barrens. The benthos at Robinson Crusoe Island (RC), in the Juan Fernández Archipelago, comprised a diverse mix of macroalgae and invertebrates, a number of which are endemic to the region. The biomass of commercially targeted fishes was >2 times higher in remote sites around RC compared to sheltered locations closest to port, and overall biomass was 35% higher around SA compared to RC, likely reflecting fishing effects around RC. The number of endemic fish species was extremely high at both islands, with 87.5% of the species surveyed at RC and 72% at SA consisting of regional endemics. Remarkably, endemics accounted for 99% of the numerical abundance of fishes surveyed at RC and 96% at SA, which is the highest assemblage-level endemism known for any individual marine ecosystem on earth. Our results highlight the uniqueness and global significance of these biodiversity hotspots exposed to very different fishing pressures.

Long-term effectiveness of a multi-use marine protected area on reef fish assemblages and fisheries landings.

The Loreto Bay National Park (LBNP) is a large, multi-use marine protected area in the Gulf of California, Mexico, where several types of small-scale commercial and recreational fishing are allowed, but where less than 1% of the park is totally protected from fishing. The LBNP was created in 1996; its management plan was completed in 2000, but it was not effectively implemented and enforced until 2003. Between 1998 and 2010, we monitored reef fish populations annually at several reefs inside and outside the LBNP to measure the effects of the park on fish assemblages. We also evaluated reported fisheries landings within the LBNP for the same time series. Our results show that reef fish biomass increased significantly after protection at a small no-take site at LBNP relative to the rest of the park. However, the multi-use part of LBNP where fishing is allowed (99% of its surface) has had no measurable effect on reef fish biomass relative to open access sites outside the park boundaries. Reported fisheries landings have decreased within the park while increasing in nearby unprotected areas. Although the current partial protection management regime has not allowed for reef fish populations to recover despite 15 years as a "protected area," we conclude that LBNP's regulations and management have maintained the conditions of the ecosystem that existed when the park was established. These results suggest that community livelihoods have been sustained, but a re-evaluation of the multi-use management strategy, particularly the creation of larger no-take zones and better enforcement, is needed to improve the reef fish populations in the park in order to ensure sustainable fisheries far into the future. These recommendations can be applied to all multi-use MPAs in Mexico where ecosystem recovery is not occurring despite maintenance of fish stocks.

Deepwater fish assemblages at Isla del Coco National Park and Las Gemelas Seamount, Costa Rica / Conjuntos de peces de aguas profundas en el Parque Nacional Isla del Coco y el Monte Submarino Las Gemelas, Costa Rica

The deepwater faunas of oceanic islands and seamounts of the Eastern Tropical Pacific are poorly known. From 11-22 September 2009 we conducted an exploration of the deepwater areas of the Isla del Coco Marine Conservation Area, Costa Rica and a nearby seamount using a manned submersible. The goal of the exploration was to characterize the habitats and biota, and conduct quantitative surveys of the deepwater portions of Isla del Coco National Park and Las Gemelas Seamount, located about 50km southwest of Isla del Coco. We completed a total of 22 submersible dives, spanning more than 80hr underwater, and collected a total of 36hr of video. We surveyed habitats from 50-402m and observed more than 45 species of fishes, some of which have not yet been described and are likely new to science. The diversity of fish species in deep water at Isla del Coco National Park was lower than the diversity of fishes in shallow water, and eight species groups accounted for more than 95% of the total fish biomass. The combined density of all fish species was higher at Las Gemelas Seamount (253 fishes/100m²) than at Isla del Coco National Park (138 fishes/100m²). The combined density of fishes in habitats comprised primarily of bedrock or large boulders outcrops was more than three times as high at Las Gemelas Seamount as it was at Isla del Coco National Park. This discrepancy was caused by the extremely high concentration of Anthiinae fishes in rocky habitats at Las Gemelas Seamount. Densities of fishes in the other habitats were similar between the two sites. Similarly, when estimates of fish density were plotted by slope categories the density was much greater on steep slopes, which were usually comprised of rock habitats. Also, the density of fishes was greatest on high rugosity habitats. Results of these submersible surveys indicate that seamounts in the tropical eastern Pacific Ocean may be an important source of biodiversity and that more quantitative surveys are needed to characterize the fauna of the region.

