Marine algal flora of São Miguel Island, Azores

Ana Isabel Neto1, Ignacio  Moreu1, Edgar F.  Rosas-Alquicira2, Karla León-Cisneros3, Eva Cacabelos4, Andrea Z. Botelho5, Joana Micael6, Ana C. Costa5, Raul M. A. Neto7, José M. N. Azevedo1, Sandra Monteiro5, Roberto  Resendes8, Pedro Afonso9, Afonso C. L. Prestes1, Rita F. Patarra10, Nuno Vaz Álvaro11, David Milla-Figueras9, Enric Ballesteros12, Robert Lawson Fletcher 13, William Frank Farnham14, Ian  Tittley15, Manuela I. Parente5, Ignacio Moreu1

1 Universidade dos Açores, Faculdade de Ciências e Tecnologia, Departamento de Biologia and Grupo de Investigação Aquática Insular, IBBC-GBA/cE3c, Rua da Mãe de Deus, 13-A, 9500-321, Ponta Delgada, Portugal; 2 Lane Community College, 4000 East 30th Ave., 97405, Eugene, United States; 3 Universidad Autónoma de Baja California Sur, Programa de Investigación en Botánica Marina, Departamento Académico de Ciencias Marinas y Costeras , Carretera al Sur Km. 5.5, colonia el Mezquitito, 23080, La Paz, Mexico; 4 MARE – Marine and Environmental Sciences Centre, Agência Regional para o Desenvolvimento da Investigação Tecnologia e Inovação (ARDITI), Edif. Madeira Tecnopolo, Piso 2, Caminho da Penteada, 9020-105, Funchal, Portugal; 5 CIBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, InBIO Laboratório Associado, Pólo dos Açores, Universidade dos Açores, Rua da Mãe de Deus, 13-A, 9500-321, Ponta Delgada, Portugal; 6 Southwest Iceland Nature Research Centre, Gardvegi 1, 245, Suðurnesjabær, Iceland; 7 NA, , Portugal; 8 Universidade dos Açores, Faculdade de Ciências e Tecnologia, Departamento de Biologia, Rua da Mãe de Deus, 13-A, 9500-321, Ponta Delgada, Portugal; 9 IMAR/Okeanos – University of the Azores, Departamento de Oceanografia e Pescas, Universidade dos Açores, Rua Prof. Doutor Frederico Machado, 9901-862, Horta, Portugal; 10 Expolab - Ciência Viva Science Centre, Avenida da Ciência - Beta, n.º 8 , 9560-421, Lagoa, Portugal; 11 Universidade dos Açores, Faculdade de Ciências Agrárias, CCMMG (Centro do Clima Meteorologia e Mudanças Globais), IITA-A (Instituto de Investigação e Tecnologias Agrárias e do Ambiente), Campus de Angra do Heroísmo Rua Capitão João d’Ávlia – Pico da Urze, 9700-042, Angra do Heroísmo, Portugal; 12 Centre d'Estudis Avançats de Blanes-CSIC , Acc. Cala Sant Francesc 14, 17320, Blanes, Spain; 13 University of Portsmouth, Institute of Marine Sciences, Department of Biological Sciences , Ferry Road, Eastney, PO4 9LY , Portsmouth, United Kingdom; 14 University of Portsmouth, Ferry Road, Eastney, PO4 9LY, Portsmouth, United Kingdom; 15 Natural History Museum, Cromwell Road, SW7 5BD, London, United Kingdom

Corresponding author(s): Ana Isabel Neto (ana.im.neto@uac.pt)Ignacio  Moreu (nacho.moreu.badia@gmail.com)Edgar F.  Rosas-Alquicira (rosasalquicirae@lanecc.edu)Karla León-Cisneros (kleon@uabcs.mx)Eva Cacabelos (ecacabelos@mare-centre.pt )Andrea Z. Botelho (andrea.zc.botelho@uac.pt)Joana Micael (joana@natturustofa.is)Ana C. Costa (ana.cm.costa@uac.pt)Raul M. A. Neto (raulmaneto@gmail.com)José M. N. Azevedo (jose.mv.azevedo@uac.pt)Sandra Monteiro (sandra.cs.monteiro@uac.pt)Roberto  Resendes (roberto.resendes@uac.pt)Pedro Afonso (pafonsopim@gmail.com)Afonso C. L. Prestes (afonso.cl.prestes@uac.pt)Rita F. Patarra (rita.patarra@expolab.pt)Nuno Vaz Álvaro (nuno.ms.alvaro@uac.pt)David Milla-Figueras (davidmfigueras@gmail.com)Enric Ballesteros (kike@ceab.csic.es)Robert Lawson Fletcher  (phaeophyta@hotmail.com)William Frank Farnham (farnham2@gmail.com)Ian  Tittley (mmit@waitrose.com)Manuela I. Parente (manuela.ip.cardoso@uac.pt)

Received {date}; Revised {date}; Accepted {date}; Published {date}

Citation: Combination of authors, year of data paper publication (in parentheses), Title, Journal Name, Volume, Issue number (in parentheses), and doi of the data paper.

