Abstract
Background
Knowledge about cyanobacteria diversity in the Azores is spread over several publications, dating from 1874, with some of them not generally available to the scientific community due to their restricted access. The dispersion and sometimes inaccessibility of this information hinder a deeper analysis and a better understanding of the biodiversity of the Azores Islands and more general ecological processes in oceanic islands. Here we present the first checklist of cyanobacteria for the Azores Archipelago with updated taxonomy of all recorded taxa.
New information
This work provides a compiled and annotated checklist of all known cyanobacteria from the Azores Archipelago with morphological identification from preserved samples and cultures, based on published literature. All records of taxa known to occur in the Azores were taxonomically updated. The present checklist comprises 225 taxa distributed by six orders (Chroococcales, Nostocales, Oscillatoriales, Pleurocapsales, Spirulinales and Synechococcales). Our literature review reveals that the Azores Archipelago hosts a high diversity of cyanobacteria, despite several overlooked habitats that may present great potential regarding cyanobacteria diversity. Increasing efforts to study these neglected habitats could contribute to the knowledge of cyanobacteria taxonomy. This checklist provides the basis for future works on the taxonomy and taxa richness of cyanobacteria in the Azores and the Atlantic Islands, as also for understanding and monitoring non-indigenous and invasive species.
Keywords: Oceanic islands, biodiversity, Cyanophyceae, Macaronesia, Atlantic Ocean
Introduction
Cyanobacteria are gram-negative photosynthetic prokaryotes that developed around 3500 million years ago (Schirrmeister et al. 2015). As one of the most primitive organisms on earth (Mareš et al. 2013), they successfully occupy various habitats in terrestrial and aquatic ecosystems, both marine and freshwater (Whitton and Potts 2012). Cyanobacteria diversity amongst these systems is unbalanced, being larger in freshwater and terrestrial ecosystems (Komárek and Johansen 2015). They persist in almost all types of illuminated habitats, with optimum growing temperatures generally higher than microalgae, which enable them to support a wide array of stress conditions, including extreme habitats (Komárek and Johansen 2015).
Freshwater cyanobacteria are commonly present in wetlands, lakes, rivers and streams, both in benthic (Scott and Marcarelli 2012) and planktonic (e.g. Stockner et al. 2000, Oliver et al. 2012) habitats. Benthic cyanobacteria are commonly found solitary or forming mats in the various stream and river substrates, such as rocks, sand, plants and many others (Casamatta and Hašler 2016). In shallow lakes and littoral zones of deep lakes, benthic species of cyanobacteria can also occur if enough light reaches the substrates (Scott and Marcarelli 2012). However, cyanobacteria are mostly known from the plankton of lentic waters, where they can grow in high abundance, usually known as blooms, especially in eutrophic lakes (Paerl et al. 2010, Carmichael and Boyer 2016). Cyanobacteria blooms negatively affect the ecosystems and services they provide (Carmichael and Boyer 2016) as most bloom-forming species produce toxins that can be accumulated at the water surface, causing unpleasant surface scums.
Extreme habitats, such as thermal springs, are successfully occupied by Cyanobacteria, where they are often the main and/or sole autotrophic organisms inhabiting these environments (Komárek and Johansen 2015). In marine systems, cyanobacteria are found in a wide array of habitats, including benthos, plankton, associated with other organisms, amongst others (Golubic et al. 2010, Konstantinou et al. 2018).
In the last ten years, cyanobacteria taxonomy has changed dramatically with the use of new techniques, mainly through 16S rRNA sequencing, contributing to a taxonomic re-assessment of the group (e.g. Wacklin et al. 2009, Komárek et al. 2011, Zapomělová et al. 2011, Strunecký et al. 2013, Komárek et al. 2014, Strunecky et al. 2017, Mai et al. 2018).
In the Azores, a remote oceanic archipelago located in the middle of the North Atlantic Ocean, the first work to be published on cyanobacteria taxonomy came from the Challenger expedition that occurred from 1872 to 1876 and had a brief passage in São Miguel Island from 3 July to 9 July 1873 (Brock and Brock 1967). Some members of the Challenger expedition visited the Furnas Village and later Moseley (1874) and Archer (1874), who received samples from Moseley, published the first records. Later, Trelease (1897) and Bohlin (1901) contributed with several cyanobacteria records from several islands. In the 20th century, more biologists visited the Islands contributing considerably to the knowledge of the microalgae and cyanobacteria of the Azores. First by Krieger (1931), with a small contribution and after with the valuable works of Cedercreutz (1941), Bourrelly and Manguin (1946) and Johansson (1977), contributing with several detailed descriptions of the cyanobacteria flora in several islands of the Azores. The later works contributed with the highest number of known species for the Azores. Bourrelly and Manguin (1946) also describes a new form Oscillatoriageitlerif.major Bourrelly in Bourrelly and Manguin (1946), which is the first and only known endemic cyanobacteria to the Azores. After 1980, works were mainly focused on planktonic freshwater species due to the rise of lake eutrophication signs. Important contributions to the known cyanobacteria flora have been provided after the implementation of the Water Framework Directive (WFD) in the Azores, with regular monitoring programmes since 1994 (Santos et al. 2005, Santos et al. 2012, Luz et al. 2020b). The more recent works on cyanobacteria were based on cultured strains isolated from freshwater lakes (Cordeiro 2015), which provide the addition of new species. Several works performed on thermal, terrestrial, brackish and marine habitats, which were overlooked in previous studies, contributed to several new cyanobacteria species records (Luz 2018, Cordeiro et al. 2020b).
Despite the increased research efforts, especially in the last decade, the knowledge of the diversity and distribution of Cyanobacteria in the Azores Archipelago is not consistently organised and a local checklist has never been published. This study aims to present an updated checklist of cyanobacteria present in the Azores, based on a taxonomically updated list of previously reported species from preserved samples and based on cultured strains.
Materials and methods
Study Area
The Azores are an oceanic group of islands located in the middle of the North Atlantic Ocean, roughly 1500 km from Europe and 1900 km from America (Fig. 1). The Archipelago is made up of nine islands roughly aligned along 615 km in a WNW-ESE trend, that are divided into three groups according to their geographical position. Although they are in geographical proximity, the Islands present unique features differentiating themselves from each other (Table 1), with different amounts of annual rainfall (Secretaria Regional do Ambiente e do Mar 2011) and distinct geological settings (e.g. Moore 1990, Azevedo and Portugal Ferreira 2006, Cole et al. 2008).
Figure 1.
Azores Archipelago location with an indication of the most represented aquatic habitats on each island.
Table 1.
Island characterisation. [1] Ávila et al. 2016, [2] Secretaria Regional do Ambiente e do Mar 2011, [3] Porteiro 2000, [4] Morton 2014, [5] Cruz and França 2006, [6] Secretaria Regional da Agricultura e Ambiente – Direção Regional do Ambiente 2015, [7] Ramsar 2013, [8] Morton et al. 1997.
| Group | Island | Age (Ma)[1] | Area (Km2)[2] | Freshwater lakes (N)[3] | Permanent Streams (N)[6] | Coastal lakes (N)[4, 7, 8] | Thermal waters (N)[5] | Total lake area (km2)[3, 4, 7, 8] | Annual precipitation (mm)[2] | 
| Eastern | São Miguel | 4.00 | 744,6 | 33 | 6 | - | 14 | 8.34 | 1027.1 | 
| Santa Maria | 6.30 | 96,9 | - | 1 | - | - | - | 775.2 | |
| Central | Terceira | 0.40 | 400,3 | 18 | - | 3 | - | 0.36 | 1125.6 | 
| Pico | 0.27 | 444,8 | 28 | - | - | - | 0.16 | 956.3 | |
| Faial | 0.85 | 173,1 | - | - | - | 2 | - | 974.0 | |
| São Jorge | 1.32 | 243,7 | - | - | 2 | - | 0.86 | 1194.3 | |
| Graciosa | 0.70 | 60,7 | - | - | - | 1 | - | 918.4 | |
| Western | Flores | 2.16 | 141,0 | 8 | 2 | - | 2 | 0.72 | 1716.1 | 
| Corvo | 1.50 | 17,1 | 1 | - | - | - | 0.24 | 1144.6 | 
The western group includes the Islands of Flores and Corvo, which are amongst the smallest islands of the Archipelago. Corvo and Flores are very rich in aquatic habitats despite their small size due to their higher annual precipitation (Secretaria Regional do Ambiente e do Mar 2011). The central group (Graciosa, Faial, Pico, São Jorge and Terceira Islands) comprise the youngest in the Archipelago (Ávila et al. 2016). The islands in the central group include a high diversity of inland aquatic habitats, including freshwater and saline lakes, streams and thermal waters (Morton et al. 1997, Porteiro 2000, Cruz and França 2006, Morton 2014). The eastern group, Santa Maria and São Miguel, includes the Archipelago’s oldest (Santa Maria) and the largest (São Miguel) islands. São Miguel is the island with most of the lakes and the larger area of water bodies (Porteiro 2000), whereas Santa Maria is the driest island of the Archipelago, with only 775.2 mm mean annual precipitation and no significant inland water habitats (Secretaria Regional do Ambiente e do Mar 2011).
The Azores are particularly rich in freshwater systems, with 88 lakes (Porteiro 2000), nine permanent streams, five saline lakes and several thermal springs (Table 1). Lakes are located between 230 and 1,050 m altitude and, according to Gonçalves 2008, could be classified into two main lake types: shallow lakes, with a maximum depth below five metres; and (ii) deep lakes, with maximum depths, greater than five metres. The insular lotic systems are small, narrow, with steep watersheds and are fed by lakes or springs, most of them having torrential or seasonal flowing regimes (Raposeiro et al. 2013).
Checklist Production
The checklist was based on all known literature mentioning cyanobacteria from the Azores with morphological identification, published until 2020. The nomenclature was revised according to Guiry and Guiry (2022). The complete taxonomic list (taxon data table and occurrence data table) is published in DwC (Suppl. material 1) in GBIF, the Global Biodiversity Information Facility (Luz et al. 2022). Taxa identified only to the family level or above were not included in the discussed taxonomic list.
Data resources
Cyanobacteria occurrence in the Azores
Data set name: Cyanobacteria Checklist of the Azores Archipelago, Portugal - Occurrence data table
Data format: Darwin Core
Description: Cyanobacteria occurrence records in the Azores Archipelago, dating from 1874 to 2020, with 2838 records (Luz et al. 2022). Used Darwin Core terms are described in Table 2.
Table 2.
