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. 2021 Apr 16;9:e64969. doi: 10.3897/BDJ.9.e64969

Marine algal flora of São Miguel Island, Azores

Ana I Azevedo Neto 1,, Ignacio Moreu 1, Edgar F Rosas Alquicira 2, Karla León-Cisneros 3, Eva Cacabelos 4,1, Andrea Z Botelho 5, Joana Micael 6, Ana C Costa 5, Raul M A Neto 7, José M N Azevedo 1, Sandra Monteiro 5, Roberto Resendes 8, Pedro Afonso 9, Afonso C L Prestes 1, Rita F Patarra 10,1, Nuno V Álvaro 11, David Milla-Figueras 9, Enric Ballesteros 12, Robert L Fletcher 13, William Farnham 13, Ian Tittley 14, Manuela I Parente 5
PMCID: PMC8065013  PMID: 33911918

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. Initial studies were mainly taxonomic, resulting in the publication of a checklist of the Azorean benthic marine algae. Later, the establishment of the University of the Azores on the Island permitted 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.

Prior to the present paper, the known macroalgal flora of São Miguel Island comprised 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), one an Azorean endemic (Predaea feldmannii subsp. azorica Gabriel), five are Macaronesian endemisms (the red algae 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 green alga Codium elisabethiae O.C.Schmidt), 19 are introduced species (15 Rhodophyta, two Chlorophyta and two Ochrophyta) and 32 are of uncertain status (21 Rhodophyta, five Chlorophyta and six Ochrophyta).

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

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, Neto 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 organisation (Schmidt 1931). Ever since the first half of the 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, Neto 2000, Neto 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 well 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, Haroun et al. 2019 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, Cacabelos et al. 2020, Freitas et al. 2019, Kellaris et al. 2019, Martins et al. 2019, Parente et al. 2019, Parente et al. 2020, Patarra et al. 2017, Patarra et al. 2019, Patarra et al. 2020, Sousa et al. 2019, Faria et al. 2020a, Faria et al. 2020b, 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, 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 (Leon-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, Leon-Cisneros et al. 2012). Some species, relatively common in the region a few years ago, have become uncommon or even very rare, for example, Scytosiphon lomentaria (Lyngbye) Link, Schimmelmannia schousboei (J.Agardh) J.Agardh. In contrast, there has been an increase in 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, Cacabelos et al. 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 itself to have a mixed nature, with species shared with Macaronesia, North Africa, the Mediterranean Sea, Atlantic Europe and America (Tittley and Neto 1995, Tittley and Neto 2005, Tittley and Neto 2006, Tittley 2003, 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 re-definition 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, opportunistic 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 organisations and the general public.

General description

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).

Project description

Title

Marine algal flora of São Miguel Island, Azores

Personnel

Collections were conducted and occurrence data recorded over several years (1989 - 2019). The main collectors were Adriá Pajares, Afonso C. L. Prestes, Alexandra Pacheco, Amine Sebti, Ana Bettencourt, Ana Carreiro, Ana Cristina Costa, Ana F. Ferreira, Ana Filipa Sousa, Ana I. Neto, Ana Leonado, Ana Rita Carreiro, Ana Rodriguez, André Amaral, André Gillon, Andrea Salamanca, Andrea Z. Botelho, Andreia Levi, Andreia Tracana, Anna Lloveras, Antalova Janouchová, Artur Oliveira, Brigida Garcia, Bruno Magalhães, Bruno Sérgio, Camille Fontaine, Carlos Campos, Carlos Mir, Carlos Rius, Carolina Moreira, Catarina Santos, Célia Albuquerque, Clara Gaspar, Cláudia Hipólito, Cristiana Figueredo, Cristina Seijo, Dálida Pereira, Daniel Torrão, Daniela Gabriel, David Milla-Figueras, Délia Cravo, Dinis Geraldes, Dolores Campos, Edgar F. Rosas-Alquicira, Emanuel Xavier, Enric Ballesteros, Eunice Nogueira, Eva Cacabelos, Fernando Feiteira, Filipe Parreira, Flávio Rodrigues, Francisco Wallenstein, Gloria Cantos, Gustavo Martins, Heather Baldwin, Helena Abreu, Hélio Dias, Hugo Lopes, Ian Tittley, Ignacio Moreu, Isadora Moniz, Joana Duarte, Joana Matzen, Jana Verdura, Joana Michael, João Brum, João Faria, João Feijó, José M. N. Azevedo, José Medeiros, Juan Garcia Marino, Juan Izaguirre, Juliana Dal Molin, Juliane Bernardi, Karla León-Cisneros, Laura Rovira, M. Canto, Marco Enoch, Margarida Leonardo, Manuela I. Parente, Marc Balcells, Marc Fernandez, Marco Henrique, Marco Santos, Maria Ana Dionísio, Maria Inês Pavão, Maria Machín-Sánchez, Maria Vale, Mariano Rego, Marisa Toste, Marlene Terra, Marta Coca, Miguel Frada, Mikel Mendizabal, Miguel Furtado, Miguel Matias, Miriam Gutierrez, Mutue Toyota Fujii, Natália Jardim, Nikola Zic, Nil Alvarez Segura, Nuno Vaz Álvaro, Núria Vila, O. Laclaustra, Olaia Morán, Olalla Torrontegi, Olivie Laroche, Patrícia Madeira, Patrícia Pereira, Paula Avelar, Paulo Azevedo, Paulo Custódio, Paulo Torres, Pedro Cavazin, Pedro Cerqueira, Pedro Raposeiro, Pedro Rodrigues, Rafael Fraga, Raquel Torres, Renato Calado, Ricardo Lacerda, Rita F. Patarra, Rita Grilo, Rita Norberto, Robert Fletcher, Rocio Sanchez, Roger Fuste, Ruben Couto, Rui Costa, Rui Jesus, Rui Moreira, Rui Patrício, Rui Sousa, Sabrina Garcia, Sandra Monteiro, Sara Peres, Sérgio Ávila, Silvia Escarduça, Sofia Carreiro, Susan Clayden, Tarso Costa, Valeria Cassano and William Farnham.

Preliminary in situ identifications were undertaken by: Ana I Neto, Andrea Z. Botelho, Andreia Levi, Daniela Gabriel, David Milla-Figueras, Edgar F. Rosas-Alquicira, Enric Ballesteros, Eva Cacabelos, Francisco Wallenstein, Heather Baldwin, Ian Tittley, Ignacio Moreu, Karla León-Cisneros, Manuela I. Parente, Maria Machín-Sanchez, Marlene Terra, Mutue Toyota Fujii, Nuno Vaz Álvaro, Raquel Torres, Robert Fletcher, Ruben Couto, Valeria Cassano and William Farnham.

Final species identification were undertaken by Ana I. Neto, Daniela Gabriel, Edgar F. Rosas-Alquicira, Enric Ballesteros, Eva Cacabelos, Ian Tittley, Ignacio Moreu, Karla León-Cisneros, Manuela I. Parente, Maria Machín-Sanchez, Marlene Terra, Mutue Toyota Fujii, Robert Fletcher, Valeria Cassano and William Farnham.

Voucher specimen management was mainly undertaken by Afonso C.L. Prestes, Ana I. Neto, Eunice Nogueira, Manuela I. Parente, Natália Cabral, Rita Patarra and Roberto Resendes.

Study area description

The Azores Archipelago (38°43′49″N, 27°19′10″W, Fig. 1), 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).

Figure 1.

Figure 1.

The Azores, its location in the Atlantic and São Miguel Island highlighted in black (by Nuno V. Álvaro).

The Archipelago comprises nine volcanic Islands and several small Islets in three separate groups (eastern, central and western).

São Miguel (in black in Fig. 1), approximately 750 km² in size, is the largest and most volcanically-active Island (Gaspar et al. 2015). Located in the eastern group of the Archipelago (37°54'58''N, -25°51'52''W, Fig. 2), its formation followed a series of volcanic events, with different parts of the Island having different ages. The oldest portion (4 M years old) is the eastern side, Nordeste, where Pico da Vara (the highest mountain of the Island with 1103 m a.s.l.) is located. The Island was then progressively formed to the west: Povoação (2 M years); Furnas (750,000 years); Serra de Água de Pau (250,000 years). The Sete Cidades complex appeared 500,000 years ago and only later (50,000 years ago) was connected to Serra de Agua de Pau through the Serra Gorda and its succeeding line of peaks (Zbyszewski et al. 1958, Zbyszewski and Ferreira 1959).

Figure 2.

Figure 2.

São Miguel Island with indication of the sampling locations (by Nuno V. Álvaro).

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 a.s.l. (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 exhibit a mosaic and/or zoned distribution pattern and have a predominance of algal turfs that cover the rocks as a carpet (Wallenstein et al. 2009), 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 (Fig. 3), while the mid-level zone is usually characterised by a fringe of chthamalid barnacles (Fig. 4), in which sometimes algae and limpets can occur (Fig. 5), followed by a lower area, in which either algal turf (generally monospecific and usually composed of Caulacanthus ustulatus (Turner) Kützing) dominates (Fig. 6) or patches of the brown alga Fucus spiralis Linnaeus and the red agarophyte Gelidium microdon Kützing (Fig. 7) 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, Fig. 8) or by various species of frondose algae growing in bands (e.g. the brown alga Gongolaria abies-marina (S.G.Gmelin) Kuntze, Fig. 9) 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, Fig. 10). 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).

Figure 3.

Figure 3.

Littorinids, a characteristic gastropod species of the Azorean high intertidal level (by the Island Aquatic Ecology Subgroup of cE3c-ABG).

Figure 4.

Figure 4.

Chthamalid barnacles on São Miguel mid-intertidal level (by the Island Aquatic Ecology Subgroup of cE3c-ABG).

Figure 5.

Figure 5.

Chthamalid barnacles, algal turf and limpet on São Miguel mid-intertidal level (by the Island Aquatic Ecology Subgroup of cE3c-ABG).

Figure 6.

Figure 6.

Algal turf, dominated by the red alga Caulacanthus ustulatus, on São Miguel mid-intertidal level (by the Island Aquatic Ecology Subgroup of cE3c-ABG).

Figure 7.

Figure 7.

The brown alga Fucus spiralis and the red agarophyte Gelidium microdon on São Miguel mid-intertidal level (by the Island Aquatic Ecology Subgroup of cE3c-ABG).

Figure 8.

Figure 8.

Multispecific algal turf and the coralline red alga Ellisolandia elongata on São Miguel low intertidal level (by the Island Aquatic Ecology Subgroup of cE3c-ABG).

Figure 9.

Figure 9.

The brown alga Gongolaria abies-marina growing in bands at the low shore level (by the Island Aquatic Ecology Subgroup of cE3c-ABG).

Figure 10.

Figure 10.

Patches of the agarophyte Pterocladiella capillacea and the calcareous Ellisolandia elongata at the low intertidal level (by the Island Aquatic Ecology Subgroup of cE3c-ABG).

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).