Resource Citation

Neto A I, Moreu I, Rosas-Alquicira E F, León-Cisneros K, Cacabelos E, Botelho A Z, Micael J, Costa A C, Neto R M A, Azevedo J M N, Monteiro S, Resendes R, Afonso P, Prestes A C L, Patarra R F, Álvaro N V, Milla-Figueras D, Ballesteros E, Fletcher R L, Farnham W F, Tittley I, Parente M I (2021): Marine algal flora of São Miguel Island, Azores. v1.3. Universidade dos Açores. Dataset/Samplingevent. http://ipt.gbif.pt/ipt/resource?r=sao_miguel_macroalgal_flora&amp;v=1.0 http://ipt.gbif.pt/ipt/resource?r=sao_miguel_macroalgal_flora&amp;v=1.0

Abstract

Background
The macroalgal flora of the Island of São Miguel (eastern group of the Azores archipelago) has attracted the interest of many researchers in the past, the first publications going back to the nineteenth century (see summary in Neto et al. 2014). Initial studies were mainly taxonomic, resulting in the publication of species lists, which were compiled by Neto (1994) in the first checklist of the Azorean benthic marine algae. Later, the establishment of the University of the Azores on the Island permited the logistic conditions to develop both temporal studies and long-term research, and this resulted in a significant increase on research directed at the benthic marine algae and littoral communities of the Island and consequent publications (see revision in Neto et al. 2014 and Haroun et al. 2019). Prior to the present paper, the known macroalgal flora of São Miguel Islandcomprised around 260 species. Despite this richness, a significant amount of the research was never made public, notably Masters and PhD theses encompassing information regarding presence data recorded at littoral and sublittoral levels down to a depth of approximately 40 m around the Island, and the many collections made, which resulted in vouchers deposited in the AZB Herbarium Ruy Telles Palhinha and the LSM- Molecular Systematics Laboratory at the Faculty of Sciences and Technology of the University of the Azores.
The present publication lists the macroalgal taxonomic records together with information on their ecology and occurrence around São Miguel Island, improving the knowledge of the Azorean macroalgal flora at local and regional scales.

