New Records of Rhodolith-Forming Species (Corallinales, Rhodophyta) from Deep Water in Espırito Santo State, Brazil
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Helgol Mar Res (2012) 66:219–231 DOI 10.1007/s10152-011-0264-1 ORIGINAL ARTICLE New records of rhodolith-forming species (Corallinales, Rhodophyta) from deep water in Espı´rito Santo State, Brazil Maria Carolina Henriques • Alexandre Villas-Boas • Rafael Riosmena Rodriguez • Marcia A. O. Figueiredo Received: 20 August 2010 / Revised: 30 May 2011 / Accepted: 5 July 2011 / Published online: 17 July 2011 Ó Springer-Verlag and AWI 2011 Abstract Little is known about the diversity of non- Introduction geniculate coralline red algae (Rhodophyta, Coral- linophycidae) from deep waters in Brazil. Most surveys ‘‘Rhodolith’’ is the name given to free-living structures undertaken in this country have been carried out in shallow composed mostly ([50%) of non-geniculate coralline red waters. In 1994, however, the REVIZEE program surveyed algae (Rhodophyta, Corallinophycidae). The communities the sustainable living resources potential of the Brazilian they denominate are called ‘‘rhodolith beds’’ (Foster 2001; exclusive economic zone to depths of 500 m. In the present Harvey and Woelkerling 2007). However, there is no study, the rhodolith-forming coralline algae from the con- consensus on use of the terms (Foster 2001). In this study, tinental shelf of Espı´rito Santo State were identified. we follow Foster (2001) and Harvey and Woelkerling Samples were taken from 54 to 60 m depth by dredging (2007). during ship cruises in 1997. Three rhodolith-forming spe- The deepest known alga is a rhodolith-forming coralline cies were found: Spongites yendoi (Foslie) Chamberlain, red algal species discovered on an uncharted seamount at Lithothamnion muelleri Lenormand ex Rosanoff and 268 m depth in the Bahamas (Littler et al. 1985). Little is Lithothamnion glaciale Kjellman. These records extend the known about the diversity of non-geniculate coralline red distribution ranges of these species into Brazilian waters algae from Brazil deep water (Amado-Filho et al. 2007; and extend the depth distribution of non-geniculate coral- Villas-Boas et al. 2009) in general and of rhodolith-form- line red algae into Brazilian water to 58 m. ing species in particular. The Brazilian continental shelf contains extensive rhodolith beds from 20 to 100 m depth Keywords Corallinales Á Taxonomy Á Spongites Á (Kempf et al. 1969; Kempf 1980; Amado-Filho et al. 2007) Lithothamnion Á Brazil distributed from 3° to 22° S and is considered the largest area supporting rhodolith beds in the world (Foster 2001). The continental shelf of Espı´rito Santo State is consid- Communicated by H.-D. Franke. ered to be the region with highest marine algal species richness in Brazil (Amado-Filho et al. 2007) and is a tran- & M. C. Henriques ( ) sitional area from tropical to temperate floras (Guimara˜es Museu Nacional/Universidade Federal do Rio de Janeiro, Av General Herculano Gomes s/n, Quinta da Boa Vista, 2003). Despite the urgent need to understand and preserve Sa˜o Cristova˜o, Rio de Janeiro, RJ, Brazil the marine biodiversity within rhodolith beds, which are a e-mail: [email protected] critical habitat for conservation, shallow rhodolith beds from Espirito Santo State have only recently been studied A. Villas-Boas Á M. A. O. Figueiredo Instituto de Pesquisas Jardim Botaˆnico do Rio de Janeiro, (Amado-Filho et al. 2007, 2010; Villas-Boas et al. 2009). Rua Pacheco Lea˜o 915, Rio de Janeiro, RJ 22460-030, Brazil Five rhodolith-forming genera (Lithothamnion, Lithophyl- lum, Hydrolithon, Neogoniolithon and Sporolithon) were R. R. Rodriguez reported from the region (Amado-Filho et al. 2007). Villas- Programa de Investigacio´n en Bota´nica Marina, Departamento de Biologı´a Marina, Universidad Auto´noma de Baja California Boas et al. (2009) report three rhodolith-forming species Sur, Apartado postal 19-B, 23080 La Paz, BCS, Mexico from the genus Lithophyllum [L. corallinae (Crouan and 123 220 Helgol Mar Res (2012) 66:219–231 Crouan) Heydrich, L. johansenii Woelkerling and Camp- degraded, and it was difficult finding intact reproductive bell, L. stictaeforme (Areschoung in J. Agardh) Hauck], structures. The material was exhaustively sectioned, but in already known to science, and one rhodolith-forming spe- one case (Fig. 1d), we could not find a representative cies (L. depressum Villas-Boas, Figueiredo et Riosmena- reproductive structure. Histological techniques for optical Rodriguez) new to science. Besides these, two other species microscopy follow Moura et al. (1997). Microslides were were reported from the region (Lithothamnion muelleri prepared, and identification of material to species level Lenormand ex Rosanoff and Mesophyllum engelhartii followed descriptions of Chamberlain (1993), Chamberlain (Foslie) Adey) (Amado-Filho et al. 2010). and Irvine (1994), and Wilks and Woelkerling (1995). A major survey evaluating the sustainable potential of Type specimens were not examined. Identified samples living resources of the Brazilian Exclusive Economic Zone were incorporated in the herbarium collection of the (REVIZEE program) was carried out in 1997. Its aim was Botanical Garden of Rio de Janeiro (RB). The herbarium to address potential fisheries resources and to identify the code follows the Index Herbariorum (Holmgren and main benthic taxonomic groups on the outer shelf and Holmgren 2011, continuously updated). Conceptacle continental slope from 50 to 500 m depth. Today, the measurements follow Adey and Adey (1973), thallus ana- REVIZEE program is the most widespread marine project tomical terminology follows Woelkerling (1988), growth carried out on the Brazilian coast generating biological, form terminology follows Woelkerling et al. (1993) and geological, physical and chemical data (Lavrado 2006). typification data follow Woelkerling (1993). Rhodolith beds were recorded during fieldwork, but cor- alline algal species composition was not investigated. The material has been stored for later study (Yoneshigue- Results Valentin et al. 2006). Thus, Brazilian deep-water marine ecosystems still remain poorly described. Taxonomic accounts The current study aims to contribute to the knowledge of coralline red algae in Espı´rito Santo state, Brazil. The study Three species were identified: Spongites yendoi (Foslie) reports previously unrecorded rhodolith-forming species in Chamberlain (Fig. 1); Lithothamnion muelleri Lenormand Brazilian deep water, discussing the implications of these ex Rosanoff (Figs. 2, 3); and Lithothamnion glaciale new records for rhodolith diversity in Brazil. Kjellman (Figs. 4, 5). Spongites yendoi (Foslie) Chamberlain, 1993: 102 (Fig. 1) Materials and methods Basionym: Goniolothon yendoi Foslie, 1900a: 25 Field work Further references: Penrose (1996) Synonyms: See Chamberlain (1993) Rhodolith samples were collected during field trips con- ducted from October to November of 1997 by the REVI- Lectotype: TRH (Yendo no. 66) Foslie (1904) ZEE program (Lavrado 2006). The material was collected Type locality: Shimoda Harbour, Izul, Japan, K. Yendo, offshore along the Espı´rito Santo State continental shelf April 1899 at three studies sites: 34R (20°4500000S–40°1000000W); 28R (19°4900000S–37°5600000W); and 29R (19°4800100S– Description: Plants non-geniculate, thallus-forming free- 37°4602200W). Depths ranged from 55 to 58 m depth (see living rhodoliths with warty growth form (Fig. 1a) con- Yoneshigue-Valentin et al. 2006 for site descriptions). The sisting of several layers of plants overgrowing one another. region is influenced by the Brazilian current that runs along Spongites yendoi compose the most external layer. Cylin- the South Atlantic Ocean and is characterized by sea sur- drical protuberances measuring 1–2 mm in height and face temperatures ranging from 21 to 27°C (Lavrado 2006). 1 mm in diameter. Thallus pseudoparenchymatous with At 50 m depth, the region is influenced by the South dorsiventral internal organization in encrusting portions Atlantic Central Waters (SACW) and is characterized by and radial internal organization in protuberances. Mono- temperatures from 6 to 20°C (Silveira et al. 2000). merous thallus construction in encrusting portions (Fig. 1b) consisting of a single system of branched filaments that Laboratory work form a core running more or less parallel to the substratum and a more peripheral region in which portions of core Since 1997, the samples had been preserved in 4% formalin filaments or their derivatives curve outwards toward the seawater (Yoneshigue-Valentin et al. 2006). As a conse- thallus surface terminating in one layer of flattened quence of this long preservation time, specimens were epithallial cells (Fig. 1c). The core region is plumose 123 Helgol Mar Res (2012) 66:219–231 221 Fig. 1 Spongites yendoi. a Morphology of a rhodolith with warty initials cells. Scale bar 10.5 lm. (RB 458711). d Longitudinal section growth form. Scale bar 2.5 mm. (RB 458711). b Longitudinal section through a uniporate tetrasporangial conceptacle with roof protruding showing the internal monomerous thallus construction. P peripheral above the surrounding thallus surface. Note the tetrasporangium region and C core region. Scale bar 52 lm. (RB 458711). c Longi- (arrow). Scale bar 45.7 lm. (RB 458711). e Longitudinal section tudinal section of the outer perithallium showing the flattened, but showing a tetrasporangium in detail. Numbers 1–4 indicate the four rounded epithallial cells (black arrow) and cell fusions between cells spores in tetrasporangium. Scale bar 9 lm. (RB 458711) of adjacent peripheral filaments (white arrow), S = subepithallial 123 222 Helgol Mar Res