Biodiversity Data Journal :
Data Paper (Biosciences)
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Corresponding author: Pedro M. Raposeiro (pedro.mv.raposeiro@uac.pt)
Academic editor: Yasen Mutafchiev
Received: 18 Jan 2021 | Accepted: 18 Feb 2021 | Published: 17 Mar 2021
© 2021 Martin Souto, Vítor Gonçalves, Xabier Pontevedra-Pombal, Pedro Raposeiro
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Souto MS, Gonçalves V, Pontevedra-Pombal X, Raposeiro PM (2021) Distribution of testate amoebae in bryophyte communities in São Miguel Island (Azores Archipelago). Biodiversity Data Journal 9: e63290. https://doi.org/10.3897/BDJ.9.e63290
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Testate amoebae are a polyphyletic group of protists living preferentially in soils, freshwaters and wetlands. These Protozoa have a worldwide distribution, but their presence and diversity in the Azores (a remote oceanic archipelago) is poorly known, with only twelve taxa recorded so far. The published information reflects occasional collections from sporadic field visits from naturalists to São Miguel Island, mainly in the nineteenth century. To overcome this limitation, a standardised survey was carried out on the Island, sampling different types of habitats from several localities to provide the distribution and information on species ecology of testate amoebae.
In this study, 43 species of testate amoebae were recorded (within a total of 499 occurrences), belonging to two orders of Protista (26 Arcellinida and 17 Euglyphida). The most frequently occurring testate amoebae were Euglypha strigosa, Trinema lineare, Euglypha rotunda, Assulina muscorum and Cyclopyxis eurystoma. The most diverse genus was Euglypha (six species). A total of 38 species are new records for the Azores Archipelago. These data help to improve knowledge of the geographical distribution of testate amoebae in the northern hemisphere and their diversity in the Azores Archipelago.
biodiversity, community ecology, island, moss, Protozoa, substratum specificity
Testate amoebae are a polyphyletic group of small size [ranging from 7 to 500 µm long (
Despite their great importance, current knowledge of testate amoebae in the Azores Archipelago is limited when compared to other groups (e.g.
Later, with the Challenger expedition that took place from 1872 to 1876 and which had a brief passage to São Miguel Island, the Irish naturalist Archer (
Taxa (Current name) |
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Arcella vulgaris Ehrenberg, 1830 |
X |
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Centropyxis aculeata (Ehrenberg, 1830) Stein, 1857 |
as Echinopyxis aculeata |
X |
X |
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Centropyxis constricta (Ehrenberg, 1841) Penard, 1890 |
X |
X |
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Centropyxis sp. |
X |
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Difflugia acuminata Ehrenberg, 1838 |
X |
X |
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Difflugia globulosa Dujardin, 1837 |
as D. globularis |
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Difflugia mitriformis Wallich, 1864 |
X |
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Difflugia oligodon Ehrenberg, 1844 |
X |
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Difflugia pyriformis Perty, 1849 |
as D. azorica |
X |
X |
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Euglypha sp. |
X |
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Euglypha acanthophora (Ehrenberg, 1841) Perty, 1849 |
as E. alveolata |
as E. alveolata |
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Nebela collaris (Ehrenberg, 1848) Leidy, 1879 s.l. |
X |
X |
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Quadrulella symmetrica (Wallich, 1863) Kosakyan et al., 2016 |
as Difflugia symmetrica |
X |
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Trinema enchelys (Ehrenberg, 1838) Leidy, 1879 |
X |
as T. acinus |
X |
X |
Almost 100 years later,
The main objective of this data paper is to provide a record of the diversity and detailed distribution of testate amoebae in São Miguel Island (Azores Archipelago, Portugal). Additional information on species ecology is also discussed. Our purpose is to release this valuable dataset since no similar datasets have been previously published for Azores and it constitutes a relevant tool for comparison for ecologists studying, for example, biogeographic patterns or climate change and as modern analogues for environmental reconstructions on oceanic islands in paleoecological studies.