New information
A total of 12,781 specimens (including some identified only to genus) belonging to 431 taxa of macroalgae are registered, comprising 284 Rhodophyta, 59 Chlorophyta and 88 Ochrophyta (Phaeophyceae). Of these, 323 were identified to species level (212 Rhodophyta, 48 Chlorophyta and 63 Ochrophyta), of which 61 are new records for the Island (42 Rhodophyta, 9 Chlorophyta and 10 Ochrophyta), 1 an Azorean endemic (Predaea feldmannii subsp. azorica Gabriel), 5 are Macaronesian endemisms (the Rhodophyta Botryocladia macaronesica Afonso-Carrillo, Sobrino, Tittley & Neto, Laurencia viridis Gil-Rodríguez & Haroun, Millerella tinerfensis (Seoane-Camba) S.M.Boo & J.M.Rico, Phyllophora gelidioides P.Crouan & H.Crouan ex Karsakoff and the Chlorophyta Codium elisabethiae O.C.Schmidt), 19 are introduced species (15 Rhodophyta, 2 Chlorophyta and 2 Ochrophyta), and 32 are of uncertain status (21 Rhodophyta, 5 Chlorophyta and 6 Ochrophyta).
Introduction
Research on the marine algae from the Azores started in the mid nineteenth century (1838) when Guthnick and the two Hochstetters, father and son, visited the archipelago (Neto 1994). Since then, many other researchers and naturalists have visited the archipelago, resulting in several publications on the marine algal flora of this region (see summary in Neto 1994; 1997). Most initial studies were taxonomic focusing on the production of species lists. Almost a century later, the German botanist Otto Christian Schmidt visited several islands, including São Miguel, and initiated a more comprehensive ecological approach describing species associations and their spatial organization (Schmidt, 1931). Ever since the first half of last century, several studies have focused more widely on intertidal and shallow subtidal communities providing information on the vertical distribution of macroalgae and invertebrates and their trophic relations (see Neto 1992, 2000, 2001 for a review on this subject). Taxonomic investigations have continued and the first checklist of the Azorean benthic marine algae published by Neto (1994) brought together the existing published information, provided distributional records within the archipelago and reported 307 species, indicating a moderately rich flora given its isolated mid-Atlantic position. A revision of this first checklist was made by Parente (2010), increasing the number of algae species to 327, but without providing their distributional information on the archipelago. Later, Rosas-Alquicira et al. (2011) published a catalogue of non-fossil geniculate coralline red algae (Corallinales, Rhodophyta) of the Macaronesia in which they made both a critical review of species and infraspecific taxa as ell as an assessment of species diversity in the region. Research by local teams was also dedicated to the Azorean littoral communities and biota conservation (see, for example, Abecasis et al. 2015, Amorim et al. 2015, Chainho et al. 2015). Taxonomic, ecological and biotechnological investigations have continued generating knowledge on the Azorean macroalgae flora, its biotechnological potential and also on the structure and functioning of littoral communities (see revisions on Neto et al. 2014 and Haroun et al. 2019). Recently, several additional studies have been published with important information on the Azorean algae biodiversity, biogeography, conservation, ecology, and taxonomy (see, for example, Bruno de Sousa et al. 2019, Cacabelos et al. 2019, 2020, Freitas et al. 2019, Kellaris et al. 2019, Martins et al. 2019, Parente et al. 2019, 2020, Patarra et al. 2017, 2019, 2020, Sousa et al. 2019, Faria et al. 2020a-b, Vieira et al. 2020).
The paper by Freitas et al. (2019) increased the number of macroalgae species occurring in the Azores to 405 and reported that amongst the mid-Atlantic archipelagos, the Azores is second in species richness after the Canary Islands, with 689 species, and followed by Madeira (396), Cabo Verde (333) and Selvagens (295 species). For some species the Azores archipelago forms a boundary in their distribution. Codium effusum (Rafinesque) Delle Chiaje, for example, is as its western distribution limit in the archipelago (León-Cisneros et al. 2012), whereas for Dudresnaya crassa M.Howe, a western Atlantic warm-water species, the Azores extends its known distributional range to the east. Some northern species such as the red alga Schizymenia dubyi (Chauvin ex Duby) J.Agardh and Lomentaria orcadensis (Harvey) Collins come close to their southern limit of distribution in the Azores while some southern warm-water species such as green alga Anadyomene stellata (Wulfen) C.Agardh and the red alga Sebdenia rodrigueziana (Feldmann) Codomier ex Athanasiadis reach their Atlantic northern limit of distribution on the islands (Neto et al. 2005, León-Cisneros et al. 2012). Some species, relatively common in the region a few years ago, have become uncommon or even very rare, e.g. Scytosiphon lomentaria (Lyngbye) Link, Schimmelmannia schousboei (J.Agardh) J.Agardh. In contrast, there has been an increase of unexpected macroalgae in the Azores, with the arrival and establishment of several non-native species (see Cardigos et al. 2006, Micael et al 2014, Vaz-Pinto et al. 2014, Parente et al. 2019, Cacabelos et al. 2019, 2020, Martins et al. 2019).
Within the spread of the archipelago there are no marked differences between floras of individual Islands or Island groups, and biogeographically the Azores algal flora reveals to have a mixed nature, with species shared with Macaronesia, North Africa, the Mediterranean Sea, Atlantic Europe and America (Tittley 2003, Tittley & Neto 1995, 2005, 2006, Wallenstein et al. 2009b). This nature of the Azorean marine algal flora was reinforced by the work of Freitas et al. (2019), who using an extensive analysis encompassing data on coastal fishes, brachyurans, polychaetes, gastropods echinoderms and macroalgae, suggested that the Azores should be a biogeographical entity on its own and proposed a redefinition of the Lusitanian biogeographical province, in which they included four ecoregions: the South European Atlantic Shelf, the Saharan Upwelling area, the Azores ecoregion, and a new ecoregion they named Webbnesia, which comprises the archipelagos of Madeira, Selvagens and the Canary Islands.
Not all the Azorean Islands have received the same attention regarding the studies on macroalgae. Furthermore, many species may have been overlooked due to their small size, opportunisctic nature or ephemeral life span.
To overcome this and gain a better and up to date knowledge of the archipelago’s macroalgae flora, an effort was made by resident teams to undertake a considerable amount of research over the past three decades on several Islands. The present paper is the last one of a series and presents physical, occurrence data, and information gathered from macroalgal surveys undertaken on São Miguel Island between 1989 and 2019 mainly by the Island Aquatic Research Group of the Azorean Biodiversity Centre of the University of the Azores (Link: https://ce3c.ciencias.ulisboa.pt/sub-team/island-aquatic-ecology), the BIOISLE, Biodiversity and Islands Research Group of CIBIO-Açores at the University of the Azores (Link: https://cibio.up.pt/research-groups-1/details/bioisle), and the OKEANOS Centre of the University of the Azores (Link: http://www.okeanos.uac.pt). In these surveys particular attention was given to the small filamentous and thin sheet-like forms that are often short-lived and fast-growing, and usually very difficult to identify in the field, without the aid of a microscope and specialised literature in the laboratory.
This paper aims to provide a valuable marine biological tool to aid research on the systematics, diversity and conservation, biological monitoring, climate change, ecology and more applied studies, such as biotechnological applications, which will be of assistance to a wide range of focal groups including academics, students, governments, private organizations and the general public.
Purpose: This paper presents taxonomic records of macroalgae for São Miguel Island and provides general information on their occurrence and distribution. By doing this, it will contribute to address several biodiversity shortfalls (see Cardoso et al. 2011, Hortal et al. 2015), namely the need to catalogue the Azorean macroalgae (Linnean shortfall) to improve current information on their local and regional geographic distribution (Wallacean shortfall), as well as to provide a better understanding of species abundance and dynamics in space (Prestonian shortfall).

Keywords: Macroalgae, Azores, São Miguel Island, new records, endemism, native, uncertain, introduced, occurrence data, ecology.