Darwin Core terms used in the occurrence data table.
| Column label | Column description | 
|---|---|
| id | Identifier. | 
| type | The nature of the resource. | 
| collectionCode | Acronym identifying the collection from which the record was derived. | 
| basisOfRecord | Specific nature of the data record. | 
| occurrenceID | Occurrence identifier. | 
| catalogNumber | Identifier for the record within the collection. | 
| associatedReferences | Literature associated with the occurrence. | 
| eventDate | Date-time or interval during which the event was recorded. | 
| continent | Name of the continent in which the occurrence location occurs. | 
| waterBody | Name of the water body in which the occurrence location occurs. | 
| islandGroup | Name of the island group in which the occurrence location occurs. | 
| island | Name of the island on which the occurrence location occurs. | 
| country | Name of the country in which the occurrence location occurs. | 
| countryCode | Standard code for the country in which the occurrence location occurs. | 
| municipality | Name of the municipality in which the occurrence location occurs. | 
| locality | Name of the locality in which the occurrence location occurs. | 
| decimalLatitude | Geographic latitude in which the occurrence location occurs. | 
| decimalLongitude | Geographic longitude in which the occurrence location occurs. | 
| geodeticDatum | Geodetic datum upon which the geographic coordinates given are based. | 
| taxonID | Taxon identifier. | 
| scientificName | The full scientific name including author. | 
| acceptedNameUsage | The full scientific name including author currently accepted. | 
| kingdom | Kingdom name in which the taxon is classified. | 
| phylum | Phylum name in which the taxon is classified. | 
| class | Class name in which the taxon is classified. | 
| taxonRank | Lowest taxonomic rank of the taxon. | 
Cyanobacteria checklist from the occurrence
Data set name: Cyanobacteria Checklist of the Azores Archipelago, Portugal - Taxon data table
Data format: Darwin Core
Description: Cyanobacteria taxa recorded in the Azores Archipelago, based on the occurrrence data table, with a total of 229 taxa (Luz et al. 2022). Used Darwin Core terms are described in Table 3.
Table 3.
Darwin Core terms used in the taxon data table.
| Column label | Column description | 
|---|---|
| id | Identifier. | 
| taxonID | Taxon identifier. | 
| scientificName | The full scientific name including author. | 
| kingdom | Kingdom name in which the taxon is classified. | 
| phylum | Phylum name in which the taxon is classified. | 
| class | Class name in which the taxon is classified. | 
| order | Order name in which the taxon is classified | 
| family | Family name in which the taxon is classified. | 
| genus | Genus name in which the taxon is classified. | 
| specificEpithet | Species epithet name in which the taxon is classified. | 
| infraspecificEpithet | Infraspecific epithet name in which the taxon is classified. | 
| taxonRank | Lowest taxonomic rank of the taxon. | 
| scientificNameAuthorship | Authorship information for the scientific name. | 
Checklists
Cyanobacteria checklist from Azores islands
Anabaena
Bory ex Bornet & Flahault, 1886
AD225C70-E126-51B1-92A0-EB32FCA1E43B
Distribution
Corvo (INOVA 1996), Flores (Bourrelly and Manguin 1946), Pico (Luz et al. 2020), São Miguel (Bohlin 1901), Terceira (Luz et al. 2020)
Notes
Freshwater (lake), thermal (pool)
Anabaena aspera
Frémy, 1930
0D2AD3D2-D028-58BE-A590-48FA64D17004
Distribution
Flores (Bourrelly and Manguin 1946)
Notes
Freshwater (lake)
Anabaena augstumalis
Schmidle, 1900
A08E012F-C9DA-5AF1-BB70-C235C1D41E16
Distribution
Flores (Bourrelly and Manguin 1946)
Notes
Freshwater (lake)
Anabaena cylindrica
Lemmermann, 1896
4111C797-D7D8-5993-AFA8-0D5E4BD049C8
Distribution
São Miguel (Oliveira 1989)
Notes
Freshwater (lake)
Anabaena inaequalis
Bornet & Flahault, 1886
1FAF86B2-90A8-535C-A933-4D1A13F04C1B
Distribution
Flores (Luz et al. 2020), Pico (Luz et al. 2020), São Miguel (Luz et al. 2020)
Notes
Freshwater (lake)
Anabaena torulosa
Lagerheim ex Bornet & Flahault, 1886
C1B36671-29CE-56F0-8A6A-081ECE6D1BBF
Distribution
Corvo (Trelease 1897)
Notes
Freshwater
Anabaenopsis
V.V.Miller, 1923
B645D965-F87A-5BF0-A39E-AEEF5BA728D8
Distribution
Corvo (Cordeiro et al. 2020b), Flores (Luz et al. 2020)
Notes
Freshwater (lake)
Anabaenopsis circularis
(G.S.West) Woloszynska & V.Miller in V.Miller, 1923
6B3731E2-A5FF-5165-9ECF-1BEBC72481B2
Distribution
Flores (Gonçalves 2008)
Notes
Freshwater (lake)
Anagnostidinema amphibium
(C.Agardh ex Gomont) Strunecký, Bohunická, J.R.Johansen & J.Komárek, 2017
3346370C-7D38-5534-A00C-B9D4226E09EB
Distribution
Flores (Bourrelly and Manguin 1946), São Miguel (Cedercreutz 1941), Terceira (Cedercreutz 1941)
Notes
Brackish (lake), freshwater (lake)
Anathece clathrata
(W.West & G.S.West) Komárek, Kastovsky & Jezberová, 2011
D865FE01-3303-5AB1-8080-DB04348ACF89
Distribution
São Miguel (Santos and Santana 2004)
Notes
Freshwater (lake)
Anathece minutissima
(West) Komárek, Kastovsky & Jezberová, 2011
CB0697C9-5FF4-5FD6-B3AB-DE2875CED4A4
Distribution
São Miguel (Luz et al. 2020)
Notes
Freshwater (lake)
Aphanizomenon
A.Morren ex É.Bornet & C.Flahault, 1886
F2B889F6-8C7A-5CC4-9D65-27039A0847C5
Distribution
São Miguel (Santos et al. 2001)
Notes
Freshwater (lake)
Aphanizomenon flos-aquae
Ralfs ex Bornet & Flahault, 1886
746F96B3-B418-564F-A851-BC6EC5320709
Distribution
Corvo (INOVA 1996), Flores (INOVA 1996), Graciosa (Azevedo et al. 2005), Pico (INOVA 1996), São Miguel (Krieger 1931)
Notes
Freshwater (lake)
Aphanizomenon gracile
Lemmermann, 1907
B4275013-C555-5491-A49B-7FA2E31BA156
Distribution
Flores (Luz et al. 2020), Pico (Luz et al. 2020), São Miguel (Luz et al. 2020)
Notes
Freshwater (lake)
Aphanizomenon manguinii
Bourrelly in Bourrelly & Manguin, 1952
8EBDF4DD-1806-5534-8691-5E77650B43E5
Distribution
Pico (Luz 2018)
Notes
Freshwater (lake)
Aphanocapsa
Nägeli, 1849
1E05D72B-365C-5C3C-BFE2-9569B08DBC37
Distribution
São Miguel (Santos and Santana 2004)
Notes
Freshwater (lake)
Aphanocapsa delicatissima
West & G.S.West, 1912
87DBC12D-68F6-5815-AA71-6FD82440167F
Distribution
Pico (Santos and Santana 2009b), São Miguel (Santos and Santana 2004)
Notes
Freshwater (lake)
Aphanocapsa elachista
West & G.S.West, 1894
8FD0214C-D1F9-5BCC-B9E7-DCF30A00BFA1
Distribution
São Jorge (Cedercreutz 1941), São Miguel (Santos and Santana 2004)
Notes
Freshwater (lake), terrestrial
Aphanocapsa grevillei
(Berkeley) Rabenhorst, 1865
39825678-DC02-5F4D-AAE9-7A5D6C688C1D
Distribution
São Jorge (Johansson 1977)
Notes
Freshwater
Aphanocapsa incerta
(Lemmermann) G.Cronberg & Komárek, 1994
54E2F8B0-827F-526B-863B-2652DB96E871
Distribution
São Miguel (Santos and Santana 2009a)
Notes
Freshwater (lake)
Aphanothece
Nägeli, 1849
BA39489C-B663-5554-A246-5D4BEC62B1EA
Distribution
Flores (Luz et al. 2020), São Miguel (Luz et al. 2020)
Notes
Freshwater (lake)
Aphanothece castagnei
(Kützing) Rabenhorst, 1865
497B852B-F6D3-5A40-9BF8-65C6038B0BFF
Distribution
Flores (Bourrelly and Manguin 1946)
Notes
Terrestrial
Aphanothece microscopica
Nägeli, 1849
E5C56523-31FD-5A31-91B1-3EF75A823619
Distribution
São Miguel (Bohlin 1901)
Notes
Freshwater (lake), terrestrial
Aphanothece naegelii
Wartmann in Rabenhorst, 1865
6DCFDD27-026B-589E-A429-349A6ED23A87
Distribution
São Miguel (Bohlin 1901)
Notes