Records of testate amoebae in São Miguel Island (Azores Archipelago)
Collections were undertaken and occurrence data recorded during 2020 in São Miguel Island. The collectors were Martin Souto, Vitor Gonçalves and Pedro Miguel Raposeiro. Identification was done by Martin Souto and Xabier Pontevedra-Pombal. Production and analysis of scanning electron microscopy images was done by Xabier Pontevedra-Pombal.
The Azores is an oceanic archipelago located in the middle of the North Atlantic, about 1500 and 2100 km off the coast of Portugal (Europe) and North America, respectively (Fig.
Native forests cover less than 10% of the total area, mostly at elevations > 800 m a.s.l. (
This work was funded by FCT– Foundation for Science and Technology, the European Union, QREN, FEDER, COMPETE programmes (PMR - DL57/2016/ICETA/EEC2018/25; MSS - ICETA/EEC2018/25; DiscoverAzores project - PTDC/CTA-AMB/28511/2017; CIBIO/InBIO - UID/BIA/50027/2013 and POCI-01-0145-FEDER-006821) and Consolidation and Structuring project 2018 GRC-ED431C 2018/32 of the government of the Xunta Galicia.
This study covers 16 sampling locations on São Miguel Island (Fig.
Habitat characteristics and location of the sixteen studied localities in São Miguel
Cod. | Locality | Latitude (Nº) | Longitude (Wº) | Alt. (m) | Date | Habitat |
Loc. 1 | Lagoa Azul |
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260 | 04- Feb. | Lake shore with degraded aquatic vegetation |
Loc. 2 | Lagoa Verde |
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260 | 04- Feb. | Strongly altered and degraded aquatic vegetation |
Loc. 3 | Lagoa do Canário |
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755 | 04- Feb. | Aquatic vegetation with Sphagnum and Thuidium communities |
Loc. 4 | Lagoa do Caldeirão Norte |
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775 | 04- Feb. | Lake shore and shrub communities rich in Sphagnum |
Loc. 5 | Lagoa do Carvão |
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700 | 15- May. | Lake shore and shrub communities |
Loc. 6 | Lagoa da Prata |
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520 | 15- May. | Peatlands with Sphagnum and Cryptomeria forest |
Loc. 7 | Alto da Barrosa |
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800 | 27- May. | Shrub communities of Calluna with epiphytic bryophytes |
Loc. 8 | Lagoa do Fogo |
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580 | 22-Jun. | Lake shore and Cryptomeria forest |
Loc. 9 | Ribeira da Praia |
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180 | 05- May. | Fissidens communities |
Loc. 10 | Lombadas |
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600 | 11-May. | Hydrothermal, native and Cryptomeria forest |
Loc. 11 | Lagoa de São Brás |
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600 | 27-Feb. | Aquatic vegetation surrounded by Cryptomeria forest |
Loc. 12 | Lagoa do Areeiro |
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580 | 13-Feb. | Aquatic vegetation surrounded by Cryptomeria forest |
Loc. 13 | Lagoa do Congro |
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700 | 15-May. | Lake shore and Fissidens communities |
Loc. 14 | Ribeira do Caldeirão |
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600 | 27-May. | Native and Cryptomeria forest |
Loc. 15 | Planalto dos Graminhais |
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900 | 10-Jun. | Native forest and peatlands |
Loc. 16 | Ribeira do Guilherme |
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580 | 09- March. | Native forest |
Testate amoebae collections were taken at the beginning of the growing season, between spring (February - March) and summer 2020 (May - June). In each location, several types of vegetation were chosen for their homogeneity and abundance (e.g. forest, lowland shrub, shrubland, wetland, peatland, riparian communities (Fig.
Testate amoebae were sorted by fragmenting, washing and stirring of 10 × 10 cm of a wet mass of moss material into 1 litre of distilled water and then sieved through a 300 μm mesh size to remove large moss particles. The samples were concentrated by sedimentation and stored in vials with 50% alcohol at 4°C. A small aliquot of each sample was stored as a reference collection.