At cobble locations, the zonation pattern of macroalgae species is not clear (Costa 1994). The many microhabitats and substrate instability tend to mask and attenuate the limits of the biological zones. Nevertheless, in locations where cobbles are large and their size enlarges towards the sea (e.g. Fenais da Luz, north shore), the profile is steeper and usually the mid-intertidal level is dominated by the green macroalgae Ulva linza Linnaeus, U. clathrata (Roth) C.Agardh and U. rigida C.Agardh; the lower level is characterised by the presence of algal turf, mainly composed of Jania crassa J. V. Lamouroux and Corallina officinalis Linnaeus, with epiphytic Rhodophyta, such as Asparagopsis armata Harvey, phase Falkenbergia rufolanosa (Harvey) F.Schmitz, Centroceras clavulatum (C.Agardh) Montagne, Ceramium ciliatum (J.Ellis) Ducluzeau, C. deslongchampsii Chauvin ex Duby and Polysiphonia atlantica Kapraun & J.N.Norris. In locations with small cobbles (e.g. Caloura, south coast) or where there is a mixture of large and small cobbles (e.g. Povoação, south coast), the mid-intertidal level is usually characterised by fast growing algae, such as Cyanobacteria and the green algae Ulva spp., whereas the lowest level is also dominated by algal turf, but here mainly composed of C. officinalis and C. clavulatum (Caloura) or by C. clavulatum, Chondracanthus acicularis (Roth) Fredericq, Jania sp. and Lophosiphonia sp. (Povoação).

Important habitats at the shore level in bedrock locations are rock pools (Fig. 11). Differing in shape and size, they recreate a shallow subtidal habitat which may contain a rich diversity of marine algae and other marine organisms (Neto et al. 2005, Wallenstein et al. 2010).

Figure 11.

Figure 11.

Low shore pool (by the Island Aquatic Ecology Subgroup of cE3c-ABG).

The macroalgae diversity varies according to the pool location on the shore. Pools in the upper shore region are dominated by green algae, whilst those lower on the shore are dominated by red and brown algae. Similarly, faunal diversity in rock pools is greater at lower intertidal levels.

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 (Fig. 12). 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 (Fig. 13) and the brown species Padina pavonica (Linnaeus) Thivy can also be locally common, mainly in locations with sand influence (Neto 2001).

Figure 12.

Figure 12.

The frondose brown algae Zonaria tournefortii and Dictyota spp. at the deepest level sampled (by the Island Aquatic Ecology Subgroup of cE3c-ABG).

Figure 13.

Figure 13.

The Macaronesian endemic Codium elisabethiae on the shallow bottoms of São Miguel Island (by the Island Aquatic Ecology Subgroup of cE3c-ABG).

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.

Funding

This study was mainly financially supported by the following projects/scientific expeditions:

  • Projects:
    • ABLA/MAC – “Associações Biológicas do Litoral Açoreano/Moluscos, Algas e Crustáceos”, funded by the Portuguese Science and Technology Foundation (1987-1991);
    • Azorean Algal Flora – “Studies on algal communities of São Miguel, Azores”, partially funded by CIRN/DB/UAc (1992-1996);
    • BIA - “Biodiversity of Azores Archipelago”, funded by the Portuguese Science and Technology Foundation. PRAXIS/2/2.1/BIA/169/94 (1996-1999);
    • BIOTOPE – “ Classification, mapping and modelling of Azorean littoral biotopes”, funded by the Portuguese Science and Technology Foundation, POCTI MGS/45319/2002 (2003-2006);
    • 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);
    • GESMAR – “ Sustainable management of marine Resources”, funded by the EU Funding Programme III B 2000-2006, Açores-Madeira-Canárias, GESMAR/MAC/2/C068 (2009-2012);
    • 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);
    • MACROBIOMOL – “Macroalgae biodiversity under a molecular view - for a better understanding of North Atlantic Biogeography”, funded by PTDC/MAR/114613/2009 (2011-2015);
    • 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);
    • 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);
    • 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);
    • 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);
    • 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);
    • 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);
    • 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);
    • 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:
    • “Campaign Macaronesia 2000”, under the project Macaronesia 2000 (2000-2001);
    • “Waitt Foundation”, under the projects BALA and PIMA (2016);
    • “BALA/PIMA”, under the projects BALA and PIMA (2018);
    • “PORBIOTA/2019” under the project ACORES-01-0145-FEDER-000072 - AZORES BIOPORTAL – PORBIOTA (2019);
  • Other funds:
    • 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;
    • European Regional Development (ERD) funds through the Operational Programme for Competitiveness Factors (COMPETE);
    • 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;
    • CIRN/DB/UAc (Research Centre for Natural Resources, Universidade dos Açores, Departamento de Biologia);
    • CIIMAR (Interdisciplinary Centre of Marine and Environmental Research, Porto, Portugal).

Sampling methods

Study extent

The present publication includes sampling performed over a relatively large area, covering littoral and sublittoral levels down to approximately 40 m around the Island (Table 1, Fig. 2).

Table 1.