Freshwater
Aphanothece nidulans
P.Richter, 1884
DC09DA52-E952-5BBD-BFCB-DB537B81C2A3
Distribution
São Miguel (Bourrelly and Manguin 1946)
Notes
Freshwater (lake)
Aphanothece pallida
(Kützing) Rabenhorst, 1863
2E7D654E-447C-5597-8F66-C5D97900A1C2
Distribution
São Miguel (Bourrelly and Manguin 1946)
Notes
Freshwater (lake)
Aphanothece saxicola
Nägeli, 1849
8411CDF5-C0CE-5678-BB2F-697D7C02E1E8
Distribution
São Miguel (Bohlin 1901)
Notes
Terrestrial
Aphanothece stagnina
(Sprengel) A.Braun in Rabenhorst, 1863
92387C66-DD74-547C-AC27-D22A52C37C46
Distribution
Pico (Johansson 1977)
Notes
Freshwater
Arthrospira
Sitzenberger ex Gomont, 1892
3D71333A-E7E2-5B03-B290-633181D35210
Distribution
São Miguel (Cordeiro et al. 2020b)
Notes
Freshwater (lake)
Calothrix
C.Agardh ex Bornet & Flahault, 1886
7EE8080E-33F4-5D0D-B13F-F56B3F7F90D5
Distribution
Flores (Cordeiro et al. 2020b), São Jorge (Luz 2018), Santa Maria (Cordeiro et al. 2020b), São Miguel (Cordeiro et al. 2020b)
Notes
Freshwater (lake, stream)
Calothrix breviarticulata
West & G.S.West, 1897
7F2A9670-72FA-5282-A8A1-657C0CB5CA08
Distribution
Flores (Luz 2018)
Notes
Freshwater (lake)
Calothrix castellii
Bornet & Flahault, 1886
EEB25B65-A92B-5B2A-A6A6-07FB71990651
Distribution
Notes
Freshwater (lake)
Calothrix parietina
Thuret ex Bornet & Flahault, 1886
7421F633-1861-5DE5-99F0-3F466F984AC4
Distribution
São Miguel (Bohlin 1901)
Notes
Terrestrial
Chlorogloeopsis fritschii
(A.K.Mitra) A.K.Mitra & D.C.Pandey, 1967
61D91FBD-E047-5ACB-A99A-516EF09A6401
Distribution
São Miguel (Luz 2018)
Notes
Thermal (stream)
Chroococcus
Nägeli, 1849
6EA6AF4F-B38D-53ED-B0E8-4BAE0D8371AB
Distribution
São Miguel (Moseley 1874)
Notes
Freshwater (lake), thermal (spring)
Chroococcus dispersus
(Keissler) Lemmermann, 1904
E13877D8-7240-563C-961C-FB00B97DEBD4
Distribution
Pico (Santos and Santana 2009b), São Miguel (Santos and Santana 2009a)
Notes
Freshwater (lake)
Chroococcus membraninus
(Meneghini) Nägeli, 1849
D0F29FFA-1B90-5FF8-BEA8-6331700F7430
Distribution
São Miguel (Bohlin 1901)
Notes
Thermal (stream)
Chroococcus minor
(Kützing) Nägeli, 1849
6FCC9259-D6DF-5AC4-8A82-A2004AFACA42
Distribution
São Miguel (Archer 1874)
Notes
Freshwater (lake)
Chroococcus minutus
(Kützing) Nägeli, 1849
4F1AED92-AD9C-5A81-B4E8-4FC65E7E21A8
Distribution
Flores (Luz et al. 2020), Pico (Johansson 1977), São Miguel (Bourrelly and Manguin 1946)
Notes
Freshwater (lake)
Chroococcus tenax
(Kirchner) Hieronymus, 1892
C0A6C736-2475-581E-9B95-A841D97CBEA0
Distribution
São Miguel (Oliveira 1989)
Notes
Freshwater (lake)
Chroococcus turgidus
(Kützing) Nägeli, 1849
51399019-4C66-5844-9523-0BFE6A161DFC
Distribution
Corvo (Cedercreutz 1941), Faial (Johansson 1977), Flores (Cedercreutz 1941), Pico (Johansson 1977), São Jorge (Johansson 1977), São Miguel (Bohlin 1901), Terceira (Cedercreutz 1941)
Notes
Freshwater (lake), terrestrial
Chroococcus turicensis
(Nägeli) Hansgirg, 1887
F5B2773F-B33D-5A1B-802A-071C23288BE3
Distribution
Flores (Bourrelly and Manguin 1946)
Notes
Freshwater
Chroococcus westii
J.B.Petersen, 1923
45271513-58C3-51E7-BBD4-2E5F56FDCC4D
Distribution
São Miguel (Oliveira 1989)
Notes
Terrestrial
Coelosphaerium
Nägeli, 1849
327A108E-1D20-5FF7-A77F-34556379954C
Distribution
São Miguel (Santos et al. 2001)
Notes
Freshwater (lake)
Coelosphaerium kuetzingianum
Nägeli, 1849
732E25B0-EA89-59F3-8178-3D047DF4DF90
Distribution
São Miguel (Santos and Santana 2004)
Notes
Freshwater (lake)
Coleospermum
Kirchner ex Frank, 1886
ADD1E5D6-BA23-5557-8C20-25144AF9EADD
Distribution
Notes
Freshwater (lake), thermal (pool, spring)
Coleospermum goeppertianum
Kirchner ex Frank, 1886
9ACACF4D-5D2C-5DA9-B3E0-C89AF2C18D73
Distribution
São Miguel (Krieger 1931)
Notes
Freshwater (lake)
Cyanobacterium synechococcoides
Komárek 1999
4D270238-0B1E-5853-9CBB-2AFCEC985639
Distribution
São Miguel (Cordeiro et al. 2020b)
Notes
Freshwater (lake)
Cyanobium
R.Rippka & G.Cohen-Bazire, 1983
BC7204EE-5549-518E-B23E-F522E5A9E588
Distribution
São Miguel (Cordeiro et al. 2020b)
Notes
Freshwater (lake)
Cyanobium plancticum
(G.Drews, H.Prauser & D.Uhlmann) Komárek, J.Kopecký & Cepák, 1999
6510DFA6-A64B-5CAF-8366-EF20536A93DB
Distribution
São Miguel (Xavier et al. 2018)
Notes
Freshwater (lake)
Cyanosaccus
K.J.Lukas & S.Golubic, 1981
490DA872-E818-5B86-86C6-4DFE8D7740EC
Distribution
Faial (Wisshak et al. 2011)
Notes
Marine (intertidal)
Cylindrospermum
Kützing ex Bornet & Flahault, 1886
76D98B8A-1394-5587-B60D-DE3B1F3481A3
Distribution
Pico (Cedercreutz 1941), São Miguel (Cordeiro et al. 2020b)
Notes
Freshwater (lake, stream)
Cylindrospermum licheniforme
Kützing ex Bornet & Flahault, 1886
CD0072DE-A493-58B1-B413-2BF2DD2535A2
Distribution
São Miguel (Trelease 1897)
Notes
Freshwater
Cylindrospermum majus
Kützing ex Bornet & Flahault, 1886
CBB9E118-DC1A-5F6A-9EDF-A06F2BD09469
Distribution
Faial (Johansson 1977), Flores (Bourrelly and Manguin 1946), São Jorge (Johansson 1977), São Miguel (Trelease 1897)
Notes
Freshwater (lake)
Dichothrix baueriana
Bornet & Flahault, 1886
605545D7-3ADB-54CE-ACCB-E1ED74D5C226
Distribution
Corvo (Trelease 1897)
Notes
Freshwater
Dichothrix orsiniana var. africana
Frémy, 1924
4520628B-ECB1-566B-8BAA-9415AAF283FA
Distribution
Flores (Bourrelly and Manguin 1946)
Notes
Freshwater (lake)
Dolichospermum
(Ralfs ex Bornet & Flahault) P.Wacklin, L.Hoffmann & J.Komárek, 2009
C646515E-D412-5E81-A619-B70D9690425B
Distribution
São Miguel (Cordeiro et al. 2020b)
Notes
Freshwater (lake)
Dolichospermum affine
(Lemmermann) Wacklin, L.Hoffmann & Komárek, 2009
608983D2-24B1-5FB3-AEC7-518B93CCE7A8
Distribution
Pico (Santos and Santana 2009b), São Miguel (Oliveira 1989)
Notes
Freshwater (lake)
Dolichospermum circinale
(Rabenhorst ex Bornet & Flahault) Wacklin, L.Hoffmann & Komárek, 2009
B91AFB58-208C-5199-834C-428612019756
Distribution
São Miguel (Cordeiro 2015)
Notes
Freshwater (lake)
Dolichospermum delicatulum
(Lemmermann) Wacklin, L.Hoffmann & Komárek, 2009
DCB702F8-E4B4-51D9-9E3C-BB0BA69AA83F
Distribution
Corvo (Luz et al. 2020), Flores (Luz et al. 2020), Pico (Luz et al. 2020), São Miguel (Luz et al. 2020)
Notes
Freshwater (lake)
Dolichospermum flos-aquae
(Brébisson ex Bornet & Flahault) Wacklin, L.Hoffmann & Komárek, 2009
9AAD69CE-9507-5D0A-BF8D-4DCEEC859E4F
Distribution
São Miguel (INOVA 1996)
Notes
Freshwater (lake)
Dolichospermum planctonicum
(Brunnthaler) Wacklin, L.Hoffmann & Komárek, 2009
BD58B186-0E34-5D9C-AC6E-6456585EBFD1
Distribution
Pico (Luz et al. 2020), São Miguel (Luz et al. 2020)
Notes
Freshwater (lake)
Dolichospermum scheremetieviae
(Elenkin) Wacklin, L.Hoffmann & Komárek, 2009
1221A41B-8B2A-5882-83A8-EC38D2C68492
Distribution
Corvo (Luz et al. 2020), Flores (Luz et al. 2020), Pico (Santos and Santana 2009b), São Miguel (Oliveira 1989)
Notes
Freshwater (lake)
Dolichospermum sigmoideum
(Nygaard) Wacklin, L.Hoffmann & Komárek, 2009
D088D6B9-11F2-5244-BDE2-4DABC9EDEC1E
Distribution
Pico (Santos and Santana 2009b), São Miguel (Cordeiro 2015)
Notes
Freshwater (lake)
Dolichospermum solitarium
(Klebahn) Wacklin, L.Hoffmann & Komárek, 2009
E4A9E0C6-7AC7-5C97-A7A3-C9BF9608CE35
Distribution
Flores (Luz et al. 2020), Pico (Luz et al. 2020), São Miguel (Bourrelly and Manguin 1946), Terceira (Luz et al. 2020)
Notes
Freshwater (lake)
Dolichospermum spiroides
(Klebhan) Wacklin, L.Hoffmann & Komárek, 2009
5456697E-14D1-5872-9A51-463C92738403
Distribution
São Miguel (Luz et al. 2020)
Notes
