The resources available for research are always limited. When setting priorities for research funding, governments, industry, and funding agencies must balance the demands of human health, food supply, and standards of living, against the less-tangible benefits of discovering more about the planet's biodiversity. Scientists have discovered almost 2 million species indicating that we have made great gains in our knowledge of biodiversity. However, this knowledge may distract attention from the estimated four-fifths of species on Earth that remain unknown to science, many of them inhabiting our oceans [1], [2]. The world's media still find it newsworthy when new species are discovered [1]. However, the extent of this taxonomic challenge no longer appears to be a priority in many funding agencies, perhaps because society and many scientists believe we have discovered most species, or that doing so is out of fashion except when new technologies are employed. Another symptom of this trend may be that the increased attention to novel methods available in molecular sciences is resulting in a loss of expertise and know-how in the traditional descriptive taxonomy of species [3]. The use of molecular techniques complements traditional methods of describing species but has not significantly increased the rate of discovery of new species (at least of fish), although it may help classify them [4]. At least in Europe, there was a mismatch between the number of species in a taxon and the number of people with expertise in it [5]. Unfortunately, because most species remain to be discovered in the most species-rich taxa [2], [5], [6], [7], there are then few experts to appreciate that this work needs to be done. Evidently, a global review of gaps in marine biodiversity knowledge and resources is overdue.

History of discovering marine biodiversity
Although the economic exploitation of marine resources dates back to prehistoric times, and historical documentation has existed since the third century B.C. with Aristotle's contributions in the Mediterranean Sea (e.g. [8]), the establishment of systematic collections of marine organisms began only during the seventeenth and eighteenth centuries. Global marine biodiversity investigations at these times depended not only on the availability of expertise, but also on foreign policies of the colonial powers of the time. For those reasons, the specimens collected from several regions (e.g., Caribbean, Japan, South America, Africa) were mostly brought to Europe, where they were described, deposited in museum collections, and used for the production of marine biological monographs. These early publications contained descriptions and checklists of many marine species, such as molluscs, crustaceans, fishes, turtles, birds, and mammals (e.g. [9], [10], [11]).

The history of research on marine biodiversity can generally be divided into three periods: early exploratory studies, local coastal “descriptive” studies, and large-scale multidisciplinary investigations and syntheses. These periods vary in timing by different seas and countries. The first exploratory studies in several regions (e.g., South America, Caribbean, South Africa, Pacific Ocean) took place from the mid-1700s until the late-1800s, in association with mainly European, North American, and Russian exploration expeditions, such as the Kamchatka Expedition in the 1740s, James Cook's voyages in the 1770s, the cruise of HMS Beagle in the 1830s, the voyage of HMS Challenger in the 1870s, and the first deep-sea investigations in the Mediterranean Sea [8], [9], [12], [13]. Pioneer investigations on deep-sea organisms were conducted in the Aegean Sea, where Forbes [14] noticed that sediments became progressively more impoverished in terms of biodiversity with increasing sampling depth. The azoic hypothesis proposed by Forbes suggested that life would be extinguished beyond 500 m depth, although a work published 68 years earlier provided indisputable evidence of the presence of life in the Gulf of Genoa at depths down to 1,000 m [15].

The taxonomists who described marine species at these times seldom collected specimens themselves in the field and, therefore, had only second-hand information about the distribution and ecology of the samples they received [4], [8]. Some of the early descriptions of tropical species thus do not even have the locality where the holotype or voucher material was collected (some examples in Chenu 1842–1853). The second period of regional studies was initiated by enhanced availability of research resources (experts, institutes, and vessels) in developing countries around the mid-1900s. The earliest institutions and research stations, many of which continue to operate, were founded in some areas as early as the late 1800s and early 1900s (e.g. [11], [16], [17]). Wide-scale establishment of laboratories in several continents (Europe, New Zealand, North and South America) have only been operational since the 1950s–1960s. The third stage, large-scale multidisciplinary investigations, has evolved since the 1990s, and is related to development and application of modern technologies and implementation of large, multinational research projects. Perhaps the largest of such investigations was the Census of Marine Life (Census).

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A census of marine biodiversity knowledge, resources, and future challenges.