General description

Additional information: Additional information on this study may also be requested to the first author

Project details

Project title: Marine algal flora of São Miguel Island, Azores
Personnel: Ana Isabel Neto, Manuela I. Parente
Funding: This study was mainly finantially supported by the following projects/scientific expeditions:
•	Projects:
o	ABLA/MAC – “Associações Biológicas do Litoral Açoreano/Moluscos, Algas e Crustáceos”, funded by the Portuguese Science and Technology Foundation (1987-1991);
o	Azorean Algal Flora – “Studies on algal communities of São Miguel, Azores”, partially funded by CIRN/DB/UAc (1992-1996);
o	BIA - “Biodiversity of Azores Archipelago”, funded by the Portuguese Science and Technology Foundation. PRAXIS/2/2.1/BIA/169/94 (1996-1999);
o	BIOTOPE – “Classification, mapping and modelling of Azorean littoral biotopes”, funded by the Portuguese Science and Technology Foundation, POCTI MGS/45319/2002 (2003-2006);
o	CAMAG/ORI – “Characterization of coastal water bodies on the islands of Santa Maria and São Miguel”, funded by the Regional Government of the Azores, Regional Secretariat for the Environment and the Sea, Regional Directorate for Planning and Water Resources (2008-2012);
o	GESMAR – “Sustainable management of marine Resources”, funded by the EU Funding Program III B 2000-2006, Açores-Madeira-Canárias, GESMAR/MAC/2/C068 (2009-2012);
o	PATELGENE – “Genetic Structure of Azorean Limpets: Implications for Conservation and Marine Protected Areas”, funded by the Portuguese Science and Technology Foundation, PTDC/BIA-BIC/115837/2009 (2011-2014);
o	MACROBIOMOL – “Macroalgae biodiversity under a molecular view - for a better understanding of North Atlantic Biogeography”, funded by PTDC/MAR/114613/2009 (2011-2015);
o	ASMAS – “Açores: Stop-over for Marine Alien Species?”, funded by the Government of the Azores - Regional Secretariat for the Sea, Science and Technology, M2.1.2/I/032/2011 (2012-2016);
o	BUS – “Urban Structures: a driver of biodiversity change in coastal ecosystems?”, funded by the Portuguese Science and Technology Foundation, PTDC/MAR-EST/2160/2012 (2013-2015);
o	ECOSUBVEG – “Changes in submersed vegetation: assessing loss in ecosystems services from frondose to depauperate systems dominated by opportunistic vegetation”, funded by the Voluntary Scheme for Biodiversity and Ecosystem Services in Territories of the EU Outermost Regions and Oversees Countries and Territories, BEST 07.032700/2012/635752/SUB/B2 (2013-2016);
o	LAUMACAT - “Diversity and phylogenetic relationships on the benthic marine algae with pharmacological potential: the Laurencia complex (Rhodophyta) in Macaronesian archipelagos, tropical and subtropical Atlantic”, funded by the Ministerio de Ciencia e Innovación, Dirección General de Investigación y Gestión del Plan Nacional de R+D+i, Subdirección General de Proyectos de Investigación, Gobierno de España (2010-2013) and by the São Paulo State Research Support Foundation (FAPESP), Brazil, Proc. 2014 / 00012-1 (2013 a 2016);
o	BALA – “Elaboration of the implementation program of the marine strategy framework directive - biodiversity of the coastal environments of the Azores” (2 /DRAM /2015), funded by the Government of the Azores - Regional Secretariat for the Sea, Science and Technology, Regional Directorate for Sea Affairs, GRA /SRMCT-DRAM, (2015);
o	PIMA – “Elaboration of the implementation program of the Marine Strategy Framework Directive - Marine Invasion Program in the Azores” (3/DRAM /2015), funded by the Government of the Azores - Regional Secretariat for the Sea, Science and Technology, Regional Directorate for Sea Affairs, GRA /SRMCT-DRAM, (2015);
o	ASPAZOR – “Ecosystem impacts and socioeconomic benefits of Asparagopsis armata in the Azores”, funded by the Regional Direction for Science, Technology. ACORES-01 -0145-FEDER-000060 (2016-2020);
o	PORBIOTA - “ACORES-01-0145-FEDER-000072 - AZORES BIOPORTAL”, funded by the Operational Programme Azores 2020 (85% ERDF and 15% regional funds) (2019-2021);
•	Scientific Expeditions and campaigns:
o	“Campaign Macaronesia 2000”, under the project Macaronesia 2000 (2000-2001);
o	“Waitt Foundation”, under the projects BALA and PIMA (2016);
o	“BALA/PIMA”, under the projects BALA and PIMA (2018);
o	“PORBIOTA/2019” under the project ACORES-01-0145-FEDER-000072 - AZORES BIOPORTAL – PORBIOTA (2019);
•	Other funds:
o	Portuguese National Funds, through FCT – the Portuguese Science and Technology Foundation, within the projects UID/BIA/00329/2013, 2015-2019, UID/BIA/00329/2020-2023 and UID/BIA/50027/2019, UID/BIA/50027/2013-2020, UID/Multi/04423/2013, PEst-C/MAR/LA0015/2013 and POCI-01-0145-FEDER-006821;
o	European Regional Development (ERD) funds through the Operational Programme for Competitiveness Factors (COMPETE);
o	Portuguese Regional Funds, through DRCT - Regional Directorate for Science and Technology, within several projects, 2019 and 2020 and SRMCT /DRAM - Regional Secretariat for the Sea, Science and Technology, Regional Directorate for Sea Affairs;
o	CIRN/DB/UAc (Research Centre for Natural Resources, Universidade dos Açores, Departamento de Biologia);
o	CIIMAR (Interdisciplinary Centre of Marine and Environmental Research, Porto, Portugal).
Study area descriptions/descriptor: The Azores archipelago (38°43′49″N, 27°19′10″W), isolated in the mid-Atlantic Ocean, comprises nine volcanic Islands and several islets spread over 500 km in a WNW–ESE direction, emerging from the Azores Plateau and located above an active triple junction between three of the world's largest tectonic plates (the North American Plate, the Eurasian Plate and the African Plate, Hildenbrand et al. 2014). The archipelago comprises nine volcanic Islands and several small Islets in three separate groups (eastern, central and western).