Freshwater (lake)
Eucapsis alpina
F.E.Clements & H.L.Schantz, 1909
754423E1-3256-547E-B07F-C607CCC76555
Distribution
Corvo (Luz et al. 2020), Flores (Luz et al. 2020), Pico (Luz et al. 2020), São Miguel (Luz et al. 2020), Terceira (Luz et al. 2020)
Notes
Freshwater (lake)
Eucapsis minuta
F.E.Fritsch, 1912
CB370973-6823-52D0-97DC-D726C2358396
Distribution
São Miguel (Luz et al. 2020)
Notes
Freshwater (lake)
Fischerella
(Bornet & Flahault) Gomont, 1895
40F27740-A289-594B-909C-D13CFFACF8F2
Distribution
São Miguel (Cordeiro et al. 2020b)
Notes
Thermal (pool)
Fortiea
De Toni, 1936
D306D1CC-AAAE-5C46-B132-4F957B23002B
Distribution
Pico (Luz 2018)
Notes
Freshwater (lake)
Fortiea striatula
(F.C.Hy) De Toni, 1936
B6B3BC2F-0524-52FA-819A-1166708AEBF1
Distribution
Pico (Cordeiro et al. 2020b)
Notes
Freshwater (lake)
Geitlerinema ionicum
(Skuja) Anagnostidis, 1989
FC04E65F-29F9-5D3D-8396-2652EDB8D803
Distribution
Santa Maria (Bourrelly and Manguin 1946)
Notes
Terrestrial
Geitlerinema splendidum
(Greville ex Gomont) Anagnostidis, 1989
8390719B-3AA3-5717-8F40-68E97BD9EC77
Distribution
Pico (Luz et al. 2020), São Miguel (Cedercreutz 1941)
Notes
Freshwater (lake)
Gloeocapsa
Kützing, 1843
A37A7C2F-E744-5BFF-BE9D-BB661590FE41
Distribution
São Miguel (Luz et al. 2020)
Notes
Freshwater (lake)
Gloeocapsa atrata
Kützing, 1843
7BE6EE57-67E8-514B-914F-859004B4C25D
Distribution
São Miguel (Cedercreutz 1941), Terceira (Johansson 1977)
Notes
Freshwater, terrestrial
Gloeocapsa caldariorum
Rabenhorst, 1865
D4BAF589-2FDC-5142-8E17-AFA299A8F4AB
Distribution
Terceira (Johansson 1977)
Notes
Freshwater
Gloeocapsa compacta
Kützing, 1847
30649D81-E503-5960-BC44-04FEA3407185
Distribution
Flores (Bourrelly and Manguin 1946)
Notes
Freshwater (lake)
Gloeocapsa gelatinosa
Kützing, 1843
D3D4627F-5A99-5CCE-AC62-8B448724F8E3
Distribution
São Jorge (Johansson 1977), Terceira (Johansson 1977)
Notes
Freshwater
Gloeocapsa quaternata
Kützing, 1846
CFEC0C70-C9A3-5820-9EF4-D4149633E7F7
Distribution
São Miguel (Johansson 1977)
Notes
Freshwater
Gloeocapsa rupestris
Kützing, 1847
98995BAA-76E2-5DD9-97FE-2A19E17DB7FC
Distribution
São Jorge (Johansson 1977), Terceira (Johansson 1977)
Notes
Freshwater
Gloeocapsa thermalis
Kützing, 1843
C4DB2CC3-FED3-5ED7-A966-F96D11B9E93D
Distribution
São Miguel (Johansson 1977)
Notes
Thermal
Gloeocapsopsis dvorakii
(Novácek) Komárek & Anagnostidis ex Komárek 1993
0D8C7E2E-A879-5D1F-A7BA-4C4A5ECAAC3E
Distribution
São Miguel (Cordeiro et al. 2020b)
Notes
Thermal (pool)
Gloeocapsopsis magma
(Brébisson) Komárek & Anagnostidis ex Komárek, 1993
DFC47F20-3A6B-553C-9036-7665DF7A6678
Distribution
São Miguel (Bohlin 1901)
Notes
Terrestrial
Gloeothece cystifera
(Hassall) Rabenhorst, 1865
A7C4A689-2C1C-57CB-AAC5-CBA2D7318CC6
Distribution
São Miguel (Bohlin 1901)
Notes
Terrestrial
Gloeothece rupestris
(Lyngbye) Bornet in Wittrock & Nordstedt, 1880
DC2E50FA-F6D1-5806-B78F-E6B8FDD54616
Distribution
Flores (Bourrelly and Manguin 1946), São Miguel (Cedercreutz 1941)
Notes
Freshwater (lake)
Gloeotrichia pisum
Thuret ex Bornet & Flahault, 1886
09E5ADCA-D25A-5C28-AFEC-37D61E8AA44D
Distribution
São Miguel (Bohlin 1901)
Notes
Terrestrial
Goleter apudmare
Miscoe & J.R.Johansen, 2016
E2CF2F03-F5E2-5849-A65E-A8F40B75C549
Distribution
Flores (Cordeiro et al. 2020b)
Notes
Freshwater (lake)
Gomphosphaeria
Kützing, 1836
349C4809-0663-5289-BBAB-6BDF0F1F9C77
Distribution
São Miguel (Santos et al. 2001)
Notes
Freshwater (lake)
Hapalosiphon hibernicus
West & G.S.West, 1896
C653B3E8-D5BE-54F8-ABB2-4ACB7EB84967
Distribution
Corvo (Cedercreutz 1941), Flores (Cedercreutz 1941), São Miguel (Bourrelly and Manguin 1946)
Notes
Freshwater (lake)
Hapalosiphon intricatus
West & G.S.West, 1894
32DF9264-1C08-5D22-A516-FC43697EDAFE
Distribution
São Jorge (Cedercreutz 1941), Terceira (Johansson 1977)
Notes
Freshwater (lake, stream)
Hapalosiphon pumilus
Kirchner ex Bornet & Flahault, 1887
47DFBC51-FC9C-56AB-9315-55CB3780F404
Distribution
Flores (Bourrelly and Manguin 1946), Santa Maria (Luz 2018)
Notes
Freshwater (stream), terrestrial
Hapalosiphon
Nägeli ex É.Bornet & C.Flahault, 1886
98E15AF8-C4EF-5A52-AC93-EB2168A2EB78
Distribution
São Miguel (Cordeiro et al. 2020b)
Notes
Terrestrial
Heteroleibleinia kuetzingii
(Schmidle) Compère, 1985
803EECD2-B18B-510A-89DE-7524AED43D2E
Distribution
São Jorge (Cedercreutz 1941)
Notes
Freshwater
Homoeothrix africana
G.S.West, 1912
C5EB2BAA-E51F-589A-B07F-B43890A79542
Distribution
São Jorge (Johansson 1977)
Notes
Freshwater
Hydrocoryne spongiosa
Schwabe ex Bornet & Flahault 1887
03DF7F50-C3A3-5112-94B5-AF68ADD1DAF0
Distribution
São Miguel (Cordeiro et al. 2020b)
Notes
Freshwater (lake)
Hyella
É.Bornet & C.Flahault, 1888
6588D41C-AF0E-5D30-BACF-2FFFB3C665AC
Distribution
Faial (Wisshak et al. 2011)
Notes
Marine (intertidal)
Hyella caespitosa
Bornet & Flahault, 1888
AFDB46F8-C22A-5062-BE0B-912F4391D908
Distribution
Faial (Wisshak et al. 2011)
Notes
Marine (intertidal)
Hyella gigas
Lukas & Golubic, 1983
85CAA134-6978-55EB-A7B6-BD600287FE89
Distribution
Faial (Wisshak et al. 2011)
Notes
Marine (intertidal)
Isocystis planctonica
Starmach 1962
92F4F0E2-A517-5298-8771-8F6BCA36A2B0
Distribution
Flores (Cordeiro et al. 2020b)
Notes
Freshwater (lake)
Kamptonema
O.Strunecký, J.Komárek & J.Smarda, 2014
8BED7D41-3DC7-5DA1-B982-E08CB245EEF8
Distribution
Flores (Cordeiro et al. 2020b)
Notes
Freshwater (lake)
Kamptonema formosum
(Bory ex Gomont) Strunecký, Komárek & J.Smarda, 2014
9939C7C1-000A-537D-9296-D8C2CFAA4EBE
Distribution
Santa Maria (Trelease 1897), São Miguel (Bohlin 1901), Terceira (Cedercreutz 1941)
Notes
Freshwater, brackish (lake), thermal (stream)
Kyrtuthrix dalmatica
Ercegovic, 1929
FC2C266E-CC53-529F-A8B7-E8A794B80998
Distribution
Faial (Wisshak et al. 2011)
Notes
Marine (intertidal)
Leptodesmis
Raabová, Kovacik & Strunecký, 2019
416BE2FB-5390-5BF0-A207-6DC44FBF4BCB
Distribution
São Miguel (Cordeiro et al. 2020b)
Notes
Freshwater (lake)
Leptolyngbya
Anagnostidis & Komárek, 1988
541015A2-AB83-51AD-81AC-793EA01FDF72
Distribution
Notes
Freshwater (lake), marine (lake), thermal (pool, stream)
Leptolyngbya gelatinosa
(Woronichin) Anagnostidis & Komárek, 1988
4B830626-D8B2-53B7-8938-4135F4D35596
Distribution
São Miguel (Luz 2018)
Notes
Thermal (stream)
Leptolyngbya granulifera
(J.J.Copeland) Anagnostidis 1936
C2EBE813-7B38-5D80-B1C1-703CFBD03695
Distribution
São Miguel (Cordeiro et al. 2020b)
Notes
Thermal (pool, spring)
Leptolyngbya laminosa
(Gomont ex Gomont) Anagnostidis & Komárek, 1988
4E808AF6-FFC4-5188-85E6-FA70E21E70C7
Distribution
São Miguel (Trelease 1897)
Notes
Thermal (spring)
Leptolyngbya nostocorum
(Bornet ex Gomont) Anagnostidis & Komárek, 1988
81E64AEC-EA8C-56D1-B716-42ABE80F86D4
Distribution
São Miguel (Bohlin 1901)
Notes
Freshwater (lake)
Leptolyngbya ochracea
(Thuret ex Gomont) Anagnostidis & Komárek, 1988
FA3EA5D5-6F68-5FD1-ACEA-21B97685FB6B
Distribution
São Miguel (Bohlin 1901)
Notes
Thermal (pool)
Leptolyngbya rivulariarum
(Gomont) Anagnostidis & Komárek, 1988
AFDF8AF7-C5A4-5379-89A8-27FED5A8133B
Distribution
São Miguel (Bohlin 1901)
Notes
Freshwater (lake)
Leptolyngbya subuliformis
(Gomont) Anagnostidis 2001
039FE232-94C8-5E4F-833C-5DCE69EB496A
Distribution
São Miguel (Cordeiro et al. 2020b)
Notes
Thermal (spring)
Leptolyngbya valderiana
(Gomont) Anagnostidis & Komárek, 1988
42D201A5-4778-5DD8-AFD1-2BDDB89BD077
Distribution
São Miguel (Bourrelly and Manguin 1946)
Notes
