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Non-Coralline Crustose Algae Associated with Maerl Beds in Portugal: a Reappraisal of Their Diversity in the Atlantic Iberian Beds

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Non-Coralline Crustose Algae Associated with Maerl Beds in Portugal: a Reappraisal of Their Diversity in the Atlantic Iberian Beds DOI 10.1515/bot-2013-0083 Botanica Marina 2013; 56(5-6): 481–493 Viviana Peña* and Ignacio Bárbara Non-coralline crustose algae associated with maerl beds in Portugal: a reappraisal of their diversity in the Atlantic Iberian beds Abstract: In recent years (2007–2009), the distribution Introduction and associated flora of maerl beds in southern Portugal (Algarve) were investigated by dredging and SCUBA diving Crustose algae are considered an important component in (12–30 m depth). The present work provides the first data the diversity and structure of marine communities. They on the non-coralline crustose flora associated with maerl also constitute an interesting subject for biogeographic beds in Portugal. Peyssonnelia bornetii is a new record studies due to their wide distribution, slow growth rates, for Atlantic European coasts, and new records for Portu- and persistence in the face of mechanical and biological gal are “Rhododiscus pulcherrimus” (sporophyte phase of disturbances (Maggs 1990). In addition, studies of crus- Atractophora hypnoides), Contarinia peyssonneliaeformis, tose algae have contributed to the discovery of new phases “Cruoria rosea” (sporophyte phase of Halarachnion ligu- in the life histories of species with alternating heteromor- latum), Peyssonnelia armorica, and “Aglaozonia chilosa” phic generations (Sauvageau 1899, Maggs and Guiry 1982, (sporophyte phase of Cutleria chilosa). In addition, the Maggs et al. 1983). European maerl beds are coastal habi- records of three other species (Hildenbrandia crouanio- tats that provide a wide range of ecological niches because rum, Peyssonnelia dubyi, and Peyssonnelia harveyana) of their three-dimensional structure (Barberá et al. 2003). completed the distribution gap between the North Ibe- They harbor a high number of algal species, including rian Peninsula and the Macaronesian region. The female crustose taxa and phases in heteromorphic life histories, and carposporangial structures of C. peyssonneliaeformis some of them mostly restricted to this habitat (Jacquotte and P. bornetii are described for the first time. The diver- 1962, Maggs and Irvine 1983, Maggs and Guiry 1989, Soto sity and species composition observed are compared with 1990, Ballesteros 1992, Birkett et al. 1998, Mannino et al. previous studies from maerl beds in the northwestern 2002, Peña and Bárbara 2010a). Iberian Peninsula (Galicia) and neighboring areas in the In the Atlantic Iberian Peninsula, a previous study Atlantic Ocean and Mediterranean Sea. In order to facili- of the diversity of the non-coralline crustose flora associ- tate and promote further studies of the crustose flora of ated with the Galician maerl and gravel beds brought the subtidal habitats along European coasts, an identification total number of species up to 23 taxa (Peña and Bárbara key is provided for the 26 crustose taxa associated with 2010a). Recently, we were able to record subtidal maerl maerl and gravel beds of the Atlantic coast of the Iberian beds in two areas of southern Portugal (Peña et al. 2009). Peninsula. Up to this date, the flora (and particularly the crustose species) associated with maerl beds of Portugal had not Keywords: crustose seaweeds; Iberian Peninsula; maerl; been studied in detail. The present study aims to contrib- Portugal; Rhodophyta. ute to a better knowledge of the subtidal benthic flora of the Portuguese coast, which is poorly known in the litera- *Corresponding author: Viviana Peña, BioCost Research Group, ture. We provide here the first data on the non-coralline University of A Coruña. Facultade de Ciencias, Campus da Zapateira crustose red algae associated with these maerl beds. We S/N. 15071 A Coruña, Spain; Phycology Research Group, Ghent also compare the crustose flora of the Atlantic Iberian University, Krijgslaan 281, Building S8, 9000 Ghent, Belgium; and maerl beds with those reported from adjacent beds of the UMR 7138 Systématique, Adaptation et Evolution, Muséum National d’Histoire Naturelle, 75231 Cedex 05 Paris, France, Atlantic Europe and the Mediterranean Sea. Given the dif- e-mail: [email protected] ficulty of identifying crustose species and the scattered Ignacio Bárbara: BioCost Research Group, University of A Coruña. information for the species recorded, we provide an iden- Facultade de Ciencias, Campus da Zapateira S/N. 15071 A Coruña, tification key, which encompasses all non-coralline crus- Spain tose species recorded in the maerl and gravel beds of the Atlantic Iberian Peninsula. Brought to you by | CSIC - Unidad Coordinacion Bibliotecas Authenticated | 150.214.192.170 Download Date | 12/7/13 5:35 