São Miguel, located in the eastern group of the archipelago (37°54'58''N, -25°51'52''W), and approximately 750 km²in size, is the largest and most volcanically active Island (Gaspar et al. 2015).
As in the other Azorean Islands, the climate is considerably influenced by the surrounding ocean and is characterised by regular rainfall, medium levels of relative humidity and persistent winds, mainly during the winter and autumn seasons (Morton et al. 1998). The tidal range is small (<2 m) and the coastal extension is restricted, with deep waters occurring within a few kilometres offshore (Hidrográfico 1981). Most sea-shores are subject to swell and surge most of the year and few are sheltered, except for some bays and harbours. Extremely heavy seas occur during winter (Neto et al. 2005).
São Miguel has the longest coastline in the archipelago, about 155 km, corresponding to 25.3% of the whole Azorean coastline. The coastal topology, resulting from the effect of the maritime agitation, responsible for the predominance of erosive morphologies, is mainly composed of high, steep cliffs with a variety of stack, arch and gully formations, and is mostly difficult to access by land. Most of the cliffs and coastal slopes are less than 50 m in altitude (Borges 2003) and fall directly into the sea. The coastline is mainly composed of irregular compact, bedrock platforms, alternating with boulder and cobble locations. On some shores boulders entrap coarse sand and gravel and there are a few sandy beaches (Wallenstein et al. 2009b).
Intertidal communities of São Miguel Island, as on the other islands of the archipelago, are primarily dominated by macroalgae, which mainly exhibits a mosaic and /or zoned distribution pattern, and have a predominance of algal turfs that cover the rocks as a carpet (Wallenstein et al. 2009a), best seen when rocks are uncovered at low tide. There is a very distinct horizontal pattern of species distribution, with three major zones commonly found on bedrock and boulder shores (Neto 2000, Neto et al. 2005; Wallenstein et al. 2009b). The uppermost intertidal level is dominated by littorinids, while the mid-level zone is usually characterised by a fringe of chthamalid barnacles, in which sometimes algae and limpets can occur, followed by a lower area in which either algal turf (generally monospecific and usually composed of Caulacanthus ustulatus (Turner) Kützing) dominates, or patches of the brown alga Fucus spiralis Linnaeus and the red agarophyte Gelidium microdon Kützing grow interspaced with barnacles and algal turf. The lowest intertidal zone, representing the transition to the sublittoral envrironment, is either dominated by algal turf (generally multispecific and commonly dominated by coralline algae, or by various species of frondose algae growing in bands (e.g. the brown alga Gongolaria abies-marina (S.G.Gmelin) Kuntze), or forming patches amongst and over turf species (e.g. the agarophyte Pterocladiella capillacea (S.G.Gmelin) Santelices & Hommersand, and the calcareous Ellisolandia elongata (J.Ellis & Solander) K.R.Hind & G.W.Saunders). The brown alga Colpomenia sinuosa (Mertens ex Roth) Derbès & Solier is very common at this level, growing epiphytically on several other algae. Seasonally, the red algae Porphyra/ Neopyropia and/ or Nemalion elminthoides (Velley) Batters can be seen growing in patches at the mid-intertidal level. In some locations the brown crust Nemoderma tingitanum Schousboe ex Bornet can be common at this shore level (Neto et al. 2005, Wallenstein et al. 2009b). In spring and summer, considerable amounts of the introduced red alga Asparagopsis armata Harvey can be seen at the lower intertidal level, normally as an epiphyte on other algae (Neto, personal observation).
The adjacent submerged zone is also dominated by algal vegetation, with the rocky bottoms covered by more frondose macrophytes (Neto 2001, Wallenstein et al. 2009b) such as the red algae Asparagopsis taxiformis (Delile) Trevisan, Ellisolandia elongata, Jania spp., Plocamium cartilagineum (Linnaeus) P.S.Dixon, Pterocladiella capillacea, and Sphaerococcus coronopifolius Stackhouse, and the brown algae Dictyopteris polypodioides (A.P.De Candolle) J.V.Lamouroux, Dictyota spp., Gongolaria abies marina (S.G.Gmelin) Kuntze, Halopteris scoparia (Linnaeus) Sauvageau, and Zonaria tournefortii (J.V.Lamouroux) Montagne. The introduced red alga Symphyocladia marchantioides (Harvey) Falkenberg can be locally abundant below 15 m depth, usually as an ephiphyte on calcareous crusts; Hypnea musciformis (Wulfen) J.V.Lamouroux and Dasya spp. are other red algal species that can be locally abundant. The green species Codium elisabethiae and the brown species Padina pavonica (Linnaeus) Thivy can also be locally common, mainly in locations with sand influence (Neto 2001).
Design description: The sampling referred to in this study was performed across littoral and sublittoral levels down to approximately 40 m. Each sampling location was visited several times and, on each occasion, a careful and extensive survey was undertaken to provide a good coverage of the area. Both presence recording and physical collections were made by walking over the intertidal shores during low tides or by SCUBA diving in the subtidal. The specimens collected were taken to the laboratory for identification and preservation and the resulting vouchers were deposited at the AZB Herbarium Ruy Telles Palhinha and the LSM- Molecular Systematics Laboratory at the Faculty of Sciences and Technology of the University of the Azores.