Terrestrial
Limnothrix
M.-E.Meffert, 1988
01B69F67-4BB1-583A-979C-6098D2CFA6F1
Distribution
Flores (Cordeiro et al. 2020b), São Miguel (Cordeiro et al. 2020b)
Notes
Freshwater (lake)
Limnothrix planctonica
(Woloszynska) Meffert, 1988
422E794B-09DC-5C26-8098-9E60113DFD28
Distribution
São Miguel (Santos and Santana 2009a)
Notes
Freshwater (lake)
Lyngbya
C.Agardh ex Gomont, 1892
1394AE00-73BF-5720-9E8D-4C50AB36D3C5
Distribution
São Miguel (Santos and Santana 2004)
Notes
Freshwater (lake)
Lyngbya lutea
Gomont ex Gomont, 1892
F36215C1-C130-51A9-BF4A-56A2B165F739
Distribution
Terceira (Neto et al. 2009)
Notes
Marine
Lyngbya martensiana
Meneghini ex Gomont, 1892
B2589F32-A84E-5CAE-8803-BA40FA5DB33F
Distribution
São Miguel (Bohlin 1901), Terceira (Johansson 1977)
Notes
Freshwater, thermal, terrestrial
Mastigocladus
Cohn ex Kirchner, 1898
ADD76FAA-B7B9-51CC-AC79-70D1CA76489E
Distribution
São Miguel (Cordeiro et al. 2020b)
Notes
Thermal (spring, stream)
Mastigocladus laminosus
Cohn ex Kirchner, 1898
DC1E7734-3205-5857-BE27-D9A58E52D9CC
Distribution
São Miguel (Bohlin 1901)
Notes
Thermal (spring, pool, stream)
Mastigocoleus testarum
Lagerheim ex Bornet & Flahault, 1886
7D733C69-72DF-58F1-8E82-624DD0DAEBC8
Distribution
Faial (Wisshak et al. 2011)
Notes
Marine (intertidal)
Merismopedia
Meyen, 1839
3664D887-87A5-5ACF-8D81-350E4491F435
Distribution
São Miguel (Archer 1874), Pico (Luz et al. 2020)
Notes
Freshwater (lake)
Merismopedia glauca
(Ehrenberg) Kützing, 1845
06B94FB9-EA53-5880-A2EC-11F2DF98BB7A
Distribution
São Miguel (Cedercreutz 1941), Terceira (Cedercreutz 1941)
Notes
Freshwater (lake)
Merismopedia tenuissima
Lemmermann, 1898
8D51F50A-BD13-5ED3-92DA-76C84678A48D
Distribution
Graciosa (Azevedo et al. 2005), São Miguel (Oliveira 1989), Terceira (Luz et al. 2020)
Notes
Freshwater (lake)
Microchaete bulbosa
J.Copeland, 1936
CEA81422-B88D-507A-964C-D6F4BF0C8342
Distribution
São Miguel (Luz 2018)
Notes
Thermal (spring)
Microchaete tenera
Thuret ex Bornet & Flahault, 1886
1C3B2104-118D-5F9F-8BF2-DF992186E19D
Distribution
Pico (Luz 2018), São Miguel (Bohlin 1901)
Notes
Freshwater (lake), terrestrial
Microcoleus amoenus
(Gomont) Strunecky, Komárek & J.R.Johansen, 2013
8EB10850-E428-50B4-97E0-31EA76A3BCDD
Distribution
Flores (Bourrelly and Manguin 1946), São Miguel (Cedercreutz 1941)
Notes
Freshwater (lake)
Microcoleus autumnalis
(Gomont) Strunecky, Komárek & J.R.Johansen, 2013
1BD74790-908A-5C96-8B7E-AEA77CCD7732
Distribution
Flores (Cedercreutz 1941), São Miguel (Cedercreutz 1941)
Notes
Freshwater (stream), terrestrial
Microcoleus lyngbyaceus
Kützing ex Forti, 1907
2D1AAA68-E16F-54D3-A17F-5CAE600F154A
Distribution
Terceira (Neto et al. 2009)
Notes
Marine
Microcystis
Lemmermann, 1907
D0BD10AF-5680-54F2-8D50-79431D9E342A
Distribution
São Miguel (INOVA 1996)
Notes
Freshwater (lake)
Microcystis aeruginosa
(Kützing) Kützing, 1846
2DEBF0D2-749E-5B17-8DA6-A07B3DCE594B
Distribution
Corvo (Luz et al. 2020), Flores (INOVA 1996), São Miguel (Vasconcelos et al. 1994)
Notes
Freshwater (lake)
Microcystis flos-aquae
(Wittrock) Kirchner, 1898
861390ED-F70D-574A-A12D-8469593AC2BF
Distribution
Flores (Luz et al. 2020), São Miguel (Oliveira 1989)
Notes
Freshwater (lake)
Microcystis pulverea
(H.C.Wood) Forti, 1907
C73B7D99-7A17-5D65-AE82-8E4E3B389155
Distribution
Pico (Santos and Santana 2009b), São Miguel (Santos and Santana 2004)
Notes
Freshwater (lake)
Microcystis robusta
(H.W.Clark) Nygaard, 1925
A32BC5F3-66D5-54FC-87A8-41A428764EB6
Distribution
São Miguel (Santos and Santana 2009a), Terceira (Santos and Santana 2009b)
Notes
Freshwater (lake)
Nodularia harveyana
Thuret ex Bornet & Flahault, 1886
C9A5F77F-12BC-5F0D-8FA2-9846A59B9BF1
Distribution
Corvo (Trelease 1897), Terceira (Cedercreutz 1941)
Notes
Brackish (lake), marine
Nostoc
Vaucher ex Bornet & Flahault, 1886
0D64A689-B759-5FD5-8F43-E84FDC561634
Distribution
Corvo (Cedercreutz 1941), Flores (Cedercreutz 1941), Santa Maria (Bourrelly and Manguin 1946), São Miguel (Moseley 1874)
Notes
Freshwater (lake, stream), terrestrial
Nostoc carneum
C.Agardh ex Bornet & Flahault, 1886
3504816D-643E-5E8A-BC52-BA57396FBA60
Distribution
Faial (Johansson 1977)
Notes
Freshwater
Nostoc commune
Vaucher ex Bornet & Flahault, 1886
2ABBBB94-EA1F-5222-8F7C-6CE439EE2578
Distribution
Pico (Johansson 1977)
Notes
Freshwater (lake)
Nostoc ellipsosporum
Rabenhorst ex Bornet & Flahault, 1886
FE19FC1D-AD73-5DE3-B0DE-5B54BC182A77
Distribution
Corvo (Trelease 1897), São Miguel (Bohlin 1901)
Notes
Freshwater
Nostoc paludosum
Kützing ex Bornet & Flahault, 1886
EA708117-914D-55F6-B68B-0AA4D4836EA2
Distribution
Corvo (Cordeiro et al. 2020b), Santa Maria (Bourrelly and Manguin 1946), São Miguel (Bohlin 1901)
Notes
Freshwater (lake)
Nostoc punctiforme
Hariot, 1891
E1749B75-D37F-5E43-9D4F-764E93EA3339
Distribution
São Miguel (Bohlin 1901)
Notes
Freshwater (lake), terrestrial
Nostoc sphaericum
Vaucher ex Bornet & Flahault, 1886
C7A90830-003D-5439-BA48-09318FE82DB4
Distribution
Faial (Johansson 1977), Flores (Bourrelly and Manguin 1946), São Jorge (Johansson 1977), São Miguel (Bourrelly and Manguin 1946)
Notes
Freshwater (lake), terrestrial
Nostoc sphaeroides
Kützing ex Bornet & Flahault, 1886
BF7A8666-B84A-5336-9C50-A8562CE303E7
Distribution
Faial (Johansson 1977), São Jorge (Johansson 1977)
Notes
Freshwater
Nostoc verrucosum
Vaucher ex Bornet & Flahault, 1886
9081A85F-45F8-52FF-8A28-2CE46A1BB116
Distribution
São Miguel (Cedercreutz 1941), Terceira (Trelease 1897)
Notes
Freshwater (stream)
Nostochopsis lobatus
H.C.Wood ex Bornet & Flahault, 1886
8A5D8CAA-A884-5E5E-8181-EDD72490185E
Distribution
São Miguel (Bohlin 1901)
Notes
Terrestrial
Oscillatoria
Vaucher ex Gomont, 1892
DCBF69E6-84DD-51C8-BD2F-AD4D08A29F46
Distribution
Flores (Luz et al. 2020), Pico (INOVA 1996), São Miguel (Moseley 1874), Terceira (Neto et al. 2009)
Notes
Freshwater (lake), marine
Oscillatoria geitleri f. major
Bourrelly in Bourrelly & Manguin, 1946
624C3A67-32CB-5903-9830-A1F1B72051A6
Distribution
Flores (Bourrelly and Manguin 1946)
Notes
Freshwater (stream)
Oscillatoria planctonica
Woloszynska, 1912
F448449E-96CF-5022-AB0C-4B4696B4C372
Distribution
São Miguel (Oliveira 1989)
Notes
Freshwater (lake)
Oscillatoria princeps
Vaucher ex Gomont, 1892
DC340E66-4A89-5F9A-8D52-AAE724E171D7
Distribution
Terceira (Neto et al. 2009)
Notes
Marine
Oscillatoria sancta
Kützing ex Gomont, 1892
0EBB4815-E988-5846-AE6D-F85788925D50
Distribution
São Miguel (Bohlin 1901)
Notes
Terrestrial
Oscillatoria tenuis
C.Agardh ex Gomont, 1892
50EEEE63-B5C9-5190-8602-C0D3B894A7D4
Distribution
Corvo (Cedercreutz 1941), Faial (Cedercreutz 1941), Flores (Cedercreutz 1941), Graciosa (Cedercreutz 1941), Pico (Luz et al. 2020), São Miguel (Bohlin 1901), Terceira (Luz et al. 2020)
Notes
Freshwater (lake, stream), terrestrial
Pegethrix
Mai, J.R.Johansen & Bohunická, 2018
A2B01D08-5109-5AAA-8195-C13EA19C31A2
Distribution
São Miguel (Cordeiro et al. 2020b)
Notes
Freshwater (lake)
Petalonema velutinum
Migula, 1907
1FDA5E34-D314-5044-A091-E398FB8682D1
Distribution
Flores (Cedercreutz 1941)
Notes
Freshwater
Phormidium
Kützing ex Gomont, 1892
6D106293-F8FA-5AD2-9EAA-0BCA8DCCBF7E
Distribution
Corvo (Luz et al. 2020), Faial (Cedercreutz 1941), Pico (Luz et al. 2020), São Jorge (Luz 2018), São Miguel (Fish and Codd 1994)
Notes
Freshwater (lake), marine (lake), thermal
Phormidium aerugineo-caeruleum
(Gomont) Anagnostidis & Komárek, 1988