A drop of each sample (three subplots) was mounted on a semi-permanent slide and all testate amoebae were identified at 200x and 400x magnification using a compound microscope Leica DM2500. All measurements were made on photomicrographs (Leica DFC495 camera) of at least than 10 specimens, using image analysis software (Leica Application Suite version 3.8.0).
In order to obtain Scanning Electronic Microscopy (SEM) images, an aliquot was dried on aluminium supports with a carbon film. They were metallised with Iridium (40 nm) in a BioRad Microscience ion plating system and examined in a Zeiss Ultra Plus field emission microscope, at 5 kV high electric voltage.
The identification of testate amoebae was based on
The Azores is an oceanic archipelago located in the middle of the North Atlantic, about 1500 and 2100 km off the coast of Portugal (Europe) and North America, respectively (Fig.
37.704 and 37.917 Latitude; -25.857 and -25.125 Longitude.
Testate amoebae found on São Miguel Island
Rank | Scientific Name |
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order | Arcellinida |
family | Arcellidae |
species | Arcella arenaria Greeff, 1866 |
species | Arcella catinus Penard, 1890 |
family | Netzeliidae |
species | Cyclopyxis eurystoma Deflandre, 1929 |
species | Cyclopyxis kahli Deflandre, 1929 |
infraorder | Incertae sedis Sphaerothecina |
species | Trigonopyxis arcula (Leidy, 1879) Penard, 1912 |
family | Difflugiidae |
species | Difflugia bacillifera Penard, 1890 |
species | Difflugia glans Penard, 1902 |
species | Difflugia elegans Penard, 1890 |
family | Centropyxidae |
species | Centropyxis aerophila Deflandre, 1929 |
species | Centropyxis constricta (Ehrenberg, 1841) Penard, 1890 |
species | Centropyxis discoides (Penard, 1890) Deflandre, 1929 |
species | Centropyxis elongata (Penard, 1890) Tomas, 1959 |
family | Hyalospheniidae |
species | Alabasta militaris (Penard, 1890) Duckert, Blandenier, Kosakyan & Singer, 2018 |
species | Nebela collaris (Ehrenberg, 1848) Leidy, 1879 |
species | Padaungiella lageniformis (Penard, 1890) Lara & Todorov, 2012 |
species | Padaungiella tubulata (Brown, 1911) Lara & Todorov, 2012 |
species | Planocarina carinata (Archer, 1867) Kosakyan et al., 2016 |
species | Quadrulella symmetrica (Wallich, 1863) Kosakyan et al., 2016 |
family | Heleoperidae |
species | Heleopera rosea Penard, 1890 |
species | Heleopera sphagni Leidy, 1874 |
family | Microchlamyidae |
species | Pyxidicula cymbalum Penard, 1902 |
family | Phryganellidae |
species | Phryganella acropodia (Hertwig & Lesser, 1874) Hopkinson, 1909 |
family | Cryptodifflugiidae |
species | Cryptodifflugia oviformis Penard, 1890 |
order | Incertae sedis Arcellinida |
species | Argynnia caudata (Leidy, 1879) |
species | Argynnia dentistoma (Penard, 1890) |
species | Physochila griseola Penard, 1911 |
order | Euglyphida |
family | Euglyphidae |
species | Euglypha acanthophora (Ehrenberg, 1841) Perty,1849 |
species | Euglypha cristata Leidy, 1874 |
species | Euglypha filifera Penard, 1890 |
species | Euglypha laevis (Ehrenberg, 1845) Perty, 1849 |
species | Euglypha rotunda Wailes, 1911 |
species | Euglypha strigosa (Ehrenberg, 1871) Leidy, 1879 |
species | Tracheleuglypha dentata (Vejdovsky, 1882) Deflandre, 1928 |
family | Assulinidae |
species | Assulina muscorum Greeff, 1888 |
family | Cyphoderiidae |
species | Cyphoderia ampulla (Ehrenberg, 1840) Leidy, 1879 |
family | Sphenoderiidae |
species | Sphenoderia fissirostris Penard, 1890 |
species | Trachelocorythion pulchellum (Penard, 1890) Bonnet, 1979 |
family | Trinematidae |
species | Corythion constricta (Certes, 1889) Jung, 1942 |
species | Corythion dubium Taranek, 1881 |
species | Playfairina valkanovi Golemansky, 1966 |
species | Trinema complanatum Penard, 1890 |
species | Trinema enchelys (Ehrenberg, 1838) Leidy, 1879 |
species | Trinema lineare Penard, 1890 |
04-02-2020 through to 22-06-2020
This work is licensed under a Creative Commons Attribution (CC-BY) 4.0 Licence.