Data published through GBIF: http://ipt.gbif.pt/ipt/resource?r=sao_miguel_macroalgal_flora

Taxonomic coverage

General taxonomic coverage description: All macroalgae were identified to genus or species level. In total, 431 taxa were identified belonging to 36 orders and 83 families, distributed among the phyla Rhodophyta (20 orders and 50 families), Chlorophyta (5 orders and 14 families), and Ochrophyta (11 orders and 19 families).
Taxonomic ranks
Phylum: Rhodophyta, Chlorophyta, Ochrophyta
Common names: Red algae, Green algae, Brown algae

Spatial coverage

General spatial coverage: São Miguel Island Description: Azores, Portugal (approximately 37°54'58''N, -25°51'52''W)
Coordinates: 37°42'8.26''N and 37°54'57.72''N Latitude; 25°51'52.43''W and 25°7'47.47''W Longitude 

Natural collections description

Parent collection identifier: AZB Herbarium Ruy Telles Palhinha, Faculty of Sciences and Technology of the University of the Azores.
Collection name: AZB | Marine macroalgae collection of São Miguel Island (Azores)-Campaign Macaronesia 2000
Collection identifier: 3cee8546-66d5-49c1-b63f-8efd4227ccc9
Natural collections description

Parent collection identifier: AZB Herbarium Ruy Telles Palhinha, Faculty of Sciences and Technology of the University of the Azores.
Collection name: AZB | Marine macroalgae collection of São Miguel Island (Azores)-Occasional sampling
Collection identifier: 5f198d55-a6ad-42f8-9342-3d96513fe808
Natural collections description

Parent collection identifier: AZB Herbarium Ruy Telles Palhinha, Faculty of Sciences and Technology of the University of the Azores.
Collection name: AZB | Marine macroalgae collection of São Miguel Island (Azores)-Project ABLA/MAC
Collection identifier: 15c76196-2b68-40cf-b49a-9392237f8d4d
Natural collections description

Parent collection identifier: AZB Herbarium Ruy Telles Palhinha, Faculty of Sciences and Technology of the University of the Azores.
Collection name: AZB | Marine macroalgae collection of São Miguel Island (Azores)-Project ASPAZOR
Collection identifier: c063f27f-5530-4e08-b50a-03843d61fb77
Natural collections description

Parent collection identifier: AZB Herbarium Ruy Telles Palhinha, Faculty of Sciences and Technology of the University of the Azores.
Collection name: AZB | Marine macroalgae collection of São Miguel Island (Azores)-Project Azorean Algal Flora
Collection identifier: 996ba65a-d07a-4f4f-a17f-511a40983710
Natural collections description

Parent collection identifier: AZB Herbarium Ruy Telles Palhinha, Faculty of Sciences and Technology of the University of the Azores.
Collection name: AZB | Marine macroalgae collection of São Miguel Island (Azores)-Project BIA (Biodiversity of Azores Archipelago)
Collection identifier: 445ca45c-3ba5-4a95-9289-39b944d6894e
Natural collections description

Parent collection identifier: AZB Herbarium Ruy Telles Palhinha, Faculty of Sciences and Technology of the University of the Azores.
Collection name: AZB | Marine macroalgae collection of São Miguel Island (Azores)-Project BIOTOPE
Collection identifier: 90fb970e-9be1-4caf-89ea-d3823235908a
Natural collections description

Parent collection identifier: AZB Herbarium Ruy Telles Palhinha, Faculty of Sciences and Technology of the University of the Azores.
Collection name: AZB | Marine macroalgae collection of São Miguel Island (Azores)-Project BUS
Collection identifier: 21215ae4-0e1f-44f0-95a8-dbed8022cfb0
Natural collections description

Parent collection identifier: AZB Herbarium Ruy Telles Palhinha, Faculty of Sciences and Technology of the University of the Azores.
Collection name: AZB | Marine macroalgae collection of São Miguel Island (Azores)-Project ECOSUBVEG
Collection identifier: 3dee61eb-0b56-4794-9d4e-75f2dce26918
Natural collections description

Parent collection identifier: AZB Herbarium Ruy Telles Palhinha, Faculty of Sciences and Technology of the University of the Azores.
Collection name: AZB | Marine macroalgae collection of São Miguel Island (Azores)-Project GESMAR
Collection identifier: 2840e5e1-4353-40f1-81c7-9c29ade05d2c
Natural collections description

