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Rebentos Rhodoliths Beds Review Article / Artigo de Revisão Horta et al.: BrazilianReBentos rhodoliths beds Rhodoliths in Brazil: Current knowledge and potential impacts of climate change Paulo Antunes Horta1*, Pablo Riul2, Gilberto M. Amado Filho3, Carlos Frederico D. Gurgel1, Flávio Berchez4, José Marcos de Castro Nunes5, Fernando Scherner6, Sonia Pereira6, Tito Lotufo7, Letícia Peres1, Marina Sissini1, Eduardo de Oliveira Bastos1, João Rosa1, Pamela Munoz1, Cintia Martins1, Lidiane Gouvêa1, Vanessa Carvalho1, Ellie Bergstrom1, Nadine Schubert8, Ricardo G. Bahia3, Ana Claudia Rodrigues1, Leonardo Rörig1, José Bonomi Barufi1, Marcia Figueiredo3 1 Departamento de Botânica, Centro de Ciências Biológicas Universidade Federal de Santa Catarina (CEP:88040-970, Florianópolis, SC, Brasil) 2 Departamento de Engenharia e Meio Ambiente, Universidade Federal da Paraíba (CEP: 58297-000, Rio Tinto, PB, Brasil) 3 Instituto de Pesquisas Jardim Botânico do Rio de Janeiro (CEP: 22460-030, Rio de Janeiro, RJ, Brasil) 4 Laboratório de Algas Marinhas, Instituto de Biociências, Universidade de São Paulo (CEP: 05508-090, São Paulo, SP, Brasil) 5 Laboratório de Algas Marinhas, Instituto de Biologia, Universidade Federal da Bahia (CEP: 40210-730, Salvador, BA, Brasil) 6 Laboratório de Ficologia, Departamento de Biologia, Universidade Federal Rural de Pernambuco (CEP: 52171-900, Recife, PE, Brasil) 7 Instituto de Oceanografia, Universidade de São Paulo (CEP: 05508-090, São Paulo, SP, Brasil) 8 Programa de Pós-Graduação em Oceanografia, Departamento de Geociências, Universidade Federal de Santa Catarina(UFSC) (CEP: 88040-900, Florianópolis, SC, Brasil) *Corresponding author: [email protected] Financial Support: CNPQ, CAPES, FAPESP. AbstrAct resumo Rhodolith beds are important marine benthic ecosystems, Bancos de rodolitos formam oásis de alta biodiversidade em representing oases of high biodiversity among sedimentary ambientes marinhos de fundo arenoso. Os rodolitos são formados seabed environments. They are found frequently and abundantly, por espécies bioconstrutoras, que fornecem abrigo e substrato acting as major carbonate ’factories’ and playing a key role in para diversas e abundantes comunidades bentônicas. No Brasil the biogeochemical cycling of carbonates in the South Atlantic. esses ambientes são frequentes, representando grandes “fábricas” Rhodoliths are under threat due to global change (mainly related de carbonato com um papel fundamental no ciclo biogeoquímico to ocean acidification and global warming) and local stressors, do carbono no Atlântico Sul. Estes organismos e ambientes such as fishing and coastal run-off. Here, we review different estão ameaçados pelas mudanças climáticas (principalmente aspects of the biology of these organisms, highlighting the a acidificação dos oceanos e o aquecimento global) e pelos predicted effects of global change, considering the additional estressores locais, tais como os impactos causados pela pesca e impact of local stressors. Ocean acidification (OA) represents as descargas costeiras de efluentes. Neste trabalho fazemos uma a particular threat that can reduce calcification or even promote revisão da taxonomia, filogenia e biologia desses organismos, the decalcification of these bioengineers, thus increasing the eco- com destaque para os efeitos previstos das mudanças climáticas physiological imbalance between calcareous and fleshy algae. OA e suas relações com estressores locais. A maioria dos estudos should be considered, but this together with extreme events such acerca dos efeitos das mudanças climáticas e da acidificação as heat waves and storms, as main stressors of these ecosystems dos oceanos em algas calcárias relatam respostas negativas não at the present time, will worsen in the future, especially if só no crescimento e na calcificação, mas também no processo possible interactions with local stressors like coastal pollution de fotossíntese, espessura da parede celular, reprodução e are taken into consideration. Thus, in Brazil there is a serious sobrevivência das algas. Este cenário reforça a necessidade em need for starting monitoring programs and promote innovative estabelecer uma consistente rede de trabalho para proporcionar experimental infrastructure in order to improve our knowledge um programa de monitoramento amplo e de longo prazo, bem of these rich environments, optimize management efforts and como infraestrutura para avaliações experimentais de impactos enhance the needed conservation initiatives. locais e regionais das mudanças climáticas e dos estressores locais em bancos de rodolitos. Descriptors: Algae, Conservation, Brazil, Global warming, Pollution, Ocean acidification. Descritores: Algas, Conservação, Brasil, Mudanças climáticas, Poluição, Acidificação dos oceanos