Information and location of the sampling sites on São Miguel Island

Location N0 Location ID Municipality Locality Latitude / Longitude Littoral zone
1 SMG_L_APs Lagoa Água de Pau | Subtidal 37°43'08''N, 25°27'45''W Subtidal
2 SMG_L_APsE Lagoa Água de Pau | Subtidal E 37°43'10''N, 25°27'44''W Subtidal
3 SMG_L_APsW Lagoa Água de Pau | Subtidal W 37°43'27''N, 25°27'16''W Subtidal
4 SMG_L_Avb Lagoa Atalhada | Viteleiro | baía 37°44'38''N, 25°23'23''W Subtidal
5 SMG_L_Avem Lagoa Atalhada | Viteleiro | Entre-marés 37°44'43''N, 25°23'24''W Intertidal
6 SMG_L_Cbab Lagoa Caloura | Baixa da Areia | baía 37°42'50''N, 25°28'11''W Subtidal
7 SMG_L_Ccb Lagoa Caloura | Cerco | baía 37°42'24''N, 25°29'30''W Subtidal
8 SMG_L_CcbW Lagoa Caloura | Cerco | baía W 37°42'21''N, 25°29'07''W Subtidal
9 SMG_L_Ccem Lagoa Caloura | Cerco | Entre-marés 37°42'27''N, 25°29'27''W Intertidal
10 SMG_L_Cepgb Lagoa Caloura | Entre Porto e Galera | baía 37°42'24''N, 25°29'52''W Subtidal
11 SMG_L_Cgb Lagoa Caloura | Galera | baía 37°42'10''N, 25°29'27''W Subtidal
12 SMG_L_Chem Lagoa Caloura | Hotel | Entre-marés 37°42'50''N, 25°28'59''W Intertidal
13 SMG_L_Cpb Lagoa Caloura | porto | baía 37°42'45''N, 25°30'15''W Subtidal
14 SMG_L_Cpem Lagoa Caloura | porto | Entre-marés 37°42'47''N, 25°30'15''W Intertidal
15 SMA_L_Crcb Lagoa Caloura | Ribeira Chã | baía 37°42'45''N, 25°30'16''W Subtidal
16 SMG_L_Lbp Lagoa Lagoa | Bairro dos Pescadores 37°44'23''N, 25°25'00''W Subtidal
17 SMG_L_Lcrb Lagoa Lagoa | Cruzeiro | baía 37°44'34''N, 25°25'56''W Subtidal
18 SMG_L_Lcrem Lagoa Lagoa | Cruzeiro | Entre-marés 37°44'32''N, 25°25'49''W Intertidal
19 SMG_L_Lovem Lagoa Lagoa | Observatório vulcanológico | Entre-marés 37°44'31''N, 25°24'44''W Intertidal
20 SMG_L_Lpib Lagoa Lagoa | Piscina | baía 37°44'26''N, 25°25'33''W Subtidal
21 SMG_L_Lpiem Lagoa Lagoa | Piscina | Entre-marés 37°44'29''N, 25°25'34''W Intertidal
22 SMG_L_Lpb Lagoa Lagoa | porto | baía 37°44'26''N, 25°25'25''W Subtidal
23 SMG_L_Pepem Lagoa Pisão | Entre praias | Entre-marés 37°43'02''N, 25°31'12''W Intertidal
24 SMG_N_AN Nordeste Achada do Nordeste 37°51'34''N, 25°43'23''W Intertidal
25 SMG_N_LGpsem Nordeste Lombo Gordo | Ponta do Sossego | Entre-marés 37°47'18''N, 25°51'27''W Intertidal
26 SMG_N_Nb6 Nordeste Nordeste | baía 6 37°50'21''N, 25°52'15''W Subtidal
27 SMG_N_Nb7 Nordeste Nordeste | baía 7 37°48'55''N, 25°53'22''W Subtidal
28 SMG_N_Npinb Nordeste Nordeste | Piscinas Naturais | baía 37°50'39''N, 25°51'17''W Subtidal
29 SMG_N_Npmb Nordeste Nordeste | Ponta da Madrugada | baía 37°49'30''N, 25°52'25''W Subtidal
30 SMG_N_Npmqb Nordeste Nordeste | Ponta da Marquesa | baía 37°47'59''N, 25°51'35''W Subtidal
31 SMG_N_Npaem Nordeste Nordeste | Ponta do Arnel | Entre-marés 37°49'21''N, 25°51'49''W Intertidal
32 SMG_PD_Bab Ponta Delgada Bretanha | Ajuda | baía 37°54'02''N, 25°14'56''W Subtidal
33 SMG_PD_Baem Ponta Delgada Bretanha | Ajuda | Entre-marés 37°54'00''N, 25°15'00''W Intertidal
34 SMG_PD_Cb Ponta Delgada Candelária | baía 37°49'15''N, 25°10'04''W Subtidal
35 SMG_PD_CPpb Ponta Delgada Capelas | porto | baía 37°50'32''N, 25°18'46''W Subtidal
36 SMG_PD_FLem Ponta Delgada Fenais da Luz | Entre-marés 37°49'54''N, 25°22'23''W Intertidal
37 SMG_PD_Fb Ponta Delgada Ferraria | baía 37°51'26''N, 25°08'51''W Subtidal
38 SMG_PD_Fem Ponta Delgada Ferraria | Entre-marés 37°51'30''N, 25°08'50''W Intertidal
39 SMG_PD_Ftb Ponta Delgada Feteiras | baía 37°46'57''N, 25°13'24''W Subtidal
40 SMG_PD_Ftem Ponta Delgada Feteiras | Entre-marés 37°48'13''N, 25°11'47''W Intertidal
41 SMG_PD_Ftpfgs Ponta Delgada Feteiras | Ponta da Fonte Grande | Subtidal 37°46'12''N, 25°13'48''W Subtidal
42 SMG_PD_Ftpem Ponta Delgada Feteiras | Porto | Entre-marés 37°48'12''N, 25°11'49''W Intertidal
43 SMG_PD_Mib Ponta Delgada Mosteiros | Ilhéus | baía 37°53'20''N, 25°10'00''W Subtidal
44 SMG_PD_Mpinb Ponta Delgada Mosteiros | piscinas naturais | baía 37°54'02''N, 25°10'31''W Subtidal
45 SMG_PD_Mpinemb Ponta Delgada Mosteiros | piscinas naturais | Entre-marés | Blocos 37°53'56''N, 25°10'58''W Intertidal
46 SMG_PD_Mpinemc1 Ponta Delgada Mosteiros | piscinas naturais | Entre-marés | Calhau 1 37°53'58''N, 25°10'45''W Intertidal
47 SMG_PD_Mpinemc2 Ponta Delgada Mosteiros | piscinas naturais | Entre-marés | Calhau 2 37°53'57''N, 25°10'56''W Intertidal
48 SMG_PD_Mpineme1 Ponta Delgada Mosteiros | piscinas naturais | Entre-marés | Escoada 1 37°54'01''N, 25°10'50''W Intertidal
49 SMG_PD_Mpineme2 Ponta Delgada Mosteiros | piscinas naturais | Entre-marés | Escoada 2 37°53'59''N, 25°10'44''W Intertidal
50 SMG_PD_Mpineme3 Ponta Delgada Mosteiros | piscinas naturais | Entre-marés | Escoada 3 37°53'59''N, 25°10'46''W Intertidal
51 SMG_PD_Mpineme4 Ponta Delgada Mosteiros | piscinas naturais | Entre-marés | Escoada 4 37°54'02''N, 25°10'54''W Intertidal
52 SMG_PD_Mpinpo Ponta Delgada Mosteiros | piscinas naturais | Poças 37°54'01''N, 25°10'48''W Intertidal
53 SMG_PD_Mpem Ponta Delgada Mosteiros | porto | Entre-marés 37°53'33''N, 25°10'42''W Intertidal
54 SMG_PD_Mpremw Ponta Delgada Mosteiros | Praia | Entre-marés (W) 37°53'20''N, 25°10'27''W Intertidal
55 SMG_PD_Pdcacb Ponta Delgada Ponta Delgada | Calheta | atrás da cadeia | baía 37°44'30''N, 25°20'46''W Subtidal
56 SMG_PD_PDeb Ponta Delgada Ponta Delgada | Etar | baía 37°44'27''N, 25°21'02''W Subtidal
57 SMG_PD_PDeem Ponta Delgada Ponta Delgada | Etar | Entre-marés 37°44'29''N, 25°21'02''W Intertidal
58 SMG_PD_PDeem Ponta Delgada Ponta Delgada | Etar | Entre-marés 37°44'29''N, 25°21'02''W Subtidal
59 SMG_PD_PDmle Ponta Delgada Ponta Delgada | marina | lado externo 37°44'22''N, 25°20'35''W Subtidal
60 SMG_PD_Pdmleem Ponta Delgada Ponta Delgada | marina | lado externo | Entre-marés 37°44'22''N, 25°20'29''W Intertidal
61 SMG_PD_PDmli Ponta Delgada Ponta Delgada | marina | lado interno 37°44'26''N, 25°20'29''W Subtidal
62 SMG_PD_PDmn Ponta Delgada Ponta Delgada | marina nova 37°44'20''N, 25°20'05''W Intertidal
63 SMG_PD_PDmn Ponta Delgada Ponta Delgada | marina nova 37°44'20''N, 25°20'05''W Subtidal
64 SMG_PD_PDpe Ponta Delgada Ponta Delgada | pesqueiro 37°44'21''N, 25°20'18''W Subtidal
65 SMG_PD_PDple Ponta Delgada Ponta Delgada | porto | lado externo 37°44'06''N, 25°20'28''W Subtidal
66 SMG_PD_PDpli Ponta Delgada Ponta Delgada | porto | lado interno 37°44'11''N, 25°20'32''W Subtidal
67 SMG_PD_PDscb Ponta Delgada Ponta Delgada | Santa Clara | baía 37°43'53''N, 25°18'59''W Subtidal
68 SMG_PD_PDscfem Ponta Delgada Ponta Delgada | Santa Clara | Farol | Entre-marés 37°43'58''N, 25°18'46''W Intertidal
69 SMG_PD_PDscpoem Ponta Delgada Ponta Delgada | Santa Clara | pontão | Entre-marés 37°43'58''N, 25°18'45''W Intertidal
70 SMG_PD_Popmem Ponta Delgada Pópulo | Praia das milícias | Entre-marés 37°44'58''N, 25°22'38''W Intertidal
71 SMG_PD_Poppem Ponta Delgada Pópulo | Praia pequena | Entre-marés 37°44'56''N, 25°22'59''W Intertidal
72 SMG_PD_Rb Ponta Delgada Relva | baía 37°45'57''N, 25°15'21''W Subtidal
73 SMG_R_bes Ponta Delgada Relva | Baixa do Espelho | Subtidal 37°45'41''N, 25°15'29''W Subtidal
74 SMG_PD_SAACem Ponta Delgada Santo António Além Capelas | Entre-marés 37°51'52''N, 25°18'00''W Intertidal
75 SMG_PD_SRd Ponta Delgada São Roque | Dori 37°44'42''N, 25°22'22''W Subtidal
76 SMG_PD_SRfcem Ponta Delgada São Roque | Forno da Cal | Entre-marés 37°44'39''N, 25°21'34''W Intertidal
77 SMG_PD_SRpb Ponta Delgada São Roque | Pranchinha | baía 37°44'37''N, 25°21'08''W Subtidal
78 SMG_PD_SRpem Ponta Delgada São Roque | Pranchinha | Entre-marés 37°44'38''N, 25°21'07''W Intertidal
79 SMG_PD_SRrcb Ponta Delgada São Roque | Rosto do Cão | baía 37°44'36''N, 25°21'39''W Subtidal
80 SMG_PD_SRrcem1 Ponta Delgada São Roque | Rosto do Cão | Entre-marés 1 37°44'40''N, 25°21'37''W Intertidal
81 SMG_PD_SRrcem2 Ponta Delgada São Roque | Rosto do Cão | Entre-marés 2 37°44'38''N, 25°21'40''W Intertidal
82 SMG_PD_SRrcem3 Ponta Delgada São Roque | Rosto do Cão | Entre-marés 3 37°44'40''N, 25°21'39''W Intertidal
83 SMG_PD_SVpobe Ponta Delgada São Vicente | Poços | baía (E) 37°50'06''N, 25°20'04''W Subtidal
84 SMG_PD_SVpobw Ponta Delgada São Vicente | Poços | baía (W) 37°50'04''N, 25°19'55''W Subtidal
85 SMG_PD_SVpoeme Ponta Delgada São Vicente | Poços | Entre-marés (E) 37°50'02''N, 25°19'53''W Intertidal
86 SMG_PD_SVpoemw Ponta Delgada São Vicente | Poços | Entre-marés (W) 37°50'03''N, 25°19'53''W Intertidal
87 SMG_PD_SVpoi Ponta Delgada São Vicente | Poços | ilhéu 37°50'03''N, 25°20'02''W Subtidal
88 SMG_P_bls Povoação Baixa da Lobeira | Subtidal 37°43'14''N, 25°40'52''W Subtidal
89 SMG_P_brqs Povoação Baixa da Ribeira Quente | Subtidal 37°43'26''N, 25°41'22''W subtidal
90 SMG_P_FTem Povoação Faial da Terra | Entre-marés 37°44'20''N, 25°48'12''W Intertidal
91 SMG_P_RQbf Povoação Ribeira Quente | baía | Fumarolas 37°43'36''N, 25°41'22''W Subtidal
92 SMG_P_RQbr Povoação Ribeira Quente | Baixa da Ribeira 37°43'59''N, 25°42'16''W Subtidal
93 SMG_P_RQborb Povoação Ribeira Quente | Boca da Ribeira | baía 37°43'59''N, 25°42'23''W Subtidal
94 SMG_P_RQrcem Povoação Ribeira Quente | Rua do Castelo | Entre-marés 37°43'43''N, 25°41'32''W Intertidal
95 SMG_RG_apgrpcpm21s Ribeira Grande Área Protegida de Gestão de Recursos da Ponta do Cintrão – Ponta da Maia (SMG21) | Subtidal 37°50'34''N, 25°30'51''W Subtidal
96 SMG_RG_FAem Ribeira Grande Fenais da Ajuda | Entre-marés 37°51'56''N, 25°41'01''W Intertidal
97 SMG_RG_Mamem Ribeira Grande Maia | Alameda do Mar | Entre-marés 37°50'03''N, 25°36'17''W Intertidal
98 SMG_RG_M_cms Ribeira Grande Maia | Cabecinhos da Maia | Subtidal 37°50'44''N, 25°36'00''W Subtidal
99 SMG_RG_Mfmb Ribeira Grande Maia | Frade da Maia | baía 37°50'17''N, 25°36'07''W Subtidal
100 SMG_RG_Mlmb Ribeira Grande Maia | Lomba da Maia | baía 37°50'56''N, 25°38'43''W Subtidal
101 SMG_RG_Mpinpb Ribeira Grande Maia | Piscinas naturais | pontas | baía 37°50'08''N, 25°36'20''W Subtidal
102 SMG_RG_Mpinpem Ribeira Grande Maia | Piscinas naturais | pontas | Entre-marés 37°50'10''N, 25°36'23''W Intertidal
103 SMG_RG_Mpem Ribeira Grande Maia | porto | Entre-marés 37°50'02''N, 25°36'47''W Intertidal
104 SMG_RG_PFbE Ribeira Grande Porto Formoso | baía E 37°50'00''N, 25°33'58''W Subtidal
105 SMG_RG_PFpb Ribeira Grande Porto Formoso | porto | baía 37°49'29''N, 25°34'22''W Subtidal
106 SMG_RG_PFpem Ribeira Grande Porto Formoso | porto | Entre-marés 37°49'27''N, 25°34'27''W Intertidal
107 SMG_RG_Pfprem Ribeira Grande Porto Formoso | praia | Entre-marés 37°49'25''N, 25°33'25''W Intertidal
108 SMG_RG_PFpmb Ribeira Grande Porto Formoso | Praia dos moinhos | baía 37°50'13''N, 25°33'05''W Subtidal
109 SMG_RG_PFsbb Ribeira Grande Porto Formoso | São Brás | baía 37°49'46''N, 25°35'08''W Subtidal
110 SMG_RG_Pfztem Ribeira Grande Porto Formoso | Zona termal | Entre-marés 37°49'36''N, 25°31'38''W Intertidal
111 SMG_RG_RPcem Ribeira Grande Rabo de Peixe | Calhetas | Entre-marés 37°49'28''N, 25°23'42''W Intertidal
112 SMG_RG_RGbem Ribeira Grande Ribeira Grande | Bandejo | Entre-marés 37°49'17''N, 25°27'51''W Intertidal
113 SMG_RG_Rfpb Ribeira Grande Ribeirinha | Furna da Pataca | baía 37°50'35''N, 25°30'52''W Subtidal
114 SMG_RG_Rpcb Ribeira Grande Ribeirinha | Ponta do Cintrão | baía 37°50'11''N, 25°31'20''W Subtidal
115 SMG_RG_Rpsib Ribeira Grande Ribeirinha | Porto de Santa Iria | baía 37°50'08''N, 25°30'59''W Subtidal
116 SMG_RG_RpsibW Ribeira Grande Ribeirinha | Porto de Santa Iria | baía W 37°50'34''N, 25°30'06''W Subtidal
117 SMG_RG_Rpsiem Ribeira Grande Ribeirinha | Porto de Santa Iria | Entre-marés 37°50'08''N, 25°30'56''W Intertidal
118 SMG_PD_SVpoemw Ribeira Grande São Vicente | Poços | Entre-marés (W) 37°50'03''N, 25°19'53''W Intertidal
119 SMG_VF_AA_bgs Vila Franca do Campo Água de Alto | Baixa da Garoupa | Subtidal 37°42'31''N, 25°31'47''W Subtidal
120 SMG_VF_AAbr Vila Franca do Campo Água de Alto | Boca da Ribeira 37°42'56''N, 25°32'48''W Intertidal
121 SMA_VF_AAtm Vila Franca do Campo Água de Alto | Três Marias 37°42'29''N, 25°31'56''W Subtidal
122 SMG_VF_PGb Vila Franca do Campo Ponta Garça | baía 37°42'56''N, 25°36'41''W Intertidal
123 SMG_VF_PGb Vila Franca do Campo Ponta Garça | baía 37°42'56''N, 25°36'41''W Subtidal
124 SMG_VF_PGbE Vila Franca do Campo Ponta Garça | baía E 37°42'46''N, 25°37'36''W Subtidal
125 SMG_VF_PGem1 Vila Franca do Campo Ponta Garça | Entre-marés 1 37°42'58''N, 25°36'48''W Intertidal
126 SMG_VF_PGem2 Vila Franca do Campo Ponta Garça | Entre-marés 2 37°42'59''N, 25°36'41''W Intertidal
127 SMG_VF_RPem Vila Franca do Campo Ribeira da Praia | Entre-marés 37°42'54''N, 25°34'43''W Intertidal
128 SMG_VF_RTem Vila Franca do Campo Ribeira das Tainhas | Entre-marés 37°42'55''N, 25°35'52''W Intertidal
129 SMG_VF_VFile Vila Franca do Campo Vila Franca do Campo | Ilhéu | lado externo 37°42'14''N, 25°33'27''W Subtidal
130 SMG_VF_VFilep Vila Franca do Campo Vila Franca do Campo | Ilhéu | lado externo | picos 37°42'00''N, 25°33'22''W Subtidal
131 SMG_VF_VFileE Vila Franca do Campo Vila Franca do Campo | Ilhéu | lado externo E 37°42'18''N, 25°33'32''W Subtidal
132 SMG_VF_VFileS Vila Franca do Campo Vila Franca do Campo | Ilhéu | lado externo S 37°42'14''N, 25°33'20''W Subtidal
133 SMG_VF_VFileSW Vila Franca do Campo Vila Franca do Campo | Ilhéu | lado externo SW 37°42'15''N, 25°33'19''W Subtidal
134 SMG_VF_VFmem Vila Franca do Campo Vila Franca do Campo | marina | Entre-marés 37°42'54''N, 25°34'16''W Intertidal
135 SMG_VF_VFmli Vila Franca do Campo Vila Franca do Campo | marina | lado interno 37°42'50''N, 25°34'12''W Subtidal
136 SMG_VF_VFpbw Vila Franca do Campo Vila Franca do Campo | porto | baía (W) 37°42'45''N, 25°33'53''W Subtidal
137 SMG_VF_VFpemw Vila Franca do Campo Vila Franca do Campo | porto | Entre-marés (W) 37°42'50''N, 25°33'58''W Intertidal
138 SMG_VF_VFpem Vila Franca do Campo Vila Franca do Campo | praia | Entre-marés 37°42'59''N, 25°34'22''W Intertidal