The freshwater and marine fish faunas of South America are the most diverse on Earth, with current species richness estimates standing above 9100 species. In addition, over the last decade at least 100 species were described every year. There are currently about 5160 freshwater fish species, and the estimate for the freshwater fish fauna alone points to a final diversity between 8000 and 9000 species. South America also has c. 4000 species of marine fishes. The mega-diverse fish faunas of South America evolved over a period of >100 million years, with most lineages tracing origins to Gondwana and the adjacent Tethys Sea. This high diversity was in part maintained by escaping the mass extinctions and biotic turnovers associated with Cenozoic climate cooling, the formation of boreal and temperate zones at high latitudes and aridification in many places at equatorial latitudes. The fresh waters of the continent are divided into 13 basin complexes, large basins consolidated as a single unit plus historically connected adjacent coastal drainages, and smaller coastal basins grouped together on the basis of biogeographic criteria. Species diversity, endemism, noteworthy groups and state of knowledge of each basin complex are described. Marine habitats around South America, both coastal and oceanic, are also described in terms of fish diversity, endemism and state of knowledge. Because of extensive land use changes, hydroelectric damming, water divergence for irrigation, urbanization, sedimentation and overfishing 4-10% of all fish species in South America face some degree of extinction risk, mainly due to habitat loss and degradation. These figures suggest that the conservation status of South American freshwater fish faunas is better than in most other regions of the world, but the marine fishes are as threatened as elsewhere. Conserving the remarkable aquatic habitats and fishes of South America is a growing challenge in face of the rapid anthropogenic changes of the 21st century, and deserves attention from conservationists and policy makers.

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Fish biodiversity and conservation in South America.

Large rivers create major gaps in reef distribution along tropical shelves. The Amazon River represents 20% of the global riverine discharge to the ocean, generating up to a 1.3 × 106–km2 plume, and extensive muddy bottoms in the equatorial margin of South America. As a result, a wide area of the tropical North Atlantic is heavily affected in terms of salinity, pH, light penetration, and sedimentation. Such unfavorable conditions were thought to imprint a major gap in Western Atlantic reefs. We present an extensive carbonate system off the Amazon mouth, underneath the river plume. Significant carbonate sedimentation occurred during lowstand sea level, and still occurs in the outer shelf, resulting in complex hard-bottom topography. A permanent near-bottom wedge of ocean water, together with the seasonal nature of the plume’s eastward retroflection, conditions the existence of this extensive (~9500 km2) hard-bottom mosaic. The Amazon reefs transition from accretive to erosional structures and encompass extensive rhodolith beds. Carbonate structures function as a connectivity corridor for wide depth–ranging reef-associated species, being heavily colonized by large sponges and other structure-forming filter feeders that dwell under low light and high levels of particulates. The oxycline between the plume and subplume is associated with chemoautotrophic and anaerobic microbial metabolisms. The system described here provides several insights about the responses of tropical reefs to suboptimal and marginal reef-building conditions, which are accelerating worldwide due to global changes.

https://advances.sciencemag.org/content/advances/2/4/e1501252.full.pdf

An extensive reef system at the Amazon River mouth

The European, Canadian, and Latin American seaweed industries rely on the sustainable harvesting of natural resources. As several countries wish to increase their activity, the harvest should be managed according to integrated and participatory governance regimes to ensure production within a long-term perspective. Development of regulations and directives enabling the sustainable exploitation of natural resources must therefore be brought to the national and international political agenda in order to ensure environmental, social, and economic values in the coastal areas around the world. In Europe, Portugal requires an appraisal of seaweed management plans while Norway and Canada have developed and implemented coastal management plans including well-established and sustainable exploitation of their natural seaweed resources. Whereas, in Latin America, different scenarios of seaweed exploitation can be observed; each country is however in need of long-term and ecosystem-based management plans to ensure that exploitation is sustainable. These plans are required particularly in Peru and Brazil, while Chile has succeeded in establishing a sustainable seaweed-harvesting plan for most of the economically important seaweeds. Furthermore, in both Europe and Latin America, seaweed aquaculture is at its infancy and development will have to overcome numerous challenges at different levels (i.e., technology, biology, policy). Thus, there is a need for regulations and establishment of “best practices” for seaweed harvesting, management, and cultivation. Trained human resources will also be required to provide information and education to the communities involved, to enable seaweed utilization to become a profitable business and provide better income opportunities to coastal communities.

https://link.springer.com/content/pdf/10.1007%2Fs10811-014-0304-8.pdf

Seaweeds: an opportunity for wealth and sustainable livelihood for coastal communities

Pelagic provinces of the world: A biogeographic classification of the world’s surface pelagic waters