Las faunas de aguas profundas de islas oceánicas del Pacífico Tropical Oriental se conocen poco y de los montes submarinos nada. Del 11 al 22 de septiembre de 2009 llevamos a cabo una exploración de zonas profundos del Área de Conservación Marina Isla del Coco utilizando un submarino tripulado. El objetivo del estudio fue caracterizar los hábitats y las comunidades, y cuantificar las poblaciones de peces de profundidad en la Isla del Coco y los montes submarinos Las Gemelas, situados a 50km al suroeste de la Isla del Coco. Realizamos 22 inmersiones con el submarino, con más de 80 horas de observación submarina, y filmamos 30 horas de video. Investigamos hábitats entre 50-402m de profundidad y observamos más de 45 especies de peces, algunas de las cuales son especies nuevas para la ciencia. La diversidad de peces profundos en la Isla del Coco fue menor que en aguas someras, y ocho grupos de especies representaron más del 95% de la biomasa total de peces. La densidad combinada de peces fue 253 peces/100m² en Las Gemelas y 138 peces/100m² en la Isla del Coco.

The shallow-water fish assemblage of Isla del Coco National Park, Costa Rica: structure and patterns in an isolated, predator-dominated ecosystem / El conjunto de peces de aguas poco profundas del Parque Nacional Isla del Coco, Costa Rica: estructura y patrones en un ecosistema aislado dominado por depredadores

Fishes at Isla del Coco National Park, Costa Rica, were surveyed as part of a larger scientific expedition to the area in September 2009. The average total biomass of nearshore fishes was 7.8 tonnes per ha, among the largest observed in the tropics, with apex predators such as sharks, jacks, and groupers accounting for nearly 40% of the total biomass. The abundance of reef and pelagic sharks, particularly large aggregations of threatened species such as the scalloped hammerhead shark (up to 42 hammerheads ha-1) and large schools of jacks and snappers show the capacity for high biomass in unfished ecosystems in the Eastern Tropical Pacific. However, the abundance of hammerhead and reef whitetip sharks appears to have been declining since the late 1990s, and likely causes may include increasing fishing pressure on sharks in the region and illegal fishing inside the Park. One Galapagos shark tagged on September 20, 2009 in the Isla del Coco National Park moved 255km southeast towards Malpelo Island in Colombia, when it stopped transmitting. These results contribute to the evidence that sharks conduct large-scale movements between marine protected areas (Isla del Coco, Malpelo, Galápagos) in the Eastern tropical Pacific and emphasize the need for regional-scale management. More than half of the species and 90% of the individuals observed were endemic to the tropical eastern Pacific. These high biomass and endemicity values highlight the uniqueness of the fish assemblage at Isla del Coco and its importance as a global biodiversity hotspot.

La biomasa promedio de peces costeros en el Parque Nacional Isla del Coco en septiembre de 2010 fue de 7,8 toneladas por hectárea, entre las más elevadas halladas jamás en zonas tropicales. Los grandes depredadores representaron el 40% de la biomasa total. La abundancia de tiburones costeros y pelágicos, particularmente las enormes agregaciones de tiburón martillo (hasta 42 individuos por hectárea) y los extensos bancos de carángidos y lutjánidos, muestran la capacidad que tienen los ecosistemas marinos no pescados para albergar elevadas biomasas de peces, y hacen de la Isla del Coco un lugar único en el mundo. No obstante, la abundancia de tiburones parece estar decreciendo desde 1999, probablemente a causa de la creciente presión pesquera en la región y la pesca ilegal en el interior del Parque. Un tiburón de Galápagos marcado se dirigió 255km en dirección a la Isla de Malpelo, Colombia. Estos resultados sugieren que los tiburones realizan importantes movimientos entre áreas marinas protegidas (Isla del Coco, Malpelo, Galápagos) en el Pacífico Tropical Oriental y remarcan la necesidad de una gestión regional de estos animales. Más del 50% de las especies y el 90% de individuos observados en los contajes eran endémicos del Pacífico Tropical Oriental.