This paper presents data distribution of testate amoebae in São Miguel Island (Azores Archipelago) collected during 2020. The dataset has been published as a Darwin Core Archive (DwC-A), which is a standardised format for sharing biodiversity data as a set of one or more data tables (
Column label | Column description |
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id | Identifier of the record, coded as a global unique identifier |
locality | Name of the locality where the event occurred |
continent | Continent of the sampling site |
country | Country of the sampling site |
island | Island from the Island Group of the sampling site |
islandGroup | Island group of the sampling site |
eventID | Identifier of the event, unique for the dataset |
occurrenceID | Identifier of the occurrence, coded as a global unique identifier |
type | The nature of the resource |
Habitat | Habitat sampled |
basisOfRecord | The specific nature of the data record |
samplingProtocol | Sampling protocol |
recordedBy | Person who collected the specimens |
identifiedBy | Person who identified the specimens |
eventDate | Time interval when the event occurred |
taxonID | The identifier for the set of taxon information (data associated with the Taxon class). Specific identifier to the dataset |
scientificName | The name with authorship applied on the first identification of the specimen |
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 |
scientificNameAuthorship | The specimen accepted name, with authorship |
taxonRank | The taxonomic rank of the most specific name in the scientificName |
minimumElevationInMetres | Elevation in metres |
decimalLatitude | The geographic latitude of the sampling site |
decimalLongitude | The geographic longitude of the sampling site |
coordinateUncertaintylnMetres | The indicator for the accuracy of the coordinate location in metres, described as the radius of a circle around the stated point location. |
geodeticDatum | The spatial reference system upon which the geographic coordinates are based |
countryCode | Code of the country where the event occurred |
This study presents 499 testate amoebae (Protista) occurrences in 46 sampled sites (16 localities) in São Miguel Island, belonging to 43 species from 25 genera, 14 families and two orders. The order Euplyphida, represented by five families, accounted for 58.5% of the total occurrences and the order Arcellinida 41.5% of the total occurrences.
The families with the highest number of occurrences were Euglyphidae and Trinematidae (121), Hyalospheniidae (54), Heleoperidae (38), Centropyxidae (35) and Assulinidae (32). Additionally, the families with the highest number of taxa were Euglyphidae and Trinematidae (7), followed by Hyalospheniidae (6) and Centropyxidae (5). The families with lower occurrences (< 5) were Cyphoderiidae (4) and Microchlamyidae (1). The genera with the highest number of occurrences were Euglypha (106), Trinema (82), Heleopera (38) and Centropyxis (35). The other 21 genera had less than 35 occurrences. The genera with the highest number of taxa were Euglypha (6) and Centropyxis (4). Euglypha strigosa, Trinema lineare and Euglypha rotunda were the most frequent species occurring in 39, 37 and 34 sites, respectively. Assulina muscorum (32 sites), Cyclopyxis eurystoma (27 sites), Corythion dubium (26 sites), Trinema complanatum (25 sites) and Euglypha laevis (24 sites) were amongst the most ubiquitous testate amoebae (Figs
A total of six taxa occurring at only one sampling site were considered rare (Fig.