Sampling description

Sampling involved species presence recording and/or specimen collecting at each sampling location. Species recording data were gathered by registering all species present in the sampled locations (Fig. 14). Destructive samples were obtained by scraping and/or manually collecting one or two specimens of every species found (Fig. 15). Intertidal collections were made during low tide by walking over the shores. Subtidal collections were made by SCUBA diving.

Figure 14.

Figure 14.

Quantitative recording of the presence and coverage of macroalgal species at the intertidal rocky habitat (by the Island Aquatic Ecology Subgroup of cE3c-ABG).

Figure 15.

Figure 15.

Collecting macroalgae in the subtidal of São Miguel Island (by the Island Aquatic Ecology Subgroup of cE3c-ABG).

Quality control

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 literature.

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, Taylor 1978, Levring 1974, Dixon and Irvine 1977, Lawson and John 1982, Irvine 1983, Gayral and Cosson 1986, Fletcher 1987, Afonso-Carrillo and Sansón 1989, Burrows 1991, Boudouresque et al. 1992, Cabioc'h et al. 1992, Maggs and Hommersand 1993, Irvine and Chamberlain 1994, Brodie et al. 2007, Lloréns et al. 2012, 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 a 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 and 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 and Guiry 2021). The database was organised on FileMaker Pro.

Geographic coverage

Description

São Miguel Island Description: Azores, Portugal (approximately 37°54'58''N, 25°51'52''W).

Coordinates

37°42'45''N and 37°54'57''N Latitude; 25°52'10''W and 25°08'06''W Longitude.

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 amongst the phyla Rhodophyta (20 orders and 50 families), Chlorophyta (5 orders and 14 families) and Ochrophyta (11 orders and 19 families).

Taxa included

Rank Scientific Name Common Name
phylum Rhodophyta Red algae
phylum Chlorophyta Green algae
phylum Ochrophyta Brown algae

Temporal coverage

Notes

The sampling was performed on several occasions in the period between 1989 and 2019.

Collection data

Collection name

AZB | Marine macroalgae collection of São Miguel Island (Azores)-Campaign Macaronesia 2000; AZB | Marine macroalgae collection of São Miguel Island (Azores)-Occasional sampling; AZB | Marine macroalgae collection of São Miguel Island (Azores)-Project ABLA/MAC; AZB | Marine macroalgae collection of São Miguel Island (Azores)-Project ASPAZOR; AZB | Marine macroalgae collection of São Miguel Island (Azores)-Project Azorean Algal Flora; AZB | Marine macroalgae collection of São Miguel Island (Azores)-Project BIA (Biodiversity of Azores Archipelago); AZB | Marine macroalgae collection of São Miguel Island (Azores)-Project BIOTOPE; AZB | Marine macroalgae collection of São Miguel Island (Azores)-Project BUS; AZB | Marine macroalgae collection of São Miguel Island (Azores)-Project ECOSUBVEG; AZB | Marine macroalgae collection of São Miguel Island (Azores)-Project GESMAR; AZB | Marine macroalgae collection of São Miguel Island (Azores)-Project LAUMACAT; AZB | Marine macroalgae collection of São Miguel Island (Azores)-Project PATELGENE; LSM | Marine macroalgae collection of São Miguel Island (Azores)-Nordeste Expedition; LSM | Marine macroalgae collection of São Miguel Island (Azores)-Occasional sampling; LSM | Marine macroalgae collection of São Miguel Island (Azores)-Postdoc Manuela I Parente; LSM | Marine macroalgae collection of São Miguel Island (Azores)-Project LusoMarBol; LSM | Marine macroalgae collection of São Miguel Island (Azores)-Sabrina Expedition; LSM | Marine macroalgae collection of São Miguel Islands (Azores)-Master Project Artur Oliveira; LSM | Marine macroalgae collection of São Miguel Island (Azores)-Occasional sampling; LSM | Marine macroalgae collection of São Miguel Island (Azores)-Project MACROBIOMOL; AZB | Marine macroalgae occurrence of São Miguel Island (Azores)-Project BIA (Biodiversity of Azores Archipelago); AZB | Marine macroalgae occurrence of São Miguel Island (Azores)-Project BIOTOPE; AZB | Marine macroalgae occurrence of São Miguel Island (Azores)-Project BUS; AZB | Marine macroalgae occurrence of São Miguel Island (Azores)-Project GESMAR; AZB | Marine macroalgae occurrence of São Miguel Island (Azores)-Occasional sampling; AZB | Marine macroalgae occurrence of São Miguel Island (Azores)-Project Azorean Algal Flora; AZB | Marine macroalgae occurrence of São Miguel Island (Azores)-Project CAMAG-ORI-SMG; LSM | Marine macroalgae occurrence of São Miguel Island (Azores)-Occasional sampling; LSM | Marine macroalgae occurrence of São Miguel Island (Azores)-Campaign Waitt Foundation/2016; LSM | Marine macroalgae occurrence of São Miguel Island (Azores)-Campaign Waitt Foundation-BALA/PIMA/2016; LSM | Marine macroalgae occurrence of São Miguel Island (Azores)-PhD Andrea Z Botelho; LSM | Marine macroalgae occurrence of São Miguel Island (Azores)-PIMA/2016; LSM | Marine macroalgae occurrence of São Miguel Island (Azores)-PIMA/BALA/2019

Collection identifier

3cee8546-66d5-49c1-b63f-8efd4227ccc9; 5f198d55-a6ad-42f8-9342-3d96513fe808; 15c76196-2b68-40cf-b49a-9392237f8d4d; c063f27f-5530-4e08-b50a-03843d61fb77; 996ba65a-d07a-4f4f-a17f-511a40983710; 445ca45c-3ba5-4a95-9289-39b944d6894e; 90fb970e-9be1-4caf-89ea-d3823235908a; 21215ae4-0e1f-44f0-95a8-dbed8022cfb0; 3dee61eb-0b56-4794-9d4e-75f2dce26918; 2840e5e1-4353-40f1-81c7-9c29ade05d2c; 7e12ab6e-a568-48aa-8ec3-0697ca734dac; ee3acc7e-abcf-4dec-9c08-961e15d4c029; c4d4cb43-19d4-4633-b252-0e73e6600800; 1fbe2045-3ccb-4e38-9058-d7927e76db62; 22941d45-0678-49fb-bdfe-8b0052ceb298; ef2b1875-8520-4520-b6e0-48a1039f9b1f; 64a1cad8-6242-4075-b762-64b84350864c; 4a97a5db-4970-4f54-b598-d1125a6b8c63; 494c9846-f867-4203-b476-42eb0789ca39; 54b8c165-ab17-4489-9382-e19f7a6af090; 3353c5f1-a12d-4c21-a0a2-c6c080d5d9fc; 2d7e1c20-23be-4e81-b349-4dba060ef8de; 5b048916-da07-4366-97f6-31741d804e51; 7a3f16c8-fb0e-4118-bf69-54dd20024146; 15799ce3-106e-48ff-933d-3d6a0a5b079d; bd4dddc7-c708-4b0d-b912-e9a0a14a87f8; f9a307a6-1137-4265-baae-85924ef72ae7; b252cbd3-385a-4808-bd39-05add8d8eca0; 5efb9cd5-d89c-4ee9-b29d-9c4ce401e186; fd958ad3-143a-4be7-bc89-0b58879c36ef; 52e0ab57-3fe9-436e-90d8-5ea2e1f4899e; 57fc6968-c1f3-4802-9c73-0609a61b8b10; f5b2f2cc-58c8-4bdf-9ab3-1fe3e5feea65

Parent collection identifier

AZB Herbarium Ruy Telles Palhinha, Faculty of Sciences and Technology of the University of the Azores; AZB Herbarium Ruy Telles Palhinha, Faculty of Sciences and Technology of the University of the Azores; AZB Herbarium Ruy Telles Palhinha, Faculty of Sciences and Technology of the University of the Azores; AZB Herbarium Ruy Telles Palhinha, Faculty of Sciences and Technology of the University of the Azores; AZB Herbarium Ruy Telles Palhinha, Faculty of Sciences and Technology of the University of the Azores; AZB Herbarium Ruy Telles Palhinha, Faculty of Sciences and Technology of the University of the Azores; AZB Herbarium Ruy Telles Palhinha, Faculty of Sciences and Technology of the University of the Azores; AZB Herbarium Ruy Telles Palhinha, Faculty of Sciences and Technology of the University of the Azores; AZB Herbarium Ruy Telles Palhinha, Faculty of Sciences and Technology of the University of the Azores; AZB Herbarium Ruy Telles Palhinha, Faculty of Sciences and Technology of the University of the Azores; AZB Herbarium Ruy Telles Palhinha, Faculty of Sciences and Technology of the University of the Azores; AZB Herbarium Ruy Telles Palhinha, Faculty of Sciences and Technology of the University of the Azores; LSM - Molecular Systematics Laboratory, Faculty of Sciences and Technology of the University of the Azores; LSM - Molecular Systematics Laboratory, Faculty of Sciences and Technology of the University of the Azores; LSM - Molecular Systematics Laboratory, Faculty of Sciences and Technology of the University of the Azores; LSM - Molecular Systematics Laboratory, Faculty of Sciences and Technology of the University of the Azores; LSM - Molecular Systematics Laboratory, Faculty of Sciences and Technology of the University of the Azores; LSM - Molecular Systematics Laboratory, Faculty of Sciences and Technology of the University of the Azores; LSM - Molecular Systematics Laboratory, Faculty of Sciences and Technology of the University of the Azores; LSM - Molecular Systematics Laboratory, Faculty of Sciences and Technology of the University of the Azores; Not applicable; Not applicable; Not applicable; Not applicable; Not applicable; Not applicable; Not applicable; Not applicable; Not applicable; Not applicable; Not applicable; Not applicable; Not applicable

Specimen preservation method

Air dry, Dried and pressed; Wet (Formalin; fixing agent Kew), Silica gel

Curatorial unit

AZB Herbarium Ruy Telles Palhinha, Faculty of Sciences and Technology of the University of the Azores

Usage licence

Usage licence

Creative Commons Public Domain Waiver (CC-Zero)

Data resources

Data package title

Marine algal flora of São Miguel Island, Azores

Resource link

http://ipt.gbif.pt/ipt/resource?r=sao_miguel_macroalgal_flora

Alternative identifiers

https://www.gbif.org/dataset/322b5629-997c-4986-ada9-7d9d078d8648; https://doi.org/10.15468/xtuzd3

Number of data sets

1

Data set 1.