Large recovery of fish biomass in a no-take marine reserve.

No-take marine reserves are effective management tools used to restore fish biomass and community structure in areas depleted by overfishing. Cabo Pulmo National Park (CPNP) was created in 1995 and is the only well enforced no-take area in the Gulf of California, Mexico, mostly because of widespread support from the local community. In 1999, four years after the establishment of the reserve, there were no significant differences in fish biomass between CPNP (0.75 t ha(-1) on average) and other marine protected areas or open access areas in the Gulf of California. By 2009, total fish biomass at CPNP had increased to 4.24 t ha(-1) (absolute biomass increase of 3.49 t ha(-1), or 463%), and the biomass of top predators and carnivores increased by 11 and 4 times, respectively. However, fish biomass did not change significantly in other marine protected areas or open access areas over the same time period. The absolute increase in fish biomass at CPNP within a decade is the largest measured in a marine reserve worldwide, and it is likely due to a combination of social (strong community leadership, social cohesion, effective enforcement) and ecological factors. The recovery of fish biomass inside CPNP has resulted in significant economic benefits, indicating that community-managed marine reserves are a viable solution to unsustainable coastal development and fisheries collapse in the Gulf of California and elsewhere.

Mangroves in the Gulf of California increase fishery yields.

Mangroves are disappearing rapidly worldwide despite their well documented biodiversity and the ecosystem services they provide. Failure to link ecological processes and their societal benefits has favored highly destructive aquaculture and tourism developments that threaten mangroves and result in costly "externalities." Specifically, the potentially irreparable damage to fisheries because of mangrove loss has been belittled and is greatly underestimated. Here, we show that, in the Gulf of California, fisheries landings are positively related to the local abundance of mangroves and, in particular, to the productive area in the mangrove-water fringe that is used as nursery and/or feeding grounds by many commercial species. Mangrove-related fish and crab species account for 32% of the small-scale fisheries landings in the region. The annual economic median value of these fisheries is US $37,500 per hectare of mangrove fringe, falling within the higher end of values previously calculated worldwide for all mangrove services together. The ten-year discounted value of one hectare of fringe is >300 times the official cost set by the Mexican government. The destruction of mangroves has a strong economic impact on local fishing communities and on food production in the region. Our valuation of the services provided by mangroves may prove useful in making appropriate decisions for a more efficient and sustainable use of wetlands.

Viability analysis of reef fish populations based on limited demographic information.

Marine protected areas (MPAs) that allow some degree of artisanal fishing have been proposed to control the overexploitation of marine resources while allowing extraction by local communities. Nevertheless, the management of MPAs is often impaired by the absence of data on the status of their resources. We devised a method to estimate population growth rates with the type of data that are usually available for reef fishes. We used 7 years of spatially explicit abundance data on the leopard grouper (Mycteroperca rosacea) in an MPA in the Gulf of California, Mexico, to construct a matrix population model that incorporated the effects of El Niño/La Niña Southern Oscillation on population dynamics. An environmental model that estimated different demographic estimates for El Niño and La Niña periods performed better than a single-environment model, and a single-habitat model performed better than a model that considered different depths as different habitats. Our results suggest that the population of the leopard grouper off the main island of the MPA is not viable under present conditions. Although the impact of fishing on leopard grouper populations in the MPA has not yet been established, fishing should be closed as a precautionary measure at this island if a priority of the MPA is to ensure the sustainability of its fish populations.