170CD658-E933-5F78-8EED-5BBDA9A07FC5
Distribution
São Miguel (Bohlin 1901)
Notes
Freshwater (lake)
Phormidium allorgei
(Frémy) Anagnostidis & Komárek, 1988
8745AAB7-6F51-581A-86C6-791A2403BC9B
Distribution
Terceira (Johansson 1977)
Notes
Freshwater
Phormidium breve
(Kützing ex Gomont) Anagnostidis & Komárek, 1988
F1939C00-6610-5FCC-993F-E00C5F8BA537
Distribution
São Miguel (Bohlin 1901)
Notes
Marine, thermal
Phormidium durum
N.L.Gardner, 1927
976B2EF6-DAE6-5169-AC4B-D2F22ACA0425
Distribution
São Jorge (Johansson 1977)
Notes
Freshwater
Phormidium irriguum
(Kützing ex Gomont) Anagnostidis & Komárek, 1988
F1217E6A-93E8-56C4-BD3A-35B2E2702A22
Distribution
São Jorge (Johansson 1977), Terceira (Bohlin 1901)
Notes
Freshwater (stream)
Phormidium pachydermaticum
Frémy, 1930
542492AB-F5B9-571F-8D6F-77150A2458DD
Distribution
São Jorge (Johansson 1977), Terceira (Johansson 1977)
Notes
Freshwater
Phormidium retzii
Kützing ex Gomont, 1892
5F2C1CBB-3DF1-5E29-9F40-213BE887C7D8
Distribution
Faial (Johansson 1977), São Jorge (Johansson 1977), São Miguel (Cedercreutz 1941)
Notes
Freshwater
Phormidium rotheanum
Itzigsohn in Rabenhorst, 1865
EEEAF295-3AEF-5C6F-8EA4-5ECF20493ED5
Distribution
São Jorge (Johansson 1977)
Notes
Freshwater
Phormidium terebriforme
(C.Agardh ex Gomont) Anagnostidis & Komárek, 1988
91BE4B1D-9F31-5A08-824C-1BE87AA1C733
Distribution
São Miguel (Bohlin 1901)
Notes
Thermal
Planktolyngbya
Anagnostidis & Komárek, 1988
424EF739-A621-5534-B535-EF66951A6898
Distribution
Flores (Luz et al. 2020), Pico (Luz et al. 2020), São Miguel (Cordeiro 2015)
Notes
Freshwater (lake)
Planktolyngbya limnetica
(Lemmermann) Komárková-Legnerová & Cronberg, 1992
081964B8-8A08-5C9E-8267-DB5602DA1713
Distribution
Pico (Luz et al. 2020), São Miguel (Cedercreutz 1941)
Notes
Freshwater (lake)
Planktothrix agardhii
(Gomont) Anagnostidis & Komárek, 1988
39DA6936-415F-5ADD-A16A-3AB4B772982D
Distribution
São Miguel (Santos and Santana 2009a)
Notes
Freshwater (lake)
Plectonema endolithicum
Ercegovic, 1932
BECB3156-D1B4-57DF-80EF-A214C58D8FE2
Distribution
Faial (Wisshak et al. 2011)
Notes
Marine (intertidal)
Plectonema terebrans
Bornet & Flahault ex Gomont, 1892
6A2CAE44-AE45-5789-B5C1-9EBCB17D6BA1
Distribution
Faial (Wisshak et al. 2011)
Notes
Marine (intertidal)
Pseudanabaena
Lauterborn, 1915
D7A53FFE-E282-5537-890B-7A77C1F7165F
Distribution
Corvo (Luz et al. 2020), Flores (Luz et al. 2020), Pico (Luz et al. 2020), São Miguel (Santos et al. 2005)
Notes
Freshwater (lake)
Pseudanabaena catenata
Lauterborn, 1915
C84CFF8A-311B-5950-A45C-3DD3C2489636
Distribution
São Miguel (Oliveira 1989)
Notes
Freshwater (lake)
Pseudanabaena limnetica
(Lemmermann) Komárek, 1974
DDD3F482-CBB4-57C6-87EF-B67D916046CD
Distribution
Corvo (Luz et al. 2020), Flores (Luz et al. 2020), Pico (Luz et al. 2020), São Miguel (Luz et al. 2020)
Notes
Freshwater (lake)
Pseudanabaena minima
(G.S.An) Anagnostidis, 2001
37631284-F293-59FC-9CE5-C0729E4C9404
Distribution
Pico (Luz 2018)
Notes
Freshwater (lake)
Pseudanabaena mucicola
(Naumann & Huber-Pestalozzi) Schwabe, 1964
31E35EB1-8639-5C50-A0AC-553E39D0D570
Distribution
São Miguel (Santos and Santana 2004)
Notes
Freshwater (lake)
Pseudophormidium pauciramosum
(Anissimova) Anagnostidis, 2001
87385094-7C2F-5121-9349-7C77CA85F825
Distribution
Santa Maria (Luz 2018)
Notes
Brackish
Raphidiopsis curvata
F.E.Fritsch & M.F.Rich, 1930
3A722206-9AB1-5D6E-9A17-CF13C825CC19
Distribution
Corvo (INOVA 1996), Flores (INOVA 1996), São Miguel (INOVA 1996)
Notes
Freshwater (lake)
Rivularia
C.Agardh ex Bornet & Flahault, 1886
42B20208-B89E-5807-94D2-3FBA4452FD97
Distribution
São Miguel (Cordeiro et al. 2020b), Terceira (Neto et al. 2009)
Notes
Freshwater, marine
Rivularia biasolettiana
Meneghini ex Bornet & Flahault 1886
3160A454-4A04-59F0-AFEE-655A8B61CE3E
Distribution
São Miguel (Cordeiro et al. 2020b)
Notes
Freshwater (lake)
Rivularia bullata
Berkeley ex Bornet & Flahault, 1886
9D4D8EE8-D694-5307-8DC9-9F3728882850
Distribution
São Miguel (Trelease 1897)
Notes
Freshwater
Rivularia nitida
C.Agardh ex Bornet & Flahault, 1886
A46388FB-F8D7-5479-BB64-B958C5C201BA
Distribution
Flores (Trelease 1897)
Notes
Freshwater
Schizothrix
Kützing ex M.Gomont, 1892
31F968A0-515F-5248-B3BC-8D059332E39A
Distribution
Terceira (Johansson 1977)
Notes
Freshwater
Schizothrix cuspidata
(West & G.S.West) West & G.S.West, 1896
08E0E674-48CD-5B5C-9F5B-ACCBD9BB343C
Distribution
Faial (Bourrelly and Manguin 1946)
Notes
Terrestrial
Schizothrix fuscescens
Kutzing ex Gomont, 1892
047A30CD-2CFD-5872-A650-870B836EE18D
Distribution
Terceira (Johansson 1977)
Notes
Freshwater
Schizothrix lacustris
A.Braun ex Gomont, 1892
7A9DF08F-78C6-5FAA-83B7-65DEFCB50236
Distribution
São Jorge (Johansson 1977)
Notes
Freshwater
Schizothrix pallida
(Kützing ex Forti) Geitler, 1932
72497F8B-92AA-56AA-B265-6EAFFAABA2C0
Distribution
Terceira (Neto et al. 2009)
Notes
Marine
Schizothrix symplocoides
(N.L.Gardner) Geitler, 1932
FBABED4B-739C-50F5-A2AC-9272B7C92A36
Distribution
Terceira (Johansson 1977)
Notes
Freshwater
Schizothrix telephoroides
Gomont, 1890
6E3A16CB-DF26-5765-A68D-117120FDDF50
Distribution
Faial (Johansson 1977)
Notes
Freshwater
Schizothrix vaginata
Gomont, 1890
5DCA9FDD-7E6D-56BC-AD98-E97330CBDFC2
Distribution
São Jorge (Johansson 1977)
Notes
Freshwater
Scytonema
C.Agardh ex É.Bornet & C.Flahault, 1886
70A0C04A-3976-5165-8064-FD9C492C6892
Distribution
Flores (Cedercreutz 1941)
Notes
Terrestrial
Scytonema amplum
West & G.S.West, 1895
299294E7-25B1-5DD5-8C14-8E0E2B7418CD
Distribution
São Miguel (Bourrelly and Manguin 1946)
Notes
Terrestrial
Scytonema dilatatum
Bharadwaja, 1934
3FC47D9F-74F5-501F-93D2-16148B94C535
Distribution
Terceira (Johansson 1977)
Notes
Freshwater
Scytonema guyanense
Bornet & Flahault, 1888
4D41D234-53B5-582B-BB8F-5AE76A05ABDE
Distribution
Flores (Bourrelly and Manguin 1946), São Miguel (Bourrelly and Manguin 1946)
Notes
Terrestrial
Scytonema hofmannii
C.Agardh ex Bornet & Flahault, 1886
EE7ACCD6-0733-5846-A65D-84F072549D31
Distribution
São Jorge (Johansson 1977), São Miguel (Cedercreutz 1941)
Notes
Freshwater (lake)
Scytonema javanicum
Bornet ex Bornet & Flahault, 1886
82B473F8-3801-57BC-8C6B-80DB19B69118
Distribution
Flores (Bourrelly and Manguin 1946)
Notes
Freshwater (lake)
Scytonema mirabile
Bornet, 1889
25239CE2-7AFC-5963-85D4-408D7A91CDEA
Distribution
Pico (Cedercreutz 1941), São Jorge (Cedercreutz 1941), São Miguel (Cedercreutz 1941), Terceira (Cedercreutz 1941)
Notes
Freshwater (lake, stream), terrestrial
Scytonema stuposum
Bornet ex Bornet & Flahault, 1887
64736D9B-8EFE-53DB-9CD5-BD27C912F2C4
Distribution
Flores (Bourrelly and Manguin 1946)
Notes
Freshwater (lake)
Scytonematopsis
E.I.Kiseleva, 1930
10ABBF2D-A791-5A8C-8518-8A784FEC0455
Distribution
Flores (Cordeiro et al. 2020b), Pico (Cordeiro et al. 2020b)
Notes
Freshwater (lake)
Snowella
A.A.Elenkin, 1938
DBE1C8D2-C455-537E-9828-C3AB301CE9DC
Distribution
São Miguel (Luz et al. 2020)
Notes
Freshwater (lake)
Snowella lacustris
(Chodat) Komárek & Hindák, 1988
63FC5AEE-E9C9-55CA-94C0-E9C24E7C50A7
Distribution
São Miguel (Santos and Santana 2004)
Notes
Freshwater (lake)
Sphaerospermopsis
Zapomelová, Jezberová, Hrouzek, Hisem, Reháková & Komárková, 2010
137EBCE7-E44A-5419-A1D3-ADF0E71501D8
Distribution
Pico (Luz 2018)
Notes
Freshwater (lake)
Sphaerospermopsis aphanizomenoides
(Forti) Zapomelová, Jezberová, Hrouzek, Hisem, Reháková & Komárková, 2010