Data set name

Marine algal flora of São Miguel Island, Azores

Data format

Darwin Core Archive

Number of columns

50

Download URL

http://ipt.gbif.pt/ipt/resource?r=sao_miguel_macroalgal_flora&amp;v=1.0

Data format version

1.3

Description

This data paper presents physical and occurrence data from macroalgal surveys undertaken on São Miguel Island between 1989 and 2019 (Neto et al. 2021b). The dataset submitted to GBIF is structured as a sample event dataset, with two tables: event (as core) and occurrences. The data in this sampling event resource have been published as a Darwin Core Archive (DwCA), which is a standardised format for sharing biodiversity data as a set of one or more data tables. The core data table contains 506 records (eventID). The extension data table has 12,781 occurrences. An extension record supplies extra information about a core record. The number of records in each extension data table is illustrated in the IPT link. This IPT archives the data and thus serves as the data repository. The data and resource metadata are available for downloading in the downloads section.

Data set 1.
Column label Column description
eventID Identifier of the event, unique for the dataset
country Country of the sampling site
countryCode Code of the country where the event occurred
stateProvince Name of the region
island Name of the island
municipality Name of the municipality
locality Name of the locality
locationID Identifier of the location
decimalLatitude The geographic latitude of the sampling site
decimalLongitude The geographic longitude of the sampling site
geodeticDatum The spatial reference system upon which the geographic coordinates are based
coordinateUncertaintyInMetres The horizontal distance (in metres) from the given decimalLatitude and decimalLongitude describing the smallest circle containing the whole of the Location
eventDate Time interval when the event occurred
year The year of the event
samplingProtocol Sampling method used during an event
locationRemarks Zonation level
minimumDepthInMetres The minimum depth in metres where the specimen was found
maximumDepthInMetres The maximum depth in metres where the specimen was found
eventRemarks Notes about the event
occurrenceID Identifier of the record, coded as a global unique identifier
institutionID The identifier for the institution having custody of the object or information referred to in the record
institutionCode The acronym of the institution having custody of the object or information referred to in the record
collectionID An identifier of the collection to which the record belongs
collectionCode The name of the collection from which the record was derived
datasetName The name identifying the dataset from which the record was derived
kingdom Kingdom name
phylum Phylum name
class Class name
order Order name
family Family name
genus Genus name
specificEpithet The name of the first or species epithet of the scientificName
infraspecificEpithet The name of the lowest or terminal infraspecific epithet of the scientificName, excluding any rank designation
acceptedNameUsage The specimen accepted name, with authorship
previousIdentifications Previous name of the specimen, with authorship
scientificName The name without authorship applied on the first identification of the specimen
scientificNameAuthorship The authorship information for the scientificName formatted according to the conventions of the applicable nomenclaturalCode
taxonRank The taxonomic rank of the most specific name in the scientificName
basisOfRecord The specific nature of the data record
habitat Description of the habitat where the specimen was found
organismQuantityType The type of quantification system used to quantify the organisms
organismQuantity Percentage of the organism coverage
recordedBy Person(s) responsible for sampling
catalogNumber Identifying code for a unique sample lot in a biological collection
identifiedBy Person(s) responsible for taxa identification
type The nature of the resource
preparations The preservation method used for the specimen
establishmentMeans The establishment status of the organism in the study region
occurrenceRemarks New record status assignment
licence Reference to the licence under which the record is published

Additional information

This paper accommodates the 12,781 specimens of macroalgae recorded from São Miguel Island in 431 taxa comprising 323 confirmed species (Tables 2, 3) and 108 taxa identified only to genus level. The confirmed species (Table 3) include 212 Rhodophyta, 48 Chlorophyta and 63 Ochrophyta (Phaeophyceae). Of these, 61 species are newly recorded to the Island (42 Rhodophyta, 9 Chlorophyta and 10 Ochrophyta). Most species are native, Predaea feldmannii subsp. azorica is an Azorean endemic, whereas the rhodophyta Botryocladia macaronesica, Laurencia viridis, Millerella tinerfensis, Phyllophora gelidioides and the Chlorophyta Codium elisabethiae are Macaronesian endemics. Nineteen species represent introductions to the algal flora (the Rhodophyta Antithamnion diminuatum Wollaston, Antithamnion hubbsii E.Y.Dawson, Antithamnionella spirographidis (Schiffner) E.M.Wollaston, Antithamnionella ternifolia (J.D.Hooker & Harvey) Lyle, Asparagopsis armata, Grallatoria reptans M.Howe, Gymnophycus hapsiphorus Huisman & Kraft, Laurencia brongniartii J.Agardh, Laurencia dendroidea J.Agardh, Neoizziella divaricata (C.K.Tseng) S.-M.Lin, S.-Y.Yang & Huisman, Scageliopsis patens E.M.Wollaston, Symphyocladia marchantioides, Xiphosiphonia pennata (C.Agardh) Savoie & G.W.Saunders and Xiphosiphonia pinnulata (Kützing) Savoie & G.W.Saunders; the Chlorophyta Caulerpa prolifera (Forsskål) J.V.Lamouroux and Codium fragile subsp. fragile (Suringar) Hariot; and the Ochrophyta Papenfussiella kuromo (Yendo) Inagaki and Petalonia binghamiae (J.Agardh) K.L.Vinogradova. Thirty-two species have an uncertain status (21 Rhodophyta, 5 Chlorophyta and 6 Ochrophyta).

Table 2.

Macroalgal species recorded from São Miguel Island, with information on relative abundance, origin and status.