PM 482 V. Peña and I. Bárbara: New records for crustose flora of Atlantic Iberian maerl beds Materials and methods for this species (Newton 1931, Denizot 1968, Irvine 1983, Peña and Bárbara 2010a). Together with the report of From 2007 to 2009, we carried out 74 subtidal surveys both R. pulcherrimus in the Canary Islands (Gil-Rodríguez et al. by SCUBA and dredging (12–30 m depth) in two main areas 1985), the present record constitutes the southernmost of southern Portugal (Armaçao de Pêra, 37°01′N, 8°19′W; record for Atlantic Europe. Bahia de Lagos-Portimao, 37°06′N, 8°38′W). We collected Habitat and occurrence: tetrasporangia recorded in 181 samples from maerl beds with variable maerl cover March. The species was found growing on living maerl, mixed with gravel, dead shells, and pebbles, sometimes occasionally on dead shells. Depth range: 17–19 m. The interrupted by rocky outcrops (Figure 1A–D). The samples present study constitutes the first record for Portugal. The were preserved in 4% formalin/seawater and kept in total tetrasporophyte phase (“R. pulcherrimus”) only is cited for darkness at 4°C. Sections were cut by hand using a razor the Atlantic maerl beds (Table 1). blade. Observations were made in different planes accord- ing to Denizot (1968) and Irvine (1983). The species were identified using specialized literature on crustose red Contarinia peyssonneliaeformis algae (Denizot 1968, Boudouresque and Denizot 1973, 1975, Belsher and Marcot 1975, Maggs and Guiry 1989, Kato Creeping thallus up to 200 μm in thickness, margin et al. 2005, Peña and Bárbara 2010a). Pictures of each appressed to the substratum (Figure 2A, B). Color bright species were taken under light microscopy, and the most red to brownish. Rhizoids are multicellular, unbranched significant contributions to the European crustose flora or branched. No hypobasal calcification is observed. In were illustrated in the present study. vertical section, the thallus has principal axial filaments Representative specimens were deposited in the her- composed of colorless cells that give rise to one to two barium SANT (acronyms follow Holmgren et al. 1990). dorsal cortical cells pseudodichotomously branched at The terms sorus, conceptacle, perithallus, and hypothal- the second cell. Gland cells are ovoid, immersed among lus were used according to Peña and Bárbara (2010a). the cortical filaments and also spreading on the surface. The habitat and occurrence of each species are provided. Carpogonial branches are produced in sori with multi- Present records were also compared with previous phy- cellular paraphyses. Auxiliary filaments are three to four cological studies from the Portuguese coast (Ardré 1970, celled, each 30–37 × 10 μm, with pit connections between Araújo et al. 2009, Berecibar et al. 2009, Berecibar 2011). cells (Figure 2C, D). Mature gonimocarps are external The maerl-associated crustose flora of the Atlantic Iberian globular structures, 75–150 × 50–75 μm, and contain car- Peninsula was compared with those of neighboring Atlan- pospores, each 10–15 × 7–13 μm (Figure 2E). The diagnostic tic European and Mediterranean beds according to data vegetative features of the Portuguese specimens match compiled by Peña and Bárbara (2008, 2010a). The identi- with descriptions provided from the Mediterranean and fication key for non-coralline crustose species associated from the only Atlantic record hitherto known in Galicia with Atlantic Iberian maerl and gravel beds was based on (Feldmann 1939, Denizot 1968, Athanasiadis 1987, Peña Peña and Bárbara (2010a) and the present study. and Bárbara 2010a). The female structures and gonimo- carps are described for the first time in this species. Fertile plants of the Mediterranean species Contarinia squamar- iae were also recently reported from Portugal (Berecibar Results et al. 2009). The structure of female nemathecia and the development of gonimocarps are similar in both species. “Rhododiscus pulcherrimus” (sporophyte Berecibar et al. (2009) also recorded tetrasporangial phase of Atractophora hypnoides) nemathecia in the Portuguese specimens of C. squamar- iae as dark reddish masses, composed of dense stands of Crust up to 100 μm thickness. Color bright red to carmine cylindrical tetrasporangia zonately or irregularly zonately red, mucilaginous when fertile. Rhizoids are absent. divided. By contrast, the tetrasporangia of C. peyssonne- Monostromatic basal layer composed of branched radial liaeformis are irregularly cruciate (Denizot 1968). filaments. Each basal cell gives rise to one to two erect fila- Habitat and occurrence: female structures recorded ments, unbranched or rarely branched, separable under only in December. The species was found growing on pressure. In surface view, cells are round. Tetrasporangia living maerl, occasionally epilithic on pebbles. Depth are terminal, cruciately arranged. Paraphyses are absent. range: 15–19 m. This study represents the
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