65F558B6-ED44-5180-B435-CF311BE91388
Distribution
Pico (Santos and Santana 2009b)
Notes
Freshwater (lake)
Spirulina subsalsa
Oersted ex Gomont, 1892
3EE7C9AC-5043-5CC6-9045-6AE70547876C
Distribution
São Jorge (Luz 2018)
Notes
Marine (lake)
Stenomitos
Miscoe & J.R.Johansen, 2016
B32D9844-A013-54F3-B033-E57ED65D674D
Distribution
Pico (Cordeiro et al. 2020b)
Notes
Freshwater (lake)
Stigonema hormoides
Bornet & Flahault, 1886
6BD919A7-E6FF-5A3A-B580-1D0D03ADD80F
Distribution
Flores (Bourrelly and Manguin 1946), São Jorge (Johansson 1977), São Miguel (Cedercreutz 1941), Terceira (Cedercreutz 1941)
Notes
Freshwater (lake), terrestrial
Stigonema informe
Kützing ex Bornet & Flahault, 1886
6F5FAA7E-D4FE-5EC1-A7E6-8418FB59DE6C
Distribution
São Jorge (Johansson 1977), Terceira (Johansson 1977)
Notes
Freshwater
Stigonema mamillosum
C.Agardh ex Bornet & Flahault, 1886
22169D47-4EEB-5C16-ABCD-76F02AB91E49
Distribution
São Jorge (Johansson 1977), São Miguel (Bourrelly and Manguin 1946)
Notes
Freshwater, terrestrial
Stigonema minutum
Hassall ex Bornet & Flahault, 1886
754A2D0E-CA33-5F36-8B5C-C44ED15B7BB0
Distribution
Flores (Bourrelly and Manguin 1946), São Jorge (Johansson 1977), São Miguel (Bohlin 1901), Terceira (Johansson 1977)
Notes
Freshwater (lake)
Stigonema multipartitum
N.L.Gardner, 1927
277B6A92-D8C9-512F-9388-6E4A5D9C0288
Distribution
São Jorge (Johansson 1977)
Notes
Freshwater
Stigonema ocellatum
Thuret ex Bornet & Flahault, 1886
B9F5BB81-4CFC-5CD5-8B5E-C85CD2BD0696
Distribution
Flores (Cedercreutz 1941), São Miguel (Cedercreutz 1941), Terceira (Johansson 1977)
Notes
Freshwater (lake)
Stigonema panniforme
Bornet & Flahault, 1886
1330F108-17FC-5C05-B141-A525244AD341
Distribution
São Miguel (Bourrelly and Manguin 1946)
Notes
Freshwater (stream)
Stigonema robustum
N.L.Gardner, 1927
8A74E630-C1AF-58C7-B0E9-22CBED7AACF1
Distribution
São Jorge (Johansson 1977)
Notes
Freshwater
Stigonema tomentosum
Hieronymus, 1895
1045DD00-12D4-53AA-825D-C66527124414
Distribution
São Jorge (Johansson 1977), São Miguel (Cedercreutz 1941)
Notes
Freshwater, terrestrial
Symploca dubia
Gomont, 1892
A43E25D7-0839-504E-80F4-D75E2F3E3FC9
Distribution
São Miguel (Cedercreutz 1941)
Notes
Thermal (pool)
Symploca thermalis
Gomont, 1892
2E16802F-02EF-5943-8166-25744AD2B127
Distribution
São Miguel (Bohlin 1901)
Notes
Thermal
Synechococcus
Nägeli, 1849
D0B2F7AA-BF3C-5370-8A8F-7CBAD85CAAF6
Distribution
São Miguel (Luz et al. 2020), Terceira (Luz et al. 2020)
Notes
Freshwater (lake)
Synechococcus nidulans
(Pringsheim) Komárek, 1970
C77F166C-9212-5512-A98A-58F52E052F9E
Distribution
São Miguel (Xavier et al. 2018)
Notes
Freshwater (lake)
Synechocystis
C.Sauvageau, 1892
056B3AAE-A756-5693-BE6B-956F18AA1215
Distribution
São Miguel (Luz et al. 2020), Pico (Luz et al. 2020)
Notes
Freshwater (lake)
Tildeniella torsiva
Mai, J.R.Johansen & Pietrasiak, 2018
9B23D44E-FAC3-501A-818D-EAB3F9F691E1
Distribution
São Miguel (Cordeiro et al. 2020b)
Notes
Freshwater (lake)
Tolypothrix
Kützing ex É.Bornet & C.Flahault, 1886
7520D80D-06F6-5E30-B71E-9004D6CB5EE0
Distribution
São Miguel (Archer 1874)
Notes
Freshwater
Tolypothrix distorta
Kützing ex Bornet & Flahault, 1886
2D14EF00-E006-5AE9-8B5F-C1CC1C660B3C
Distribution
São Miguel (Bohlin 1901)
Notes
Freshwater
Tolypothrix helicophila
Lemmermann, 1910
06F802E4-E334-50CB-AB11-E3583746C1BB
Distribution
Pico (Cordeiro et al. 2020b)
Notes
Freshwater (lake)
Tolypothrix lanata
Wartmann ex Bornet & Flahault, 1886
2DB9AB26-62EF-5163-9B99-0A2FEDD0EFAF
Distribution
São Miguel (Cedercreutz 1941)
Notes
Freshwater (lake)
Tolypothrix tenuis
Kützing ex Bornet & Flahault, 1886
0649CE44-C607-5C1E-A860-CA690555C2F8
Distribution
Flores (Bourrelly and Manguin 1946)
Notes
Freshwater
Trichormus variabilis
(Kützing ex Bornet & Flahault) Komárek & Anagnostidis, 1989
6CC32149-43D9-50EE-A5D7-E394851983EA
Distribution
São Miguel (Bohlin 1901)
Notes
Thermal (pool)
Tychonema
K.Anagnostidis & J.Komárek, 1988
E422F595-49ED-5FCA-98EB-F0AABA8BCE92
Distribution
São Miguel (Cordeiro et al. 2020b)
Notes
Freshwater (lake)
Westiellopsis
Janet, 1941
0ADAA7EE-9A01-588E-9516-525C8CC18002
Distribution
São Miguel (Cordeiro et al. 2020b)
Notes
Freshwater (lake), thermal (stream)
Woronichinia naegeliana
(Unger) Elenkin, 1933
4FBC75DF-F1C4-5FB8-B0A8-E83DA94C0A07
Distribution
Corvo Luz et al. 2020), Flores (Luz et al. 2020), Pico (Luz et al. 2020), São Miguel (Santos and Santana 2004)
Notes
Freshwater (lake)
Analysis
The present work comprises 225 taxa, 179 identified species and 11 only to genus level, distributed by six orders (Chroococcales, Nostocales, Oscillatoriales, Pleurocapsales, Spirulinales and Synechococcales), 30 families and 79 genera (Table 4). Most species belong to the Nostocales (43.0%) and Synechococcales (21.2%) orders. Chroococcales and Oscillatoriales orders contributed almost with the same number of species (17.3% and 16.8%, respectively), despite their different genera contributions.
Table 4.
Cyanobacteria taxa richness of the Azores Archipelago.
| Order | Taxa | Family | Genus | Species | Habitat (by taxa) | ||||
| Nº | % | Freshwater | Thermal | Brackish | Marine | ||||
| Chroococcales | 36 | 5 | 8 | 31 | 17.3 | 34 | 2 | - | - | 
| Nostocales | 95 | 11 | 34 | 77 | 43.0 | 85 | 9 | 1 | 4 | 
| Oscillatoriales | 36 | 4 | 14 | 30 | 16.8 | 27 | 7 | 2 | 2 | 
| Pleurocapsales | 4 | 1 | 2 | 2 | 1.1 | - | - | - | 4 | 
| Spirulinales | 1 | 1 | 1 | 1 | 0.6 | - | - | - | 1 | 
| Synechococcales | 53 | 8 | 20 | 38 | 21.2 | 47 | 5 | - | 2 | 
| Total: | 225 | 30 | 79 | 179 | 100 | 193 | 23 | 3 | 13 | 
A summary of cyanobacteria species richness found in the Azores and on each of the nine islands in the different types of habitats is given in Table 5. The number of recorded species was highest on São Miguel Island (115) and lowest on Graciosa Island (3). Freshwater systems were the most diverse habitats, comprising 193 taxa (85.7%), followed by thermal, with 23 species (10.2%), marine (13 species, 5.8%) and brackish systems (3 species, 1.3%).
Table 5.
Cyanobacteria taxa richness in the Azores by island.
| Island | Taxa | Taxonomy | Habitat | |||||||
| Order | Family | Genus | Species | Freshwater | Thermal | Brackish | Marine | |||
| Eastern | São Miguel | 151 | 4 | 25 | 59 | 115 | 133 | 20 | 1 | 1 | 
| Santa Maria | 7 | 2 | 5 | 6 | 5 | 5 | - | 2 | - | |
| Central | Terceira | 37 | 4 | 14 | 21 | 31 | 28 | 1 | 2 | 4 | 
| Pico | 43 | 4 | 16 | 27 | 31 | 43 | - | - | - | |
| São Jorge | 31 | 5 | 14 | 15 | 28 | 28 | - | - | 3 | |
| Graciosa | 3 | 3 | 3 | 3 | 3 | 3 | - | - | - | |
| Faial | 18 | 5 | 8 | 11 | 15 | 10 | - | - | 8 | |
| Western | Flores | 56 | 4 | 18 | 33 | 43 | 56 | - | 1 | - | 
| Corvo | 21 | 4 | 11 | 16 | 16 | 20 | - | - | 1 | |
A positive Pearson correlation coefficient (r = 0.86, n = 9, P = 0.003) was evident between species richness (S) and island area. This correlation is best described by a linear relationship (Fig. 2), where Pico and Flores seem to be outliers. Flores presented higher, while Pico has lower than expected species richness concerning its surface area.
Figure 2.
Species-area relationship. Regression line indicates a significant linear relationship with P = 0.003 and R2 = 0.86 (Pearson correlation). Dashed lines represent 95% interval confidence. COR: Corvo, FAI: Faial, FLO: Flores, GRA: Graciosa, PIC: Pico, SMR: Santa Maria, SJG: São Jorge, SMG: São Miguel, TER: Terceira.