Parent collection identifier: AZB Herbarium Ruy Telles Palhinha, Faculty of Sciences and Technology of the University of the Azores.
Collection name: AZB | Marine macroalgae collection of São Miguel Island (Azores)-Project LAUMACAT
Collection identifier: 7e12ab6e-a568-48aa-8ec3-0697ca734dac
Natural collections description

Parent collection identifier: AZB Herbarium Ruy Telles Palhinha, Faculty of Sciences and Technology of the University of the Azores.
Collection name: AZB | Marine macroalgae collection of São Miguel Island (Azores)-Project PATELGENE
Collection identifier: ee3acc7e-abcf-4dec-9c08-961e15d4c029
Natural collections description

Parent collection identifier: LSM- Molecular Systematics Laboratory, Faculty of Sciences and Technology of the University of the Azores
Collection name: LSM | Marine macroalgae collection of São Miguel Island (Azores)-Nordeste Expedition
Collection identifier: c4d4cb43-19d4-4633-b252-0e73e6600800
Natural collections description

Parent collection identifier: LSM- Molecular Systematics Laboratory, Faculty of Sciences and Technology of the University of the Azores
Collection name: LSM | Marine macroalgae collection of São Miguel Island (Azores)-Occasional sampling
Collection identifier: 1fbe2045-3ccb-4e38-9058-d7927e76db62
Natural collections description

Parent collection identifier: LSM- Molecular Systematics Laboratory, Faculty of Sciences and Technology of the University of the Azores
Collection name: LSM | Marine macroalgae collection of São Miguel Island (Azores)-Postdoc Manuela I Parente
Collection identifier: 22941d45-0678-49fb-bdfe-8b0052ceb298
Natural collections description

Parent collection identifier: LSM- Molecular Systematics Laboratory, Faculty of Sciences and Technology of the University of the Azores
Collection name: LSM | Marine macroalgae collection of São Miguel Island (Azores)-Project LusoMarBol
Collection identifier: ef2b1875-8520-4520-b6e0-48a1039f9b1f
Natural collections description

Parent collection identifier: LSM- Molecular Systematics Laboratory, Faculty of Sciences and Technology of the University of the Azores
Collection name: LSM | Marine macroalgae collection of São Miguel Island (Azores)-Sabrina Expedition
Collection identifier: 64a1cad8-6242-4075-b762-64b84350864c
Natural collections description

Parent collection identifier: LSM- Molecular Systematics Laboratory, Faculty of Sciences and Technology of the University of the Azores
Collection name: LSM | Marine macroalgae collection of São Miguel Islands (Azores)-Master Project Artur Oliveira
Collection identifier: 4a97a5db-4970-4f54-b598-d1125a6b8c63
Natural collections description

Parent collection identifier: : LSM- Molecular Systematics Laboratory, Faculty of Sciences and Technology of the University of the Azores
Collection name: LSM | Marine macroalgae collection of São Miguel Island (Azores)-Occasional sampling
Collection identifier: 494c9846-f867-4203-b476-42eb0789ca39
Natural collections description

Parent collection identifier: : LSM- Molecular Systematics Laboratory, Faculty of Sciences and Technology of the University of the Azores
Collection name: LSM | Marine macroalgae collection of São Miguel Island (Azores)-Project MACROBIOMOL
Collection identifier: 54b8c165-ab17-4489-9382-e19f7a6af090
Natural collections description

Parent collection identifier: Not applicable
Collection name: AZB | Marine macroalgae occurrence of São Miguel Island (Azores)-Project BIA (Biodiversity of Azores Archipelago)
Collection identifier: 3353c5f1-a12d-4c21-a0a2-c6c080d5d9fc
Natural collections description

Parent collection identifier: Not applicable
Collection name: AZB | Marine macroalgae occurrence of São Miguel Island (Azores)-Project BIOTOPE
Collection identifier: 2d7e1c20-23be-4e81-b349-4dba060ef8de
Natural collections description

Parent collection identifier: Not applicable
Collection name: AZB | Marine macroalgae occurrence of São Miguel Island (Azores)-Project BUS
Collection identifier: 5b048916-da07-4366-97f6-31741d804e51
Natural collections description

Parent collection identifier: Not applicable
Collection name: AZB | Marine macroalgae occurrence of São Miguel Island (Azores)-Project GESMAR
Collection identifier: 7a3f16c8-fb0e-4118-bf69-54dd20024146
Natural collections description

Parent collection identifier: Not applicable
Collection name: AZB | Marine macroalgae occurrence of São Miguel Island (Azores)-Occasional sampling
Collection identifier: 15799ce3-106e-48ff-933d-3d6a0a5b079d
Natural collections description

Parent collection identifier: Not applicable
Collection name: AZB | Marine macroalgae occurrence of São Miguel Island (Azores)-Project Azorean Algal Flora
Collection identifier: bd4dddc7-c708-4b0d-b912-e9a0a14a87f8
Natural collections description