There are very few inventories of testate amoebae in the Azores, therefore the overall species richness of testate amoebae is unknown. Here, we present the first systematic study that explored the distribution of testate amoebae in bryophyte communities mainly in forest habitats. Forty-three species are recorded for São Miguel Island, 38 of these being new records for the Azores Archipelago. However, we must take into account the numerous cryptic taxa that testate amoebae present and some taxonomic uncertainty (
The three most representative families in terms of species richness, Centropyxidae, Euglyphidae and Trinematidae, are, in general, the most commonly registered in other oceanic archipelagos (
The 43 taxa recorded to the Azores is higher than what was reported to other oceanic archipelagos, such as the Canary Islands (10 species), Balearic Islands (15) and Island of Annobón (30 species). Testate amoebae assessment in the Balearic and Canary Islands was focused on mosses under Pinus forests of drier characteristics (
The testate amoebae assemblages in São Miguel Island were composed mainly by genus with a cosmopolitan distribution which are also known from other oceanic islands. For example, in the comparable Annobón tropical rainforest, situated much further south (
Moss biotopes are very abundant in the Azores, where extant bryoflora comprises about 430 species of mosses and hepatics (
Most of the natural vegetation of the Island has been replaced by pastures and Cryptomeria forests (Fig.
The most frequent genera shared on these terrestrial and semi-terrestrial habitats are Nebela and Euglypha, which are less represented in the aquatic systems. According to
Bryophytes assemblages on lake shores presented a high diversity of testate amoebae (n = 13, 33 species). Acarcarpic mosses, like Fissidens or Campylopus in more open areas near lakes, maintain less developed testate amoebae communities (Fig. 6). However, the genera Sphagnum and Rhytidiadelphus squarrosus, especially in more waterlogged areas, maintain rich testate communities (n = 13, 33 species). Several works consider aquatic environments and sediments to be the preferred habitat for these organisms, yet they end up functioning as a data collector for the surrounding ecosystems and many species are actually terrestrial (
On hydrothermal vents, lower diversity of testate amoebae was observed (n =3, 14 species). These vents develop an abundant bryophyte extension (Fig.
Here, we presented the first study that explored the distribution of testate amoebae of different habitats from the Azores Archipelago, mainly in São Miguel Island. Moreover, this work indicates that there are typical species on the different sampled habitats. This a matter of concern on islands, where large areas of native forest have been replaced by exotic forest and changes in land uses, driven by human activities, will affect population dynamics. In order to better understand the complexity of these habitats, population dynamics and species specificity need to be carried out. Larger datasets located in different islands and habitats are required to better understand how these communities respond to environment changes. Additionally, molecular barcoding is a useful tool, not only for species identification, but also for studying evolutionary and ecological processes. The results of this study provide indications that testate amoebae assemblages are habitat specific and therefore constitute a promising group for paleoenvironmental reconstruction of Azorean ecosystems. Future studies in drier ecosystems, coastal areas and hydrothermal zones may reveal and offer us a greater diversity of these organisms.
This work was funded by FCT – Foundation for Science and Technology, the European Union, QREN, FEDER, COMPETE programmes (PMR - DL57/2016/ICETA/EEC2018/25; MSS - ICETA/EEC2018/25; DiscoverAzores Project - PTDC/CTA-AMB/28511/2017; UID/BIA/50027/2020 and POCI-01-0145-FEDER-006821), AZORESBIOPORTAL – PORBIOTA (ACORES-01-0145-FEDER-000072) and Consolidation and Structuring Project 2018 GRC-ED431C 2018/32 of Xunta de Galicia government. We would like to thank Rafael Carballeira for his collaboration in the realisation of the SEM photographs. A special thanks to Mr. Matthew James Mole for revising this article. Additionally, we thank the reviewers for their constructive and valuable comments that helped to improve this paper.
Martin Souto, Vitor Gonçalves and Pedro Miguel Raposeiro conceived the study and carried out the sampling campaign. Identification was done by Martin Souto and Xabier Pontevedra-Pombal. Production and analysis of scanning electron microscopy images was done by Xabier Pontevedra-Pombal. Martin Souto and Pedro Miguel Raposeiro wrote the paper with inputs from all authors. All authors agree with the final version of the paper.