Phylum Species (Accepted Name) Number of records Establishment means Occurrence remarks
Rhodophyta Acrosorium ciliolatum (Harvey) Kylin 826 Native
Rhodophyta Agardhinula browneae (J.Agardh) De Toni 1 Native
Rhodophyta Aglaothamnion bipinnatum (P.Crouan & H.Crouan) Feldmann & G. Feldmann 24 Native New record
Rhodophyta Aglaothamnion pseudobyssoides (P.Crouan & H.Crouan) Halos 1 Native
Rhodophyta Aglaothamnion tenuissimum (Bonnemaison) Feldmann-Mazoyer 14 Uncertain
Rhodophyta Ahnfeltiopsis devoniensis (Greville) P.C.Silva & DeCew 5 Native
Rhodophyta Amphiroa beauvoisii J.V.Lamouroux 12 Native
Rhodophyta Amphiroa cryptarthrodia Zanardini 4 Native
Rhodophyta Amphiroa rigida J.V.Lamouroux 3 Native
Rhodophyta Anotrichium barbatum (C.Agardh) Nägeli 8 Native
Rhodophyta Anotrichium furcellatum (J.Agardh) Baldock 14 Uncertain
Rhodophyta Anotrichium tenue (C.Agardh) Nägeli 1 Native
Rhodophyta Antithamnion cruciatum (C.Agardh) Nägeli 11 Native New record
Rhodophyta Antithamnion decipiens (J.Agardh) Athanasiadis 1 Native
Rhodophyta Antithamnion diminuatum Wollaston 6 Introduced
Rhodophyta Antithamnion hubbsii E.Y.Dawson 7 Introduced New record
Rhodophyta Antithamnionella boergesenii (Cormaci & G.Furnari) Athanasiadis 1 Uncertain
Rhodophyta Antithamnionella floccosa (O.F.Müller) Whittick 1 Native New record
Rhodophyta Antithamnionella spirographidis (Schiffner) E.M.Wollaston 1 Introduced New record
Rhodophyta Antithamnionella ternifolia (J.D.Hooker & Harvey) Lyle 2 Introduced
Rhodophyta Aphanocladia stichidiosa (Funk) Ardré 2 Native
Rhodophyta Apoglossum ruscifolium (Turner) J.Agardh 2 Native New record
Rhodophyta Asparagopsis armata Harvey 188 Introduced
Rhodophyta Asparagopsis armata Harvey, phase Falkenbergia rufolanosa (Harvey) F.Schmitz 34 Introduced
Rhodophyta Asparagopsis taxiformis (Delile) Trevisan 154 Native
Rhodophyta Asteromenia peltata (W.R.Taylor) Huisman & A.J.K.Millar 6 Native
Rhodophyta Balliella cladoderma (Zanardini) Athanasiadis 3 Native New record
Rhodophyta Bangia atropurpurea (Mertens ex Roth) C.Agardh 7 Native
Rhodophyta Bonnemaisonia asparagoides (Woodward) C.Agardh 8 Native
Rhodophyta Bonnemaisonia hamifera Hariot 1 Introduced
Rhodophyta Bornetia secundiflora (J.Agardh) Thuret 6 Native
Rhodophyta Bostrychia scorpioides (Hudson) Montagne 1 Native New record
Rhodophyta Botryocladia botryoides (Wulfen) Feldmann 4 Native
Rhodophyta Botryocladia macaronesica Afonso-Carillo, Sobrino, Tittley & Neto 52 Macaronesian endemism
Rhodophyta Callithamnion corymbosum (Smith) Lyngbye 7 Native
Rhodophyta Callithamnion granulatum (Ducluzeau) C. Agardh 38 Native
Rhodophyta Callithamnion tetragonum (Withering) S.F.Gray 15 Native New record
Rhodophyta Callithamnion tetricum (Dillwyn) S.F.Gray 1 Native
Rhodophyta Carradoriella denudata (Dillwyn) Savoie & G.W.Saunders 26 Uncertain
Rhodophyta Carradoriella elongata (Hudson) A.M.Savoie & G.W.Saunders 1 Native
Rhodophyta Catenella caespitosa (Withering) L.M.Irvine 5 Native
Rhodophyta Caulacanthus ustulatus (Turner) Kützing 121 Uncertain
Rhodophyta Centroceras clavulatum (C.Agardh) Montagne 82 Native
Rhodophyta Ceramium botryocarpum A.W.Griffiths ex Harvey 12 Native
Rhodophyta Ceramium ciliatum (J.Ellis) Ducluzeau 25 Native
Rhodophyta Ceramium cimbricum H.E.Petersen 8 Native New record
Rhodophyta Ceramium circinatum (Kützing) J.Agardh 3 Native
Rhodophyta Ceramium deslongchampsii Chauvin ex Duby 6 Native
Rhodophyta Ceramium diaphanum (Lightfoot) Roth 9 Native
Rhodophyta Ceramium echionotum J.Agardh 10 Native
Rhodophyta Ceramium pallidum (Kützing) Maggs & Hommersand 24 Native New record
Rhodophyta Ceramium secundatum Lyngbye 19 Native New record
Rhodophyta Ceramium tenuicorne (Kützing) Waern 11 Native New record
Rhodophyta Ceramium virgatum Roth 23 Native
Rhodophyta Ceratodictyon intricatum (C.Agardh) R.E.Norris 2 Native
Rhodophyta Champia parvula (C.Agardh) Harvey 10 Native
Rhodophyta Chondracanthus acicularis (Roth) Fredericq 203 Native
Rhodophyta Chondracanthus teedei (Mertens ex Roth) Kützing 57 Native
Rhodophyta Chondria capillaris (Hudson) M.J.Wynne 2 Native
Rhodophyta Chondria coerulescens (J.Agardh) Sauvageau 29 Uncertain
Rhodophyta Chondria dasyphylla (Woodward) C.Agardh 22 Uncertain
Rhodophyta Coelothrix irregularis (Harvey) Børgesen 2 Native
Rhodophyta Compsothamnion decompositum (J.Agardh) Maggs & L'Hardy-Halos 2 Native
Rhodophyta Corallina ferreyrae E.Y.Dawson, Acleto & Foldvik 30 Native New record
Rhodophyta Corallina officinalis Linnaeus 140 Native
Rhodophyta Cottoniella filamentosa (M.Howe) Børgesen 15 Native New record
Rhodophyta Crouania attenuata (C.Agardh) J.Agardh 5 Native
Rhodophyta Cruoria pellita (Lyngbye) Fries 2 Native New record
Rhodophyta Cryptonemia crenulata (J.Agardh) J.Agardh 1 Uncertain New record
Rhodophyta Cryptonemia seminervis (C.Agardh) J.Agardh 2 Native
Rhodophyta Cryptopleura ramosa (Hudson) L.Newton 9 Native
Rhodophyta Dasya baillouviana (S.G.Gmelin) Montagne 6 Uncertain
Rhodophyta Dasya caraibica Børgesen 1 Native New record
Rhodophyta Dasya corymbifera J.Agardh 3 Native
Rhodophyta Dasya crouaniana J.Agardh 1 Native New record
Rhodophyta Dasya hutchinsiae Harvey 6 Native
Rhodophyta Dasya ocellata (Grateloup) Harvey 3 Native
Rhodophyta Dermocorynus dichotomus (J.Agardh) Gargiulo, M.Morabito & Manghisi 67 Native
Rhodophyta Diplothamnion jolyi C.Hoek 8 Native
Rhodophyta Drachiella heterocarpa (Chauvin ex Duby) Maggs & Hommersand 5 Native New record
Rhodophyta Dudresnaya crassa M.Howe 7 Native
Rhodophyta Dudresnaya verticillata (Withering) Le Jolis 2 Native
Rhodophyta Ellisolandia elongata (J.Ellis & Solander) K.R.Hind & G.W.Saunders 67 Native
Rhodophyta Erythrocystis montagnei (Derbès & Solier) P.C.Silva 4 Native
Rhodophyta Erythrodermis traillii (Holmes ex Batters) Guiry & Garbary 10 Uncertain
Rhodophyta Erythroglossum laciniatum (Lightfoot) Maggs & Hommersand 1 Native New record
Rhodophyta Erythrotrichia carnea (Dillwyn) J.Agardh 1 Uncertain
Rhodophyta Eupogodon planus (C.Agardh) Kützing 2 Native
Rhodophyta Gaillona gallica (Nägeli) Athanasiadis 19 Native
Rhodophyta Gaillona hookeri (Dillwyn) Athanasiadis 9 Native
Rhodophyta Gastroclonium clavatum (Roth) Ardissone 1 Native
Rhodophyta Gastroclonium ovatum (Hudson) Papenfuss 17 Native
Rhodophyta Gastroclonium reflexum (Chauvin) Kützing 23 Native
Rhodophyta Gayliella flaccida (Harvey ex Kützing) T.O.Cho & L.J.McIvor 8 Native
Rhodophyta Gelidium corneum (Hudson) J.V.Lamouroux 2 Native
Rhodophyta Gelidium microdon Kützing 228 Native
Rhodophyta Gelidium pusillum (Stackhouse) Le Jolis 52 Native
Rhodophyta Gelidium spinosum (S.G.Gmelin) P.C.Silva 150 Native
Rhodophyta Gigartina pistillata (S.G.Gmelin) Stackhouse 20 Native
Rhodophyta Gracilaria gracilis (Stackhouse) Steentoft, L.M.Irvine & Farnham 3 Native New record
Rhodophyta Gracilaria multipartita (Clemente) Harvey 1 Native
Rhodophyta Gracilariopsis longissima (S.G.Gmelin) Steentoft, L.M.Irvine & Farnham 12 Native
Rhodophyta Grallatoria reptans M.Howe 2 Introduced
Rhodophyta Grateloupia filicina (J.V.Lamouroux) C.Agardh 21 Native
Rhodophyta Griffithsia corallinoides (Linnaeus) Trevisan 2 Native
Rhodophyta Griffithsia phyllamphora J.Agardh 2 Uncertain
Rhodophyta Gymnogongrus crenulatus (Turner) J.Agardh 107 Native
Rhodophyta Gymnogongrus griffithsiae (Turner) C.Martius 115 Native
Rhodophyta Gymnophycus hapsiphorus Huisman & Kraft 1 Introduced
Rhodophyta Gymnothamnion elegans (Schousboe ex C.Agardh) J.Agardh 4 Native
Rhodophyta Halarachnion ligulatum (Woodward) Kützing 26 Native
Rhodophyta Halurus flosculosus (J.Ellis) Maggs & Hommersand 4 Native
Rhodophyta Haraldia lenormandii (Derbès & Solier) Feldmann 1 Native New record
Rhodophyta Haraldiophyllum bonnemaisonii (Kylin) A.D.Zinova 17 Native New record
Rhodophyta Herposiphonia secunda (C.Agardh) Ambronn 19 Native
Rhodophyta Heterosiphonia crispella (C.Agardh) M.J.Wynne 5 Native
Rhodophyta Hildenbrandia rubra (Sommerfelt) Meneghini 8 Native
Rhodophyta Hypnea arbuscula P.J.L.Dangeard 1 Native
Rhodophyta Hypnea cervicornis J.Agardh 2 Native New record
Rhodophyta Hypnea musciformis (Wulfen) J.V.Lamouroux 148 Uncertain
Rhodophyta Hypoglossum hypoglossoides (Stackhouse) Collins & Hervey 16 Native
Rhodophyta Itonoa marginifera (J.Agardh) Masuda & Guiry 5 Native
Rhodophyta Jania crassa J.V.Lamouroux 3 Native
Rhodophyta Jania longifurca Zanardini 11 Uncertain
Rhodophyta Jania pedunculata var. adhaerens (J.V.Lamouroux) A.S.Harvey, Woelkerling & Reviers 4 Native
Rhodophyta Jania rubens (Linnaeus) J.V.Lamouroux 14 Native
Rhodophyta Jania squamata (Linnaeus) J.H.Kim, Guiry & H.-G.Choi 2 Native
Rhodophyta Jania verrucosa J.V.Lamouroux 9 Native
Rhodophyta Jania virgata (Zanardini) Montagne 9 Uncertain
Rhodophyta Kallymenia reniformis (Turner) J.Agardh 37 Native
Rhodophyta Laurencia brongniartii J.Agardh 2 Introduced New record
Rhodophyta Laurencia dendroidea J.Agardh 1 Introduced
Rhodophyta Laurencia intricata J.V.Lamouroux 2 Native New record
Rhodophyta Laurencia microcladia Kützing 1 Native New record
Rhodophyta Laurencia obtusa (Hudson) J.V.Lamouroux 23 Native
Rhodophyta Laurencia pyramidalis Bory ex Kützing 8 Native
Rhodophyta Laurencia tenera C.K.Tseng 1 Native
Rhodophyta Laurencia viridis Gil-Rodríguez & Haroun 21 Macaronesian endemism
Rhodophyta Laurenciella marilzae (Gil-Rodríguez, Sentíes, Díaz-Larrea, Cassano & M.T.Fujii) Gil-Rodríguez, Sentíes, Díaz-Larrea, Cassano & M.T.Fujii 6 Native New record
Rhodophyta Leptosiphonia brodiei (Dillwyn) Savoie & G.W.Saunders 14 Uncertain
Rhodophyta Liagora distenta (Mertens ex Roth) J.V.Lamouroux 21 Native New record
Rhodophyta Liagora viscida (Forsskål) C.Agardh 11 Native
Rhodophyta Lomentaria articulata (Hudson) Lyngbye 128 Native
Rhodophyta Meredithia microphylla (J.Agardh) J.Agardh 55 Native New record
Rhodophyta Millerella tinerfensis (Seoane-Camba) S.M.Boo & J.M.Rico 4 Macaronesian endemism New record
Rhodophyta Monosporus pedicellatus (Smith) Solier 1 Native
Rhodophyta Myriogramme minuta Kylin 12 Native
Rhodophyta Nemalion elminthoides (Velley) Batters 41 Native
Rhodophyta Neoizziella divaricata (C.K.Tseng) S.-M.Lin, S.-Y.Yang & Huisman 14 Introduced
Rhodophyta Neopyropia leucosticta (Thuret) L.-E.Yang & J.Brodie 1 Native New record
Rhodophyta Nitophyllum punctatum (Stackhouse) Greville 19 Native
Rhodophyta Osmundea hybrida (A.P.de Candolle) K.W.Nam 12 Native
Rhodophyta Osmundea oederi (Gunnerus) G.Furnari 1 Native
Rhodophyta Osmundea pinnatifida (Hudson) Stackhouse 151 Native
Rhodophyta Osmundea truncata (Kützing) K.W.Nam & Maggs 7 Native
Rhodophyta Palisada corallopsis (Montagne) Sentíes, Fujii & Díaz-Larrea 2 Native New record
Rhodophyta Peyssonnelia squamaria (S.G.Gmelin) Decaisne ex J.Agardh 71 Native
Rhodophyta Phyllophora crispa (Hudson) P.S.Dixon 37 Native