The South American region
The marine areas of the South American continent extend for almost 30,000 km of coastline and encompass three different oceanic domains—the Caribbean, the Pacific, and the Atlantic. The latitudinal and longitudinal ranges within this region are similarly wide, from 12°N to 55°S, and from 34° to 81°W. Ten countries border on these coasts, each with different research capabilities and taxonomic traditions; therefore, taxonomic knowledge differs among countries. Coastal biodiversity is strongly influenced by the physical and geological history of these coasts. The eastern tropical Pacific region, which encompasses the continental coasts of southern Central America (Costa Rica and Panama) and of northwestern South America (Colombia and Ecuador) is characterized by cliffs alternating with pocket beaches, alluvial and deltaic plains with extensive sandy beaches, well-developed mangrove forests, estuaries, lagoons, and, reefs. It also includes important offshore island systems such as the Pearl and Galapagos islands [1], [2]. The Peruvian coast also is diverse with bays, cliffs, kelp and macroalgal beds, rocky shores and sandy beaches, islands, and peninsulas, as well as wetlands, which include the southernmost limit to the tropical Pacific mangrove ecosystem [3], [4]. The Chilean coast is 4,500 km of mainly rocky shores, but does include some sandy-beach bays with channels and archipelagos toward the south (Patagonian region) [5], [6]. Some of the most diverse ecosystems in Chile are the beds of kelp (Lessonia and Macrosystis) and macroalgae (Gracillaria and Ulva). The combination of the unique oceanographic conditions and coastal heterogeneity in the Chilean coast has resulted in high levels of endemism (near 40%) in many invertebrate groups [5], and several marine invertebrate taxa show latitudinal biodiversity patterns, some of them explained by the presence of Antarctic fauna [7]–[9]. Ecuador, Peru, and Chile are under the influence of the Humboldt upwelling system and subject to high environmental variability caused by the ENSO (El Nino Southern Oscillation) and LNSO (La Nina Southern Oscillation), which cause important changes in community composition and abundance, particularly of the plankton [1], [10].

The Atlantic coast of the South American continent is distinctly different from the Pacific coast. It includes three major rivers (Orinoco, Amazon, and La Plata), which discharge enormous amounts of freshwater and sediment to the ocean, and the coast has an extensive continental platform. Argentina's coast has mostly sandy beaches [6], [11] and some rocky formations located mainly at Mar del Plata and at Peninsula Valdes. At Mar del Plata, these rocky shores are dominated by two mussel species and by a diverse macroalgal community with a clear tidal zonation [12], [13]. The Uruguayan coast is dominated by sandy beaches with a narrow portion of rocky habitats known to sustain a rich biological diversity [14]. Observed variations in community composition and distribution may be related to the salinity gradient caused by La Plata River discharge [15].

The coast of Brazil, extending almost 7,500 km, is under the influence of the warm Brazil Current, the cold Malvinas/Falklands Current, and many rivers and upwelling regions [16]. The warm northern coast, where the Amazon discharges into the ocean, is characterized by a combination of freshwater, estuarine, and marine ecosystems, with diverse but poorly known habitats [17]. The colder southern coast is characterized by a variety of ecosystems, including mangrove forests, seagrass beds, coral reefs, sandy beaches, rocky shores, lagoons, and estuaries. Because of its vastness, extensive areas of Brazil's coast remain unexplored. North of Brazil are Suriname, French Guiana, Guyana, and the Venezuelan Atlantic Front. This area, including about 1,900 km of coastline, is under the strong influence of the Amazon River. Therefore, the typical ecosystems are estuaries, mudflats, sandy beaches, and mangrove forests, which extend along most of the coastline [18]. The Venezuelan Atlantic coast is also under the influence of the Orinoco River, with coastal mudflats and extensive mangrove forests [19].

In this paper, we analyze the status of knowledge of marine biodiversity in five subregions along the Atlantic and Pacific coasts of South America. As most of the information is based in national reports, these subregions were based in the Large Marine Ecosystem boundaries as defined for South America, with a few practical adaptations, based in country political borders. The paper also provides an updated review of ecosystem threats, such as invasive species, and the marine conservation strategies employed by South American countries with access to the coast, excluding the Caribbean coasts of Venezuela and Colombia, as these are included in another paper of this collection [20].

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Felipe Artigas

Papers

Marine biodiversity in the Atlantic and Pacific coasts of South America: knowledge and gaps.