Discussion
The cyanobacteria diversity in the Azores Archipelago is understudied compared to other European regions (Gkelis et al. 2016), despite being one of the best studied archipelagos in the North Atlantic (Cordeiro et al. 2020a). With its first records of cyanobacteria in Furnas, São Miguel Island, under the Challenger expedition, Moseley (1874) identified three genera: Chroococcus Nägeli, 1849, Nostoc Vaucher ex Bornet & Flahault, 1886 and Oscillatoria Vaucher ex Gomont, 1892. After that, as seen in Fig. 3, the contributions of the Cyanobacteria flora were sporadic, but significant, mainly with Bohlin (1901), Cedercreutz (1941), Bourrelly and Manguin (1946) and Johansson (1977). Their contributions were important, but geographically restricted to the larger islands, such as São Miguel and Terceira. From thereon, the number of recorded cyanobacteria species in the Azores has risen throughout the end of the 20th century and the 21st century and presently stands at 225 taxa with 179 identified species (Fig. 3). This boost in the 21st century is mainly due to the implementation of the WFD (e.g. Santos et al. 2005, Santos et al. 2012, Luz et al. 2020) with 39 new described species, which makes 21.8% of the total described species. This programme has helped the continuous study of freshwater cyanobacteria present in the major lakes of the Azores, in Pico, Flores and São Miguel. The works of Luz (2018), Xavier et al. (2018) and Cordeiro et al. (2020b) significantly contributed to the records of cyanobacteria for the Azores, reporting 19 new species, 10.6% of all taxa recorded only through in-vitro cultivation methods. Using a cultivation approach, these studies were able to isolate strains from lakes, terrestrial and thermal habitats, increasing the ability to identify small or rare species normally not detected in regular monitoring works.
Figure 3.
The number of described species through time from 1874 to 2020. Bars represent the number of species described per year, the line represents the cumulative number of described species.
Freshwater cyanobacteria are the most represented taxa in the Azores records, mainly from lakes. Although this result may reflect the abundance of this type of habitat in the Azores, it may also denote the less effort on diversity studies in other types of habitats. A much lower percentage of cyanobacteria was identified from thermal, marine and brackish habitats (10%, 6% and 1%, respectively), probably due to low sampling efforts. The availability of freshwater habitats in the Azores favours the establishment of incoming cyanobacteria in São Miguel, Flores, Terceira and São Jorge. These Islands have permanent streams, lakes, peat bogs and wetlands, providing highly diverse habitats for incomers, while these are absent in Faial, Graciosa and Santa Maria.
Several islands of the Azores have active volcanoes and present high numbers of fumarolic fields, geysers and hot springs (Cruz and França 2006) creating conditions for the growth of thermophilic cyanobacteria. Nevertheless, the current knowledge about thermal cyanobacteria in the Azores is low with few published works focusing on this habitat (Moseley 1874, Bohlin 1901, Luz 2018, Cordeiro et al. 2020b). The recent works by Luz (2018), with morphological identifications and Cordeiro et al. (2020b), that used both morphological and genetic characters for its identification, contributed to several new cyanobacteria taxa reports for the Azores from thermal habitats in São Miguel.
The cyanobacteria diversity and distribution in lotic systems in the Azores are much less known compared to other sites (Branco et al. 2001, Casamatta and Hašler 2016). Only a few works are available addressing this type of habitat in the Azores, with contributions mainly by Cedercreutz (1941), Bourrelly and Manguin (1946) and Johansson (1977), with no relevant works in the latest years. This is unusual as, in lotic systems, cyanobacteria are easily identified and sometimes even the dominant taxa (Schultz et al. 2013, Casamatta and Hašler 2016).
One of the most accepted explanations for regional biodiversity is the species-area relationship (SAR), according to which the number of species along the spatial scale increases with the area (e.g. Rosenzweig 1995, Drakare et al. 2006). This pattern was thoroughly studied on islands (e.g. Lomolino and Weiser 2001, Whittaker and Fernandez-Palacios 2007, Triantis et al. 2012), where the number of species from different taxonomic groups increases with the increase in island size (Triantis et al. 2012). In our data, a positive relationship between the island area and the number of species was observed (Fig. 3). This increase in cyanobacteria species richness with increasing island area in the Azores is consistent with the work of Borges et al. (2005), for arthropods and bryophytes and Raposeiro et al. (2009) for chironomids with an exception from Flores and Corvo Islands. A possible explanation for these exceptions is the higher percentage that water bodies represent in the total island area (Porteiro 2000) and also the higher precipitation (Secretaria Regional do Ambiente e do Mar 2011, Reichwaldt and Ghadouani 2012, Haakonsson et al. 2017). Compared to Pico Island, the percentage of land covered with water is double in Corvo and almost six times higher in Flores (Porteiro 2000). This suggests that, for cyanobacteria in the Azores, habitat diversity is an important factor in determining the SAR, as shown for other taxonomic groups and islands (Hortal et al. 2009, Chase et al. 2019).
Compared to other North Atlantic islands, the Azores present the highest species richness (Table 6). The overall distribution of species richness in the different cyanobacteria orders on the North Atlantic archipelagos generally follows the same pattern as the total world species, with Nostocales and Oscillatoriales being the richest orders. However, Nostocales represents a much higher contribution to the regional species richness in the Azores and Madeira Archipelagos, which could reflect their longer dispersion capabilities (Ribeiro et al. 2018). Nostocacean cyanobacteria are able to produce akinetes that can resist long periods of unfavourable conditions (Sarma 2012), enabling them to survive during long dispersion routes and colonise remote oceanic archipelagos, such as the Azores and Madeira. The absence of some orders, such as the Chroococcidiopsidales and Thermostichales, in the Azores and the other Macaronesia Archipelagos, suggests that they probably have a more restricted geographic distribution. Although biological, geographical and climatic factors may contribute to cyanobacteria regional species richness (Moreira et al. 2013, Walter et al. 2017, Ribeiro et al. 2018), the differences amongst islands are most probably related to different sampling efforts (Cordeiro et al. 2020b) and, between Azorean Islands, the distribution of planktic cyanobacteria seems to be mainly related to lake typology rather than environmental parameters (Cordeiro et al. 2020c). For instance, the reduced richness of Oscillatoriales in the Azores could be related to their preference for terrestrial and benthic habitats that are less studied in this Archipelago. With the increase in sampling campaigns covering all types of habitats, the reports of new cyanobacteria in the Azores are expected to increase and the representation of the different orders can become similar to the global pattern.
Table 6.
Cyanobacteria species richness in the Azores compared to world-known species richness (World's order and species number retrieved from Guiry and Guiry (2022); Canary Islands, Madeira and Cuba numbers retrieved from Cordeiro et al. (2020a)).
| Order | Azores | Madeira | Canary Islands | Cuba | World | ||||||
| Nº | % | Nº | % | Nº | % | Nº | % | Nº | % | ||
| Chroococcales | 31 | 17.32 | 2 | 8.33 | 4 | 6.35 | 28 | 19.05 | 649 | 13.20 | |
| Chroococcidiopsidales | 0 | 0 | 0 | 0 | 0 | 0 | 2 | 1.36 | 37 | 0.75 | |
| Gloeobacterales | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 3 | 0.06 | |
| Gloeomargaritales | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0.02 | |
| Nostocales | 77 | 43.02 | 11 | 45.83 | 20 | 31.75 | 44 | 29.93 | 1547 | 31.46 | |
| Oscillatoriales | 30 | 16.76 | 8 | 33.33 | 18 | 28.57 | 30 | 20.41 | 1397 | 28.41 | |
| Pleurocapsales | 2 | 1.12 | 1 | 4.17 | 6 | 9.52 | 3 | 2.04 | 223 | 4.53 | |
| Spirulinales | 1 | 0.56 | 0 | 0 | 4 | 6.35 | 5 | 3.40 | 56 | 1.14 | |
| Synechococcales | 38 | 21.23 | 2 | 8.33 | 11 | 17.46 | 35 | 23.81 | 995 | 20.23 | |
| Thermostichales | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 10 | 0.20 | |
| Total: | 179 | 24 | 63 | 147 | 4918 | ||||||
The hereby presented taxonomic list of cyanobacteria in the Azores represents a valuable resource for biodiversity research and awareness of described cyanobacteria tracked through years that, in the future, will allow the identification of possible invader species and studies of the influence of temperature changes in the World. Besides that, knowing the biodiversity of a specific archipelago enriches its value and allows future works in ecology and, in a more practical way, in biotechnology or pharmaceutical if found to be of increased value.
Supplementary Material
Cyanobacteria checklist of the Azores Archipelago, Portugal
Luz R, Cordeiro R, Fonseca A, Gonçalves V
Data type
Darwin Core Archive (.zip) of cyanobacteria taxa and occurrence data used for the presented and analysed checklist.
Brief description
Published Darwin Core Archive with two data tables in GBIF (doi: 10.15468/bfktqo) about the reports of cyanobacteria in the Azores Archipelago. The taxon (core) data table contains 229 records of cyanobacteria (from class to species level). One extension data table exists, with a total of 2838 occurrence records of cyanobacteria found in literature. The taxon data table is constructed, based on the occurrence data table.
File: oo_694835.zip
Acknowledgements
Rúben Luz was supported by a Ph.D. grant (M3.1.a/F/002/2020) from the Fundo Regional da Ciência e Tecnologia (FRCT). This work was funded by FEDER funds through the Interreg-MAC 2014-2020 Programme under the projects REBECA—Red de excelencia en biotecnología azul (algas) de la región macaronesia (MAC1.1a/060) and REBECA-CCT—Red de Excelencia en Biotecnología Azul de la Región Macaronésica. Consolidación, Certificación y Transferencia (MAC2/1.1b/269) and by Portuguese National Funds, through FCT—Fundação para a Ciência e a Tecnologia, the European Union, QREN, FEDER, COMPETE, by funding the CIBIO/InBIO (project UID/BIA/ 50027/2013 and POCI-01-0145-FEDER-006821). The work was also funded by FEDER (85%) and by Azorean Public funds (15%) through Operational Programme Azores 2020, under the project AZORESBIOPORTAL –PORBIOTA (ACORES-01-0145-FEDER-000072). CIIMAR acknowledges funding by FCT through UIDB/04423/2020 and UIDP/04423/2020.
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Associated Data
This section collects any data citations, data availability statements, or supplementary materials included in this article.
Supplementary Materials
Cyanobacteria checklist of the Azores Archipelago, Portugal
Luz R, Cordeiro R, Fonseca A, Gonçalves V
Data type
Darwin Core Archive (.zip) of cyanobacteria taxa and occurrence data used for the presented and analysed checklist.
Brief description
Published Darwin Core Archive with two data tables in GBIF (doi: 10.15468/bfktqo) about the reports of cyanobacteria in the Azores Archipelago. The taxon (core) data table contains 229 records of cyanobacteria (from class to species level). One extension data table exists, with a total of 2838 occurrence records of cyanobacteria found in literature. The taxon data table is constructed, based on the occurrence data table.
File: oo_694835.zip