Parent collection identifier: Not applicable
Collection name: AZB | Marine macroalgae occurrence of São Miguel Island (Azores)-Project CAMAG-ORI-SMG
Collection identifier: f9a307a6-1137-4265-baae-85924ef72ae7
Natural collections description

Parent collection identifier: Not applicable
Collection name: LSM | Marine macroalgae occurrence of São Miguel Island (Azores)-Occasional sampling
Collection identifier: b252cbd3-385a-4808-bd39-05add8d8eca0
Natural collections description

Parent collection identifier: Not applicable
Collection name: LSM | Marine macroalgae occurrence of São Miguel Island (Azores)-Campaign Waitt Foundation/2016
Collection identifier: 5efb9cd5-d89c-4ee9-b29d-9c4ce401e186
Natural collections description

Parent collection identifier: Not applicable
Collection name: LSM | Marine macroalgae occurrence of São Miguel Island (Azores)-Campaign Waitt Foundation-BALA/PIMA/2016
Collection identifier: fd958ad3-143a-4be7-bc89-0b58879c36ef
Natural collections description

Parent collection identifier: Not applicable
Collection name: LSM | Marine macroalgae occurrence of São Miguel Island (Azores)-PhD Andrea Z Botelho
Collection identifier: 52e0ab57-3fe9-436e-90d8-5ea2e1f4899e
Natural collections description

Parent collection identifier: Not applicable
Collection name: LSM | Marine macroalgae occurrence of São Miguel Island (Azores)-PIMA/2016
Collection identifier: 57fc6968-c1f3-4802-9c73-0609a61b8b10
Natural collections description

Parent collection identifier: Not applicable
Collection name: LSM | Marine macroalgae occurrence of São Miguel Island (Azores)-PIMA/BALA/2019
Collection identifier: f5b2f2cc-58c8-4bdf-9ab3-1fe3e5feea65
Living time period: 1989 - 2019
Specimen preservation method: Dried and pressed
Specimen preservation method: Dried
Specimen preservation method: Formalin
Specimen preservation method: Other

Methods

Method step description: At the laboratory, specimen sorting and macroalgae identification followed standard procedures. A combination of morphological and anatomical characters and reproductive structures was used for species identification. For small and simple thalli, this required the observation of the entire thallus with the naked eye and/or using dissecting and compound microscopes. For larger and more complex algae, investigation of the thallus anatomy required histological preparations (longitudinal and transverse sections) or squashed preparations of mucilaginous thalli, sometimes after staining, to observe vegetative and reproductive structures and other diagnostic features.
The Azorean algal flora has components from several geographical regions, which implies difficulties in species identification. Floras and keys for the North Atlantic, Tropical Atlantic and Western Mediterranean were, therefore, used (e.g. Schmidt 1931, Taylor 1967, 1978, Levring 1974, Dixon & Irvine 1977, Lawson & John 1982, Irvine 1983, Gayral & Cosson 1986, Fletcher 1987, Afonso-Carrillo & Sansón 1989, Burrows 1991, Boudouresque et al. 1992, Cabioc'h et al. 1992, Maggs & Hommersand 1993, Irvine & Chamberlain 1994, Brodie et al. 2007, Lloréns et al. 2012 and Rodríguez-Prieto et al. 2013). For more critical and taxonomically difficult taxa, specimens were taken to the Natural History Museum (London) for comparison with collections there.
A reference collection was made for all collected specimens by assigning them an herbarium code number and depositing them at the AZB Herbarium Ruy Telles Palhinha and the LSM- Molecular Systematics Laboratory, University of Azores. Depending on the species and on planned further research, different methods of preservation were used, namely (i) wet collections using 5% buffered formaldehyde seawater and then replacing it by the fixing agent Kew (Bridsen & Forman 1999); (ii) dried collections, either by pressing the algae (most species) as described by Gayral and Cosson (1986), or by letting them air dry (calcareous species); and (iii) silica gel collections for molecular studies.
Nomenclatural and taxonomic status used here follow Algaebase (Guiry & Guiry, 2021). The database was organized on FileMaker Pro.
Study extent description: The present publication includes sampling performed over a relatively large area,  covering littoral and sublittoral levels down to approximately 40 m around the Island.
Sampling description: Sampling involved species presence recording and/ or specimen collecting at each sampling location. Species recording data was gathered by registering all species present in the sampled locations. Destructive samples were obtained by scraping and/ or manually collecting one or two specimens of every species found. Intertidal collections were made during low tide by walking over the shores. Subtidal collections were made by SCUBA diving.
Quality control description: Each specimen collected was identified by trained taxonomists and involved morphological and anatomical observations of whole specimens by eye, and/or of histological preparations under the microscope to determine the main diagnostic features of each species as described in the literature.

Datasets

Dataset description
Object name: Darwin Core Archive Marine algal flora of São Miguel Island, Azores
Character encoding: UTF-8
Format name: Darwin Core Archive format
Format version: 1.0
Distribution: http://ipt.gbif.pt/ipt/archive.do?r=sao_miguel_macroalgal_flora
Publication date of data: 2021-02-14
Language: English
Licences of use: Creative Commons Attribution (CC-BY) 4.0 License

Metadata language: English
Date of metadata creation: 2021-01-31
Hierarchy level: Dataset

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