http://dx.doi.org/10.1590/S1679-875920160870064sp2 BRAZILIAN JOURNAL OF OCEANOGRAPHY, 64(sp2):117-136;2016 117 Horta et al.: Brazilian rhodoliths beds INTRODUCTION Rhodoliths represent heavily calcified organisms, depositing calcium carbonate in the form of high Mg- Pink or red “rocks” on the sea floor often consist of calcite, a mineral that attains up to ~90 % of the dry weight different morphotypes of a particular kind of calcareous of these species (WOELKERLING, 1993; OLIVEIRA, red algae (Corallinophycidae). This particular kind of 1997) and ensures good preservation in fossil record. These seaweed, known as coralline red algae, present calcium features call the attention not only of phycologists, but also carbonate impregnated (i.e. calcified) cell walls. palaeontologists and geologists interested in the carbonate Crustose coralline red algae are completely calcified abundance of limestone deposits found since the Cretaceous encrusting organisms (Figure 1) that either adhere (LITTLER, 1972; BASSO, 2012). It has been estimated that tightly to hard substratum or remain unattached to the the Brazilian continental shelf carbonate stock holds 2 x 1011 seafloor (BROOM et al., 2008). Rhodoliths (including tonnes of CaCO (MILLIMANN; AMARAL, 1974). Among maërl) can be defined as calcareous nodules composed 3 the rhodolith beds along the Brazilian coast, the Abrolhos of more than 50% of coralline red algal material and shelf boasts the world’s largest expansion of rhodoliths in consisting of one to several coralline species growing the Atlantic, with a mean relative cover of rhodoliths around together (BOSELLINI; GINSBURG, 1971). They may 69.1% (± 1.7%) and a mean density of 211 ± 20 nodules per cover extensive areas of the seafloor forming large square meter (AMADO-FILHO et al., 2012a). These authors beds, often associated with high marine biodiversity estimated a mean CaCO production rate of about 0.025 Gt/ (STELLER et al., 2003; RIUL et al., 2009; PEÑA et 3 year for the Abrolhos shelf, containing approximately 5% of al., 2014). Rhodoliths may be composed entirely of the CaCO inventory of all the world’s carbonate banks. In encrusting coralline algae or may have a core of non- 3 addition to its importance in the global carbon cycle (VAN coralline material (e.g. a shell, piece of dead coral or a DER HEIJDEN; KAMENOS, 2015), the reported amount pebble), in addition to encrustations by other calcified of limestone has also called the attention of multinational organisms. They are distributed worldwide (FOSTER, companies interested in the large-scale exploitation of these 2001), from tropical to polar regions (ADEY; deposits for agronomic purposes. However, the relatively STENECK, 2001), and from the intertidal zone to low growth rate of mesophotic rhodoliths (about 1.0 mm depths of 268 m (LITTLER et al., 1986). Rhodolith per year, BLAKE; MAGGS, 2003; AMADO- FILHO et beds represent one of the largest carbonate depositional al., 2012a) confers on this resource a nearly non-renewable environments in the world (TESTA; BOSENCE, characteristic, at least on human time-scales (WILSON 1999), which makes them an important component in et al., 2004). In addition to the use of rhodolith beds as a the global carbon biogeochemical cycle and hence, in source of CaCO , other potential economic uses include the atmospheric CO balance (OLIVEIRA, 1996). 3 2 exploiting their associated biodiversity as a source of In Brazil, rhodolith beds extend over almost the bioactive compounds in the pharmacology, agriculture and entire continental shelf from Maranhão (0° 50’S) to nutraceutical industries (AMADO-FILHO; PEREIRA- Santa Catarina state (27°17’S). They predominate FILHO, 2012; MARINS et al., 2012). within the mesophotic zone (~ 30-150 m depth) along the Despite their great importance for biodiversity continental shelf, on the tops of seamounts, and around resilience and the global carbon budget, in Brazil the oceanic islands (AMADO-FILHO; PEREIRA-FILHO, functional ecology of these complex habitats, considering 2012). The mesophotic coralline ecosystems represent the intrinsic latitudinal and vertical variability, has received extensions of shallower ecosystems (HINDERSTEIN relatively little attention. In contrast, research programs and et al., 2010), however, information on biotic and abiotic initiatives in Europe (e.g. BIOMAERL, 1996-1999; United aspects of mesophotic habitats remains extremely Nations Environment Programme, UNEP, 2007, 2015) scarce, due to logistic and technological restrictions, have contributed to the knowledge of: (1) rhodolith bed when compared to shallow habitats (BRIDGE et al., diversity, (2) rhodolith-bed associated biota, (3) functional 2011). However, recent advances in mixed-gas diving roles played by key elements of
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