Rhodophyta Phyllophora gelidioides P.Crouan & H.Crouan ex Karsakoff 2 Macaronesian endemism
Rhodophyta Phyllophora sicula (Kützing) Guiry & L.M.Irvine 5 Native
Rhodophyta Platoma cyclocolpum (Montagne) F.Schmitz 139 Native
Rhodophyta Platysiphonia delicata (Clemente) Cremades 3 Native New record
Rhodophyta Pleonosporium borreri (Smith) Nägeli 7 Native
Rhodophyta Plocamium cartilagineum (Linnaeus) P.S.Dixon 173 Native
Rhodophyta Pneophyllum confervicola (Kützing) Y.M.Chamberlain 1 Native New record
Rhodophyta Polysiphonia atlantica Kapraun & J.N.Norris 6 Native
Rhodophyta Polysiphonia havanensis Montagne 1 Native
Rhodophyta Polysiphonia opaca (C.Agardh) Moris & De Notaris 1 Native
Rhodophyta Polysiphonia stricta (Mertens ex Dillwyn) Greville 3 Native
Rhodophyta Porphyra umbilicalis Kützing 14 Native
Rhodophyta Porphyrostromium ciliare (Carmichael) M.J.Wynne 3 Native
Rhodophyta Predaea feldmannii Børgesen 3 Native New record
Rhodophyta Predaea feldmannii subsp. azorica Gabriel 4 Azorean endemism
Rhodophyta Pterocladiella capillacea (S.G.Gmelin) Santelices & Hommersand 377 Native
Rhodophyta Pterothamnion crispum (Ducluzeau) Nägeli 54 Native
Rhodophyta Pterothamnion plumula (J.Ellis) Nägeli 6 Native
Rhodophyta Ptilothamnion pluma (Dillwyn) Thuret 4 Uncertain
Rhodophyta Radicilingua thysanorhizans (Holmes) Papenfuss 8 Native New record
Rhodophyta Rhodophyllis divaricata (Stackhouse) Papenfuss 7 Native
Rhodophyta Rhodymenia holmesii Ardissone 92 Native
Rhodophyta Rhodymenia pseudopalmata (J.V.Lamouroux) P.C.Silva 18 Native
Rhodophyta Scagelia pylaisaei (Montagne) M.J.Wynne 4 Native
Rhodophyta Scageliopsis patens E.M.Wollaston 1 Introduced
Rhodophyta Schimmelmannia schousboei (J.Agardh) J.Agardh 29 Native New record
Rhodophyta Schizymenia apoda (J.Agardh) J.Agardh 89 Native
Rhodophyta Schottera nicaeensis (J.V.Lamouroux ex Duby) Guiry & Hollenberg 28 Uncertain
Rhodophyta Scinaia furcellata (Turner) J.Agardh 33 Native
Rhodophyta Scinaia interrupta (A.P.de Candolle) M.J.Wynne 34 Native
Rhodophyta Sebdenia dichotoma Berthold 19 Native
Rhodophyta Sebdenia rodrigueziana (Feldmann) Codomier ex Athanasiadis 41 Native
Rhodophyta Spermothamnion repens (Dillwyn) Magnus 5 Native
Rhodophyta Sphaerococcus coronopifolius Stackhouse 68 Native
Rhodophyta Spyridia filamentosa (Wulfen) Harvey 1 Native
Rhodophyta Stichothamnion cymatophilum Børgesen 1 Native
Rhodophyta Stylonema alsidii (Zanardini) K.M.Drew 2 Native
Rhodophyta Stylonema cornu-cervi Reinsch 3 Native
Rhodophyta Symphyocladia marchantioides (Harvey) Falkenberg 85 Introduced
Rhodophyta Taenioma nanum (Kützing) Papenfuss 1 Native New record
Rhodophyta Tenarea tortuosa (Esper) Me.Lemoine 3 Native
Rhodophyta Vertebrata foetidissima (Cocks ex Bornet) Díaz-Tapia & Maggs 1 Native
Rhodophyta Vertebrata fruticulosa (Wulfen) Kuntze 5 Native
Rhodophyta Vertebrata fucoides (Hudson) Kuntze 7 Uncertain
Rhodophyta Vertebrata furcellata (C.Agardh) Kuntze 6 Native
Rhodophyta Vertebrata hypnoides (Welwitsch) Kuntze 4 Uncertain
Rhodophyta Vertebrata nigra (Hudson) Díaz-Tapia & Maggs 1 Native New record
Rhodophyta Vertebrata reptabunda (Suhr) Díaz-Tapia & Maggs 4 Uncertain
Rhodophyta Vertebrata tripinnata (Harvey) Kuntze 3 Native
Rhodophyta Wrangelia penicillata (C.Agardh) C.Agardh 6 Native New record
Rhodophyta Wurdemannia miniata (Sprengel) Feldmann & Hamel 1 Native
Rhodophyta Xiphosiphonia ardreana (Maggs & Hommersand) Savoie & G.W.Saunders 4 Native
Rhodophyta Xiphosiphonia pennata (C.Agardh) Savoie & G.W.Saunders 2 Introduced
Rhodophyta Xiphosiphonia pinnulata (Kützing) Savoie & G.W.Saunders 2 Introduced
Chlorophyta Anadyomene stellata (Wulfen) C.Agardh 4 Uncertain
Chlorophyta Blidingia marginata (J.Agardh) P.J.L.Dangeard ex Bliding 1 Native New record
Chlorophyta Blidingia minima (Nägeli ex Kützing) Kylin 2 Native
Chlorophyta Bryopsis cupressina J.V.Lamouroux 27 Native
Chlorophyta Bryopsis duplex De Notaris 1 Native
Chlorophyta Bryopsis hypnoides J.V.Lamouroux 28 Native
Chlorophyta Bryopsis pennata J.V.Lamouroux 6 Native New record
Chlorophyta Bryopsis plumosa (Hudson) C.Agardh 71 Native
Chlorophyta Caulerpa prolifera (Forsskål) J.V.Lamouroux 10 Introduced
Chlorophyta Chaetomorpha aerea (Dillwyn) Kützing 10 Native
Chlorophyta Chaetomorpha linum (O.F.Müller) Kützing 26 Native
Chlorophyta Chaetomorpha pachynema (Montagne) Kützing 2 Native
Chlorophyta Cladophora albida (Nees) Kutzing 5 Native
Chlorophyta Cladophora coelothrix Kützing 15 Native
Chlorophyta Cladophora conferta P.Crouan & H.Crouan 5 Native
Chlorophyta Cladophora dalmatica Kützing 1 Uncertain
Chlorophyta Cladophora hutchinsiae (Dillwyn) Kützing 1 Native
Chlorophyta Cladophora laetevirens (Dillwyn) Kützing 7 Uncertain
Chlorophyta Cladophora lehmanniana (Lindenberg) Kützing 5 Native
Chlorophyta Cladophora liebetruthii Grunow 1 Native New record
Chlorophyta Cladophora prolifera (Roth) Kützing 82 Native
Chlorophyta Cladophoropsis membranacea (Hofman Bang ex C.Agardh) Børgesen 1 Uncertain
Chlorophyta Codium adhaerens C. Agradh 115 Native
Chlorophyta Codium decorticatum (Woodward) M.Howe 10 Native
Chlorophyta Codium elisabethiae O.C.Schmidt 84 Macaronesian endemism
Chlorophyta Codium fragile (Suringar) Hariot 9 Native
Chlorophyta Codium fragile subsp. atlanticum (A.D.Cotton) P.C.Silva 3 Native New record
Chlorophyta Codium fragile subsp. fragile (Suringar) Hariot 15 Introduced
Chlorophyta Codium tomentosum Stackhouse 2 Native New record
Chlorophyta Codium vermilara (Olivi) Delle Chiaje 2 Native New record
Chlorophyta Gayralia oxysperma (Kützing) K.L.Vinogradova ex Scagel & al. 5 Native
Chlorophyta Lychaete pellucida (Hudson) M.J.Wynne 3 Native
Chlorophyta Microdictyon boergesenii Setchell 3 Native New record
Chlorophyta Microdictyon umbilicatum (Velley) Zanardini 3 Native New record
Chlorophyta Pseudochlorodesmis furcellata (Zanardini) Børgesen 1 Native New record
Chlorophyta Pseudorhizoclonium africanum (Kützing) Boedeker 5 Native
Chlorophyta Ulothrix flacca (Dillwyn) Thuret 2 Native
Chlorophyta Ulva clathrata (Roth) C.Agardh 12 Native
Chlorophyta Ulva compressa Linnaeus 12 Native
Chlorophyta Ulva intestinalis Linnaeus 37 Native
Chlorophyta Ulva lactuca Linnaeus 2 Uncertain
Chlorophyta Ulva linza Linnaeus 6 Native
Chlorophyta Ulva polyclada Kraft 1 Native
Chlorophyta Ulva prolifera O.F.Müller 6 Native
Chlorophyta Ulva ralfsii (Harvey) Le Jolis 1 Native
Chlorophyta Ulva rigida C.Agardh 198 Native
Chlorophyta Valonia macrophysa Kützing 3 Native
Chlorophyta Valonia utricularis (Roth) C. Agardh 7 Native
Ochrophyta Ascophyllum nodosum (Linnaeus) Le Jolis 8 Native
Ochrophyta Bachelotia antillarum (Grunow) Gerloff 2 Native
Ochrophyta Canistrocarpus cervicornis (Kützing) De Paula & De Clerck 2 Native New record
Ochrophyta Carpomitra costata (Stackhouse) Batters 16 Native
Ochrophyta Cladostephus spongiosus (Hudson) C.Agardh 33 Native
Ochrophyta Colpomenia sinuosa (Mertens ex Roth) Derbès & Solier 472 Native
Ochrophyta Compsonema saxicola (Kuckuck) Kuckuck 28 Native
Ochrophyta Cutleria multifida (Turner) Greville 27 Uncertain
Ochrophyta Cutleria multifida (Turner) Greville phase Aglaozonia parvula (Greville) Zanardini 4 Uncertain
Ochrophyta Cystoseira compressa (Esper) Gerloff & Nizamuddin 81 Native
Ochrophyta Cystoseira foeniculacea (Linnaeus) Greville 46 Native
Ochrophyta Cystoseira humilis Schousboe ex Kützing 21 Native
Ochrophyta Dictyopteris polypodioides (A.P.De Candolle) J.V.Lamouroux 21 Native
Ochrophyta Dictyota bartayresiana J.V.Lamouroux 9 Native
Ochrophyta Dictyota ciliolata Sonder ex Kützing 7 Native New record
Ochrophyta Dictyota cyanoloma Tronholm, De Clerck, A.Gómez-Garreta & Rull Lluch 7 Native
Ochrophyta Dictyota dichotoma (Hudson) J.V.Lamouroux 189 Native
Ochrophyta Dictyota dichotoma var. intricata (C.Agardh) Greville 10 Native New record
Ochrophyta Dictyota fasciola (Roth) J.V.Lamouroux 5 Native New record
Ochrophyta Dictyota implexa (Desfontaines) J.V.Lamouroux 9 Native New record
Ochrophyta Ectocarpus fasciculatus Harvey 7 Native
Ochrophyta Ectocarpus siliculosus (Dillwyn) Lyngbye 6 Uncertain
Ochrophyta Elachista flaccida (Dillwyn) Fries 1 Native New record
Ochrophyta Feldmannia irregularis (Kützing) Hamel 1 Native
Ochrophyta Feldmannia mitchelliae (Harvey) H.-S.Kim 10 Native
Ochrophyta Feldmannia paradoxa (Montagne) Hamel 3 Native
Ochrophyta Fucus spiralis Linnaeus 89 Uncertain
Ochrophyta Gongolaria abies-marina (S.G.Gmelin) Kuntze 265 Native
Ochrophyta Halopteris filicina (Grateloup) Kützing 217 Native
Ochrophyta Halopteris scoparia (Linnaeus) Sauvageau 207 Native
Ochrophyta Hapalospongidion macrocarpum (Feldmann) León-Álvarez & González-González 16 Native New record
Ochrophyta Hecatonema terminale (Kützing) Kylin 20 Native
Ochrophyta Hincksia ovata (Kjellman) P.C.Silva 4 Native
Ochrophyta Hydroclathrus clathratus (C.Agardh) M.Howe 111 Native
Ochrophyta Leathesia marina (Lyngbye) Decaisne 6 Uncertain
Ochrophyta Lobophora variegata (J.V.Lamouroux) Womersley ex E.C.Oliveira 88 Native
Ochrophyta Mesogloia vermiculata (Smith) S.F.Gray 9 Native New record
Ochrophyta Myriactula rivulariae (Suhr ex Areschoug) Feldmann 20 Native
Ochrophyta Myrionema strangulans Greville 13 Native New record
Ochrophyta Nemoderma tingitanum Schousboe ex Bornet 50 Native
Ochrophyta Padina pavonica (Linnaeus) Thivy 231 Native
Ochrophyta Papenfussiella kuromo (Yendo) Inagaki 11 Introduced
Ochrophyta Petalonia binghamiae (J.Agardh) K.L.Vinogradova 177 Introduced
Ochrophyta Petalonia fascia (O.F.Müller) Kuntze 3 Native
Ochrophyta Petrospongium berkeleyi (Greville) Nägeli ex Kützing 12 Native
Ochrophyta Pseudolithoderma adriaticum (Hauck) Verlaque 1 Native
Ochrophyta Pseudolithoderma roscoffense Loiseaux 48 Native
Ochrophyta Punctaria tenuissima (C.Agardh) Greville 4 Native
Ochrophyta Ralfsia verrucosa (Areschoug) Areschoug 92 Native
Ochrophyta Sargassum cymosum C.Agardh 20 Native
Ochrophyta Sargassum furcatum Kützing 37 Native
Ochrophyta Sargassum vulgare C.Agardh 15 Native
Ochrophyta Scytosiphon lomentaria (Lyngbye) Link 127 Native
Ochrophyta Scytosiphon lomentaria (Lyngbye) Link, phase Microspongium gelatinosum Reinke 43 Native
Ochrophyta Sphacelaria cirrosa (Roth) C.Agardh 7 Native
Ochrophyta Sphacelaria plumula Zanardini 2 Native
Ochrophyta Sphacelaria rigidula Kützing 1 Native
Ochrophyta Sphacelaria tribuloides Meneghini 5 Uncertain
Ochrophyta Sphaerotrichia divaricata (C.Agardh) Kylin 41 Uncertain
Ochrophyta Sporochnus pedunculatus (Hudson) C.Agardh 2 Native
Ochrophyta Stragularia clavata (Harvey) Hamel 1 Native New record
Ochrophyta Stypopodium zonale (J.V.Lamouroux) Papenfuss 8 Native
Ochrophyta Taonia atomaria (Woodward) J.Agardh 41 Native
Ochrophyta Zanardinia typus (Nardo) P.C.Silva 7 Native
Ochrophyta Zonaria tournefortii (J.V.Lamouroux) Montagne 248 Native

Table 3.

Summary of the macroalgal flora of the Island of São Miguel (N spec- number of specimens; N taxa- taxa; N spp- number of species) with information on the species origin and status (Introd- introduced; Uncrt- uncertain origin; Azo end- Azores endemism; Mac end- Macaronesia endemism; New rec- new record).

Phyllum Order Family N spec N taxa N spp Native Introd Uncrt Azo end Mac end New rec
Rhodophyta 20 50 7510 284 212 171 15 21 1 4 42
Chlorophyta 5 14 1103 59 48 40 2 5 1 9
Ochrophyta 11 19 4168 88 63 55 2 6 10
Total 36 83 12781 431 323 266 19 32 1 5 61

Many species were only sporadically recorded, but 10 were widely recorded around the Island and occurred quite abundantly in some locations, namely: the Rhodophyta Acrosorium ciliolatum (Harvey) Kylin, Chondracanthus acicularis, Gelidium microdon and Pterocladiella capillacea; and the Ochrophyta Colpomenia sinuosa, Gongolaria abies-marina, Halopteris filicina (Grateloup) Kützing, Halopteris scoparia, Padina pavonica and Zonaria tournefortii.

This paper increases the total of macroalgae species previously listed for São Miguel Island by 63 (44 Rhodophyta, 9 Chlorophyta and 10 Ochrophyta). When compared with the other Azorean Islands (Table 4), São Miguel has the highest number of species in all phylla, which reflects the greater amount of research undertaken on this Island on a more regular basis, involving both temporal and long-term studies.

Table 4.

Number of macroalgae species on the Azorean Islands: Santa Maria (Neto et al. 2020d); São Miguel (the present paper); Terceira (Neto et al. 2020a); Graciosa (Neto et al. 2020c); São Jorge and Faial (authors' unpublished data); Pico (Neto et al. 2020b); Flores and Corvo (Neto et al. 2021a).

Phyllum Santa Maria São Miguel Terceira Graciosa São Jorge Pico Faial Flores Corvo
Rhodophyta 103 212 73 126 35 142 59 80 22
Chlorophyta 29 48 24 31 17 41 16 22 8
Ochrophyta 44 63 16 38 10 42 8 26 13
Total 176 323 113 195 62 225 83 128 43

In general and in keeping with other warm-water areas of the North Atlantic Ocean, the Azorean macroalgae flora has a larger proportion of red seaweeds.

A relatively high number of non-native species has been recorded on São Miguel Island (see Tables 2, 3), similarly to what has been reported for Santa Maria (Neto et al. 2020d), Terceira (Neto et al. 2020a), Graciosa (Neto et al. 2020c), Pico (Neto et al. 2020b) and Flores and Corvo (Neto et al. 2021a). Research over the past 15 years (Cardigos et al. 2006, Micael et al. 2014, Vaz-Pinto et al. 2014, Chainho et al. 2015, Cacabelos et al. 2019, Cacabelos et al. 2020, Martins et al. 2019, Parente et al. 2019) has indicated that the arrival of non-native species is increasing in the Azores, which is very likely to be related to the geostrategic position of the Archipelago as a “crossroad” in the distribution of marine algae in the North Atlantic, with documented exchanges with European, Mediterranean and American coasts. This may favour the arrival of new species via maritime traffic, both commercial and recreational (hull fouling, ballast waters), which may be a high risk to local marine ecosystems, as non-indigenous species can become invasive, resulting in impacts on ecosystem services and biodiversity (e.g. Katsanevakis et al. 2014).

The discovery of the new macroalgae records on São Miguel Island (present study) and on the other Azorean Islands (Neto et al. 2020a, Neto et al. 2020b, Neto et al. 2020c, Neto et al. 2020d, Neto et al. 2021a), demonstrates the need for continuing taxonomic and floristic studies in this region of the Atlantic Ocean. The biogeographically-variable nature of the new records found confirms the overall mixed nature of the marine algal flora of the Azores with elements shared with Macaronesia, the Mediterranean Sea, Atlantic Europe and the subtropical and tropical Atlantic America.

Taxonomic mismatch

A mismatch regarding the GBIF backbone taxonomy of some of the macroalgae species names was identified as detailed in Suppl. material 1.

Supplementary Material

Supplementary material 1

DP-SMG-id_15785_normalized.csv

Ana I Neto

Data type

Macroalgae taxonomic mismatching

Brief description

GBIF does not have the more actualised nomenclature for some of the macroalgae species names. Therefore, the matching tools of its platform were applied to the species list, as required by Pensoft's data auditor, to identify the problematic taxonomic situations. The resulting file (DP-SMG-id_15785_normalized.csv) is included here, since the names will not be immediately updated in the GBIF Taxonomic Backbone. A request was already sent to GBIF helpdesk to solve this situation.

File: oo_505943.csv

bdj-09-e64969-s001.csv (66.2KB, csv)

Acknowledgements

This research was supported by several projects, expeditions and campaigns (see Funding above) and lately by the project “ACORES-01-0145-FEDER-000072” funded the Operational Programme Azores 2020 (85% ERDF and 15% regional funds). Thanks are due to the campaign teams for their critical involvement in this project (Abel Sentíes, Aina del Alcázar, Ana Alfaya, Ana Belén Villalba Lapeña, Ana Santos, Ana Sofia Carreiro, André Amaral, Andrea Tracana, Ane Laborda, Anna Lloveras Armengol, António Brigos Plafon, Berta Solé Nadal, Camille Fontaine, Carlos Rius, Carles Mir, Caroline Terral, Catarina Santos, Cláudia Hipólito, Daniela Gabriel, Edward Hehre, Emanuel Xavier, Eduardo García, Enrique Almira, Esteban Belles, Eunice Nogueira, Fátima Vaz Pinto, Francisco Wallenstein, Gustavo M Martins, Heather Baldwin, Isadora Moniz, Jana Verdura, Joana Pombo, João Brum, João Faria Santos, João Ferreira, Laura Busquier, Marco Enoch, Maria Ana Dionísio, Maria Machín-Sánchez, Maria Vale, Marlene Terra, Mónica Martínez, Mutue Toyota Fujii, Patrícia Madeira, Pedro Raposeiro, Richard Fralick, Richard Thompson, Rocío Sánchez, Ruben Couto, Rubén Mosquera, Rui Sousa, Sara Peres, Tarso Costa, Tito Silva, Valeria Cassano, Virginie Leyendecker). Edgar Rosas Alquicira and Karla León Cisneros were supported by the Programme AlBan, the European Union Programme of High Level Scholarships for Latin America (through scholarships E05D060221MX and E05D060520MX), “Consejo Nacional de Ciencia y Tecnología” (doctoral scholarships 176162 and 157904) and the UNAMUNO Programme of PhD Scholarships for Europe. Eva Cacabelos was supported by a postdoctoral grant (Project M1420-09-5369-FSE-000002) from ARDITI (Regional Agency for Development of Research, Technology and Innovation of Madeira). Andrea Z. Botelho was supported by a PhD grant (M3.1.a/F/083/2015), awarded by Fundo Regional da Ciência e Tecnologia (FRCT). Afonso C.L. Prestes was supported by a PhD grant (M3.1.a/F/083/2015), awarded by Fundo Regional da Ciência e Tecnologia (FRCT). Rita F. Patarra was supported by a Science and Technology Management Fellowship grant (SFRH/BGCT/135478/2018), awarded by Fundação para a Ciência e a Tecnologia (FCT I.P.). Manuela I. Parente was supported by a Postdoc grant (SFRH/BPD/34246/2006), awarded by Fundação para a Ciência e a Tecnologia (FCT).

Funding Statement

This work is financed by the ERDF (85%) and by regional funds (15%), through the Operational Programme Azores 2020, within the scope of the project “ACORES-01-0145-FEDER-000072”.

Author contributions

AIN: Conceptualisation; Methodology; Research (field and laboratory work); Resources; Data Curation; Formal analysis and interpretation; Paper writing

IM: Research (field and laboratory work); Data Curation; Formal analysis and interpretation; Paper writing

EFRA: Research (field and laboratory work); Data Curation; Paper writing

KLC: Research (field and laboratory work); Data Curation; Paper writing

EC: Research (field and laboratory work); Data Curation; Paper writing

AZB: Research (field and laboratory work); Data Curation

JM: Research (field and laboratory work); Data Curation

ACC: Research (field and laboratory work); Resources; Data Curation

RMAN: Data Curation; Formal analysis and interpretation

JMNA: Research (field work and laboratory work); Formal analysis and interpretation

SM: Research (field and laboratory work); Data Curation

RR: Resources; Data Curation

PA: Resources

ACLP: Research (field and laboratory work); Data Curation

RFP: Research (field and laboratory work); Data Curation; Paper writing

NVA: Research (field work); Maps elaboration

DM-F: Research (field and laboratory work); Data Curation

EB: Research (fieldwork and laboratory work); Resources; Data Curation; Paper writing

RF: Methodology; Research (fieldwork and laboratory work); Data Curation; Paper writing

WF: Methodology; Research (fieldwork and laboratory work); Data Curation; Paper writing

IT: Methodology; Research (fieldwork and laboratory work); Resources; Data Curation; Paper writing

MIP: Research (field and laboratory work); Resources; Data Curation; Formal analysis and interpretation; Paper writing

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Associated Data

This section collects any data citations, data availability statements, or supplementary materials included in this article.

Supplementary Materials

Supplementary material 1

DP-SMG-id_15785_normalized.csv

Ana I Neto

Data type

Macroalgae taxonomic mismatching

Brief description

GBIF does not have the more actualised nomenclature for some of the macroalgae species names. Therefore, the matching tools of its platform were applied to the species list, as required by Pensoft's data auditor, to identify the problematic taxonomic situations. The resulting file (DP-SMG-id_15785_normalized.csv) is included here, since the names will not be immediately updated in the GBIF Taxonomic Backbone. A request was already sent to GBIF helpdesk to solve this situation.

File: oo_505943.csv

bdj-09-e64969-s001.csv (66.2KB, csv)

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